Molecular phylogeny, divergence time, biogeography and trends in host plant usage in the agriculturally important tortricid tribe Grapholitini (Lepidoptera: Tortricidae: Olethreutinae).

The leaf-roller moth tribe Grapholitini comprises about 1200 described species and contains numerous notorious pests of fruits and seeds. The phylogeny of the tribe has been little studied using contemporary methods, and the monophyly of several genera remains questionable. In order to provide a more robust phylogenetic framework for the group, we conducted a multiple-gene phylogenetic analysis of 104 species representing 27 genera of Grapholitini and 29 outgroup species. Divergence time, ancestral area, and host plant usage were also inferred to explore evolutionary trends in the tribe. Our analyses indicate that Larisa and Corticivora, traditionally assigned to Grapholitini, are best excluded from the tribe. After removal of these two genera, the tribe is found to be monophyletic, represented by two major lineages-a Dichrorampha clade and a Cydia clade, the latter of which can be divided into seven generic groups. The genus Grapholita was found to be polyphyletic, comprising three different clades, and we propose three genera to accommodate these groups: Grapholita (sensu stricto), Aspila (formerly a subgenus of Grapholita) and Ephippiphora (formerly considered a synonym of Grapholita). We summarize each generic group, including related genera not included in our analysis, providing morphological, pheromone and food plant characters that support particular branches within the molecular hypotheses. Biogeographical analyses indicate that Grapholitini probably originated in the Nearctic, Afrotropical and Neotropical regions in the Lutetian of the middle Eocene (ca. 44.3 Ma). Our results also indicate that most groups in Grapholitini originated from Fabaceae-feeding monophagous or oligophagous ancestors, and that host plant shifts probably promoted species diversification within the tribe.

Species-Level Para- and Polyphyly in DNA Barcode Gene Trees: Strong Operational Bias in European Lepidoptera.

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.

Revision of African Neaspasia Diakonoff, 1989 and the related Conaspasia, n. gen. (Lepidoptera, Tortricidae).

The six species of Neaspasia Diakonoff present in mainland Africa are described and illustrated. Niphadophylax albonigra Razowski & Wojtusiak and N. sophrona Razowski & Wojtusiak are transferred to Conaspasia, new genus. Four new species are described: Neaspasia coronana Aarvik, new species, N. karischi Aarvik, new species, N. malamigambo Aarvik, new species, and Conaspasia congolana Aarvik, new species. Argyroploce orthacta Meyrick, Argyroploce brevisecta Meyrick, and Penthina brevibasana Walsingham are transferred to Neaspasia. Neaspasia rhodesiae Razowski & Brown is a junior synonym of Neaspasia orthacta (Meyrick), new combination. Genetancylis homalota Razowski and Rhopobota cornuta Razowski, both described from Oman, are transferred to Neaspasia. Genetancylis Razowski is synonymised with Neaspasia.

A new species of Neurothaumasia Le Marchand (Lepidoptera, Tineidae) from Crete, Greece.

A new species of fungus moths (Tineidae), Neurothaumasia cretica sp. nov., is described from specimens collected on Crete isl. (Greece). It differs from congeneric taxa by the characteristic black and white forewing pattern which is only shared with N. fasciata Petersen, 1959 from the Middle East, and the widespread western Palaearctic N. ankerella (Mann, 1867). However, the new species differs strongly from the former by several characters of male and female genitalia, and from the latter species particularly from external appearance and by the highly divergent DNA barcode (cytochrome c-oxidase subunit 1) (unknown for N. fasciata). Adult and genitalia of N. cretica sp. nov. and the only similar European species N. ankerella are shown for comparison. Finally, a complete checklist of the genus is added.

A new genus and two new species of the tribe Pexicopiini (Lepidoptera, Gelechiidae) from the Afrotropical region.

A new genus, Falcipenna gen. nov., and two species, F. irinae sp. nov. (Kenya) and F. argenteomaculata sp. nov. (Kenya, Tanzania, South Africa), are described. Based on external and genital characters the new genus is placed in the subfamily Apatetrinae, tribe Pexicopiini. Within the tribe, the genus shows affinity to Harpagidia Raganot, 1895 and Sitotrogoides Sohn, Ponomarenko & Sakamaki, 2019. The differences between the new genus and allied genera are discussed. Adults of the new species are illustrated, including details of external morphology and male and female genitalia.

Gibbalaria : A new genus of Olethreutini from the Afrotropical Region (Lepidoptera: Tortricidae: Olethreutinae), and a new combination in Cosmopoda Diakonoff.

Gibbalaria Brown and Aarvik, new genus, is described to accommodate the following species: G. divergana Brown and Aarvik, new species (TL: South Africa); G. occidentana Brown and Aarvik, new species (TL: South Africa); G. chiloanei Brown and Aarvik, new species (TL: South Africa); G. scabellana Zeller, 1852, new combination (TL: South Africa); G. mabalingwae Razowski, 2008, status revised, new combination (TL: South Africa); G. longiphallus Brown and Aarvik, new species (TL: South Africa); G. bagamoyo (Razowski & Wojtusiak, 2014), new combination (TL: Tanzania); and G. analcis (Razowski, 2015), new combination (TL: Republic of Cameroon). Gibbalaria sistrata (Meyrick, 1911), new combination (TL: South Africa), is synonymized with G. scabellana, new synonymy. Although the male genitalia of Gibbalaria form two moderately divergent groups, the two groups share extremely similar female genitalia, which feature a broad, somewhat cylindrical antrum, followed immediately anterad by a conspicuously elbowed region of the ductus bursae forming a weak diverticulum on the left side, and a signum in the form of a patch of small arrowhead-shaped sclerites, usually with a shallow pocket. Male genitalia morphology shows many similarities with the Neopotamia group (Olethreutini), Afroploce Aarvik in particular. The monophyly of Gibbalaria is supported by the raised, curled, opalescent scales on the forewing in both sexes, which occurs in a few other genera scattered through the subfamily (e.g., Thylacandra Diakonoff in Grapholitini; Astronauta Diakonoff and Cosmopoda Diakonoff in Olethreutini); the angled diverticulum in the ductus bursae in the female genitalia; the subrectangular flange from the lower margin of the valva immediately basal to the cucullus; and the setose, protruding ridge immediate above the basal cavity, which appears to be homologous with the subbasal process of the valva in many members of the Neopotamia group of genera. In addition, Diakonoffiana graziani Razowski, 2015 (TL: Republic of Cameroon) is transferred to Cosmopoda Diakonoff, resulting in C. graziani (Razowski), new combination.

New species and new country distribution records of Athrips (Lepidoptera, Gelechiidae) from the Afrotropical region.

Athrips nigribasella sp. nov., A. dorsochrella sp. nov., A. macrosignella sp. nov. and A. spinosum sp. nov., are described from Kenya, and A. asymmetrica sp. nov. is described from South Africa. The genus Parapsectris Meyrick, 1911, syn. nov., is synonymized with Athrips Billberg, 1820. Athrips sisterina (Povoln, 1989) is recorded for the first time from Kenya, Athrips savannae (Bidzilya, 2010), comb. nov. and A. ochrocosma (Meyrick, 1911) from Tanzania. Adults and genitalia of all newly described species are illustrated. The diagnoses of A. sisterina and A. bruneosparsa (Janse, 1958) are updated with descriptions and illustrations of details of male and female genitalia.

Integrative taxonomy reveals overlooked cryptic diversity in the conifer feeding Batrachedrapinicolella (Zeller, 1839) (Lepidoptera, Batrachedridae).

During efforts to generate DNA barcodes for all European Lepidoptera, Batrachedrapinicolella (Zeller, 1839) was found to comprise two genetically distinct clusters. Morphological investigation and results from two nuclear markers and ddRAD sequencing furthermore support the existence of two distinct taxa which we treat as two separate species, B.pinicolella and B.confusella sp. nov. A lectotype for B.pinicolella is designated. Available data indicate that the biology of both species also differs, with Piceaabies (L.) Karsten as a proved host-plant for B.pinicolella and Pinussylvestris L. for B.confusella sp. nov. Both species are mainly distributed on the European continent with B.pinicolella occurring in boreal parts of North and Central Europe and introduced to Canada, reflecting a boreo-montane distribution pattern. Batrachedraconfusella sp. nov. is more widely distributed in temperate Northern and Central Europe.

Two new species of the genus Thubdora Park and a new species of Ptilothyris Walsingham (Lepidoptera, Gelechioidea, Lecithoceridae) from Uganda.

Two species of Thubdora Park (T. onsemiro Park, sp. nov. and T. elgozmanyi Park, sp. nov.) and one species of Ptilothyris Walsingham (P. galbiplatella Park, sp. nov.) are described from Uganda.

A taxonomic review of the Afrotropical genus Dragmatucha Meyrick, 1908 (Lepidoptera, Gelechioidea, Lecithoceridae, Torodorinae), with descriptions of eleven new species.

The genus Dragmatucha Meyrick is reviewed and 11 species of a total of 13 are described as new. The new species originate from the Democratic Republic of Congo, Ghana, Kenya, and Uganda. Based on morphological characters, especially the color pattern of both wings, the genus is divided into two species-groups: the proaula species-group with five species (D. proaula Meyrick, 1908; D. galbinea Park, sp. nov.; D. kabarolensis Park, sp. nov.; D. saltualis Park, sp. nov. and D. vittatella Park, sp. nov.) and the hispidula species-group with eight species (D. hispidula Park, 2018; D. goniotes Park, sp. nov.; D. cochliana Park, sp. nov.; D. pedalis Park, sp. nov.; D. ghanaensis Park, sp. nov.; D. crinifrutalis Park, sp. nov.; D. dizostera Park, sp. nov. and D. kakumensis Park, sp. nov.). Adults and male genitalia of all new species are illustrated, but female genitalia are provided only for D. ghanaensis Park, sp. nov. and D. kakumensis Park, sp. nov. When fresh material was available, the analysis of COI sequences was undertaken.

DNA barcode library for European Gelechiidae (Lepidoptera) suggests greatly underestimated species diversity.

For the first time, a nearly complete barcode library for European Gelechiidae is provided. DNA barcode sequences (COI gene - cytochrome c oxidase 1) from 751 out of 865 nominal species, belonging to 105 genera, were successfully recovered. A total of 741 species represented by specimens with sequences ≥ 500bp and an additional ten species represented by specimens with shorter sequences were used to produce 53 NJ trees. Intraspecific barcode divergence averaged only 0.54% whereas distance to the Nearest-Neighbour species averaged 5.58%. Of these, 710 species possessed unique DNA barcodes, but 31 species could not be reliably discriminated because of barcode sharing or partial barcode overlap. Species discrimination based on the Barcode Index System (BIN) was successful for 668 out of 723 species which clustered from minimum one to maximum 22 unique BINs. Fifty-five species shared a BIN with up to four species and identification from DNA barcode data is uncertain. Finally, 65 clusters with a unique BIN remained unidentified to species level. These putative taxa, as well as 114 nominal species with more than one BIN, suggest the presence of considerable cryptic diversity, cases which should be examined in future revisionary studies.

The importance of correct labelling of types: an example in Tortricidae (Lepidoptera) and its rectification.

Taxonomy is the basic discipline in biology. The taxonomist defines each taxon and provides a unique name for it. Of the systematic categories, the species level is crucial. When a species is described and named, great care should be taken to ensure that all information related to the name is correct. Information on ecology, biology, distribution, etc. is linked to the name, and may be added and repeated in subsequent literature. Great confusion may arise from ambiguous descriptions and/or mislabelled figures. Therefore editors should take special care to ensure that taxonomic papers containing descriptions of new taxa receive careful refereeing and editing. Here we present a case where descriptions of new species became misleading when the labels of the type specimens were switched. If the manuscript had been more thoroughly refereed, the mistake could have been discovered and avoided.

Redefinition and revision of African Cosmetra Diakonoff, 1977 (Lepidoptera: Tortricidae) with description of six new species.

The eight species of Cosmetra Diakonoff present in mainland Africa are described and illustrated. Six new species are described: Cosmetra larseni, new species; C. fibigeri, new species; C. truncana, new species; C. multidentana, new species; C. usambarensis, new species; and C. juu; new species. Cosmetra neka Razowski & Brown is a junior synonym of Cosmetra tumulata (Meyrick), and C. mucronata Razowski & Wojtusiak is a junior synonym of C. spiculifera (Meyrick).

Additions to the fauna of plume moths (Lepidoptera, Pterophoridae) of Uganda.

An overview of 20 Pterophoridae species from Uganda is given. Three of them are described as new to science: Crassuncus agassizi Ustjuzhanin & Kovtunovich sp. nov., Hellinsia anikini Ustjuzhanin & Kovtunovich sp. nov. and Hellinsia nawrothi Ustjuzhanin & Kovtunovich sp. nov. Twelve species are recorded as new to the fauna of Uganda.

New Tortricidae (Lepidoptera) from East Africa with an account of the tortricid fauna of acacia in the Kenyan Rift Valley.

Species of Tortricidae whose larvae feed on acacia are listed, including five new species: Hystrichophora bussei Agassiz, Endotera cyaneana Agassiz, Paraeccopsis variegana Agassiz & Aarvik, Coniostola flavitinctana Agassiz & Aarvik, and C. rufitinctana Agassiz & Aarvik. Six additional species related to the aforementioned, whose life histories are not known, also are described: Paraeccopsis tanzanica Aarvik, P. addis Aarvik, P. turi Aarvik, P. botswanae Aarvik, P. pseudoinsellata Aarvik, and Coniostola laikipiana Agassiz & Aarvik. Endotera nodi Agassiz is synonymised with Endotera cyphospila (Meyrick), comb. n.; and Coniostola omistus Diakonoff is synonymised with Coniostola stereoma (Meyrick). Paraeccopsis inflicta (Meyrick) and Paraeccopsis atricapsis (Meyrick) are removed from the synonymy of Paraeccopsis insellata (Meyrick). Eucosma pharangodes Meyrick is transferred to Eucosmocydia Diakonoff. Age onychistica Diakonoff is recorded from Africa for the first time. 

Deep sympatric mtDNA divergence in the autumnal moth (Epirrita autumnata).

Deep sympatric intraspecific divergence in mtDNA may reflect cryptic species or formerly distinct lineages in the process of remerging. Preliminary results from DNA barcoding of Scandinavian butterflies and moths showed high intraspecific sequence variation in the autumnal moth, Epirrita autumnata. In this study, specimens from different localities in Norway and some samples from Finland and Scotland, with two congeneric species as outgroups, were sequenced with mitochondrial and nuclear markers to resolve the discrepancy found between mtDNA divergence and present species-level taxonomy. We found five COI sub-clades within the E. autumnata complex, most of which were sympatric and with little geographic structure. Nuclear markers (ITS2 and Wingless) showed little variation and gave no indications that E. autumnata comprises more than one species. The samples were screened with primers for Wolbachia outer surface gene (wsp) and 12% of the samples tested positive. Two Wolbachia strains were associated with different mtDNA sub-clades within E. autumnata, which may indicate indirect selection/selective sweeps on haplotypes. Our results demonstrate that deep mtDNA divergences are not synonymous with cryptic speciation and this has important implications for the use of mtDNA in species delimitation, like in DNA barcoding.