A taxonomic revision of the North American species of Stigmella (Lepidoptera: Nepticulidae)

Abstract. A revision of the fifty-one species of the genus Stigmella from North America is presented. Thirty-one species are recorded only from the U.S.A., seventeen species and one subspecies from the U.S.A. and Canada and three species and one subspecies are known only from Canada. Five species are new: Stigmella inconspicuella (U.S.A., California); S.heteromelis (California); S.sclerostylota (Arkansas); S.longisacca (California) and S.plumosetaeella (Arizona). Life histories, mine characteristics and larval habits are given where known. These are collated from field study, material examined, breeding records and the literature. A list of host plants is also provided.
A neotype is designated for Nepticula prunifoliella Clemens, previously considered to be possibly dipterous.
Although most Stigmella species have different genital morphology, some pairs of species do not. In these pairs, however, external features and characteristics of the mine often differ – some pairs also differ in distribution or season. The taxonomic implications are discussed in the individual treatments – not all the problems, however, can be resolved without further rearing studies and collection. Species-groups have been designated on the basis of morphology. Some species are, however, structurally intermediate between species-groups and others may resemble Ectoedemia Busck in aspects of the male genitalia while their female genitalia, external features and venation are typical of Stigmella.     

Monophagous leaf-mining larvae of Stigmella (Lepidoptera: Nepticulidae) on birch: patterns and differentiation in exploitation of the host

Patterns of larval coexistence, density and mortality of six birch feeding Stigmella leaf-miners were analysed in four field sites in the Netherlands. Larval feeding was further characterized by quantitative records on phenology, oviposition (per host, per tree and per leaf), larval growth rate and relative use of different mesenchyme layers. As the Stigmella guild on birch in western Europe is known to consist of three phylogenetically distinct species-groups, the specific feeding syndromes were primarily interpreted in this context. Ecological segregation among species groups tended to be larger than that within these groups. Among group segregation mainly involved differences in phenology, types of birch trees, mesenchyme layers used for feeding and larval body size and growth rate. Within group differentiation primarily concerned the distribution and clumping of eggs and the use of differently sized trees. A preliminary analysis of the frequency of mines per infested leaf produced a rather aggregated pattern, suggesting that only a limited number of the available birch leaves are considered as suitable by ovipositing females.     

Stigmella epicosma (Meyrick) (Lepidoptera: Nepticulidae): first distribution records from Chile and first host plant record

The presence of Stigmella epicosma (Meyrick) is reported for the first time from two coastal valleys from northern Chile: Azapa and Chaca. Adults examined were reared from leafminer larvae on Trixis cacalioides (Asteraceae), the first host plant known for S. epicosma.     

Challenging limits: Ultrastructure and size‐related functional constraints of the compound eye of Stigmella microtheriella (Lepidoptera: Nepticulidae)

With a body length of only 2 mm, the nepticulid Stigmella microtheriella (Stainton, 1854) is one of the smallest moths known to date. We investigated the optical design of its lemon‐shaped compound eyes, which measure 83.60 μm in anterior–posterior and 119.77 μm in dorso‐ventral direction. The eyes consist of about 123 facets, each of the latter just 9.9 μm in diameter. Transmission electron microscopy reveals an optical design with features intermediate between apposition and superposition optics similar to that known from two other small species of moths (one Nepticulid and one Gracillarid). Size‐related evolutionary adaptations of the ommatidial organization include (1) the involvement of only five rhabdomeres in the formation of the distal rhabdom (2) the complete absence of a rhabdomere of the eighth (= basal) retinula cell, (3) the “hourglass” shape of the rhabdom with a characteristic narrow waist separating distal from proximal portion, and (4) the reduction to one single layer of tracheoles as an adaptation to the overall restricted space available in this minute eye. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae)

The generic names Euptychia, Cissia and Argyreuplychia are reviewed in relation to the Cissia confusa species-group in Costa Rica and Trinidad, for which the generic name Cissia Doubleday is formally resurrected. Three new species of Cissia are described, C. confusa itself is resurrected from synonymy, three neotypes and three lectotypes are designated, and six new combinations established. On the basis of early instar morphology three subgroups are recognized and the general biology of the confusa species-group is summarized. A key to species in Costa Rica is provided and the ecology of the adults is reviewed.     

The Disphragis notabilis (Schaus) species-group in Costa Rica (Lepidoptera,Notodontidae)

The four described taxa in the Disphragis notabilis (Schaus) species-group are reviewed, including the types and their dissected genitalia. Disphragis hemicera (Schaus), stat. rev., is elevated to species rank, D. normula (Dognin) is retained as a synonym of D. notabilis, D. sobolis Miller is confirmed as distinct from D. hemicera, and D. bifurcata sp. n., is newly described. Both D. hemicera and D. bifurcata occur in Costa Rica. The known ranges of the other species are outlined. Defining characters of each species are presented and a key to species is provided. Unusual variation in the genitalia is noted.     

Three Unrecorded Species of Family Nepticulidae (Lepidoptera) from South Korea

Here, we report three previously unrecorded species of Nepticulidae, Stigmella aladina Puplesis, S. fumida Kemperman et Wilkinson, and Bohemannia nubila Puplesis from South Korea. The external characteristics of adults and genitalia are briefly redescribed with illustrations.     

Pigmy moths (Lepidoptera, Nepticulidae) from the Middle Volga Basin

Fifty-seven species of the family Nepticulidae were found in the Middle Volga Basin. The family includes four genera: Bohemannia Stainton, 1859 (1 species), Ectoedemia Busck, 1907 (15), Stigmella Schrank, 1802 (38), and Trifurcula Zeller, 1848 (3 species). The mines made by pigmy moth larvae are very specific in the study area. Larvae of the largest genus, Stigmella, make typical serpentine leaf mines. Larvae of the genera Bohemannia and Ectoedemia make blotchy mines; some larvae develop in fruits and tree bark. Most species of the genera Bohemannia, Ectoedemia, and Stigmella were recorded by their mines. At the larval stage, representatives of Trifurcula feed on stems of legumes. The pupae develop in the soil. Most species in the family Nepticulidae are leaf miners, mainly associated with Rosaceae (34.7%), Salicaceae (15.8%), Fagaceae (12.3%), and Betulaceae (12.3%). All the species investigated do not act as pests of forage plants in the area studied.     

A Global Phylogeny of Leafmining Ectoedemia Moths (Lepidoptera: Nepticulidae): Exploring Host Plant Family Shifts and Allopatry as Drivers of Speciation

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.     

Phylogenetic relationships in brown argus butterflies (Lepidoptera: Lycaenidae: Aricia) from northwestern Europe

Lycaenid butterflies of the Aricia agestisartaxerxes complex pose an unresolved taxonomic and conservation problem in northwestern Europe. Two key issues require resolution: (i) how many species of Aricia occur in northwestern Europe and what are their distributions?; (ii) how is the morphological variation observed in northwestern Europe best explained? We investigated phylogenetic relationships and phylogeographic patterns in this species group using mitochondrial and nuclear markers in comparison with morphological variation. A 325 bp fragment of the mitochondrial cytochromeb gene was sequenced from 179 individuals representing 18 populations from the UK and Scandinavia. Seventeen enzymecoding loci were analysed from 538 individuals from the same populations. Highly congruent phylogenies between mitochondrial and allozyme markers demonstrate that the sample is composed of two closely related species, A. agestis and A. artaxerxes. Both marker types also suggest that Scottish and northern Scandinavian A. artaxerxes populations are conspecific, and consequently do not support the endemic status of A. artaxerxes in the UK. The subspecies division of British populations of A. artaxerxes is also not supported by phylogenetic analyses. Allozyme and mitochondrial analyses cluster two populations from the Peak District, UK, differently. The former suggests that they are A. artaxerxes whilst the latter suggests they are A. agestis. Further research is required to find the reason for this disagreement, which could be associated with the different dynamics of nuclear and mitochondrial genes across a hybrid zone between the two species. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75 , 27–37.     

First record of a walnut shield bearer Coptodisca (Lepidoptera: Heliozelidae) in Europe

A leafminer of the Nearctic genus Coptodisca Walsingham (Lepidoptera: Heliozelidae), a species of potential economic interest, is reported for the first time from Europe, infesting the black (Juglans nigra L.) and the common walnut (Juglans regia L.). Mines were collected since September of 2010 in several sites of two Italian regions (Campania and Lazio). The species is rather similar to Coptodisca juglandella (Chambers), the only Coptodisca known to attack walnuts, but at present, an unambiguous identification cannot be provided because of the unsatisfactory characterization of this leafminer and congeneric species. Three generations were recorded per year and leafminers overwinter as mature larvae. The first adults emerged in May–June while mature larvae of the last generation started the overwintering in September. During the last generation of the year, infestation levels of leaves were 100% in all sampled localities. Several species of parasitoids were reared from infested mines, with specimens belonging to the genus Chrysocharis (Hymenoptera: Eulophidae) being the most frequent parasitoids.     

The taxonomy of the Pollenia rudis species-group in the Holarctic Region (Diptera: Calliphoridae)

Abstract. A rudis species-group is defined within Pollenia Robineau-Desvoidy, and new characters found useful in the taxonomy of this genus are presented. P.rudis (Fabricius), P.angustigena Wainwright, stat.rev. and P.pseudorudis Rognes are redescribed. P.hungarica sp.n., P.longitheca sp.n. and P.luteovillosa sp.n. are described as new to science. A key is provided, and the terminalia of both sexes are illustrated for all the species. Some features of the puparia are figured for the species where these are known. A neotype is designated for Musca rudis , and a lectotype for P.angustigena. P.angustigena, P.pseudorudis and P.rudis are Holarctic species, and the latter two have also been found in New Zealand. The remaining species are confined to the western Palaearctic. P.hungarica is known from central Europe, including southern parts of Scandinavia, P.longitheca from the eastern Mediterranean, and P.luteovillosa from Algeria and Morocco in North Africa. In the larval stages P. rudis group members are parasites of or predators on earthworms. The species have several generations each year, and normally overwinter as adults. Eisenia rosea (Savigny) serves as a host for P.hungarica, P.pseudorudis and P.rudis according to the reared material available. A previous detailed account of the immature stages and life-cycle of lsquo;rudis' from North America is tentatively assigned to pseudorudis. Keilin's (1909,1915) often cited accounts of the immature stages and life-cycle of a species called 'rudis' are rejected as a source of information for any member of the rudis group.     

Insecticidal resistance and cross-resistance in populations of Cydia pomonella (Lepidoptera: Tortricidae) in central Europe

Insecticide bioassays were used to investigate resistance of Cydia pomonella (L.) (Lepidoptera: Tortricidae) to insecticides with various types of active ingredients. The efficacy baselines of selected insect growth regulators (fenoxycarb), insect growth inhibitors (diflubenzuron and teflubenzuron), organophoshorous insecticides (phosalone), and neonicotinoids (thiacloprid) against the eggs and first and fifth instars of sensitive laboratory strains of codling moth were determined. According to concentration-mortality baseline, 50% lethality concentration values and 90% lethality concentration values were determined for all the tested insecticides. The lethal concentration ratio quantified the relation between the efficacy of selected insecticides against fifth instars found by topical application and against first instars found by diet-treated bioassay. No difference was detected when the efficacy of technical grade diflubenzuron diluted in tetrahydrofuran and diflubenzuron in the formulated product Dimilin 48 SC diluted in water was compared. However, just before the application of insecticide, the integument of larvae must be treated with acetone. Two bioassays were used to monitor the resistance of codling moths collected in 2003-2005 in two apple (Malus spp.) orchards with different intensities of chemical control. Resistance ratios (RRs) to the tested insecticides were determined for both field populations of codling moth. For the population of codling moth from a commercial apple orchard in Velké Bílovice, cross-resistance to fenoxycarb, teflubenzuron, and phosalone was detected after the topical application of insecticides to fifth instars. The population of codling moth from Prague-Ruzyne was slightly resistant to phosalone and teflubenzuron. No resistance to diflubenzuron was detected in either tested population.     

The taxonomy and phylogenetic relationships of the Pollenia semicinerea species-group (Diptera: Calliphoridae)

Abstract. Within Pollenia Robineau-Desvoidy, a semicinerea species-group is defined. It consists of the redescribed P.semicinerea Villeneuve, P.grunini sp.n., P.mystica sp.n. and P.paragrunini sp.n., and P.pectinata Grunin, for which some new characters are given. A key is provided and the known terminalia of both sexes are illustrated. A lectotype is designated for P.semicinerea Villeneuve. P.semicinerea is known from the Middle East (Israel, Syria, Lebanon), P.grunini, P.mystica and P.paragrunini from the Caucasus and neighbouring areas in the U.S.S.R. and P.pectinata from the temperate parts of the Palaearctic Region except west of Poland and U.S.S.R. It is argued that the semicinerea- and racfe-groups are monophyletic sister-groups. The tenuiforceps-group is most probably the monophyletic sister-group of the composite monophyletic group ( rudis-group+semicinerea-group ). The intermedia-group and the griseo-tomentosa-group are both monophyletic. The composite group ( rudis- group + semicinerea-group + tenuiforceps-group + intermedia-group + griseotomentosa-group ) is probably a monophyletic subgroup of Pollenia. A partly resolved synapomorphy diagram is provided, which is compared to trees generated by various computer programs in the Phylogeny Inference Package (PHYLIP) of Felsenstein. Pollenia alajensis Rodendorf (= sytshevskajae Grunin) is assigned to the tenuiforceps species-group on the basis of male and female genitalia.     

Host plant relationships of an endangered butterfly, Lopinga achine (Lepidoptera: Nymphalidae) in northern Europe

The aim of the present study was to evaluate—in a geographic perspective—the role of host plant as a determinant of habitat quality for Lopinga achine, a satyrine butterfly endangered over much of its European range. Laboratory trials were performed to record host choices made by the ovipositing females as well as by neonate larvae. In rearing experiments, growth performance and mortality on different host plants was determined. Oviposition was found to be indiscriminate but larvae were shown to be able to choose between host plants, with the choices made broadly consistent with growth performance of the larvae on particular hosts. Nevertheless, most grasses and sedges offered were found to support larval development reasonably well. No clear superiority of the previously suggested primary host plant Carex montana could be shown. Importantly, no differences in host plant relationships were found between the populations of Sweden, western Estonia and eastern Estonia. In particular, the larvae originating from eastern Estonian populations developed on C. montana equally well even if the plant is absent from their native habitat. In the context of species conservation, one should conclude that L. achine is polyphagous enough on various grasses and sedges so that the presence of any particular host species cannot be a critical component of habitat quality. Nevertheless, some preference to broad- and soft-leaved hosts, as well as sensitivity to host wilting, may partly explain the butterfly’s preference to moist forest habitats, further emphasizing the central role of habitat management in the conservation practice of this species. In turn, the absence of ecological differences between geographic populations should enable conservationists to successful transfer their experience across national boundaries.     

The genus Cleora Curtis (Lepidoptera: Geometridae) in Fiji

An account is given of the 10 species of the genus Cleora known to inhabit Fiji; two of these are described as new and two are transferred to Cleora from other genera. Species previously known from Fiji are re-described, and a key to the males is included.     

Eulophid wasps (Hymenoptera, Eulophidae), parasitoids of leaf-mining moths (Lepidoptera: Gracillariidae, Nepticulidae, Tischeriidae) on the English oak in the Middle Volga Area

Data on the parasitoid eulophid complexes reared from 6 species of leaf mining moths damaging the oak in the Middle Volga Area are reported. The reared eulophids belong to three subfamilies: Eulophinae (8 species from 6 genera): Cirrospilus diallus Walker, C. viticola (Rondani), Euplectrus liparidis Ferrière, Hyssopus geniculatus (Hartig), Sympiesis gordius (Walker), Pnigalio rotundiventris (Erdös), P. soemius (Walker), Rhicnopelte crassicornis (Nees); Entedoninae (11 species from 4 genera): Chrysocharis laomedon (Walker), Ch. prodice (Walker), Ch. pubens Delucchi, Ch. submutica Graham, Closterocerus formosus Westwood, Neochrysocharis aratus (Walker), Pediobius cassidae Erdös, P. flaviscapus (Thomson), P. metallicus (Nees), P. pyrgo Walker; and Tetrastichinae (3 species from 3 genera): Aprostocetus sp., Minotetrastichus frontalis (Nees), and Sigmophora brevicornis (Panzer). The most abundant of the moth parasitoids are M. frontalis and Ch. laomedon, while Tischeria ekebladella has the greatest number of parasitoids. The ectoparasitoids/endoparasitoids ratio in the number of species is 2: 1, and in the number of individuals reared, 1.76: 1. Thus, the host density is regulated primarily by the ectoparasitoids.