High molecular and phenotypic diversity in the Merodon avidus complex (Diptera,Syrphidae): cryptic speciation in a diverse insect taxon

This paper examines molecular and phenotypic variability in the widely spread European hoverfly species complex Merodon avidus. Herein, based on the mitochondrial DNA (mtDNA) sequences of the cytochrome c oxidase subunit I (COI) and morphometric wing parameters, M. avidus is shown to comprise a complex of cryptic species, and one variety is redefined as a valid species: M. bicolor Gil Collado, 1930 (as var. of spinipes). The species M. bicolor, M. avidus A, and M. avidus B were clearly delimited based on their wing size. A total of 29 M. avidus and M. bicolor individuals presented 20 mtDNA haplotypes, four of which were shared by M. avidus A and M. avidus B, three were confined to M. bicolor, seven to M. avidus A, and six to M. avidus B. Sequence divergences between lineages occurring in the Balkan and in Spain ranged from 4.93 to 6.0 (uncorrected p in %) whereas divergences between M. avidus A and M. avidus B were 0.26 to 1.56. Divergence among morphologically identified individuals of M. avidus A and M. avidus B species ranged from 0.13 to 1.58, and from 0.13 to 0.52, respectively. The phenotypic substructuring and observed genetic uniqueness of populations in spatially and temporally fragmented M. avidus taxa were used to identify genetic units. The early split of two allopatric lineages, Spanish M. bicolor and Balkan M. avidus, was followed by diversification in each lineage. Present‐day morphological uniformity masks much of the genetic complexity of lineages within the M. avidus complex. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 819–833.     

Amiota sinuata species group from eastern Malaysia (Diptera, Drosophilidae)

Eight species of the Amiota sinuata species group are reported from eastern Malaysia, including six new species, A. bispinula, A. cerata, A. curvibacula, A. lambirensis, A. parviserrata and A. quadrifoliolata spp. nov. A key to Asian species of the group is provided.     

Systematics and phylogeny of the tribe Paragini (Diptera: Syrphidae) based on molecular and morphological characters

We studied the phylogeny and systematics of the tribe Paragini (Diptera: Syrphidae) using morphological and molecular data. The paper presents separate parsimony analyses of both adult morphological characters and partial DNA sequence data from mitochondrial cytochrome c oxidase I and nuclear ribosomal 28S rRNA gene, as well as a combined analysis of all the data. The data set of morphological characters included some features of the male terminalia (i.e. shape of the ejaculatory apodeme; relative position of elements of the aedeagal complex; shape of surstylar apodeme; shape of the aedeagal apodeme) not previously used in the systematics of the Paragini. The trees obtained from separate parsimony analyses of molecular and morphological data produced almost identical topologies. Four lineages are supported by the combined data set, and we establish two new subgenera, i.e. Serratoparagus Vujić et Radenković subgen. nov., and Afroparagus Vujić et Radenković subgen. nov., and redefine Pandasyopthalmus Stuckenberg, 1954 stat. rev. and Paragus Latreille, 1804, stat. rev. The monophyly of the Pandasyopthalmus clade, including the species fitting neither of the current species groups ( jozanus -group) of Paragini, is established. Diagnoses of all known species groups are presented, including a new arrangement of almost all valid species of Paragini.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 507–536.     

New data on the evolutionary history of the European bison (Bison bonasus) based on subfossil remains from Southeastern Europe

The origin and evolutionary history of the European bison Bison bonasus (wisent) have become clearer after several morphological, genomic, and paleogenomic studies in the last few years, but these paleogenomic studies have raised new questions about the evolution of the species. Here, we present additional information about the population diversity of the species based on the analysis of new subfossil Holocene remains from the Balkan Peninsula. Seven ancient samples excavated from caves in Western Stara Planina in Bulgaria were investigated by mitochondrial D‐loop (HVR1) sequence analysis. The samples were dated to 3,800 years BP by radiocarbon analysis. Additionally, a phylogenetic analysis was performed to investigate the genetic relationship among the investigated samples and all mitochondrial DNA sequences from the genus Bison available in GenBank. The results clustered with the sequences from the extinct Holocene South‐Eastern (Balkan) wisent to the fossil Alpine population from France, Austria, and Switzerland, but not with those from the recent Central European (North Sea) one and the now extinct Caucasian population.In conclusion, these data indicate that the Balkan wisent that existed in historical time represented a relict and probably an isolated population of the Late Pleistocene‐Holocene South‐Western mountainous population of the wisent.     

First record of the bacterial endosymbiont Wolbachia for phytophagous hoverflies from genus Merodon (Diptera: Syrphidae)

Wolbachia is a widespread bacterial endosymbiont among arthropod species. It influences the reproduction of the host species and also mitochondrial DNA diversity. Until now there were only a few studies that detected Wolbachia infections in hoverflies (Diptera: Syrphidae), and this is the first broader study with the aim of examining the incidence of Wolbachia in the hoverfly genus Merodon. The obtained results indicate an infection rate of 96% and the presence of both Wolbachia supergroup A and B, which are characteristic for most of the infected arthropod species. Additionally, the presence of multiple Wolbachia strains in the Merodon aureus group species was detected and the mitochondrial DNA COI‐based relationships of the group are discussed in the light of infection. Finally, we discuss plant‐mediated horizontal transmission of Wolbachia strains among the studied hoverfly species.     

Morphology of the larvae,male genitalia and DNA sequences of Anopheles (Kerteszia) pholidotus (Diptera: Culicidae) from Colombia

Since 1984, Anopheles (Kerteszia) lepidotus has been considered amosquito species that is involved in the transmission of malaria in Colombia, afterhaving been incriminated as such with epidemiological evidence from a malariaoutbreak in Cunday-Villarrica, Tolima. Subsequent morphological analyses of femalescaptured in the same place and at the time of the outbreak showed that the speciesresponsible for the transmission was not An. lepidotus, but ratherAnopheles pholidotus. However, the associated morphologicalstages and DNA sequences of An. pholidotus from the foci ofCunday-Villarrica had not been analysed. Using samples that were caught recently fromthe outbreak region, the purpose of this study was to provide updated and additionalinformation by analysing the morphology of female mosquitoes, the genitalia of malemosquitoes and fourth instar larvae of An. pholidotus, which wasconfirmed with DNA sequences of cytochrome oxidase I and rDNA internal transcribedspacer. A total of 1,596 adult females were collected in addition to 37 larvalcollections in bromeliads. Furthermore, 141 adult females, which were captured fromthe same area in the years 1981-1982, were analysed morphologically. Ninety-five DNAsequences were analysed for this study. Morphological and molecular analyses showedthat the species present in this region corresponds to An.pholidotus. Given the absence of An. lepidotus, even inrecent years, we consider that the species of mosquitoes that was previouslyincriminated as the malaria vector during the outbreak was indeed An.pholidotus, thus ending the controversy.     

Morphometric study of third-instar larvae from five morphotypes of the Anastrepha fraterculus cryptic species complex (Diptera,Tephritidae)

The occurrence of cryptic species among economically important fruit flies strongly affects the development of management tactics for these pests. Tools for studying cryptic species not only facilitate evolutionary and systematic studies, but they also provide support for fruit fly management and quarantine activities. Previous studies have shown that the South American fruit fly, Anastrephafraterculus, is a complex of cryptic species, but few studies have been performed on the morphology of its immature stages. An analysis of mandible shape and linear morphometric variability was applied to third-instar larvae of five morphotypes of the Anastrephafraterculus complex: Mexican, Andean, Ecuadorian, Peruvian and Brazilian-1. Outline geometric morphometry was used to study the mouth hook shape and linear morphometry analysis was performed using 24 linear measurements of the body, cephalopharyngeal skeleton, mouth hook and hypopharyngeal sclerite. Different morphotypes were grouped accurately using canonical discriminant analyses of both the geometric and linear morphometry. The shape of the mandible differed among the morphotypes, and the anterior spiracle length, number of tubules of the anterior spiracle, length and height of the mouth hook and length of the cephalopharyngeal skeleton were the most significant variables in the linear morphometric analysis. Third-instar larvae provide useful characters for studies of cryptic species in the Anastrephafraterculus complex.     

The role of odour and visual cues in the pan-trap catching of hoverflies (Diptera: Syrphidae)

Coloured pan traps are frequently used to attract and catch insects, such as in the monitoring of populations of beneficial insects in classical or conservation biological control. They are also used in the evaluation of the recovery of insect populations after disturbance and in many other situations where an estimate of relative insect numbers is required. However, the fact that traps may be visible to the insects over a considerable distance can influence the interpretation of catch data. This difficulty may arise, for example, if traps along a transect can attract insects from some or all of the other transect positions. This study compared the effect of different coloured traps on attraction and catch of hoverflies. The hypothesis was that completely yellow traps would attract hoverflies from a distance, while traps that were green outside and yellow inside would catch fewer flies because only those from above or near the trap can see the yellow stimulus. A subsidiary hypothesis was that rose water would enhance hoverfly capture rates. For the two main hoverfly species captured, Melanostoma fasciatum (Macquart) and Melangyna novaezelandiae (Macquart), significantly more individuals were caught in completely yellow traps than in yellow and green or in completely green traps. Moreover, the addition of rose water increased the number of hoverflies caught significantly. It is suggested that if a measure of hoverfly numbers relating to a particular distance along a transect is required, consideration should be given to the ability of hoverflies to see yellow traps from a distance. The use of traps that are green outside would more accurately reflect the local abundance of hoverflies. If higher trap catches of hoverflies are needed for statistical purposes, rose water can enhance catches.     

Taxonomy of Chrysotoxum festivum Linnaeus, 1758 (Diptera: Syrphidae) – an integrative approach

An integrative taxonomic approach revealed two taxa within Chrysotoxum festivum (Linnaeus, 1758) (Diptera, Syrphidae), C. festivum A and C. festivum B. In addition to morphological differences, results also showed significant distinction in geometric morphometrics of wings and surstyli, and in DNA sequence data (nuclear ITS2 sequences) between C. festivum A, C. festivum B, and the closely related species C. elegans Loew, 1841. From examination of type material, the name C. tomentosum Giglio‐Tos, 1890 is proposed for C. festivum B, and the taxon is redefined. © 2013 The Linnean Society of London     

Systematic and morphological studies of the genus Chaetopleurophora Schmitz (Diptera: Phoridae) occurring in Japan

Japanese species of the genus Chaetopleurophora are reviewed. All belong to the C. erythronota group. The following three species from Japan are described: C. rhomboidea sp. nov., C. pygidialis Schmitz and C. dividua sp. nov. The male and female genitalia are studied and further examples of unique characters of the genus including asymmetric features are added. The male aedeagus of the genus is illustrated for the first time. The aedeagus of the species treated in this study consists of only two components, the inner core plate and the outer jacket plate. The jacket plate wraps sinistrally around the core plate. The combination of the core plate and the jacket plate forms complex, asymmetric features of the aedeagus in the Phoridae. The structure around the genital opening in the female genitalia protrudes posteriorly under the segment IX + X, and shows asymmetric features in C. rhomboidea sp. nov. with a bilaterally different degree of sclerotization, shifted genital opening to the right side and a membranous ribbon just on the left side. In addition, C. dividua sp. nov. is different from most of the species in the C. erythronota group, and very closely related to C. multiseriata (known in North America) in the male and female genitalia, wing venation and bristle formation on the scutellum. It is suggested that C. dividua sp. nov. forms a monophyletic group with C. multiseriata and the related species.     

Taxonomic notes on the genus Holepyris (Hymenoptera: Bethylidae) from Korea with descriptions of six new species

Six new species of the genus Holepyris (Hymenoptera: Bethylidae: Epyrinae) from Korea are described: H. brevicarinatus sp. nov., H. crinitus sp. nov., H. dimidiatus sp. nov., H. longicephalus sp. nov., H. mucro sp. nov. and H. multo sp. nov. They are described with illustrations of male genitalia and the biometric measurement data. A key to the Korean species of Holepyris based on males is provided.     

Towards a phylogeny of the flesh flies (Diptera: Sarcophagidae): morphology and phylogenetic implications of the acrophallus in the subfamily Sarcophaginae

The morphology of the acrophallus, the distal portion of the male phallus carrying the phallotreme, was studied in 72 exemplar species representing 56 genera and subgenera of the flesh fly subfamily Sarcophaginae. For 42 of those species, scanning electron microscopy was used to clarify the phallic morphology. Terms used to describe the male genitalia were updated based on new interpretations of homology. Male genitalic characters, combined with other morphological characters of adult males and females and of larvae, were used to construct a phylogeny. The monophyly of the subfamily was supported, and some generic‐level sister‐group relationships proposed in the literature, but without previous cladistic analyses, were also supported. The genus Blaesoxipha Loew, as currently recognized, was not monophyletic in our analysis. The genus Helicobia Coquillett is synonymized with Sarcophaga Meigen syn. nov. and treated as a subgenus of the latter. The Sarcophaga subgenera Neobellieria Blanchard and Mehria Enderlein were not monophyletic. Many of the clades in the analysis were supported primarily or exclusively by male genitalic character states, highlighting the importance of the male genitalia as a source of morphological characters for sarcophagine phylogeny. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 740–778.     

Variation of wing shape in the Drosophila virilis species group (Diptera: Drosophilidae)

Wing shape variation was analysed with geometric morphometric methods in 17 laboratory strains, representing 11 closely related species (including two subspecies) of the Drosophila virilis group: D. virilis, D. lummei, D. novamexicana, D. americana americana, D. americana texana, D. montana, D. lacicola, D. flavomontana, D. borealis, D. littoralis, D. ezoana and D. kanekoi. Overall shape estimated using Procrustes coordinates of 14 landmarks was highly variable among strains and very similar in females and males. The landmarks in the distal part of the wing showed higher variation across strains than those in the proximal part. Procrustes distances between species were not consistent with phylogenetic distances previously suggested for the virilis group. Moreover, Procrustes distances between strains within species and within two major phylads (virilis and montana) were comparable with those between species and between phylads, respectively. The most different from other members of the group was the endemic D. kanekoi species, currently viewed as separate subphylad within the montana phylad. Allometric effects were found to be partly responsible for shape differences between the strains. Three most significant shape transformations were considered using the relative warp analysis and the strains were ordinated in accordance with transformation values. The pattern of relative warp scores could be easily interpreted only for the third warp explaining about 13% of shape variation. It separated the largest species, D. montana, D. ezoana and D. kanekoi, from other ones and was mainly associated with shape changes in the proximal region of the wing. The results of the present work suggest that wing shape in the virilis species group is not related to the speciation process. The observed proximal‐distal contrasts and allometric effects are in agreement with data of other studies, in which wing shape variation was analysed within Drosophila species.     

Macroevolution of hoverflies (Diptera: Syrphidae): the effect of using higher‐level taxa in studies of biodiversity,and correlates of species richness

We test a near‐complete genus level phylogeny of hoverflies (Diptera: Syrphidae) for consistency with a null model of clade growth having uniform probabilities of speciation and extinction among contemporaneous species. The phylogeny is too unbalanced for this null model. Importantly, the degree of imbalance in the phylogeny depends on whether the phylogeny is analysed at the genus level or species level, suggesting that genera ought not to be used uncritically as surrogates for species in large‐scale evolutionary analyses. Tests for a range of morphological, life‐history and ecological correlates of diversity give equivocal results, but suggest that high species‐richness may be associated with sexual selection and diet breadth. We find no correlation between species‐richness and either body size or reproductive rate.