ALBERT

Description: 〈span〉〈div〉Abstract〈/div〉A nepticulid leaf-mine ichnofossil, 〈span〉Stigmellites janggi〈/span〉 Sohn and Nam, n. ichnosp., is described on the basis of a fossil leaf of 〈span〉Fagus〈/span〉 from the early Miocene Geumgwangdong Formation in Pohang Basin. This mine trace is characterized by a linear-blotch type with a clear centric frass trail composed of closely and randomly dispersed pellets filling the mine width in the early stage. We found traces of a possible egg case and an exit slit from the leaf. These features are most consistent with those produced by members of Nepticulidae. Our record demonstrates for the first time the trophic association of Nepticulidae with 〈span〉Fagus〈/span〉 in the Miocene and suggests the persistence of a long-term association between the insect family and the plant genus from the Miocene to the present. Other Nepticulidae leaf mines in the Miocene and leaf-mine fossils from the Geumgwangdong Formation are briefly reviewed.UUID: http://zoobank.org/fff951f6-ca82-4cca-a274-863e0d91acaa〈/span〉

Description: Antispila treitschkiella (Fischer von Röslerstamm, 1843) and A. petryi Martini, 1899, sp. rev. were regarded as synonymous since 1978, but are shown to be two clearly separated species with different hostplants, life histories, DNA barcodes and morphology. Antispila treitschkiella feeds on Cornus mas L., is bivoltine, and has, by following its ornamentally planted host, greatly expanded its range in north-western Europe. In contrast A. petryi feeds on the widespread native C. sanguinea L., is univoltine, and is one of only two Antispila species previously resident in the British Isles, the Netherlands and northern Europe. Consequently, the increase in abundance of A. treitschkiella in the Netherlands since the early 1990s and in Great Britain in recent years must be regarded as part of a recent expansion into north-western Europe, whereas the native A. petryi is hardly expanding and less abundant. In Britain, detailed surveys of parks and living collections confirmed the monophagy of these two species. A search of British herbarium samples provided no evidence for an earlier date of establishment. Information on recognition of all stages, including DNA barcodes, and distribution is provided, and these two species are compared with the third European Cornus L. leafminer, A. metallella (Denis & Schiffermüller, 1775).

Description: Heliozelidae are a widespread, evolutionarily early diverging family of small, day-flying monotrysian moths, for which a comprehensive phylogeny is lacking. We generated the first molecular phylogeny of the family using DNA sequences of two mitochondrial genes (COI and COII) and two nuclear genes (H3 and 28S) from 130 Heliozelidae specimens, including eight of the twelve known genera: Antispila, Antispilina, Coptodisca, Heliozela, Holocacista, Hoplophanes, Pseliastis, and Tyriozela. Our results provide strong support for five major Heliozelidae clades: (i) a large widespread clade containing the leaf-mining genera Antispilina, Coptodisca and Holocacista and some species of Antispila, (ii) a clade containing most of the described Antispila, (iii) a clade containing the leafmining genus Heliozela and the monotypic genus Tyriozela, (iv) an Australian clade containing Pseliastis and (v) an Australian clade containing Hoplophanes. Each clade includes several new species and potentially new genera. Collectively, our data uncover a rich and undescribed diversity that appears to be especially prevalent in Australia. Our work highlights the need for a major taxonomic revision of the family and for generating a robust molecular phylogeny using multi-gene approaches in order to resolve the relationships among clades.

Description: After finding distinct clades in a molecular phylogeny for Nepticulidae that could not be placed in any known genera and discovering clear apomorphic characters that define these clades, as well as a number of Neotropical species that could be placed in known genera but were undescribed, three new genera and nine new species are here described from the Neotropics: Stigmella gallicola van Nieukerken & Nishida, sp. n. reared from galls on Hampea appendiculata (Malvaceae) in Costa Rica, representing the first example of a gall making Stigmella; S. schinivora van Nieukerken, sp. n. reared from leafmines on Schinus terebinthifolia (Anacardiaceae) in Argentina, Misiones; S. costaricensis van Nieukerken & Nishida, sp. n. and S. intronia van Nieukerken & Nishida, sp. n. each from a single specimen collected the same night in Costa Rica, Parque Nacional Chirripó; S. molinensis van Nieukerken & Snyers, sp. n. reared from leafmines on Salix humboldtiana, Peru, Lima, the first Neotropical species of the Stigmella salicis group sensu stricto; Ozadelpha van Nieukerken, gen. n. with type species O. conostegiae van Nieukerken & Nishida, sp. n., reared from leafmines on Conostegia oerstediana (Melastomataceae) from Costa Rica; Neotrifurcula van Nieukerken, gen. n. with type species N. gielisorum van Nieukerken, sp. n. from Chile; Hesperolyra van Nieukerken, gen. n.. with type species Fomoria diskusi Puplesis & Robinson, 2000; Hesperolyra saopaulensis van Nieukerken, sp. n., reared from an unidentified Myrtaceae, Sao Paulo, Brasil; and Acalyptris janzeni van Nieukerken & Nishida, sp. n. from Costa Rica, Guanacaste. Five new combinations are made: Ozadelpha ovata (Puplesis & Robinson, 2000), comb. n. and Ozadelpha guajavae (Puplesis & Diškus, 2002), comb. n., Hesperolyra diskusi (Puplesis & Robinson, 2000), comb. n., Hesperolyra molybditis (Zeller, 1877), comb. n. and Hesperolyra repanda (Puplesis & Diškus, 2002), comb. n. Three specimens are briefly described, but left unnamed: Ozadelpha specimen EvN4680, Neotrifurcula specimen EvN4504 and Neotrifurcula specimen RH2.

Description: 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): Stigmella acerifoliella Dovnar-Zapolski, 1969 (unavailable, = S. acerna Puplesis, 1988), Stigmella nakamurai Kemperman & Wilkinson, 1985 (= S. palionisi Puplesis, 1984), Nepticula amseli Skala, 1941 (unavailable = S. birgittae Gustafsson, 1985), Stigmella cathepostis Kemperman & Wilkinson, 1985 (= S. microtheriella (Stainton, 1854)), Stigmella populnea Kemperman & Wilkinson, 1985 (= S. nivenburgensis (Preissecker, 1942)), Nepticula obscurella Braun, 1912 (revised synonymy, = S. myricafoliella (Busck, 1900)), Nepticula mandingella Gustafsson, 1972 (= S. wollofella (Gustafsson, 1972)), Stigmella rosaefoliella pectocatena Wilkinson & Scoble, 1979 (= S. centifoliella (Zeller, 1848)), Micropteryx pomivorella Packard, 1870 (= S. oxyacanthella (Stainton, 1854)), Stigmella crataegivora Puplesis, 1985 (= S. micromelis Puplesis, 1985), Stigmella scinanella Wilkinson & Scoble, 1979 (= S. purpuratella (Braun, 1917)), Stigmella palmatae Puplesis, 1984 (= S. filipendulae (Wocke, 1871)), Stigmella sesplicata Kemperman & Wilkinson, 1985 (= S. lediella (Schleich, 1867)), Stigmella rhododendrifolia Dovnar-Zapolski & Tomilova, 1978 (unavailable, = S. lediella (Schleich, 1867)), Stigmella oa Kemperman & Wilkinson, 1985 (= S. spiculifera Kemperman & Wilkinson, 1985), Stigmella gracilipae Hirano, 2014 (= S. monticulella Puplesis, 1984), Nepticula chaoniella Herrich-Schäffer, 1863 (= S. samiatella (Zeller, 1839)), Bohemannia piotra Puplesis, 1984 (= B. pulverosella (Stainton, 1849)), Bohemannia nipponicella Hirano, 2010 (= B. manschurella Puplesis, 1984), Sinopticula sinica Yang, 1989 (= Glaucolepis oishiella (Matsumura, 1931)), Trifurcula collinella Nel, 2012 (= Glaucolepis magna (A. Laštuvka & Z. Laštuvka, 1997)), Obrussa tigrinella Puplesis, 1985 (= Etainia trifasciata (Matsumura, 1931)), Microcalyptris vittatus Puplesis, 1984 and M. arenosus Falkovitsh, 1986 (both = Acalyptris falkovitshi (Puplesis, 1984)), Ectoedemia castaneae Busck, 1913, E. heinrichi Busck, 1914 and E. helenella Wilkinson, 1981 (all three = Zimmermannia bosquella (Chambers, 1878)), Ectoedemia chloranthis Meyrick, 1928 and E. acanthella Wilkinson & Newton, 1981 (both = Zimmermannia grandisella (Chambers, 1880)), Ectoedemia coruscella Wilkinson, 1981 (= Zimmermannia mesoloba (Davis, 1978)), Ectoedemia piperella Wilkinson & Newton, 1981 and E. reneella Wilkinson, 1981 (both = Zimmermannia obrutella (Zeller, 1873)), Ectoedemia similigena Puplesis, 1994 (= E. turbidella (Zeller, 1848)), Ectoedemia andrella Wilkinson, 1981 (= E. ulmella (Braun, 1912)), Nepticula canadensis Braun, 1917 (= E. minimella (Zetterstedt, 1839)), Opostega rezniki Kozlov, 1985 (= O. cretatella Chrétien, 1915), Pseudopostega cyrneochalcopepla Nel & Varenne, 2012 (= P. chalcopepla (Walsingham, 1908)). Stigmella caryaefoliella (Clemens, 1861) and Zimmermannia bosquella (Chambers, 1878) are taken out of synonymy and re-instated as full species. Lectotypes are designated for Trifurcula obrutella Zeller, 1873 and Nepticula grandisella Chambers, 1880.

Description: 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, aswell as subgenera to allowfor 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.

Description: 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.

Description: We present an interactive key that is available online through any web browser without the need to install any additional software, making it an easily accessible tool for the larger public. The key can be found at http://identify.naturalis.nl/lithocolletinae. The key includes all 86 North-West European Lithocolletinae, a subfamily of smaller moths (“micro-moths”) that is commonly not treated in field guides. The user can input data on several external morphological character systems in addition to distribution, host plant and even characteristics of the larval feeding traces to reach an identification. We expect that this will enable more people to contribute with reliable observation data on this group of moths and alleviate the work-load of taxonomic specialists, allowing them to focus on other new keys or taxonomic work.

Description: The proliferation of DNA data is revolutionizing all fields of systematic research. DNA barcode sequences, now available for millions of specimens and several hundred thousand species, are increasingly used in algorithmic species delimitations. This is complicated by occasional incongruences between species and gene genealogies, as indicated by situations where conspecific individuals do not form a monophyletic cluster in a gene tree. In two previous reviews, non-monophyly has been reported as being common in mitochondrial DNA gene trees. We developed a novel web service “Monophylizer” to detect non-monophyly in phylogenetic trees and used it to ascertain the incidence of species non-monophyly in COI (a.k.a. cox1) barcode sequence data from 4977 species and 41,583 specimens of European Lepidoptera, the largest data set of DNA barcodes analyzed from this regard. Particular attention was paid to accurate species identification to ensure data integrity. We investigated the effects of tree-building method, sampling effort, and other methodological issues, all of which can influence estimates of non-monophyly. We found a 12% incidence of non-monophyly, a value significantly lower than that observed in previous studies. Neighbor joining (NJ) and maximum likelihood (ML) methods yielded almost equal numbers of non-monophyletic species, but 24.1% of these cases of non-monophyly were only found by one of these methods. Non-monophyletic species tend to show either low genetic distances to their nearest neighbors or exceptionally high levels of intraspecific variability. Cases of polyphyly in COI trees arising as a result of deep intraspecific divergence are negligible, as the detected cases reflected misidentifications or methodological errors. Taking into consideration variation in sampling effort, we estimate that the true incidence of non-monophyly is ~23%, but with operational factors still being included. Within the operational factors, we separately assessed the frequency of taxonomic limitations (presence of overlooked cryptic and oversplit species) and identification uncertainties. We observed that operational factors are potentially present in more than half (58.6%) of the detected cases of non-monophyly. Furthermore, we observed that in about 20% of non-monophyletic species and entangled species, the lineages involved are either allopatric or parapatric—conditions where species delimitation is inherently subjective and particularly dependent on the species concept that has been adopted. These observations suggest that species-level non-monophyly in COI gene trees is less common than previously supposed, with many cases reflecting misidentifications, the subjectivity of species delimitation or other operational factors.

Description: Background: Host association patterns in Ectoedemia (Lepidoptera: Nepticulidae) are also encountered in other insect groups with intimate plant relationships, including a high degree of monophagy, a preference for ecologically dominant plant families (e.g. Fagaceae, Rosaceae, Salicaceae, and Betulaceae) and a tendency for related insect species to feed on related host plant species. The evolutionary processes underlying these patterns are only partly understood, we therefore assessed the role of allopatry and host plant family shifts in speciation within Ectoedemia. Methodology: Six nuclear and mitochondrial DNA markers with a total aligned length of 3692 base pairs were used to infer phylogenetic relationships among 92 species belonging to the subgenus Ectoedemia of the genus Ectoedemia, representing a thorough taxon sampling with a global coverage. The results support monophyletic species groups that are congruent with published findings based on morphology. We used the obtained phylogeny to explore host plant family association and geographical distribution to investigate if host shifts and allopatry have been instrumental in the speciation of these leafmining insects. Significance: We found that, even though most species within species groups commonly feed on plants from one family, shifts to a distantly related host family have occasionally occurred throughout the phylogeny and such shifts are most commonly observed towards Betulaceae. The largest radiations have occurred within species groups that feed on Fagaceae, Rosaceae, and Salicaceae. Most species are restricted to one of the seven global biogeographic regions, but within species groups representatives are commonly found in different biogeographic regions. Although we find general patterns with regard to host use and biogeography, there are differences between clades that suggest that different drivers of speciation, and perhaps drivers that we did not examine, have shaped diversity patterns in different clades.