Cuticular hydrocarbons corroborate the distinction between lowland and highland Natal fruit fly (Tephritidae,Ceratitis rosa) populations

The cuticular hydrocarbons (CHs) and morphology of two Ceratitis rosa Karsch (Diptera: Tephritidae) populations, putatively belonging to two cryptic taxa, were analysed. The chemical profiles were characterised by two-dimensional gas chromatography with mass spectrometric detection. CHs of Ceratitis rosa that originated from the lowlands and highlands of Kenya comprised of n-alkanes, monomethylalkanes, dimethylalkanes and unsaturated hydrocarbons in the range of the carbon backbone from C₁₄ to C₃₇. Hydrocarbons containing C₂₉, C₃₁, C₃₃ and C₃₅ carbon atoms predominated in these two populations. 2-Methyltriacontane was the predominant compound in both populations. Quantitative differences in the distribution of hydrocarbons of different chain lengths, mainly the C₂₂, C₃₂, C₃₃ and C₃₄ compounds of these two populations, were observed despite indistinct qualitative differences in these hydrocarbons. Morphological analyses of male legs confirmed that the flies belong to different morphotypes of Ceratitis rosa previously labelled as R1 and R2 for lowland and highland populations, respectively. A statistical analysis of the CH compositions of the putative R1 and R2 species showed distinct interspecific identities, with several CHs specific for each of the lowland and highland populations. This study supports a hypothesis that the taxon Ceratitis rosa consists of at least two biological species.     

An integrative approach to unravel the Ceratitis FAR (Diptera,Tephritidae) cryptic species complex: a review

This paper reviews all information gathered from different disciplines and studies to resolve the species status within the Ceratitis FAR (Ceratitis fasciventris, Ceratitis anonae, Ceratitis rosa) complex, a group of polyphagous fruit fly pest species (Diptera, Tephritidae) from Africa. It includes information on larval and adult morphology, wing morphometrics, cuticular hydrocarbons, pheromones, microsatellites, developmental physiology and geographic distribution. The general consensus is that the FAR complex comprises Ceratitis anonae, two species within Ceratitis rosa (so-called R1 and R2) and two putatitve species under Ceratitis fasciventris. The information regarding the latter is, however, too limited to draw final conclusions on specific status. Evidence for this recognition is discussed with reference to publications providing further details.     

Population structure and cryptic genetic variation in the mango fruit fly,Ceratitis cosyra (Diptera,Tephritidae)

The fruit fly Ceratitis cosyra is an important agricultural pest negatively affecting the mango crop production throughout Africa and also feeding on a variety of other wild and cultivated hosts. The occurrence of deeply divergent haplotypes, as well as extensive morphological variability, previously suggested possible cryptic speciation within Ceratitis cosyra. Here we provide the first large-scale characterisation of the population structure of Ceratitis cosyra with the main objective of verifying cryptic genetic variation. A total of 348 specimens from 13 populations were genotyped at 16 polymorphic microsatellite loci. Hardy-Weinberg equilibrium (HWE) deviations were observed in 40.4% of locus-population combinations and suggested the occurrence of genetic substructuring within populations. Discriminant Analysis of Principal Components (DAPC) showed genetic divergence between the vast majority of vouchers from Burundi and Tanzania (plus a few outliers from other African countries) and all other specimens sampled. Individual Bayesian assignments confirmed the existence of two main genotypic groups also occurring in sympatry. These data provided further support to the hypothesis that Ceratitis cosyra might include cryptic species. However, additional integrative taxonomy, possibly combining morphological, ecological and physiological approaches, is required to provide the necessary experimental support to this model.     

Comparative analysis of development and survival of two Natal fruit fly Ceratitis rosa Karsch (Diptera,Tephritidae) populations from Kenya and South Africa

Comparative analysis of development and survivorship of two geographically divergent populations of the Natal fruit fly Ceratitis rosa Karsch designated as Ceratitis rosa R1 and Ceratitis rosa R2 from Kenya and South Africa were studied at seven constant temperatures (10, 15, 20, 25, 30, 33, 35 °C). Temperature range for development and survival of both populations was 15–35 °C. The developmental duration was found to significantly decrease with increasing temperature for Ceratitis rosa R1 and Ceratitis rosa R2 from both countries. Survivorship of all the immature stages of Ceratitis rosa R1 and Ceratitis rosa R2 from Kenya was highest over the range of 20–30 °C (87–95%) and lowest at 15 and 35 °C (61–76%). Survivorship of larvae of Ceratitis rosa R1 and Ceratitis rosa R2 from South Africa was lowest at 35 °C (22%) and 33 °C (0.33%), respectively. Results from temperature summation models showed that Ceratitis rosa R2 (egg, larva and pupa) from both countries were better adapted to low temperatures than R1, based on lower developmental threshold. Minimum larval temperature threshold for Kenyan populations were 11.27 °C and 6.34 °C (R1 and R2, respectively) compared to 8.99 °C and 7.74 °C (R1 and R2, respectively) for the South African populations. Total degree-day (DD) accumulation for the Kenyan populations were estimated at 302.75 (Ceratitis rosa R1) and 413.53 (Ceratitis rosa R2) compared to 287.35 (Ceratitis rosa R1) and 344.3 (Ceratitis rosa R2) for the South African populations. These results demonstrate that Ceratitis rosa R1 and Ceratitis rosa R2 from both countries were physiologically distinct in their response to different temperature regimes and support the existence of two genetically distinct populations of Ceratitis rosa. It also suggests the need for taxonomic revision of Ceratitis rosa, however, additional information on morphological characterization of Ceratitis rosa R1 and Ceratitis rosa R2 is needed.     

Wing morphometrics as a possible tool for the diagnosis of the Ceratitis fasciventris,C. anonae,C. rosa complex (Diptera,Tephritidae)

Previous attempts to resolve the Ceratitis FAR complex (Ceratitis fasciventris, Ceratitis anonae, Ceratitis rosa, Diptera, Tephritidae) showed contrasting results and revealed the occurrence of five microsatellite genotypic clusters (A, F1, F2, R1, R2). In this paper we explore the potential of wing morphometrics for the diagnosis of FAR morphospecies and genotypic clusters. We considered a set of 227 specimens previously morphologically identified and genotyped at 16 microsatellite loci. Seventeen wing landmarks and 6 wing band areas were used for morphometric analyses. Permutational multivariate analysis of variance detected significant differences both across morphospecies and genotypic clusters (for both males and females). Unconstrained and constrained ordinations did not properly resolve groups corresponding to morphospecies or genotypic clusters. However, posterior group membership probabilities (PGMPs) of the Discriminant Analysis of Principal Components (DAPC) allowed the consistent identification of a relevant proportion of specimens (but with performances differing across morphospecies and genotypic clusters). This study suggests that wing morphometrics and PGMPs might represent a possible tool for the diagnosis of species within the FAR complex. Here, we propose a tentative diagnostic method and provide a first reference library of morphometric measures that might be used for the identification of additional and unidentified FAR specimens.     

Leinendera achaeta sp. n., a new species of robber fly from Brazil (Diptera,Asilidae, Asilinae)

The third species of the Neotropical genus Leinendera Carrera, 1945, Leinendera achaeta sp. n., is described from Rio Grande do Sul state, Brazil. The habitus, wing and male terminalia are described and illustrated, and a key to the three Brazilian species is provided.     

Palaearctic species of Rhamphomyia (Pararhamphomyia) anfractuosa group (Diptera,Empididae)

Palaearctic species of the Rhamphomyia (Pararhamphomyia) anfractuosa group are revised. Rhamphomyia (Pararhamphomyia) biflexata sp. n., Rhamphomyia (Pararhamphomyia) lineodorsata sp. n., Rhamphomyia (Pararhamphomyia) nudiscutellata sp. n., and Rhamphomyia (Pararhamphomyia) shatalkini sp. n. (all from Russian Far East) are described and illustrated. A key to Palaearctic species of the Rhamphomyia (Pararhamphomyia) anfractuosa group is provided.     

Nuclear ribosomal internal transcribed spacer 1 (ITS1) variation in the Anastrepha fraterculus cryptic species complex (Diptera,Tephritidae) of the Andean region

The nuclear ribosomal internal transcribed spacer 1 (ITS1) was sequenced for Anastrepha fraterculus (Wiedemann, 1830) originating from 85 collections from the northern and central Andean countries of South America including Argentina (Tucumán), Bolivia, Perú, Ecuador, Colombia, and Venezuela. The ITS1 regions of additional specimens (17 collections) from Central America (México, Guatemala, Costa Rica, and Panamá), Brazil, Caribbean Colombia, and coastal Venezuela were sequenced and together with published sequences (Paraguay) provided context for interpretation. A total of six ITS1 sequence variants were recognized in the Andean region comprising four groups. Type I predominates in the southernmost range of Anastrepha fraterculus. Type II predominates in its northernmost range. In the central and northern Andes, the geographic distributions overlap and interdigitate with a strong elevational effect. A discussion of relationships between observed ITS1 types and morphometric types is included.     

A new species of the genus Baeoura from Morocco,with a key to the West Palaearctic species (Diptera,Tipuloidea, Limoniidae)

The genus Baeoura is represented in Morocco by two species, Baeoura ebenina Starý, 1981, and Baeoura staryi sp. n. The new species is described and illustrated, and a key to the West Palaearctic species of Baeoura is presented.     

Cytogenetic and symbiont analysis of five members of the B. dorsalis complex (Diptera,Tephritidae): no evidence of chromosomal or symbiont-based speciation events

The Bactrocera dorsalis species complex, currently comprising about 90 entities has received much attention. During the last decades, considerable effort has been devoted to delimiting the species of the complex. This information is of great importance for agriculture and world trade, since the complex harbours several pest species of major economic importance and other species that could evolve into global threats. Speciation in Diptera is usually accompanied by chromosomal rearrangements, particularly inversions that are assumed to reduce/eliminate gene flow. Other candidates currently receiving much attention regarding their possible involvement in speciation are reproductive symbionts, such as Wolbachia, Spiroplasma, Arsenophonus, Rickettsia and Cardinium. Such symbionts tend to spread quickly through natural populations and can cause a variety of phenotypes that promote pre-mating and/or post-mating isolation and, in addition, can affect the biology, physiology, ecology and evolution of their insect hosts in various ways. Considering all these aspects, we present: (a) a summary of the recently gained knowledge on the cytogenetics of five members of the Bactrocera dorsalis complex, namely Bactrocera dorsalis s.s., Bactrocera invadens, Bactrocera philippinensis, Bactrocera papayae and Bactrocera carambolae, supplemented by additional data from a Bactrocera dorsalis s.s. colony from China, as well as by a cytogenetic comparison between the dorsalis complex and the genetically close species, Bactrocera tryoni, and, (b) a reproductive symbiont screening of 18 different colonized populations of these five taxa. Our analysis did not reveal any chromosomal rearrangements that could differentiate among them. Moreover, screening for reproductive symbionts was negative for all colonies derived from different geographic origins and/or hosts. There are many different factors that can lead to speciation, and our data do not support chromosomal and/or symbiotic-based speciation phenomena in the taxa under study.     

A new species group in Megaselia,the lucifrons group,with description of a new species (Diptera,Phoridae)

With 1,400 described species, Megaselia is one of the most species-rich genera in the animal kingdom, and at the same time one of the least studied. An important obstacle to taxonomic progress is the lack of knowledge concerning the phylogenetic structure within the genus. Classification of Megaselia at the level of subgenus is incomplete although Schmitz addressed several groups of species in a series of monographs published from 1956 to 1981. Another problem is the lack of molecular phylogenetic analyses to support morphology-based conclusions. As a contribution towards addressing these problems, we here circumscribe a previously unrecognized monophyletic lineage of Megaselia consisting of species similar to Megaselia lucifrons. We base this taxonomic decision on morphological study of an extensive phorid material from Sweden, complemented by molecular analyses of select exemplars using two markers (COI and 28S). We name the clade the lucifrons species group, and show that it contains three distinct species. Our results also demonstrate that Megaselia subnitida Lundbeck, 1920, previously treated as a synonym of Megaselia lucifrons Schmitz, 1918, is a separate species, and we remove it from synonymy. The third species in the group was previously unknown; we describe it here as Megaselia albalucifrons sp. n.     

Taxonomic studies of the Lygephila lubrica (Freyer, 1842) species complex with notes on other species in the genus (Lepidoptera,Erebidae, Toxocampinae)

The taxa of the Lygephila lubrica (Freyer, 1846) species complex are revised. The genital features of all known taxa are described and illustrated, with special reference to the structure of the vesica. Genitalia of Lygephila lubrica from different places in Russia, Central Asia and China are studied, illustrated and compared with different Mongolian populations. Lygephila kazachkaratavika, described as a subspecies, is raised to a species level, stat. n. Neotypes of Lygephila lubrosa (Staudinger, 1901), Lygephila lubrosa kazachkaratavika Stshetkin YuL & Stshetkin YuYu, 1994 [1997] and Lygephila lubrosa orbonaria Stshetkin YuL & Stshetkin YuYu, 1994 [1997] are designated. The female genitalia of the type of Lygephila lupina (Graeser, 1890) is described and illustrated for the first time, and Lygephila mirabilis (Bryk, 1948) treated here as a junior subjective synonym, syn. n.     

Three new genera of Neotropical Mimallonidae (Lepidoptera,Mimallonoidea, Mimallonidae) with descriptions of three new species

Three new genera of Mimallonidae are described. The monotypic genus Tostallo gen. n. is erected to contain “Perophora” albescens Jones, 1912, which was previously placed in the preoccupied genus Perophora Harris, 1841 and was never formally moved to a valid genus. Perophora is a junior homonym of Cicinnus Blanchard, 1852, but the name albescens is not appropriately placed in Cicinnus due to external and genitalia characteristics entirely unique to the species albescens. The female of Tostallo albescens comb. n. is described and both sexes are figured for the first time. Auroriana gen. n. is erected to contain Auroriana florianensis (Herbin, 2012), comb. n. previously described as Cicinnus florianensis, and two new species: Auroriana colombiana sp. n. from Colombia and Auroriana gemma sp. n. from southeastern and southern Brazil. The female of Auroriana florianensis is described and figured for the first time. Finally, the monotypic genus Micrallo gen. n. is erected to include a new species, Micrallo minutus sp. n. described from northeastern Brazil.     

Contribution to the knowledge of the Carabus (Archiplectes) satyrus Kurnakov, 1962, species complex in Abkhazia (Coleoptera,Carabidae, Carabini)

This study is based on a comparative analysis of extensive material of Carabus (Archiplectes) satyrus Kurnakov, 1962, its various forms and related taxa recently collected by the authors and some other collectors in Abkhazia. The status or specific affiliations of several subspecies are changed and a subspecies is described. Carabus (Archiplectes) besleticus Kurnakov, 1972, stat. n. is treated as a separate species housing six hitherto established subspecies in addition to the nominal type: Carabus (Archiplectes) besleticus mtsaranus Kurnakov, 1972, Carabus (Archiplectes) besleticus duripshensis Kurnakov, 1972, Carabus (Archiplectes) besleticus napraensis Belousov & Zamotajlov, 1993, Carabus (Archiplectes) besleticus dsychvensis Kurnakov, 1972, Carabus (Archiplectes) besleticus adzinbai Retezár, 2013, and Carabus (Archiplectes) besleticus resheviensis subsp. n. Carabus (Archiplectes) satyrus is treated as monotypical while the specific status of Carabus (Archiplectes) pseudopshuensis Zamotajlov, 1991, earlier proposed by Fominykh and Zamotajlov (2012), is confirmed based on the morphological and morphometric data.     

Three new species of Tricimba Lioy from the West Palaearctic region (Diptera,Chloropidae)

Tricimba rudolfi Kubík, sp. n. (Czech Republic, Portugal), Tricimba chalupi Kubík, sp. n. (Czech Republic), and Tricimba dursuni Kubík, sp. n. (Turkey) are described and illustrated. First records of Tricimba kaplanae Dely-Draskovits, 1983 from Corsica and Tricimba hungarica Dely-Draskovits, 1983 from Turkey are listed.     

Relevant genetic differentiation among Brazilian populations of Anastrepha fraterculus (Diptera,Tephritidae)

We used a population genetic approach to detect the presence of genetic diversity among six populations of Anastrepha fraterculus across Brazil. To this aim, we used Simple Sequence Repeat (SSR) markers, which may capture the presence of differentiative processes across the genome in distinct populations. Spatial analyses of molecular variance were used to identify groups of populations that are both genetically and geographically homogeneous while also being maximally differentiated from each other. The spatial analysis of genetic diversity indicates that the levels of diversity among the six populations vary significantly on an eco-geographical basis. Particularly, altitude seems to represent a differentiating adaptation, as the main genetic differentiation is detected between the two populations present at higher altitudes and the other four populations at sea level. The data, together with the outcomes from different cluster analyses, identify a genetic diversity pattern that overlaps with the distribution of the known morphotypes in the Brazilian area.     

A review of the genus Toxorhina Loew from China,with descriptions of three new species (Diptera,Limoniidae, Limoniinae)

The genus Toxorhina Loew from China is reviewed. Seven species belonging to the subgenus Ceratocheilus Wesche are recognized, of which three species, Toxorhina (Ceratocheilus) huanglica sp. n., Toxorhina (Ceratocheilus) omnifusca sp. n. and Toxorhina (Ceratocheilus) univirgata sp. n., are described as new to science, Toxorhina (Ceratocheilus) fuscolimbata Alexander is recorded from China for the first time, and three known species are redescribed and illustrated.