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A revised cladistic classification of the Nepticulidae (Lepidoptera) with descriptions of new taxa mainly from South Africa
... The first cladistic analysis and classification resulted from the PhD studies on the South African Nepticulidae fauna by Scoble (1983), here redrawn in Fig. 2A. The family was divided into two subfamilies: the Australian Pectinivalvinae and the global Nepticulinae, the latter subdivided into Nepticulini and Trifurculini. ...
... Four years later, in his unpublished PhD thesis of Australian Nepticulidae, Hoare prepared the first cladistic analysis of Nepticulidae using maximum parsimony algorithms employed in computer software (Hoare, 1998) (Fig. 2D). The weakness of this analysis was the low number of species included, but its strength was the inclusion of characters of all life stages and this resulted in a refined version of the divisions by Scoble (1983) andvan Nieukerken (1986b). Additionally from Hoare's PhD studies, analyses of the Australian Pectinivalvinae (Hoare, 2000b;Hoare & van Nieukerken, 2013) resulted in the recognition of the new genus Roscidotoga Hoare, and division of the genus Pectinivalva into three subgenera: Pectinivalva Hoare, Casanovula Hoare and Menurella Hoare. ...
... Based on our current knowledge we therefore discontinue the use of subfamily and tribe within Nepticulidae (and Opostegidae) for the time being. We also abandon the use of subgenera, which were introduced in the 1980s (Scoble, 1983;van Nieukerken, 1986b), mainly because the former large genus Ectoedemia (Ectoedemia + Zimmermannia + Etainia + Muhabbetana + Fomoria) is shown to be polyphyletic, which can only be addressed by either raising all subgenera to full genus, or including even more genera in this large polytypic entity. We reject the latter solution because that would leave no reliable or practical apomorphies. ...
Nepticulidae represent one of the early diverging Lepidoptera lineages, and the family currently comprises over 850 described species. The larvae of the vast majority of the species are leaf miners on Angiosperms and highly monophagous, which has led to persistent ideas on coevolution with their plant hosts. We present here a molecular phylogeny based on eight gene fragments from 355 species, representing 20 out of 22 extant Nepticulidae genera. Using two fossil calibration points, we performed molecular dating to place the origin of the family in the Early Cretaceous, before the main Angiosperm diversification. Based on our results we propose a new classification, abandoning all ranks between family and genus, as well as subgenera to allow for a stable classification. The position of Enteucha Meyrick within Nepticulidae remains somewhat ambiguous, and the species-rich cosmopolitan genus Stigmella Schrank, with nearly half of all described Nepticulidae, requires further study. Ectoedemia Busck, Zimmermannia Hering, Acalyptris Meyrick, Etainia Beirne, Parafomoria Borkowski, Muhabbetana Koçak & Kemal and Fomoria Beirne appear to have diversified in a relatively short evolutionary period, leading to short branches in the molecular phylogeny and unclear suprageneric relations. Otherwise support values throughout the phylogeny are mostly high and the species groups, genera and higher clades are discussed in respect of their supporting morphological and life-history characters. Wing venation characters are confirmed to be mostly reliable and relevant for Nepticulidae classification, but some other previously used characters require reinterpretation. The species groups of most genera are recovered, but only partly so in the large genus Stigmella. The molecular dating results are compared with existing knowledge on the timing of the Angiosperm radiation and reveal that the diversification of Nepticulidae could largely have been contemporaneous with their hosts, although some of the genera restricted to a single plant family appear to have begun to diversify before their hosts.
... If we have to choose, we probably adhere most to the phylogenetic or diagnostic species concept (Mallet 2001;Isaac et al. 2004), but agree with Mallet's notion that "agreement on a unified species-level taxonomy is possible, but will be forthcoming only if we accept that species lack a single, interpretable biological reality over their geographic range and across geological time". In the taxonomic literature on Nepticuloidea, only Scoble (1983) discussed how he recognised species, but also he concluded that the practice is rather different from the theory. With the explosion of genetic data we have much more information nowadays than just two decades ago, but that does not mean that it has become much easier to recognise species. ...
... African species Stigmella fluida group (Scoble, 1978b: 92) Stigmella ingens group (Scoble, 1978b: 111) Stigmella abachausi (Janse, 1948) Scoble, 1978b AFR Nepticula abachausi Janse, 1948: 162 Stigmella abutilonica Scoble, 1978b: 93 AFR Stigmella ampullata Scoble, 1978b: 108 AFR Stigmella angustivalva Scoble, 1978b: 113 AFR Stigmella caliginosa (Meyrick, 1921b) Scoble, 1983 AFR Nepticula caliginosa Meyrick, 1921b: 140 Stigmella celtifoliella Vári, 1955: 338 AFR Stigmella charistis Vári, 1963: 71 AFR Stigmella confinalis Scoble, 1978b: 111 AFR Stigmella crotonica Scoble, 1978b: 100 AFR Stigmella dombeyivora Scoble, 1978b: 107 AFR Stigmella ficivora Gustafsson, 1985: 170 AFR Stigmella fluida (Meyrick, 1911a) Scoble, 1978b AFR Nepticula fluida Meyrick, 1911a: 236 Stigmella galactacma (Meyrick, 1924b) Diškus & Puplesis, 2003 AFR Nepticula galactacma Meyrick, 1924b: 89 Stigmella grewiae Scoble, 1978b: 112 AFR Stigmella gustafssoni (Căpuşe, 1975) Diškus & Puplesis, 2003: 366 AFR Nepticula gustafssoni Căpuşe, 1975: 211 Stigmella ingens (Meyrick, 1913) Scoble, 1978b AFR Nepticula ingens Meyrick, 1913: 327 Stigmella irrorata (Janse, 1948) Scoble, 1978b AFR Nepticula irrorata Janse, 1948: 168 Stigmella letabensis Scoble, 1978b: 113 AFR Stigmella liota Vári, 1963: 73 AFR Stigmella maytenivora Gustafsson, 1985: 174 AFR Stigmella naibabi Mey, 2004: 29 AFR Stigmella nigrata (Meyrick, 1913) Scoble, 1978b AFR Nepticula nigrata Meyrick, 1913: 326 Stigmella panconista (Meyrick, 1920a) Diškus & Puplesis, 2003 AFR Nepticula panconista Meyrick, 1920a: 312 Stigmella parinarella Vári, 1955: 337 AFR Stigmella perplexa (Janse, 1948) Scoble, 1978b AFR Nepticula perplexa Janse, 1948: 172 Stigmella platyzona Vári, 1963: 67 AFR Stigmella porphyreuta (Meyrick, 1917a) Scoble, 1978b AFR Nepticula porphyreuta Meyrick, 1917a: 13 Stigmella pretoriata Scoble, 1978b: 109 AFR Stigmella protosema (Meyrick, 1921b) Scoble, 1978b AFR Nepticula protosema Meyrick, 1921b: 140 Stigmella rhomboivora Gustafsson, 1985: 167 AFR Stigmella rhynchosiella Vári, 1955: 338 AFR Stigmella urbica (Meyrick, 1913) Scoble, 1978b AFR Nepticula urbica Meyrick, 1913: 326 Stigmella uwusebi Mey, 2004: 30 AFR Stigmella varii Scoble, 1978b: 95 AFR Stigmella wollofella (Gustafsson, 1972) Gustafsson, 1985 AFR Nepticula wollofella Gustafsson, 1972: 158 Nepticula mandingella Gustafsson, 1972: 157 syn. n. 24 Stigmella mandingella (Gustafsson, 1972) Diškus & Puplesis, 2003 Stigmella worcesteri Scoble, 1983: 16 RN for S. pallida Scoble, 1978 AFR Stigmella pallida Scoble, 1978b: 105 JSH of Stigmella pallida (Braun, 1917) Australian species ...
... African species Stigmella fluida group (Scoble, 1978b: 92) Stigmella ingens group (Scoble, 1978b: 111) Stigmella abachausi (Janse, 1948) Scoble, 1978b AFR Nepticula abachausi Janse, 1948: 162 Stigmella abutilonica Scoble, 1978b: 93 AFR Stigmella ampullata Scoble, 1978b: 108 AFR Stigmella angustivalva Scoble, 1978b: 113 AFR Stigmella caliginosa (Meyrick, 1921b) Scoble, 1983 AFR Nepticula caliginosa Meyrick, 1921b: 140 Stigmella celtifoliella Vári, 1955: 338 AFR Stigmella charistis Vári, 1963: 71 AFR Stigmella confinalis Scoble, 1978b: 111 AFR Stigmella crotonica Scoble, 1978b: 100 AFR Stigmella dombeyivora Scoble, 1978b: 107 AFR Stigmella ficivora Gustafsson, 1985: 170 AFR Stigmella fluida (Meyrick, 1911a) Scoble, 1978b AFR Nepticula fluida Meyrick, 1911a: 236 Stigmella galactacma (Meyrick, 1924b) Diškus & Puplesis, 2003 AFR Nepticula galactacma Meyrick, 1924b: 89 Stigmella grewiae Scoble, 1978b: 112 AFR Stigmella gustafssoni (Căpuşe, 1975) Diškus & Puplesis, 2003: 366 AFR Nepticula gustafssoni Căpuşe, 1975: 211 Stigmella ingens (Meyrick, 1913) Scoble, 1978b AFR Nepticula ingens Meyrick, 1913: 327 Stigmella irrorata (Janse, 1948) Scoble, 1978b AFR Nepticula irrorata Janse, 1948: 168 Stigmella letabensis Scoble, 1978b: 113 AFR Stigmella liota Vári, 1963: 73 AFR Stigmella maytenivora Gustafsson, 1985: 174 AFR Stigmella naibabi Mey, 2004: 29 AFR Stigmella nigrata (Meyrick, 1913) Scoble, 1978b AFR Nepticula nigrata Meyrick, 1913: 326 Stigmella panconista (Meyrick, 1920a) Diškus & Puplesis, 2003 AFR Nepticula panconista Meyrick, 1920a: 312 Stigmella parinarella Vári, 1955: 337 AFR Stigmella perplexa (Janse, 1948) Scoble, 1978b AFR Nepticula perplexa Janse, 1948: 172 Stigmella platyzona Vári, 1963: 67 AFR Stigmella porphyreuta (Meyrick, 1917a) Scoble, 1978b AFR Nepticula porphyreuta Meyrick, 1917a: 13 Stigmella pretoriata Scoble, 1978b: 109 AFR Stigmella protosema (Meyrick, 1921b) Scoble, 1978b AFR Nepticula protosema Meyrick, 1921b: 140 Stigmella rhomboivora Gustafsson, 1985: 167 AFR Stigmella rhynchosiella Vári, 1955: 338 AFR Stigmella urbica (Meyrick, 1913) Scoble, 1978b AFR Nepticula urbica Meyrick, 1913: 326 Stigmella uwusebi Mey, 2004: 30 AFR Stigmella varii Scoble, 1978b: 95 AFR Stigmella wollofella (Gustafsson, 1972) Gustafsson, 1985 AFR Nepticula wollofella Gustafsson, 1972: 158 Nepticula mandingella Gustafsson, 1972: 157 syn. n. 24 Stigmella mandingella (Gustafsson, 1972) Diškus & Puplesis, 2003 Stigmella worcesteri Scoble, 1983: 16 RN for S. pallida Scoble, 1978 AFR Stigmella pallida Scoble, 1978b: 105 JSH of Stigmella pallida (Braun, 1917) Australian species ...
A catalogue of all named Nepticulidae and Opostegidae is presented, including fossil species. The catalogue is simultaneously published online in the scratchpad http://nepticuloidea.info/ and in Catalogue of Life (http://www.catalogueoflife.org/col/details/database/id/172). We provide a historical overview of taxonomic research on Nepticuloidea and a brief ‘state of the art’. A DNA barcode dataset with 3205 barcodes is made public at the same time, providing DNA barcodes of ca. 779 species, of which 2563 are identified as belonging to 444 validly published species. We recognise 862 extant and 18 fossil species of Nepticulidae in 22 extant genera and the fossil form genus Stigmellites. We count 192 valid Opostegidae species in 7 genera, without fossils. We also list seven dubious Nepticulidae names that cannot be placed due to absent type material and poor descriptions, 18 unavailable names in Nepticulidae that cannot be placed and we also list the 33 names (including four fossils) that once were placed as Nepticulidae or Opostegidae but are now excluded. All synonyms and previous combinations are listed. The generic classification follows the Molecular phylogeny that is published almost simultaneously. Subfamilies and tribes are not recognised, Trifurculinae Scoble, 1983 is synonymised with Nepticulidae Stainton, 1854 and Opostegoidinae Kozlov, 1987 is synonymised with Opostegidae Meyrick, 1893. The status of Casanovula Hoare, 2013, Etainia Beirne, 1945, Fomoria Beirne, 1945, Glaucolepis Braun, 1917, Menurella Hoare, 2013, Muhabbetana Koçak & Kemal, 2007 and Zimmermannia Hering, 1940 is changed from subgenus to full genus, whereas two genera are considered synonyms again: Manoneura Davis, 1979, a synonym of Enteucha Meyrick, 1915 and Levarchama Beirne, 1945, a synonym of Trifurcula Zeller, 1848. We propose 87 new combinations in Nepticulidae and 10 in Opostegidae, largely due to the new classification, and re-examination of some species. We propose the following 37 new synonymies for species (35 in Nepticulidae, 2 in Opostegidae):
Stigmellaacerifoliella Dovnar-Zapolski, 1969 (unavailable, = Stigmellaacerna Puplesis, 1988), Stigmellanakamurai Kemperman & Wilkinson, 1985 (= Stigmellapalionisi Puplesis, 1984), Nepticulaamseli Skala, 1941 (unavailable = Stigmellabirgittae Gustafsson, 1985), Stigmellacathepostis Kemperman & Wilkinson, 1985 (= Stigmellamicrotheriella (Stainton, 1854)), Stigmellapopulnea Kemperman & Wilkinson, 1985 (= Stigmellanivenburgensis (Preissecker, 1942)), Nepticulaobscurella Braun, 1912 (revised synonymy, = Stigmellamyricafoliella (Busck, 1900)), Nepticulamandingella Gustafsson, 1972 (= Stigmellawollofella (Gustafsson, 1972)), Stigmellarosaefoliellapectocatena Wilkinson & Scoble, 1979 (= Stigmellacentifoliella (Zeller, 1848)), Micropteryxpomivorella Packard, 1870 (= Stigmellaoxyacanthella (Stainton, 1854)), Stigmellacrataegivora Puplesis, 1985 (= Stigmellamicromelis Puplesis, 1985), Stigmellascinanella Wilkinson & Scoble, 1979 (= Stigmellapurpuratella (Braun, 1917)), Stigmellapalmatae Puplesis, 1984 (= Stigmellafilipendulae (Wocke, 1871)), Stigmellasesplicata Kemperman & Wilkinson, 1985 (= Stigmellalediella (Schleich, 1867)), Stigmellarhododendrifolia Dovnar-Zapolski & Tomilova, 1978 (unavailable, = Stigmellalediella (Schleich, 1867)), Stigmellaoa Kemperman & Wilkinson, 1985 (= Stigmellaspiculifera Kemperman & Wilkinson, 1985), Stigmellagracilipae Hirano, 2014 (= Stigmellamonticulella Puplesis, 1984), Nepticulachaoniella Herrich-Schäffer, 1863 (= Stigmellasamiatella (Zeller, 1839)), Bohemanniapiotra Puplesis, 1984 (= Bohemanniapulverosella (Stainton, 1849)), Bohemannianipponicella Hirano, 2010 (= Bohemanniamanschurella Puplesis, 1984), Sinopticulasinica Yang, 1989 (= Glaucolepisoishiella (Matsumura, 1931)), Trifurculacollinella Nel, 2012 (= Glaucolepismagna (A. Laštuvka & Z. Laštuvka, 1997)), Obrussatigrinella Puplesis, 1985 (= Etainiatrifasciata (Matsumura, 1931)), Microcalyptrisvittatus Puplesis, 1984 and Microcalyptrisarenosus Falkovitsh, 1986 (both = Acalyptrisfalkovitshi (Puplesis, 1984)), Ectoedemiacastaneae Busck, 1913, Ectoedemiaheinrichi Busck, 1914 and Ectoedemiahelenella Wilkinson, 1981 (all three = Zimmermanniabosquella (Chambers, 1878)), Ectoedemiachloranthis Meyrick, 1928 and Ectoedemiaacanthella Wilkinson & Newton, 1981 (both = Zimmermanniagrandisella (Chambers, 1880)), Ectoedemiacoruscella Wilkinson, 1981 (= Zimmermanniamesoloba (Davis, 1978)), Ectoedemiapiperella Wilkinson & Newton, 1981 and Ectoedemiareneella Wilkinson, 1981 (both = Zimmermanniaobrutella (Zeller, 1873)), Ectoedemiasimiligena Puplesis, 1994 (= Ectoedemiaturbidella (Zeller, 1848)), Ectoedemiaandrella Wilkinson, 1981 (= Ectoedemiaulmella (Braun, 1912)), Nepticulacanadensis Braun, 1917 (= Ectoedemiaminimella (Zetterstedt, 1839)), Opostegarezniki Kozlov, 1985 (= Opostegacretatella Chrétien, 1915), Pseudopostegacyrneochalcopepla Nel & Varenne, 2012 (= Pseudopostegachalcopepla (Walsingham, 1908)). Stigmellacaryaefoliella (Clemens, 1861) and Zimmermanniabosquella (Chambers, 1878) are taken out of synonymy and re-instated as full species. Lectotypes are designated for Trifurculaobrutella Zeller, 1873 and Nepticulagrandisella Chambers, 1880.
... The first cladistic analysis and classification resulted from the PhD studies on the South African Nepticulidae fauna by Scoble (1983), here redrawn in Fig. 2A. The family was divided into two subfamilies: the Australian Pectinivalvinae and the global Nepticulinae, the latter subdivided into Nepticulini and Trifurculini. ...
... Four years later, in his unpublished PhD thesis of Australian Nepticulidae, Hoare prepared the first cladistic analysis of Nepticulidae using maximum parsimony algorithms employed in computer software (Hoare, 1998) (Fig. 2D). The weakness of this analysis was the low number of species included, but its strength was the inclusion of characters of all life stages and this resulted in a refined version of the divisions by Scoble (1983) andvan Nieukerken (1986b). Additionally from Hoare's PhD studies, analyses of the Australian Pectinivalvinae (Hoare, 2000b;Hoare & van Nieukerken, 2013) resulted in the recognition of the new genus Roscidotoga Hoare, and division of the genus Pectinivalva into three subgenera: Pectinivalva Hoare, Casanovula Hoare and Menurella Hoare. ...
... Based on our current knowledge we therefore discontinue the use of subfamily and tribe within Nepticulidae (and Opostegidae) for the time being. We also abandon the use of subgenera, which were introduced in the 1980s (Scoble, 1983;van Nieukerken, 1986b), mainly because the former large genus Ectoedemia (Ectoedemia + Zimmermannia + Etainia + Muhabbetana + Fomoria) is shown to be polyphyletic, which can only be addressed by either raising all subgenera to full genus, or including even more genera in this large polytypic entity. We reject the latter solution because that would leave no reliable or practical apomorphies. ...
Nepticulidae are amongst the earliest radiating Lepidoptera, with a fossil record of larval leafmines from the Albian onwards. We present a phylogeny and analysis of divergence times based on eight molecular markers, a taxonset of ca 350 exemplars, representing almost all (sub)generic groupings. The support for the monophyly of most clades is high, but not all previous groupings are recovered. Ectoedemia s.l. with several subgenera is not monophyletic, but all subgenera are. The small genus Enteucha is sistergroup to all other Nepticulidae in most analyses, but close to Stigmella in others. The large genus Stigmella is divided in three well supported clades, two of which harbour most tropical species. In our new classification we abandon subgenera and subfamilial ranks. The resulting genera are not only supported by several morphological apomorphies, several have specific biologies, a limited host plant choice and a limited biogeography. Particularly the Neotropics show separate clades. Using two fossil calibration points, we estimate the divergence times of genera and discuss their congruence with other recent insights in Lepidoptera diversification dating estimates. The estimated 95% confidence interval for the origin of Nepticulidae falls almost completely within the Early Cretaceous. Almost all of the currently recognized genera are estimated to have originated before the end of the Cretaceous, 66 mya. Considering that Nepticulidae are the first larger group of Lepidoptera, with 850 named and 2000 assumed species, specialising in angiosperm feeding, we postulate that also geologically the Nepticulidae formed the first larger group of Lepidoptera, radiating on angiosperms.
... The morphology of nepticulids has been extensively discussed by, particularly, Scoble (1983), Nieukerken (1986b), Johansson et al (1990) and Puplesis (1994). Conspicuous morphological features are discussed below to provide a background to understanding the morphological particularities of the newly describing taxa and justifying amendments in terminology. ...
... Within the Trifurculinae, Bohemannia takes the lowest phylogenetic branch, i.e., it is the sister group of the remaining Trifurculinae (for details, see Puplesis, 1984bPuplesis, , 1992Puplesis, , 1994. However, an alternative classi fication proposed by Scoble (1983) is better supported by characters of the immature stages and has often been followed in recent literature with some adjust ments at generic level (Nieukerken, 1986b;Hoare et ai, 1997;Hoare, 2000). This latter classification recognizes two alternative sub families: Pectinivalvinae (including two endemic Australian genera Pectinivalva and Roscidotoga) and Nepticulinae (including all remaining genera divided between two tribes, Nepticulini and Trifurculini). ...
... Pectinivalva Scoble, 1983. Type species: Pectinivalva commoni Scoble, 1983. ...
... Opostegidae is present in two genera of Nepticulidae (Pectinivalva, Acalyptris) (Scoble, 1983;Puplesis, 1984Puplesis, , 1989Puplesis, , 1994 there is doubt as to whether these structuresare homologousin the two familiesor repre sent an instance of parallelism. The pectinifer in Pectinivalva is almost certainly homologous with that of the Opostegidae (Nieukerken, 1986) but that in Acalyptris almost certainly not on grounds of parsi mony. ...
... However, it does not appear possible to use this feature in recognizing relationships within the family. In the Nepticulidae the plesiomorphic condi tion prevails in that microtrichia are scattered over the entire forewing and hindwing (Scoble, 1983). ...
... The presence of a pedicellum bearing a cucullar lobe with a pectinifer is an important autapomorphy of the Opostegidae as a whole. Al-11 though pectinifers have beendiscovered in twogenera of the Nepticulidae (Scoble, 1982(Scoble, , 1983Puplesis, 1990, 1994, Hoare et al., 1997 they are not borne on a separate cucullar lobe. Therefore the pectinifer in Pectinivalva (Nepticulidae) may represent a more an cestral state of the structure than that of any opostegid. ...
... Bet kokiu atveju molekulinių mažųjų gaubtagalvių (Nepticulidae) duomenų dar maža ( van Nieukerken et al., 2012avan Nieukerken et al., , 2012b, ir iki šiol visos šiuolaikinės šeimos taksonominės sistemos bei rūšių identifikacija daugiausia remiasi morfologijos ir pagal galimybes biologijos duomenimis (požymiais). Apie bendrąsias Nepticulidae morfologijos ypatybes buvo ne kartą skelbta mokslinėje literatūroje (Scoble, 1983;van Nieukerken, 1986;Johansson et al., 1990;Puplesis, Diškus, 2003 ir kt.). Toliau pateikiama Nepticulidae tegmų (ir atskirų morfologinių sistemų) apžvalga, įvertinanti morfologinių požymių svarbą diagnostikoje ir taksonomijoje. ...
... Apatinių žandų (maksilių) čiuopikliai, smailėjantys tolimajame (distaliniame) gale, sudaryti iš 5-ių segmentų. Straubliukas (proboscis) labai trumpas; šiuo metu žinomos tik kelios Pietų Afrikos mažųjų gaubtagalvių rūšys, turinčios pailgėjusį straubliuką (Scoble, 1983). Čiuopikliai labai trumpi, sudaryti iš 2-ų ar 3-ų segmentų. ...
... Kitokia Nepticulidae klasifikacija buvo pasiūlyta tyrinėjant Pietų pusrutulio fauną (Scoble, 1983). M. J. Scoble sudarytoje kladogramoje buvo daugiau panaudota ankstyvojo vystymosi stadijų požymių ir šia klasifikacija (su įvairiais pakeitimais) buvo dažnai remiamasi entomologinėje literatūroje ( van Nieukerken, 1986;Johansson et al., 1990;Hoare et al., 1997;Hoare, 2000aHoare, , 2000b. ...
... In the early seventies the interest in the family increased and resulted into several accounts on the classification of Nepticulidae, such as Johansson (197 l), Borkowski (1972) and Wilkinson & Scoble (1979) who all adopted and refined Beirne's scheme. Recently, Scoble (1983) provided a revised classification, the first which strictly followed cladistic principles. Although his classification was intended to treat the family world-wide, Scoble mainly studied the South-African species and several Holarctic genera were treated by him as "incertae sedis". ...
... The work by Scoble (1983) provides an important starting point for the classification of the Holarctic Nepticulidae; in this study I was able to corroborate most of the branching points proposed by him and to refine others. I also could resolve some of the existing uncertainties in a few Holarctic genera, not studied by Scoble. ...
... Descriptions of immature stages are not given; not enough material of pupae was available (but see Scoble, 1983 for some notes) and the larvae will be subject of a future publication. However, important apomorphies in immature stages are included in this work, but will be elaborated on later ( Van Nieukerken & Jansen, in prep.). ...
A revised classification of the Holarctic genera of Nepticulidae is provided. Eight genera belonging to the nominal subfamily are recognised and redefined. They are Enteucha Meyrick (= Johanssonia Borkowski, Artaversala Davis, Oligoneura Davis), Stigmella Schrank (including Astigmella Puplesis), Simplimorpha Scoble in the Nepticulini and Acalyptris Meyrick ( = Microcalyptris Braun, Niepeltia Strand), Trifurcula Zeller, Parafomoria Van Nieukerken, Bohemannia Stainton and Ectoedemia Busck in the Trifurculini. Trifurcula is divided into the subgenera Glaucolepis Braun (= Fedalmia Beirne), Levarchama Beirne and Trifurcula s.str. Ectoedemia is divided into the subgenera Etainia Beirne, Fomoria Beirne, Laqueus Scoble, Zimmermannia Hering and Ectoedemia s.str. The genera and subgenera are (re)described and data on biology and distribution are given. The species Simplimorpha promissa (Staudinger) and Acalyptris psammophricta Meyrick are redescribed. A phylogeny of the family in cladistic sense is presented and discussed. The monophyly of Ectoedemia is uncertain, and monophyly of the subgenus Fomoria could not be demonstrated. Current opinions about the phylogenetic position of the Nepticulidae and the division of the Heteroneura are reviewed and discussed. The biogeography and hostplant relationships of the family are reviewed.
Nepticulidae-: Key-to-genera-and-subgenera, Holarctic-, Adult-males-and-females, p.-41;
Nepticulidae: Revision;
Acalyptris-Meyrick-1921 ( Nepticulidae- ) : Emended-diagnosis, P.-59;
Acalyptris-psammophricta-Meyrick-1921 ( Nepticulidae- ) : Redescription-, Female-, p.-62;
Bohemannia-Stainton-1859 ( Nepticulidae- ) : Emended-diagnosis, P.-72;
Ectoedemia-Busck-1907 ( Nepticulidae- ) : Emended-diagnosis, P.-75;
Nepticulini- ( Nepticulidae- ) : Emended-diagnosis, P.-44;
Parafomoria-van-Nieukerken-1983 ( Nepticulidae- ) : Emended-diagnosis, P.-72;
Simplimorpha-Scoble-1983 ( Nepticulidae- ) : Emended-diagnosis, P.-45;
Simplimorpha-promissa ( Staudinger-1870 ) ( Nepticulidae- ) : Redescription-, P.-46;
Stigmella-Schrank-1802 ( Nepticulidae- ) : Emended-diagnosis, P.-55;
Trifurcula-Zeller-1848 ( Nepticulidae- ) : Emended-diagnosis, P.-63;
Trifurculini- ( Nepticulidae- ) : Emended-diagnosis, P.-58;
Enteucha-Meyrick-1915 ( Lyonetiidae- ) : Emended-diagnosis, P.-49
... These two form the genus Ectoedemia in the sense of Wilkinson & Scoble (1979) and Wilkinson & Newton (1981). The concept of Ectoedemia was recently enlarged by Scoble (1983) to contain the subgenera Fomoria Beirne and Laqueus Scoble, and one more subgenus will be included in a forthcoming generic revision of Holarctic Nepticulidae ( Van Nieukerken, in preparation). An up-to-date survey of the Western Palaearctic species assigned to the subgenera of Ectoedemia not treated here, will be presented by Van Nieukerken (in press). ...
... Most other species however bear a brush of hair-scales instead, arising near the frenulum, which is believed to be homologous with the costal bristles. Following Scoble (1983) complete dorsal surface of the hindwing, as in terebinthivora or heringella (figs. 53, 62). ...
... The enlarged portion of the vagina is referred to as the vestibulum, this part has earlier been regarded as part of the ductus bursae (Scoble, 1983) and sometimes termed colliculum (Wilkinson & Scoble, 1979;Wilkinson & Newton, 1981). Here the term ductus bursae is reserved for the narrowed Dart anterior of the entrance of the ductus spermathecae. ...
The subgenera Zimmermannia Hering and Ectoedemia s.str., together forming the genus Ectoedemia Busck sensu Wilkinson & Newton (1981) are described and redefined, and the Western Palaearctic species are revised. In total 50 species are recognised, including the new species hispanica, monemvasiae, nuristanica in Zimmermannia and andalusiae, algeriensis, leucothorax, alnifoliae, contorta and two unnamed species in Ectoedemia s.str. Fifteen new synonymies and ten new combinations are established and 42 lectotypes are designated. Primary types have been examined in many cases. Data on larvae and biology are included and keys to all species are provided. The monophyly and the sister group relationships of both subgenera are demonstrated. The subgenus Ectoedemia can be divided into the populella group, suberis group, subbimaculella group and occultella group, being monophyletic entities, and the possibly paraphyletic angulifasciella group. Two alternative hypotheses of the phylogeny within Ectoedemia s.str. are presented. Decisions on species discrimination have in many cases been corroborated by study of allozymes.
... The genus Etainia Beirne, 1945(¼Obrussa Braun, 1915, a small taxon of the family Nepticulidae (pygmy moths), has been described by B. P. Beirne based on the European Lyonetia sericopeza Zeller possessing two large apodemes in the male genitalia (Beirne, 1945). Because of striking morphological automorphies, as well as the unique biology of the species (Puplesis, 1994;Puplesis and Di skus, 1996), the genus was usually accepted and treated as a "good" genus (Braun, 1915;Wilkinson and Scoble, 1979;Scoble, 1983;Puplesis and Ivinskis, 1985;Puplesis, 1994;Puplesis and Di skus, 1996, 1997, 2003Sinev, 2008;Navickaite et al. 2011). Nevertheless, some researchers treated Etainia as a subgenus of Ectoedemia Busck ( van Nieukerken, 1986;Johansson et al. 1990; van Nieukerken and La st uvka, 2002;Hirano, 2013) because the species of Etainia share some characteristics with other genera (Laqueus, Fomoria, Zimmermannia) previously considered a subgenus of Ectoedemia and their larvae are similar to other Ectoedemia species. ...
... In total, 17 species of Etainia have been known worldwide to date, which have been well investigated in the West Palearctic region (Johansson et al. 1990;Puplesis, 1994), as well as recorded in Central Asia (Puplesis and Di skus, 2003), East Asia (Puplesis and Ivinskis, 1985;Puplesis, 1994;Yagi and Hirowatari, 2017), North America (Wilkinson and Scoble, 1979), and South Africa (Meyrick, 1918;Scoble, 1983). So far, the knowledge of the ecology of Etainia has been based on the European and North American fauna: larvae of these species mine in fruits (samaras) and buds of Acer (Aceraceae) and Arctostaphylos (Ericaceae) (Jäckh, 1951;Emmet, 1976;Johansson et al. 1990;Puplesis and Di skus, 2003). ...
The genus Etainia is reported from Korea for the first time, with four newly recorded species: Etainia capesella (Puplesis), E. peterseni (Puplesis), E. sabina (Puplesis), and E. trifasciata (Matsumura). The images and identification key of the adults, and male genitalia of the species are provided, and DNA barcodes for precise identification of the species are also given.
... In the specialized literature on Lepidoptera, the superfamily Nepticuloidea comprises some of the smallest Lepidoptera known (Davis 1999;Davis & Stonis 2007;Stonis et al. 2020d). Moreover, in Nepticulidae, or pygmy moths, their size has been emphasized, i.e., containing "some of the smallest lepidopterans" (Scoble 1983) or calling nepticulids "the smallest lepidopterans in the world" (Puplesis 1994;Diškus & Stonis 2012;Stonis et al. 2020c). ...
... The actual data were retrieved from our exhaustive species database with measurements, as well as from published references. A few of them are listed further in the text, and a small, freely selected portion of further uncited references are provided here as principally important papers for the current study: Wilkinson (1979); Wilkinson & Scoble (1979); Newton & Wilkinson (1982); Scoble (1983); Kemperman & Wilkinson (1985); Donner & Wilkinson (1989); van Nieukerken (1985avan Nieukerken ( , 2007avan Nieukerken ( , 2007bvan Nieukerken ( , 2019; Hoare et al. (1997); Hoare (2000aHoare ( , 2000b; Puplesis & Robinson (2000); van Nieukerken & Liu (2000); ; ; ; and Hoare & van Nieukerken (2013). Recently, based on new discoveries of Nepticulidae in the Neotropics, many new measurement data became available because Stonis et al. (2013aStonis et al. ( , 2013bStonis et al. ( , 2016Stonis et al. ( , 2017aStonis et al. ( , 2017bStonis et al. ( , 2017cStonis et al. ( , 2018aStonis et al. ( , 2018bStonis et al. ( , 2018cStonis et al. ( , 2020b; ; , and many others (a few are cited further in the text). ...
The world’s smallest moths in Lepidoptera (Insecta) and the complexity in making such a determination are examined and discussed. The forewing length and wingspan of 650 species were measured and the same data were retrieved from published papers to determine which species and family have the smallest moths in the world. The minimal recorded forewing length was found to be around 1.2–1.3 mm and the wingspan around 2.6–2.8 mm in two families, the Gracillariidae and Nepticulidae. Among Lepidoptera, the following species have the smallest moths globally: the European Johanssoniella acetosae (Stainton), the Peruvian Simplimorpha kailai Stonis & Diškus, the Mexican Stigmella maya Remeikis & Stonis, the Mediterranean S. diniensis (Klimesh), the Mediterranean Parafomoria liguricella (Klimesh) (Nepticulidae), the South East Asian Porphyrosela alternata Kumata, and the Central African P. desmodivora De Prins (Gracillariidae). Additionally, in the Nepticulidae, we provide a measurement update for Stigmella maya Remeikis & Stonis, one of the tiniest species with a forewing length of 1.3 mm and wingspan of 2.8 mm, and describe a new species, Stigmella incaica Diškus & Stonis, sp. nov., with a forewing length of 1.75 to 1.95 mm and a wingspan of 3.8 to 4.3 mm.
... Male genitalia (Figs. 2,(12)(13)(14)(15)(16)(17)(18)(19)(20)(21)(22)(23)(24)(25)(26)(27)(28)(29)(30). Capsule longer (285-380 μm) than wide (140-280 μm). ...
... This distinctive, highly apomorphic species was recently described and illustrated by Stonis et al., 2017: Figs. 2, 9, 33, 95, 96 (note the unusually extended cathrema, illustrated in Fig. 81 Discussion. The genus is characterized by less reduced forewing venation (see Scoble, 1983: Fig. 12), deeply divided valva, elongated cathrema, and the absence of cornuti in the male genitalia. Female genitalia without vaginal sclerites. ...
We describe one new genus (Brachinepticula Stonis & Diškus, gen. nov.) with two new species (B. plurilobata Diškus & Stonis, sp. nov., B. elongata Remeikis & Stonis, sp. nov.), and one species with uncertain taxonomic position (Johanssoniella bina Remeikis & Stonis, sp. nov.). We also provide diagnostic characters and an updated annotated catalogue of the following related genera: newly restored Johanssoniella Koçak, Brachinepticula gen. nov., Enteucha Meyrick (s. str.) and Manoneura Davis. In the Annotated Catalogue, we provide five new combinations and new data on morphology, biology, and distribution of some species, including the first photographic documentation of Manoneura basidactyla Davis discovered in the Amazon rainforest, results of re-examination of the male genitalia of the European Johanssonia acetosae (Stt.), also the first documentation of the male genitalia, host plant and leaf mines of the little known Johanssoniella diplocosma (Meyrick) from the Himalayas. All new taxa treated in the paper are illustrated with drawings and (or) photographs of the adults and genitalia; photographs of the leaf mines of Brachinepticula plurilobata Diškus & Stonis, sp. nov. and Johanssoniella diplocosma (Meyrick) are also provided.
... not as a subgenus), e.g. by Wilkinson & Scoble (1979); Puplesis (1985Puplesis ( , 1992Puplesis ( , 1994; Puplesis et al. (1996); Puplesis & Robinson (2000); ; ; Navickaitė et al. (2011Navickaitė et al. ( , 2014a; Stonis et al. (2016Stonis et al. ( , 2017; Remeikis (2017). In some other publications Glaucolepis (=Fedalmia Beirne) was treated as a subgenus of Trifurcula Zeller, 1848 (e.g., Scoble 1983;van Nieukerken 1986a, b;Johansson et al. 1990;Laštůvka & Laštůvka, 2000;van Nieukerken et al. 2004;Laštůvka et al. 2007Laštůvka et al. , 2013Ivinskis et al. 2012). Recently, Neotrifurcula van Nieukerken, 2016, was erected (van Nieukerken et al. 2016b) based on the species N. gielisorum. ...
... Fedalmia Beirne, 1945. The synonymy suggested by R. Johansson (unpublished), discussed in Scoble (1983) and provided by Puplesis 1985: 11. Type species: Nepticula headleyella Stainton, 1854: 298. ...
We provide diagnostic characters for the genus Glaucolepis Braun, re-examine the type series of the type species of the North American G. saccharella Braun, describe two new species from Chile and Argentina (G. flagellata Remeikis & Stonis, sp. nov. and G. pseudoflagellata Remeikis & Stonis, sp. nov.), and provide the first photographic documentation of the central Andean G. aerifica (Meyrick). We synonymize Neotrifurcula van Nieukerken, 2016, syn. nov. with Glaucolepis and provide one new combination for the south Andean G. gielisorum (van Nieukerken, 2016), comb. nov. All species treated in the paper are illustrated with drawings and (or) photographs of the adults and genitalia.
... They are a specialized but phylogenetically primitive Lepidoptera family which comprises the smallest moth on Earth. The family was extensively characterized in monographic reviews by Scoble (1983), Johansson et al. (1990), Puplesis (1994), and Puplesis, Diškus (2003), with special reference to South America also by Puplesis, Robinson (2000) and Stonis et al. (2016c). One of the most prominent characteristics of the Nepticulidae is their larval biology. ...
... Male (Figs. 10,11). Forewing length about 2.3 mm; wingspan about 5.1 mm. ...
The paper presents the first documentation of Urticaceae-feeding Nepticulidae species in South America and describes four new species: two species feeding on Phenax Wedd. (Stigmella singularia Diškus & Stonis, sp. nov. and S. lata Diškus & Stonis, sp. nov.), one species on Boehmeria Jacq. (S. boehmeriphaga Diškus & Stonis, sp. nov.), and one species on Pilea Lindl. (S. auripurpurata Diškus & Stonis, sp. nov.); all from the equatorial Andes. In addition, leaf-mines of an unknown Stigmella taxa feeding on Phenax are documented. The newly discovered Urticaceae-feeding Nepticulidae exhibit some morphological and taxonomical diversity: two species groups, Stigmella singularia and S. marmorea, are revealed (the latter is designated in the current paper).
... The nepticulid subgenus Etainia (in the genus Ectoedemia), often regarded as a separate genus (Scoble 1983; Puplesis 1994; Puplesis & Diškus 1996) is one of the best characterized monophyletic entities within the family, best characterized by the unique dorsal apodeme on the valve in the male genitalia. It is also rather peculiar in its biology , since the species are not leaf-miners, but – as far as known in the Holarctic fauna – feed in shoots, petioles or fruits, most on Acer (Aceraceae) and one species on Arctostaphylos (Ericaceae). ...
... Previously only one species was recorded from this family: Ectoedemia (Fomoria) oleivora Vári, feeding in Olea chrysophylla Lamk. (Vári 1955; Scoble 1983); it is not closely related. Most species of Etainia, where the biology is known, feed on Acer species (Aceraceae or Sapindaceae in the system of Bremer et al. 1998). ...
Ectoedemia (Etainia) obtusa (Puplesis & Diškus), described from Turkmenistan, is for the first time recorded from Europe: Spain, France, Italy and Croatia. It has been reared from cocoons, partly found on trunks of Fraxinus ornus L., which is considered to be its probable host. The female is described here for the first time and the male redescribed and illustrated. A checklist and key of the seven Western Palaearctic species of the subgenus are provided.
... The cosmopolitan plant-mining lepidopteran family Nepticulidae (commonly called pygmy moths) comprises over 1005 species worldwide (Dobrynina et al. 2022;Stonis et al. 2022) and is characterized by a highly pronounced stenophagy, i.e., the vast majority of species are monophagous feeding on a single plant species or are true oligophagous feeders trophically associated with a few plant genera of the same plant family (for a schematic visualization of phytophagous feeders, see Stonis et al. 2016). So far, only one species of Nepticulidae, a species from the genus Simplimorpha Scoble (Scoble 1983), is known trophically associated with Cotinus coggygria Scop. (Puplesis & Diškus 2003). ...
In this paper, the results of a recent molecular study of the Cotinus-feeding, leaf-mining Simplimorpha promissa (Staudinger) (Lepidoptera: Nepticulidae) are discussed for the first time on the basis of samples collected in Ukraine and Armenia. Newly obtained mtDNA CO1-5' sequences from these countries were compared with previously known sequences from France, Italy, Croatia and Greece. A mitotype network and a Neighbor-Joining tree of twenty-two specimens of S. promissa are provided. They show that the studied specimens from Ukraine and the rest of the European countries are genetically closer to each other than to the examined specimens from Armenia, thereby suggesting the Armenian specimens could represent a sister taxon to the European S. promissa. The study also revealed a significantly large genetic diversity of S. promissa in Ukraine.
... Brachinepticula Diškus & Stonis, 2018 is a small and still poorly investigated genus of pygmy moths (Nepticulidae) (for the updated generic composition of the Neotropical fauna of pygmy moths see Stonis et al. 2022 and some general information on the family Nepticulidae can be found in Scoble 1983;Johansson et al. 1989). ...
This paper describes two new species of Brachinepticula Stonis & Diškus: B. melania Remeikis, Mey & Stonis, sp. nov. and B. colombica Remeikis, Mey & Stonis, sp. nov. Both the new species were collected in the Northern Andean Páramo (Cundinamarca, Colombia). Since the specimens were caught at night-time with a light trap, the host plants remain unknown. The examination of the morphology of the male genitalia of B. melania and female genitalia of B. colombica revealed a highly distinctive new taxa of pygmy moths and broadened our knowledge of the morphology and distribution of the recently described endemic genus Brachinepticula Diškus & Stonis. The examination of the new findings also allowed us updating the diagnosis of Brachinepticula. The new species were illustrated with photographs of the genitalia, adults, and habitats.
... Esta familia, con cerca de 862 especies descritas en 22 géneros, incluye los lepidópteros más pequeños, para la región Neotropical se registran 123 especies y 9 géneros (van Nieukerken, Doorenweerd, Hoare, Davis, 2016). Los adultos son polillas más bien pardas, fuscas u ocres o típicamente metálicas con una o más bandas brillantes plateadas y un fondo iridiscente dorado o purpúreo (Scoble, 1983). En promedio, la envergadura alar alcanza cerca de 5-6 mm, pero en algunas especies está por debajo de los 4 mm. ...
Se brinda un inventario preliminar con datos de distribución, hospederos y hábitat de los lepidópteros acuáticos y semiacuáticos registrados en Argentina. El mismo se basa en un relevamiento intensivo de datos bibliográficos,especímenes depositados en la Colección Instituto – Fundación Miguel Lillo (IFML) y registros propios de relevamientos de campo. El objetivo del presente trabajo es presentar la diversidad de familias, géneros y especies de Lepidópteros acuáticos que se registran hasta el presente en Argentina aportando información de la distribución geográfica, hospederos,hábitos y de biología. Comprende datos de 11 familias (Coelophoridae,Cosmopterigidae,Crambidae, Erebidae,Gracillaridae, Momphidae, Pterophoridae, Pyralidae, Nepticulidae, Sphingidae y Tortricidae), 26 géneros y 44 especies. La subfamilia Acentropinae (Crambidae) es la más diversa con 11 géneros y 19 especies para Argentina. La mayor diversidad registrada fueron: el género semiacuático Paracles (Erebidae) con 9 especies registradas y Parapoynx (Crambidae) con 5 especies acuáticas. Se citan por primera vez para Argentina los géneros y especies de: Aulacodes sp. (Guenée, 1854), Argyractoides sp. (Lange, 1956), Oligostigmoides sp. (Lange, 1956), Oxyelophila sp. (Forbes, 1922), Petrophila cronialis (Druce, 1896), Petrophila longipennis (Hampson,1906) y Usingeriessa sp. Lange 1956 (Crambidae). También se brinda una actualización de los géneros de Acentropinae registrados para la región Neotropical y Argentina.
... In the male genitalia, there is a usually well-developed, wide, and strongly chitinized phallus possessing distinctive, often large cornuti. General characterization of this highly specialized, predominantly leaf-mining (occasionally gall-making, e.g., see van Nieukerken et al. 2016b) family was published by several researchers, including Newton & Wilkinson (1982), Scoble (1983), van Nieukerken (1986avan Nieukerken ( , 1986b, Johansson et al. (1990), Puplesis (1994), Puplesis & Robinson (2000), Puplesis & Diškus (2003), Diškus & Stonis (2012), and van Nieukerken et al. (2016avan Nieukerken et al. ( , 2016b. Nepticulidae consists of 18 globally distributed or endemic genera, some well differentiated and easily recognizable, some scarcely diagnosed, and some with their status still in dispute ( van Nieukerken et al. 2016b;Stonis et al. 2017cStonis et al. , 1918cStonis et al. , 2018dStonis et al. , 2019bStonis et al. , 2019c. ...
This study identifies the number of named and described species of three monotrysian, plant-mining lepidopteran families worldwide: Nepticulidae and Opostegidae (Nepticuloidea), and Tischeriidae (Tischerioidea). At the end of 2021, we estimated that a total of 1000 Nepticulidae species, 197 Opostegidae species, and 170 Tischeriidae species have been described since the taxonomic practice of describing species began in the 18th century. We examine and discuss the history of descriptions and authorship of species worldwide for each of the three families. We found that the total (accumulative) number of species described increased with each time period delineated. About five new species were described per year on average, or about 22 new species were described per year in the 21st century. We recognize researchers with the most number of described species in these three taxa.
... Its members occur in very different habitats of the Western Hemisphere, and are particularly common in lowland, tropical, or subtropical premontane habitats of the Neotropics, including the lush tropical forests of Central America, tropical slopes of the Andes, and steamy rainforests of the Amazon. General characterization of this genus was provided by several authors, notably Wilkinson & Scoble (1979), Wilkinson (1979), Scoble (1983), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis & Robinson (2000), and Puplesis & Diškus (2003). The following genitalic characters are of diagnostic importance for recognition of Acalytris: in the male genitalia the capsule usually posseses lateral apodemes, the phallus has large carinae, and usually with minute spine-like cornuti or none cornuti; in the female genitalia the vestibulum usually has a distinct vaginal sclerite (or sclerites and spines). ...
We list all 56 currently known Acalyptris Meyrick species from North and South America, designate five new species groups, and provide pictorial diagnostics for all nine revised species groups of the American fauna. We describe seven new species: A. marmor Stonis & Diškus, sp. nov., A. barbudo Stonis & Remeikis, sp. nov., A. jareki Stonis & Diškus, sp.
nov., A. hilli Stonis & Diškus, sp. nov., A. mortalis Diškus & Stonis, sp. nov., A. hyacinthum Stonis & Vargas, sp. nov., and A. extremus Stonis & Diškus, sp. nov. We provide new data on morphology, biology or distribution for the following species: A. murex Diškus & Stonis, A. hispidus Puplesis & Robinson, A. trifidus Puplesis & Robinson, A. bifidus Puplesis
& Robinson, A. terrificus Šimkevičiūtė & Stonis, and particularly A. yucatani Remeikis & Stonis. We transfer Fomoria miranda Diškus & Stonis to Acalyptris and provide the first photographic documentation of A. novenarius Puplesis & Robinson, A. fortis Puplesis & Robinson, A. martinheringi Puplesis & Robinson, A. basihastatus Puplesis & Diškus, A.
pseudohastatus Puplesis & Diškus, A. articulosus Puplesis & Diškus, A. bovicorneus Puplesis & Diškus, and A. insolentis Puplesis & Diškus. We also comment on the re-deposition of some type series to the collection of the Zoological Museum of the Natural History Museum of Denmark, Copenhagen.
... Some species have been included on lists of cultivated plant pests (e.g., Kuznetzov and Puplesis 1994). A general characterization of this family was provided by several authors, notably Scoble (1983), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis and Robinson (2000), Puplesis and Diškus (2003), Diškus and Stonis (2012), and recently van Nieukerken et al. (2016b). Nepticulidae are distributed worldwide and occur in almost all terrestrial habitats. ...
We describe a new pest of guava ( Psidium guajava L.), Hesperolyra guajavifoliae Stonis & Vargas, sp. nov. , that was recently discovered in western Colombia. Hesperolyra van Nieukerken is a small, Neotropical genus of pygmy moths (Nepticulidae). We re-examine and document the complex morphology of the male genitalia of the generic type species, H. diskusi (Puplesis & Robinson). We discuss the diagnostics and composition of the genus and provide a simple pictorial differentiation scheme for all currently known representatives of the genus. The new species is illustrated with photographs of the adults, some of the immature stages, male and female genitalia, and leaf mines. A link to the COI barcodes of H. guajavifoliaesp. nov. is provided and the relationship of Hesperolyra to other genera is discussed.
... Nepticulid moths are a phylogenetically basal lepidopteran family and are famous for being the world's smallest, with a wingspan often less than 4-5 mm. Useful and extensive characterizations of this family were provided in several monographic reviews, notably those by Newton and Wilkinson (1982), Scoble (1983), Kemperman and Wilkinson (1985), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis and Robinson (2000), Puplesis and Di skus (2003), Di skus and Stonis (2012), and Stonis et al. (2016e). ...
Pygmy moths (Nepticulidae) associated with Asteraceae are poorly known and very rare worldwide. Recently, we discovered many leaf-mining nepticulids in South America feeding on Asteraceae. We review all known records of Asteraceae-feeding Nepticulidae, which in the Neotropics (including the Andes and Patagonia) previously included only Stigmella Schrank. We describe six new species of Stigmella from equatorial South America: S. jungiae Diškus and Stonis, n. sp. (feeding on Jungia L.f.), S. aeneola Diškus and Stonis, n. sp., S. violea Diškus and Stonis, n. sp., S. bracteata Diškus and Stonis, n. sp. (feeding on Liabum Adans.), S. spatiosa Diškus and Stonis, n. sp. (feeding on Ageratina Spach), and S. auripennata Diškus and Stonis, n. sp. (feeding on Baccharis L.). All new taxa are illustrated with photographs of the adults, their genitalia, and leaf mines. Additionally, leaf mines of three unknown nepticulid taxa on Jungia (aff. J. polita Griseb.), Piptocoma discolor (Kunth) Pruski, Gynoxys laurifolia (Kunth) Cass., and G. acostae Cuatrec. are documented for the first time. We diagnose and designate two new species groups and one new species complex in Stigmella. We discuss the origin and diversity of Asteraceae in the Neotropics and illustrate the distribution of Stigmella species feeding on representatives of Asteraceae at the tribal level.
... The family Nepticulidae is characterized in several monographic reviews, notably by Newton & Wilkinson (1982), Scoble (1983), Kemperman & Wilkinson (1985), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis & Robinson (2000), Puplesis & Diškus (2003), Diškus & Stonis (2012) and Stonis et al. (2016d). ...
We provide a report on Nepticulidae feeding on Lamiaceae plants in South America and describe two new species from the Andes: Stigmella lamiacifoliae Remeikis & Stonis, sp. nov., feeding on Salvia palifolia in Colombia, and S. scutellariae Remeikis & Stonis, sp. nov., feeding on Scutellaria volubilis in Ecuador. The leaf mines, adults, genitalia, and habitats of the new species are illustrated.
... The family Nepticulidae was characterized in monographic reviews by Scoble (1983), Johansson et al. (1990), Puplesis (1994), and Puplesis, Diškus (2003), with special reference to South America also by Puplesis, Robinson (2000) and Stonis et al. (2016). ...
The paper reviews the most recent findings of the Rosaceae-feeding Nepticulidae species along with previous records of these tropically specialized leaf-miners in South America and describes three new species: one species on Hesperomeles obtusifolia (Pers.) Lindl (Stigmella circinata Diškus & Stonis, sp. nov.) and two species on Rubus spp. (S. rubiphagiella Diškus & Stonis, sp. nov. and Ectoedemia morae Diškus & Stonis, sp. nov.); all from the equatorial Andes. Additionally, leaf-mines of the unknown Stigmella taxa feeding on Acaena L., Alchemilla L., Rubus L., Prunus L., and Hesperomeles Lindl are documented. Description of previously unknown females of Stigmella nubimontana Puplesis & Diškus and photographic documentation of leaf-mines of S. nubimontana and S. rubeta Puplesis & Diškus are provided for the first time. The discovered Rosaceae-feeding Nepticulidae exhibit morphological and taxonomical diversity: two new species groups, Stigmella imperatoria and S. circinata, are designated; the latter is also compared with the most similar and probably closely related Holarctic S. hemargyrella and S. sorbi groups.
... Microtrichia also show a progressive reduction within this clade (Simonsen, 2001). This result contradicts the earlier division of Nepticulidae into the Australian subfamily Pectinivalvinae versus Nepticulinae (Scoble, 1983;van Nieukerken, 1986;Hoare, 2000). Although a suite of characters supports the combination of the two pectinivalvine genera, Pectinivalva and Roscidotoga Hoare, morphological support for Nepticulinae was not very strong and is partly based on reductions, e.g. of the valval pectinifer and of the number of antennal segments in the larva (two or three in Pectinivalva, one in other Nepticulidae). ...
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.
... Many species are (strict) monophages or oligophages (sensu Menken & Roessingh 1998), and some genera or species groups are specialized on a single plant family. Several studies provide a phylogeny of the family or its subgroups (Scoble 1983, van Nieukerken 1986, Puplesis 1994, Hoare & van Nieukerken 2013, Doorenweerd et al. 2015. ...
With phylogenetic knowledge of Lepidoptera rapidly increasing, catalysed by increasingly powerful molecular techniques, the demand for fossil calibration points to estimate an evolutionary timeframe for the order is becoming an increasingly pressing issue. The family Nepticulidae is a species rich, basal branch within the phylogeny of the Lepidoptera, characterized by larval leaf-mining habits, and thereby represents a potentially important lineage whose evolutionary history can be established more thoroughly with the potential use of fossil calibration points. Using our experience with extant global Nepticulidae, we discuss a list of characters that may be used to assign fossil leaf mines to Nepticulidae, and suggest useful methods for classifying relevant fossil material. We present a checklist of 79 records of Nepticulidae representing adult and leaf-mine fossils mentioned in literature, often with multiple exemplars constituting a single record. We provide our interpretation of these fossils. Two species now are included in the collective generic name Stigmellites: Stigmellites resupinata (Krassilov, 2008) comb. nov. (from Ophiheliconoma) and Stigmellites almeidae (Martins-Neto, 1989) comb. nov. (from Nepticula). Eleven records are for the first time attributed to Nepticulidae. After discarding several dubious records, including one possibly placing the family at a latest Jurassic position, we conclude that the oldest fossils likely attributable to Nepticulidae are several exemplars representing a variety of species from the Dakota Formation (USA). The relevant strata containing these earliest fossils are now dated at 102 Ma (million years ago) in age, corresponding to the latest Albian Stage of the Early Cretaceous. Integration of all records in the checklist shows that a continuous presence of nepticulid-like leaf mines preserved as compression-impression fossils and by amber entombment of adults have a fossil record extending to the latest Early Cretaceous.
... Aedeagus with single, very large dorsal carina (Figs 151, 152). : 69 (originally described as Ectoedemia wilkinsoni Puplesis, 1984a: 122, a junior homonym of Ectoedemia wilkinsoni Scoble, 1983. Morphology update (Figs 153-160): Caudal process of gnathos strongly sclerotized and appearing truncate at apex (Fig. 155). ...
East Asia is famous for its tremendous overall biodiversity, and the Nepticulidae are no exception. The majority of the currently known fauna of Nepticulidae was described in 1984-1987 by several authors, but R. Puplesis is the author of the largest number of new species from the region. Unfortunately, the genitalia of all species from Primorskiy Kray, Sakhalin, and the Kuril Islands were described from temporary slides in glycerol, and therefore, the drawings do not always show all details of genital armature or may be confusing. Without the baseline data providing morphology of genital armature of type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae. In this paper we reexamine and document for the first time with photographs the types of 56 species described by R. Puplesis from the Russian Far East. Details of the morphology is updated for most of the re-examined species.
... Aedeagus with single, very large dorsal carina (Figs 151, 152). : 69 (originally described as Ectoedemia wilkinsoni Puplesis, 1984a: 122, a junior homonym of Ectoedemia wilkinsoni Scoble, 1983. Morphology update (Figs 153-160): Caudal process of gnathos strongly sclerotized and appearing truncate at apex (Fig. 155). ...
East Asia is famous for its tremendous overall biodiversity, and the Nepticulidae are no exception. The majority of the currently known fauna of Nepticulidae was described in 1984–1987 by several authors, but R. Puplesis is the author of the largest number of new species from the region. Unfortunately, the genitalia of all species from Primorskiy Kray, Sakhalin, and the Kuril Islands were described from temporary slides in glycerol, and therefore, the drawings do not always show all details of genital armature or may be confusing. Without the baseline data providing morphology of genital armature of type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae. In this paper we re-examine and document for the first time with photographs the types of 56 species described by R. Puplesis from the Russian Far East. Details of the morphology is updated for most of the re-examined species.
... In the western part of the Amazon basin, this genus comprises about 50% of the fauna (Puplesis et al. 2002a, b). A great diversity of Acalyptris is also known from South Africa (Scoble 1983). The genus is also abundant in desert and steppe regions of Central Asia (Puplesis , 1994Puplesis & Diškus 1995, 2003. ...
Based on the material collected by the authors in the Crimea (Karadag Nature Reserve), Acalyptris platani (Müller-Rutz, 1934) is recorded in Eastern Europe (Ukraine) for the first time. It represents the easternmost record of this Sub-mediterranean nepticulid species in Europe, once again confirming faunal links of southern Crimea with Mediterranean region. Adults, male and female genitalia, and leaf-mines of A. platani are illustrated and provided together with a distribution map.
... Outside the Holarctic an impressive, albeit still very incomplete, amount of information on micro-moth biology/immatures has been gathered in Australia (much of it only published in outline, see Common 1990) and New Zealand (Hudson 1928/39, Donner & Wilkinson 1989. Important progress in the study of some groups of micro-moth leafminers has been made in Southern Africa (Vari 1961, Scoble 1983), but otherwise the knowledge of micro-moth immatures/biology in the Afrotropical as well as Neotropical regions remains grossly inadequate; the same is true for the Oriental region, where the major publications by T. B. Fletcher in the early decades of the 20 th century only mark a beginning. The remaining Ditrysia, which include some very species-rich superfamilies, are mostly medium-sized to large, frequently rather broad-winged insects with predominantly exophagous larvae. ...
The currently recognized robust support for the monophyly of the Lepidoptera (and the superorder Amphiesmenoptera comprising Lepidoptera + Trichoptera) is outlined, and the phylogeny of the principal lineages within the order is reviewed succinctly. The state of the taxonomic inventory of Lepidoptera is discussed separately for 'micro-moths', 'macro-moths' and butterflies, three assemblages on which work has followed historically somewhat different paths. While currently there are about 160,000 described species of Lepidoptera, the total number of extant species is estimated to be around half a million. On average, just over one thousand new species of Lepidoptera have been described annually in recent years. Allowing for the new synonyms simultaneously established, the net increase in species numbers still
... The family Nepticulidae occurs worldwide on all major landmasses, exceptions being only the cold Antarctica and Greenland. Although our knowledge of these tiny leafmining moths is still biased considerably towards the Holarctic region (see references in Hoare et al. 1997), more information is becoming available for (sub)tropical regions (Scoble 1983;Hoare et al. 1997;Hoare 2000aHoare , 2000bvan Nieukerken and Liu 2000;Puplesis and Robinson 2000). A paper summarising our knowledge for the Oriental region is in preparation by the senior author. ...
Stigmella ebbenielseni, sp. nov. is described from Guam, where it was reared from leafmines on Pipturus argenteus (G. Forst.) Wedd. (Urticaceae). Vacated leafmines of this or a related species have also been recorded from Tinian and Alamagan. These provide the first records of the Nepticulidae from Micronesia. Similar mines, probably of a related species, have been recorded from herbarium material of Maoutia australis Wedd. originating from Tahiti; these confirm the occurrence of the family in Polynesia. Adult and mines of S. ebbenielseni are described and illustrated. The literature on Microlepidoptera of Pacific islands is reviewed and previous records of Nepticulidae are discussed. We conclude that Stigmella is probably widespread in the Pacific area, with at least four species, currently all associated with Urticaceae, tribe Boehmeriae.
... In the male genitalia viridissimella shows more resemblance to some South African species, placed in Fomoria (see Scoble, 1983), than to any other. This supports the placement in the present subgenus, previously based on females only ( Van Nieukerken, 1986b). ...
The two Nepticulidae, described by A. Caradja, are reexamined. The following synonymies are established: Ectoedemia (Ectoedemia) rufifrontella (Caradja, 1920) comb. nov. as senior synonym of Nepticula nigrosparsella Klimesch, 1940 (syn. nov.); Ectoedemia (Fomoria) viridissimella (Caradja, 1920) comb. nov. as senior synonym of Nepticula nowakowskii Toll, 1957 (syn. nov.). E. viridissimella is redescribed, and the genitalia are figured.
Ectoedemia- ( Ectoedemia- ) rufifrontella- ( Caradja-1920 ) ( Nepticulidae- ) : Comb-nov, Transferred-from, Trifurcula-, p.-142, Syn-nov, Nepticula-nigrosparsella-Klimesch-1940, p.-142;
Ectoedemia- ( Fomoria- ) viridissimella- ( Caradja-1920 ) ( Nepticulidae- ) : Redescription-, P.-142, Comb-nov, Transferred-from, Nepticula-, p.-142, Syn-nov, Nepticula-nowakowskii-Toll-1957, p.-142;
Nepticula-nigrosparsella-Klimesch-1940 ( Nepticulidae- ) : New-synonym, Of-Ectoedemia ( Ectoedemia- ) rufifrontella- ( Caradja-1920 ) , p.-142;
Nepticula-nowakowskii-Toll-1957 ( Nepticulidae- ) : New-synonym, Of-Ectoedemia ( Fomoria- ) viridissimella- ( Caradja-1920 ) , p.-142;
Nepticula-viridissimella-Caradja-1920 ( Nepticulidae- ) : Referred-to, Ectoedemia- ( Fomoria- ) , p.-142;
Trifurcula-rufifrontella-Caradja-1920 ( Nepticulidae- ) : Referred-to, Ectoedemia- ( Ectoedemia- ) , p.-142
In the launched electronic Nepticulidae species identification tool (https://www.nepticulidaespecies.info), nine of ten sections of this tool are dedicated to morphology: one section is about adult forewing pattern and eight other about various structures of the male genitalia. This guide omits such characters like the adult’s head scaling, wing venation, morphology of the female genitalia, as well various other morphological characters, because
they were purposely unincluded into the launched identification tool.
The publication deals with 228 species of Nepticulidae (or pygmy moths) from the Neotropics (i.e., the Neotropical and Ando-Patagonian regions) and adjacent areas. The species are illustrated with photographs or drawings of adult forewings, the male genitalia and, if available, leaf mines. In addition to the illustrated species review, the monograph announces about the launching of the first electronic diagnostic system (a free electronic tool on the internet) designed for the identification of species of Nepticulidae and includes chapters on the global biodiversity inventory, the collecting of adults and larvae of pygmy moths, preparation of micro-mounts of genitalia structures of Nepticulidae, global generic composition of the family, some particularities of Nepticulidae biology, a unique morphology guide based on the forewing pattern and male genitalia, descriptions of four newly named species, and a taxonomic checklist of the species currently known from the Neotropics and adjacent areas.
The genus Simplimorpha Scoble is recognized for the first time from South America. We describe a subgenus, Myrtinepticula Stonis & Diškus, subgen. nov., for three new species from the southern Andes (Chile and Argentina): Simplimorpha (Myrtinepticula) cercaria Diškus & Stonis, sp. nov., S. (M.) nielseni Remeikis & Stonis, sp. nov., S. (M.) sapphirella Remeikis & Stonis, sp. nov.; and one new species from the southwestern Amazon (Peru): S. (M.) kailai Stonis, sp. nov. We provide a pictorial differentiation scheme for Simplimorpha Scoble and Pectinivalva Scoble. We synonymize the recently erected, predominantly Australian Menurella Hoare, syn. nov. and Cosanovula Hoare, syn. nov. with Pectinivalva Scoble. We also revise the taxonomic status of the Australian Roscidotoga Hoare as a subgenus of Simplimorpha which now exhibits a Gondwanan distribution in the Southern Hemisphere, with the presence of a single species in the Mediterranean. All new taxa are illustrated with photographs of the adults and the genitalia; the leaf mines of Simplimorpha (Myrtinepticula) nielseni sp. nov. are also provided.
The monograph treats 29 species of leaf-mining pygmy moths (Insecta, Lepidoptera, Nepticulidae) discovered in the northern Andean bush and grass páramo and the central Andean puna at altitudes above 3700 m. They represent the world’s highest-altitudinal Nepticulidae fauna known. The height record belongs to Stigmella nivea sp. nov. from Peru collected at an elevation of 4700 m. Except for one species, all belong to Stigmella Schrank. Twenty-two of the species recorded are new and are named and described in the current paper: Stigmella paramica Diškus & Stonis, sp. nov.; S. lachemillae Diškus & Stonis, sp. nov.; S. gynoxyphaga Diškus & Stonis, sp. nov.; S. calceolariae Diškus & Stonis, sp. nov.; S. rigida Diškus & Stonis, sp. nov.; S. altiplanica Diškus & Stonis, sp. nov.; S. robusta Remeikis & Stonis, sp. nov.; S. pseudorobusta Remeikis & Stonis, sp. nov.; S. auriargentata Remeikis & Stonis, sp. nov.; S. altimontana Remeikis & Stonis, sp. nov.; S. pandora Remeikis & Stonis, sp. nov.; S. ampla Diškus & Stonis, sp. nov.; S. evanida Diškus & Stonis, sp. nov.; S. mustelina Remeikis & Stonis, sp. nov.; S. angusta Diškus & Stonis, sp. nov.; S. alticosma Remeikis & Stonis, sp. nov.; S. nivea Remeikis & Stonis, sp. nov.; S. kristenseni Diškus & Stonis, sp. nov.; S. lobata Remeikis & Stonis, sp. nov.; S. ageratinae Diškus & Stonis, sp. nov.; S. clinopodiella Diškus & Stonis, sp. nov.; and S. calceolarifoliae Diškus & Stonis, sp. nov. Some of these species are leaf-miners on Asteraceae (Pentacalia, Baccharis, Gynoxys, and Ageratinaplants), Calceolariaceae (Calceolaria), Lamiaceae (Clinopodium), and Rosaceae (Lachemilla). Twenty species are known only from adults with no data on their biology and host-plants. In addition, we present data and discuss recently discovered nepticulid taxa associated with Polylepis forests that is the natural vegetation in much of the High Andes. All High-Andean Stigmella species treated are illustrated with photographs of the adults and genitalia, distribution maps, including some with photographs of the leaf-mines and habitats.
Previous information on the external galea morphology of primitive Lepidoptera-Glossata is reviewed. Approximately 40 species are newly examined, and scanning electron micrographs are now available for all families of non-ditrysian glossatan moths with a functional proboscis. Principal objectives of the study are the reconstruction of the groundplan features of the proboscis of Glossata, and of the evolution of this organ in higher Lepidoptera. Plesiomorphic characters of the galea of Glossata include a microtrichiated external galeal wall, a non-microtrichiated food groove composed of finely fluted plates, dorsal galeal linking structures ('dorsal legulae') arranged horizontally and multipointed ventral legulae extending from the lower anterior margins of the food groove plates. Sensory equipment comprises at least a few sensilla trichodea (restricted to the external galea wall), and uniporous sensilla basiconica on both the external galea surface and the median food groove. Evolutionary trends are primarily identifiable in the linking structures and the sensory organs of the galeae. A third type of sensory organ, sensilla styloconica, an additional ('secondary') set of ventral legulae (arising below the food groove), and pilifers bearing bristles are apparently groundplan autapomorphies of a clade comprising all non-nepticuloid Heteroneura. A tightening of the linkage of the galeae and elaborated sensory equipment, including three types of proboscis sensilla, are interpreted as key innovations in the evolution of a long proboscis adapted for flower-probing.
Genera and previously described species of Nearctic Scythrididae are revised for the first time, based on the study of adult structures. About 90 percent of the Nearctic fauna known in collections consists of undescribed species. The supraspecific taxa treated in this work encompass less than half of the Nearctic species diversity. Only six new species are described, all within the largest and structurally most diverse genus. The status of all nominal species is revised. Valid species are redescribed and their features illustrated. General problems in the systematics of the Scythrididae are discussed. A description of adult features of the family Scythrididae is providad. Extra-limital genera are briefly reviewed. A key to the Nearctic genera and informal supraspecific lineages is provided.Six genera, including three new, are treated: Areniscythris Powell, 1976, Arotrura Walsingham, 1888, Asymmetrura gen. nov., Neoscythris gen. nov., Rhamphura gen. nov., and Scythris s. str. Hübner, [1825]. Areniscythris includes a single described species, Areniscythris brachypteris Powell, but is defined more broadly to account for a number of undescribed species. Arotrura is divided into nine informal species groups with the following included species: Arotrura atascosa sp. nov., Arotrura balli sp. nov., Arotrura divaricata (Braun) comb, nov., Arotrura eburnea Walsingham, Arotrura formidabilis sp. nov., Arotrura hymenata sp. nov., Arotrura longissima sp. nov., Arotrura oxyplecta (Meyrick) comb, nov., Arotrura powelli sp. nov., and Arotrura sponsella (Busck) comb. nov. Asymmetrura includes: Asymmetrura albilineata (Walsingham) comb. nov., Asymmetrura graminivorella (Braun) comb. nov., Asymmetrura impositella (Zeller) comb. nov. and type species, Asymmetrura matutella (Clemens) comb, nov., Asymmetrura reducta (Braun) comb, nov., and Asymmetrura scintillifera (Braun) comb. nov. Neoscythris includes: Neoscythris confinis (Braun) comb, nov., Neoscythris euthia (Walsingham) comb. nov., Neoscythris fissirostris (Meyrick) comb. nov. and type species, and Neoscythris planipenella (Chambers) comb. nov. Rhamphura includes: Rhamphura altisierrae (Keifer) comb, nov., Rhamphura ochristriata (Walsingham) comb. nov. and type species, Rhamphura perspicillella (Walsingham) comb. nov., Rhamphura suffusa (Walsingham) comb. nov., and the extra-limital Rhamphura immunis (Meyrick) comb. nov. from Peru. Scythris s. str. includes: Scythris immaculatella (Chambers) rev. stat., Scythris limbella (Fabricius), Scythris mixaula Meyrick, Scythris trivinctella (Zeller), and Scythris ypsilon Braun. A further eight species are phylogenetically distinct from Scythris s. str. but provisionally are only assigned to five informal monophyletic lineages until their cladistic relationships are more firmly established. These are: the Scythris basilaris lineage including Scythris basilaris (Zeller), Scythris eboracensis (Zeller), and Scythris fuscicomella (Clemens); the Scythris interrupta lineage including Scythris interrupta Braun; the Scythris inspersella lineage including Scythris inspersella (Hübner) and Scythris noricella (Zeller); the Scythris anthracina lineage including Scythris anthracina Braun; and the Scythris charon lineage including Scythris charon Meyrick. Three species are incertae sedis: Scythris inornatella (Chambers) comb, nov., Scythrispilosella (Zeller), and Scythris piratica Meyrick. Coleophora albacostella Chambers and Coleophora inornatella Chambers are transferred from the Coleophoridae. Scythris arizoniella (Kearfott) is transferred to the Coleophoridae [ Coleophora arizoniella (Kearfott) comb. nov.].The following new synonymy is proposed: Colinita Busck, 1907 = Arotrura Walsingham, 1888; Gelechia aterrimella Walker, 1864 and Scythris epilobiella McDunnough, 1942 = Scythris inspersella [Hübner, (1817)]; Scythris magnatella Busck, 1904 = Scythris noricella (Zeller, 1843); Scythris pacifica McDunnough, 1927 = Scythris immaculatella (Chambers, 1875); Coleophora albacostella Chambers, 1875 and Scythris hemidictyas Meyrick, 1928 = Neoscythris planipenella (Chambers, 1875).A cladistic definition of the family is presented for the first time. The monophyly of the Scythrididae is supported by the following synapomorphies: very narrow ductus bursae, broad ductus seminalis anastomosed with the oviduct and the corpus bursae, lack of signum, unique shape of the apophyses of the metathoracic furca, tarsomeres 1–4 with two subapical spurs, aedeagus ankylosed, and origin of forewing veins R4 and R5 on a common stalk with R4 extended to the costa and R5 to the termen. Relationships of the Scythrididae within the Gelechioidea are discussed. Based on the cladistic analysis of 52 structural characters, phylogenetic relationships of supraspecific taxa are inferred. Two cladograms, one for the genera and one for the species groups of Arotrura , are presented and used in deriving the classification.
Over the few couple of decades, the Nepticulidae of East Asia have been the subject of continuing investigation. Male genitalia of the nepticulid species described from Primorskiy Kray (the Russian Far East) were re-examined in the preceding paper in this journal. The present paper continues our study on the diversity and systematics of the Nepticulidae of East Asia based on a sample of specimens collected in 1974-1990 at various sites of the southern part of Primorskiy Kray and treats 35 species: two new taxa (Ectoedemia ortiva sp. nov. and E. species 219) and 33 other species. Seventeen of them are briefly discussed and illustrated with photographs of male genitalia. Two new synonyms are proposed, and three new distribution records are provided. We also provide an updated checklist of the Nepticulidae of East Asia, which comprises 105 species; 67 species occur in the Russian Far East and 53 in Japan (20 of which occur both in Japan and Russia). Species with Euro-East Asiatic distribution currently comprise 11% of the Japanese fauna and 16% of the continental fauna of the Russian Far East, Primorskiy Kray.
The Ectoedemia (Fomoria) vannifera species-group is reviewed. Three species are recognized from South Africa (E. vannifera (Meyrick), E. fuscata (Janse) and E. hobohmi (Janse)), one from central Asia (E. asiatica (Puplesis)), and one from India (E. glycystrota (Meyrick) comb. n., here redescribed); three new species are described and named from Australia (E. pelops sp. n., E. squamibunda sp. n., and E. hadronycha sp. n.). All species share a striking synapomorphy in the male genitalia: a pin-cushion-like lobe at the apex of the valva. Two of the Australian species and one of the South African species have been reared from larvae mining the leaves of Brassicaceae sensu lato. A phylogeny of all currently recognized species is presented: this taken together with known distribution suggests either that the group is very ancient and antedates the split between the African and Indian parts of Gondwana (ca. 120 million years ago), or that it has dispersed more recently and has been overlooked in large parts of its range.
The simple matching coefficient (per cent similarity) and the coefficients of taxonomic distance are calculated for 47 european Notodontidae basing upon 50 characters. For this a calculator program in basic is given. The obtained dendrograms are used for discussing classification and systematics of the european Notodontidae.
Clearly the ranks of genera and subfamilies are shown in the dendrograms.
Ectoedemia (Etainia) obtusa (Puplesis & Diškus), described from Turkmenistan, is for the first time recorded from Europe: Spain, France, Italy and Croatia. It has been reared from cocoons, partly found on trunks of Fraxinus ornus L., which is considered to be its probable host. The female is described here for the first time and the male redescribed and illustrated. A checklist and key of the seven Western Palaearctic species of the subgenus are provided. Ectoedemia (Etainia) obtusa (Puplesis & Diškus), beschrieben aus Turkmenistan, wird zum erstenmal aus Europa gemeldet, namentlich aus Spanien, Frankreich, Italien und Kroatien. Die Art wurde aus Puppen gezüchtet, die teilweise auf Stämmen von Fraxinus ornus L. gefunden wurden; diese Pflanze wird daher als die wahrscheinliche Futterpflanze angesehen. Das Weibchen wird zum erstenmal beschrieben, und das Männchen aufs neue beschrieben und abgebildet. Eine Checkliste und Bestimmungsschlüssel der sieben westpaläarktischen Arten der Untergattung Etaina werden angegeben und Anmerkungen zum taxonomischen Status von Etaina gemacht. Ectoedemia (Etainia) obtusa (Puplesis & Di škus), décrit de Turkmenistan, est rapportée de l’Europe pour la première fois: provenant d’Espagne, France, Italie et Croatie. Quelques exemplaires étaient élevés des cocons trouvés sur des troncs de Fraxinus ornus L.; cette plante est regardée comme plante-hôte possible. La femelle est décrit pour la première fois, et le mâle est décrit de nouveau et figuré en détail. Nous donnons aussi un liste des sept espèces Ouest-Paléarctiques et un table d’identification.
Xanadoses nielseni, gen. nov., sp. nov. is described from New Zealand, where it is the only native member of the superfamily Incurvarioidea. The larva is a bark-miner, making long tortuous galleries on the trunks of various species of smooth-barked tree and pupating under a raised silk-lined 'cap' of bark. The moth is assigned to the Cecidosidae on the basis of five apomorphies shared with this family, but is considered to represent the basal lineage within the family as it lacks at least four apomorphies shared by all other genera. The distribution of this concept of Cecidosidae (South America, South Africa and New Zealand) confirms it is an ancient group that originated before the break-up of Gondwana in the late Cretaceous. A checklist of cecidosid taxa is provided.
Houdinia flexilissima, gen. nov., sp. nov. is described from peatlands in the North Island of New Zealand. The extremely narrow larvae mine and pupate in the living stems of Sporadanthus ferrugineus de Lange, Heenan & Clarkson, (Restionaceae), a large endemic rush. Eggs, larvae, pupae and adults are fully described and illustrated. The systematic placement of this autapomorphic taxon is discussed in detail, and it is assigned within Gelechioidea to Batrachedridae on the basis of characters shared with a taxon currently placed in Batrachedra. Because of its morphological distinctiveness, as well as threats to the habitat of its very local host plant, H. flexilissima, gen. nov., sp. nov. is considered a species of high conservation status.
Adult and larval characters place the monophyletic Ectemnorhinus-group of genera firmly within the former Adelognatha and new concept of Brachycerinae. The group, previously with tribal or subfamily status, is most closely related to the genera Heterexis Broun and Oclandius Blanchard from the Auckland and Campbell Islands. A cladistic analysis of the genera based on 35 adult and 11 larval characters, resulted in the following six genera being recognised: Christensenia Brinck, Canonopsis C. O. Waterhouse, Disker Dreux & Voisin, Palirhoeus Kuschel, Bothrometopus Jeannel and Ectemnorhinus G. R. Waterhouse. Ectemnorhinus angusticollis (C. O. Waterhouse) is transferred to Bothrometopus, Mesembriorrhinus parvulus Jeannel, also transferred to Bothrometopus, is reinstated to species level. Ectemnorhinus curtus Jeannel is placed in synonymy with Ectemnorhinus fuscus Enderlein, which is removed from synonymy with Ectemnorhinus viridis G. R. Waterhouse and reinstated as valid. Ectemnorhinus hibernantium (Dreux & Voisin) is placed in synonymy with E. geniculatus Jeannel.
In 1957, some officers of the Forest Insect Survey of Canada discovered a species of
Nepticula
mining the leaves of white birch (
Betula papyrifera
Marsh.) and yellow birch (
Betula lutea
Michx. f.) in Ontario. At that time the species was considered to be undescribed, but to confirm this it was necessary to study the life-history. This has been done by Mr. O. H. Lindquist, Forest Insect Laboratory, Sault Ste. Marie, Ontario, and the results of his studies are being presented in a companion paper. His investigations confirmed that the species had not been named and the following description is presented.
The phylogeny of the mainly Australian nepticulid genus Pectinivalva Scoble, 1983 is investigated on the basis of morphology, and a division into three monophyletic subgenera is proposed on the basis of these results. These subgenera (Pectinivalva, Casanovula Hoare, subgen. n. and Menurella Hoare, subgen. n. ) are described and diagnosed, the described species of Pectinivalva are assigned to them, and representative new species are described in each: Pectinivalva (Pectinivalva) mystaconota Hoare, sp. n., Pectinivalva (Casanovula) brevipalpa Hoare, sp. n., Pectinivalva (Casanovula) minotaurus Hoare, sp. n., Pectinivalva (Menurella) scotodes Hoare, sp. n., Pectinivalva (Menurella) acmenae Hoare, sp. n., Pectinivalva (Menurella) xenadelpha Van Nieukerken & Hoare, sp. n., Pectinivalva (Menurella) quintiniae Hoare & Van Nieukerken, sp. n., and Pectinivalva (Menurella) tribulatrix Van Nieukerken & Hoare, sp. n.
Pectinivalva (Menurella) quintiniae (from Quintinia verdonii, Paracryphiaceae) is the first known member of the genus with a host-plant not belonging to Myrtaceae. Pectinivalva (Menurella) xenadelpha from Mt Gunung Lumut, Kalimantan, Borneo, is the first pectinivalvine reported from outside Australia. Keys to the subgenera of Nepticulidae known from Australia, based on adults, male and female genitalia, and larvae, are presented. Host-plant relationships of Pectinivalva are discussed with relation to the phylogeny, and a list of known host-plants of Pectinivalva, including hosts of undescribed species, is presented. DNA barcodes are provided for most of the new and several unnamed species.
Two new species are described in the monophyletic Ectoedemia (Fomoria) weaveri group: E. (F.) festivitatis spec. nov. from mountains in Nepal, China (Yunnan) and northern Vietnam, feeding on shrubby Hypericum species, and E. (F.) degeeri spec. nov. from Turkey, food plant unknown. The species group is briefly reviewed and a checklist with information on distribution and food plants is provided; the weaveri group comprises 14 species. E. ruwenzoriensis (Bradley, 1965) comb. nov. is added to the group. E. hypericella (Kuroko, 1982) and E. permira (Puplesis, 1984) are recorded for the first time from China and Hypericum is confirmed as food plant for the latter. The reported occurrence of the Canarian endemic E. variicapitella (Chrétien, 1908) outside the Canary Islands is based on an error.
Abstract The six species of Nepticulidae feeding on Cistaceae are placed in Parafomoriagen.n.: the type-species helianthemella (Herrich-Schäffer), cistivora (Peyerimhoff), pseudocistivora sp.n., liguricella (Klimesch), ladaniphila (Mendes) and tingitella (Walsingham). The genus occurs in central Europe and the mediterranean region. Full descriptions, including figures of genitalia, and details of biology are given and a key to the species is provided. A tentative phylogeny is discussed. Primary types have been checked, and five lecto-types have been designated. Stigmella diniensis (Klimesch) is for the first time reported as feeding on Cistaceae, and redescribed.
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