SYSTEMATIC ENTOMOLOGY

Genomics-based higher classification of the species-rich Hairstreaks (Lepidoptera: Lycaenidae: Eumaeini)
Robbins RK, Cong Q, Zhang J, Shen J, Busby RC, Faynel C, Duarte M, Martins ARP, Prieto C, Lamas G and Grishin NV
We propose a higher classification of the lycaenid hairstreak tribe Eumaeini - one of the youngest and most species-rich butterfly tribes - based on autosome, Lepidopteran Z sex chromosome, and mitochondrial protein-coding genes. The subtribe Neolycaenina Korb is a of Callophryidina Tutt, and subtribe Tmolusina Bálint is a of Strephonotina K. Johnson, Austin, Le Crom, & Salazar. Proposed names are Rhammina Prieto & Busby, ; Timaetina Busby & Prieto, ; Atlidina Martins & Duarte, ; Evenina Faynel & Grishin, ; Jantheclina Robbins & Faynel, ; Paiwarriina Lamas & Robbins, ; Cupatheclina Lamas & Grishin, ; Parrhasiina Busby & Robbins, ; Ipideclina Martins & Grishin, ; and Trichonidina Duarte & Faynel, . Phylogenetic results from the autosome and Z sex chromosome analyses are similar. Future analyses of datasets with hundreds of terminal taxa may be more practical time-wise by focussing on the smaller number of sex chromosome sequences (2.6% of nuclear protein-coding sequences). The phylogenetic classification and biological summaries for each subtribe suggest that a variety of factors affected Eumaeini diversification. About a dozen kinds of male secondary sexual organs with frequent evolutionary gains and losses occur in Atlidina, Evenina, and Jantheclina (141 species combined). Females have been shown to use these organs to discriminate between conspecific and non-conspecific males, facilitating sympatry among close relatives. Eumaeina, Rhammina, and Timaetina (140 species combined) are overwhelmingly montane with some evidence for a higher incidence of sympatric diversification. Seven Neotropical lineages in five subtribes invaded the temperate parts of the Nearctic Region with a diversification increase in the Callophryidina (262 species). North American and then invaded the Palearctic at least once each, with a major species-richness increase in . The evolution of litter feeding detritivores within Calycopidina (172 species) resulted in an increase in diversification rate compared with its flower-feeding sister lineage. Atlidina, Strephonotina, Parrhasiina, and Strymonina (562 species combined) each contain a mixture of genera that specialize on one or two caterpillar food plant families and genera that are polyphagous. These would be appropriate subtribes to assess how the breadth of caterpillar food plants and the frequency of host shifts affected diversification.
Speciation in North American from a genomic perspective
Cong Q, Zhang J, Shen J, Cao X, Brévignon C and Grishin NV
Delineating species boundaries in phylogenetic groups undergoing recent radiation is a daunting challenge akin to discretizing continuity. Here, we propose a general approach exemplified by American butterflies from the genus Hübner notorious for the variety of similar phenotypes, ease of hybridization, and the lack of consensus about their classification. We obtain whole-genome shotgun sequences of about 200 specimens. We reason that discreteness emerges from continuity by means of a small number of key players, and search for the proteins that diverged markedly between sympatric populations of different species, while keeping low polymorphism within these species. Being 0.25% of the total number, these three dozen 'speciation' proteins indeed partition pairs of populations into two clusters with a prominent break in between, while all proteins taken together fail to reveal this discontinuity. Populations with larger divergence from each other, comparable to that between two sympatric species, form the first cluster and correspond to different species. The other cluster is characterized by smaller divergence, similar to that between allopatric populations of the same species and comprise conspecific pairs. Using this method, we conclude that (Cramer), Brévignon, (Cramer), and C. & R. Felder are restricted to South America. We find that six species of are present in the United States, one of which is new: Grishin, (i), found in south Texas and phenotypically closest to W. Barnes & McDunnough (ii) in its dark appearance. In the pale nudum of the antennal club, these two species resemble C. & R. Felder (iii) from Florida and the Caribbean Islands. The pair of sister species, Austin & J. Emmel (iv) and Hübner (v), represent the classic west/east U.S.A. split. The mangrove feeder (as caterpillar), dark nudum Brévignon (vi) enters south Texas as a new subspecies Grishin characterized by more extensive hybridization with and introgression from , and, as a consequence, more variable wing patterns compared with the nominal in Florida. Furthermore, a new mangrove-feeding species from the Pacific Coast of Mexico is described as Grishin Finally, genomic analysis suggests that may be a hybrid species formed by the ancestors of and
Unveiling one of the rarest 'butterflies' ever (Lepidoptera: Hesperiidae, Noctuidae)
Zilli A and Grishin NV
Many insect species named by the Danish entomologist J.C. Fabricius remain enigmatic due to loss of the original type specimens, sketchy descriptions and lack of illustrations, but even some well-illustrated taxa remain unrecognized. This is the case for , a 'butterfly' illustrated by W.J. Jones, the identity of which has puzzled experts for 225 years. Here we argue that the description and illustrations of this species are a perfect fit to a colourful moth later described by F. Walker as . Furthermore, we present evidence that Walker unwittingly based his name on the same specimen as Fabricius, and that this is the only known example of this species. An extraordinary sequence of misconceptions led the geographic origin of this specimen to become thoroughly confused, so that it is currently unknown where on Earth this species may occur (although a substantial body of evidence points to West Africa) and if it is even still extant.
Phylogenetic relationships of North American Gomphidae and their close relatives
Ware JL, Pilgrim E, May ML, Donnelly TW and Tennessen K
Intrafamilial relationships among clubtail dragonflies (Gomphidae) have been the subject of many morphological studies, but have not yet been systematically evaluated using molecular data. Here we present the first molecular phylogeny of Gomphidae. We include six of the eight subfamilies previously suggested to be valid, and evaluate generic relationships within them. We have included examples of all genera reported from the Nearctic except . This sample includes all North American species of , which has allowed us to explore intrageneric relationships in that genus. Our particular focus is on the closest relatives of the genus , especially those North American species groups that have been commonly treated as subgenera of . The complex is split into additional genera, supported by molecular and morphological evidence: and are here considered to be valid genera. The genus , in our restricted sense, does not occur in the western hemisphere; in addition, is transferred to .
Revision of the Bicyclus sciathis species group (Lepidoptera: Nymphalidae) with descriptions of four new species and corrected distributional records
Brattström O, Aduse-Poku K, Collins SC, DI Micco DE Santo T and Brakefield PM
In this paper we present a thorough revision of the sciathis species group of the butterfly genus Bicyclus (Kirby). Type materials are discussed and in several cases lectotypes are assigned to specimens from original type series. Four new, and morphologically distinct, species are described (B. elishiae Brattström , B. heathi Brattström , B. sigiussidorum Brattström and B. subtilisurae Brattström ), along with a comprehensive molecular phylogeny that includes exemplar taxa of all currently recognized species. We also investigate the types of all previously synonymized taxa and in the process invalidate the name B. ewondo Libert. This was done after finding the previously missing holotype of B. makomensis (Strand), which clearly belongs to the same species and thereby gives the older name priority. The phylogeny showed that some distinctly different species were surprisingly closely related, suggesting a high rate of morphological evolution in parts of the sciathis group. The distributional records for the group are updated after investigating over 1700 specimens kept in a range of museum collections. Many species previously thought to be broadly sympatric were found to have much more restricted ranges, with the previous overestimations probably based on misidentified specimens. The higher level of allopatry now established will make identification of many morphologically similar species easier. The fact that species often have smaller ranges than previously known, meaning that the level of endemism for African butterflies is likely to be higher than current estimates, has important implications for conservation management. An identification key for males of all 13 currently recognized species in the species group is included. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:837A9D4C-779A-4497-8176-7151D409DFA5.
A tale of two haplotype groups: Evaluating the New World ring species hypothesis using the distribution of divergent haplotypes
Gemmell AP and Marcus JM
The New World butterflies are a possible ring species with a circum-Caribbean distribution. Previous reports suggest a steady transition between North and South American forms in Mesoamerica, but in Cuba the forms were thought to co-exist without interbreeding representing the overlapping ends of the ring. Three criteria establish the existence of a ring species: a ring-shaped geographic distribution, gene flow among intervening forms, and genetic isolation in the region of range overlap. We evaluated mitochondrial haplotypes in from 9 species in the Western Hemisphere to test the ring species hypothesis. species are generally not monophyletic with respect to haplotypes, which are shared across species. However, two major haplotype groups exist. Group A predominates in South America, and Group B predominates in North and Central America. Therefore, haplotypes can be used to assess the degree of genetic influence a population receives from each continent. shows a ring-shaped distribution around the Caribbean, and evidence is consistent with gene flow among forms of including those from Mesoamerica. However, we detected no discontinuity in gene flow in Cuba or elsewhere in the Caribbean consistent with genetic isolation in the region of overlap. Though sampling is still very limited in the critical region, the only remaining possiblity for a circum-Caribbean discontinuity in gene flow is at the Isthmus of Panama, where there may be a transition from 98% Group B haplotypes in Costa Rica to 85-100% Group A haplotypes in South America.
Morphometric analysis and taxonomic revision of Ruschka (Hymenoptera: Chalcidoidea: Pteromalidae) - an integrative approach
Baur H, Kranz-Baltensperger Y, Cruaud A, Rasplus JY, Timokhov AV and Gokhman VE
We use an integrative taxonomic approach to revise the genus . In particular, we apply multivariate ratio analysis (MRA), a rather new statistical method based on principal component analysis (PCA) and linear discriminant analysis (LDA), to numerous body measurements and combine the data with those from our molecular analysis of and genetic markers (on a subset of species) and all available published data on morphology, karyology, behaviour, host associations and geographic distribution. We demonstrate that the analysis of quantitative characters using MRA plays a major role for the integration of name-bearing types and thus for the association of taxa with names. Six species are recognized, of which two are new: Baur and Gokhman & Baur For (Howard), a well-known, cosmopolitan parasitoid of stored-product pests, we have selected a neotype to foster continuity and stability in the application of this important name. The species was sometimes confused with the related , another cosmopolitan species that is frequently encountered in similar environments. We also show that several species originally described or later put under actually belong to different genera: (Risbec) ; (Szelényi) ; (Roomi, Khan & Khan) Ayyar & Mani is confirmed as a junior synonym of (Ferrière) and (Domenichini) must be considered as a valid but doubtful taxon. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:BDFE96D3-D0F4-4012-90F5-9A087F7F5864.
Identification and discrimination of species by numerical methods (Ephemeroptera: Heptageniidae)
Belfiore C
Nine species of from thirty-one populations were investigated on the basis of five meristic and five ratio characters of the larvae. The attribution of populations to species was checked by a cluster analysis based on the generalized distances between pairs. Groups representing species were then subjected to a multiple discriminant analysis; discriminant functions and loadings were calculated. The correct attribution of individuals to species by the discriminant functions ranged from 97% to 100%. The analysis of discriminant loadings shows which characters contribute more to the discrimination of species. Although many species could be identified with some confidence by a sequential monothetic key based on meristic and qualitative characters, the discriminant analysis improves the effectiveness of identification of all species.
A revision of the western Palaearctic species of Robineau-Desvoidy (Diptera: Tephritidae)
White IM and Korneyev VA
A key is provided to twenty-four western Palaearctic species of Robineau-Desvoidy. The hosts of twenty-three species which attack Asteraceae are listed, including those being used or investigated as possible weed biocontrol agents. The species are divided into four species groups and the differing host relationships and types of galls induced by these groups are discussed. sp.n. and ssp. ssp.n., associated with spp. and spp. respectively, are described. (Hering) and (Rohdendorf) are both treated as subspecies of (Meigen). is given full specific status and is removed from synonymy. The following new synonymies are made (junior synonyms in parentheses): (Hering) (= Hering); (Linnaeus) (= Robineau-Desvoidy, lectotype designated); (Hering) (= Hering); (Frauenfeld) (= Hering); Macquart (= Rondani, Loew); (Linnaeus) (= Hering, ); (Fabricius) (= Hering, Rondani); (Loew) (= Hering, Loew, Hendel). The possibility that is a species complex is discussed; it is also suggested that and represent the morphological extremes of a complex. The misuse of the name Fabricius in the genus Blot is noted.
Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea)
Woolley JB
A data set consisting of twenty-eight anatomical characters scored for twenty-eight terminal taxa representing the world fauna of Signiphoridae was analysed using parsimony and compatibility methods. The Coccophaginae (Aphelinidae) and the Azotinae (Aphelinidae) were used as outgroups to establish polarity of character state changes. Relationships of Signiphoridae to other Chalcidoidea are discussed. Several multistate characters were treated in the parsimony analyses either as unordered or as ordered into transformation series using additive binary coding, which in some cases drastically reduced the number of equally parsimonious solutions. Monophyly of Signiphoridae is supported by seven synapomorphies. Four genera, and , are recognized within Signiphoridae based on synapomorphies. . and . are synonymized under Species of are further assigned to four species groups, three of which are demonstrably monophyletic. Nine species or subspecies are transferred to from ( , , comb.n., , , , , , , one species to from , and one species to from ( . A key to genera of Signiphoridae and species groups of is presented. A diagnosis, relevant nomenclatural history, and a list of included species are given for each genus and species group, and the biology and distribution of each is summarized.