Further observations on some blackflies (Diptera: Simuliidae) associated with mayflies (Ephemeroptera: Baetidae and Heptageniidae) in Cameroon

An account is given of the infestation rates, attachment sites, larval size distribution and dispersion of Simulium baetiphilum on nymphs of Baetidae. The mechanisms of choice shown by S. afronuri and S. lumbwanum infesting Afronurus spp. are discussed.     

Towards a new paradigm in mayfly phylogeny (Ephemeroptera): combined analysis of morphological and molecular data

This study represents the first formal morphological and combined (morphological and molecular) phylogenetic analyses of the order Ephemeroptera. Taxonomic sampling comprised 112 species in 107 genera, including 42 recognized families (all major lineages of Ephemeroptera). Morphological data consisted of 101 morphological characters. Molecular data were acquired from DNA sequences of the 12S, 16S, 18S, 28S and H3 genes. The Asian genus Siphluriscus (Siphluriscidae) was supported as sister to all other mayflies. The lineages Carapacea, Furcatergalia, Fossoriae, Pannota, Caenoidea and Ephemerelloidea were supported as monophyletic, as were many of the families. However, some recognized families (for example, Ameletopsidae and Coloburiscidae) and major lineages (such as Setisura, Pisciforma and Ephemeroidea among others) were not supported as monophyletic, mainly due to convergences within nymphal characters. Clade robustness was evaluated by multiple methods and approaches.     

An electrophoretic and morphological study of three Ecdyonurus species (Ephemeroptera: Heptageniidae) occurring in the British Isles

Abstract The objective was to investigate the validity of three closely-related British species: Ecdyonurus dispar, E. venosus and E. torrentis . The species were characterized by eleven enzyme-substrates and fifteen different enzyme-loci, comparisons being made not only between species but also between five populations of E.dispar (three from Britain, one from France, one from Switzerland), two populations of E.venosus and two populations of E. torrentis (one from Britain, one from Switzerland for both species).
Four monomorph enzyme-loci (aldolase, mannose phosphate isomerase, arginine phosphokinase, glutamate-oxaloacetate transaminase-2) exhibited interspecific differences in their mobilities and therefore validated the conclusion that E. dispar, E. venosus and E. torrentis are distinct species. There were no monomorph enzyme-loci that were different between populations of the same species. There were, however, some intraspecific differences revealed by the presence of polymorphic enzyme-loci: seven in E.dispar (retinol dehydrogenase, hexokinase-1 and 2, glutamate-oxaloacetate transaminase-!, malate dehydrogenase-1, phospho-glucomutase, indophenol oxidase-2), three in E. venosus (glutamate-oxaloacetate transaminase-1, malate dehydrogenase-1, phospho-glucomutase) and three in E. torrentis (hexokinase-1, glutamate-oxaloacetate transaminase-1, malate dehydrogenase-1).
The morphological characters of larvae and adults were examined and some were used in new keys to larvae and adults.     

Swarm behaviour and mate competition in mayflies (Ephemeroptera)

Although mayfly swarms are frequently cited as an example of lekking by insects, little is known about the behaviour of individuals within a swarm, or how mate-selection takes place. A study of five species of mayfly over a period of 10 consecutive years has revealed species-specific differences in the flight pattern of swarming males and in the ability of males to recognize swarms of their own species. Males of four of the five species jostle other males in the swarm at all times except when mating: mating pairs are not jostled. The pattern of jostling varies with the species. Measurements of the sperm content of the vesicula seminalis and of the wing length of members of individual swarms show that larger wing size is positively correlated with the presence of less sperm. The vesicula seminalis is always filled with sperm at the beginning of the imaginal stage and the testes regress before the beginning of the imaginal stage. If the volume of sperm in the vesicula seminalis is a valid index of mating success then males with larger wings have the highest success. Large wings may bestow an advantage during jostling. The males of Ephemera danica , which do not jostle, glide with outspread wings; these outspread wings may attract females, the largest wings being the most attractive. Females of all five species enter the swarm a few at a time, although many females may be resting beneath the swarm. This phased entry may decrease the attraction of the swarm for predators. The number of females in a swarm is not correlated with swarm size, and the factors which enable females to regulate their entry into a swarm remain obscure.     

Molecular and morphological phylogeny of Menispermaceae (Ranunculales)

The phylogeny of Menispermaceae (Ranunculales) is reconstructed here using both morphological and molecular data from a broad sample of species. Morphological data include characters of leaves, wood, flowers, fruits, seeds, pollen and phytochemistry from 73 species representing the different subgroups recognized within the family. This dataset allowed us to study the morphological evolutionary trends in Menispermaceae. The molecular data focused on cpDNA sequences: rbcL and atpB. Maximum parsimony (MP) and Bayesian methods were used to reconstruct the phylogeny of Menispermaceae. The results obtained from the three datasets are partly incongruent. The morphology indicates that Menispermaceae can be divided into two major groups, one including the tribes Fibraureae and Tinosporeae, the other one including the tribes Anomospermeae, Pachygoneae and Menispermeae. Only Burasaia Thouars was not included in any of these groups. The characters of fruits and seeds appeared to be most useful to differentiate these groups whereas convergences were found for the androecium.     

Gondwanan Gymnochilini (Coleoptera: Trogossitidae): generic concepts,review of New Zealand species and long‐range Pacific dispersal

Members of Leperina Erichson (Trogossitidae: Gymnochilini) from New Zealand, New Caledonia and Lord Howe Island are morphologically similar to members of the endemic Juan Fernandez Island genus Phanodesta Reitter, sharing at least one obvious character, elytral carinae that are beaded and contain well‐defined punctures. To test the monophyly of Leperina and Phanodesta, we reconstructed phylogenetic relationships of the genera of the tribe Gymnochilini by a cladistic analysis of 22 terminals and 47 adult characters rooted with one genus of trogossitine. Leperina is rendered paraphyletic by the placements of Seidlitzella Jakobson and Phanodesta. Kolibacia n.gen. (type species Leperina tibialis Reitter) is described for east Palaearctic species included formerly in Leperina (two new combinations); New Zealand Leperina and other species from New Caledonia and Lord Howe Island (Ostoma pudicum Olliff) are transferred to Phanodesta (six new combinations); and the remaining species are retained in Leperina. The following species are described as new: Phanodesta carinata n.sp., P. manawatawhi n.sp., P. oculata n.sp. and P. tepaki n.sp. Leperina ambiguum Broun is transferred to Grynoma Sharp resulting in a new combination and three new synonymies for New Zealand trogossitines: Leperina interrupta Brookes n.syn. and Leperina sobrina (White) n.syn. [= Phanodesta farinosa (Sharp)], and Trogosita affinis White n.syn. (= Tenebroides mauritanicus Linnaeus). A key to the New Zealand species and a checklist for the species of the Kolibacia, Leperina and Phanodesta are provided. The derived placement of Juan Fernandez Phanodesta in the phylogeny is evidence for long‐distance dispersal from Australasia. A tally of all Juan Fernandez Islands Coleoptera shows derivation mostly from Chile and South America, with few from the southern Pacific region, rarely from Australasia.     

Croatian mayflies (Insecta,Ephemeroptera): species diversity and distribution patterns

Knowledge of the mayfly biodiversity in the Balkan Peninsula is still far from complete. Compared to the neighbouring countries, the mayfly fauna in Croatia is very poorly known. Situated at the crossroads of central and Mediterranean Europe and the Balkan Peninsula, Croatia is divided into two ecoregions: Dinaric western Balkan and Pannonian lowland. Mayflies were sampled between 2003 and 2013 at 171 sites, and a total of 66 species was recorded. Combined with the literature data, the Croatian mayfly fauna reached a total of 79 taxa. Of these, 29 species were recorded for the first time in Croatia while 15 species were not previously recorded in Dinaric western Balkan ecoregion. Based on the mayfly assemblage, sampling sites were first structured by ecoregion and then by habitat type. In comparison with the surrounding countries, the Croatian mayfly fauna is the most similar to the Hungarian and Bosnian fauna. Some morphologically interesting taxa such as Baetis cf. nubecularis Eaton, 1898 and Rhithrogena from the diaphana group were recorded. Ephemera cf. parnassiana Demoulin, 1958, the species previously recorded only from Greece, was also recorded.     

Toward a DNA taxonomy of Alpine Rhithrogena (Ephemeroptera: Heptageniidae) using a mixed Yule-coalescent analysis of mitochondrial and nuclear DNA

Aquatic larvae of many Rhithrogena mayflies (Ephemeroptera) inhabit sensitive Alpine environments. A number of species are on the IUCN Red List and many recognized species have restricted distributions and are of conservation interest. Despite their ecological and conservation importance, ambiguous morphological differences among closely related species suggest that the current taxonomy may not accurately reflect the evolutionary diversity of the group. Here we examined the species status of nearly 50% of European Rhithrogena diversity using a widespread sampling scheme of Alpine species that included 22 type localities, general mixed Yule-coalescent (GMYC) model analysis of one standard mtDNA marker and one newly developed nDNA marker, and morphological identification where possible. Using sequences from 533 individuals from 144 sampling localities, we observed significant clustering of the mitochondrial (cox1) marker into 31 GMYC species. Twenty-one of these could be identified based on the presence of topotypes (expertly identified specimens from the species' type locality) or unambiguous morphology. These results strongly suggest the presence of both cryptic diversity and taxonomic oversplitting in Rhithrogena. Significant clustering was not detected with protein-coding nuclear PEPCK, although nine GMYC species were congruent with well supported terminal clusters of nDNA. Lack of greater congruence in the two data sets may be the result of incomplete sorting of ancestral polymorphism. Bayesian phylogenetic analyses of both gene regions recovered four of the six recognized Rhithrogena species groups in our samples as monophyletic. Future development of more nuclear markers would facilitate multi-locus analysis of unresolved, closely related species pairs. The DNA taxonomy developed here lays the groundwork for a future revision of the important but cryptic Rhithrogena genus in Europe.     

Identification and discrimination of Electrogena species by numerical methods (Ephemeroptera: Heptageniidae)

Abstract. Nine species of Electrogena 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.     

Molecular phylogeny of hinge‐beak shrimps (Decapoda: Caridea: Rhynchocinetes and Cinetorhynchus) and allies: a formal test of familiar and generic monophyly using a multilocus phylogeny

The Rhynchocinetidae (‘hinge‐beak’ shrimps) is a family of marine caridean decapods with considerable variation in sexual dimorphism, male weaponry, mating tactics, and sexual systems. Thus, this group is an excellent model with which to analyse the evolution of these important characteristics, which are of interest not only in shrimps specifically but also in animal taxa in general. Yet, there exists no phylogenetic hypothesis, either molecular or morphological, for this taxon against which to test either the evolution of behavioural traits within the Rhynchocinetidae or its genealogical relationships with other caridean taxa. In this study, we tested (1) hypotheses on the phylogenetic relationships of rhynchocinetid shrimps, and (2) the efficacy of different (one‐, two‐, and three‐phase) methods to generate a reliable phylogeny. Total genomic DNA was extracted from tissue samples taken from 17 species of Rhynchocinetidae and five other species currently or previously assigned to the same superfamily (Nematocarcinoidea); six species from other superfamilies were used as outgroups. Sequences from two nuclear genes (H3 and Enolase) and one mitochondrial gene (12S) were used to construct phylogenies. One‐phase phylogenetic analyses (SATé‐II) and classical two‐ and three‐phase phylogenetic analyses were employed, using both maximum likelihood and Bayesian inference methods. Both a two‐gene data set (H3 and Enolase) and a three‐gene data set (H3, Enolase, 12S) were utilized to explore the relationships amongst the targeted species. These analyses showed that the superfamily Nematocarcinoidea, as currently accepted, is polyphyletic. Furthermore, the two major clades recognized by the SATé‐II analysis are clearly concordant with the genera Rhynchocinetes and Cinetorhynchus, which are currently recognized in the morphological‐based classification (implicit phylogeny) as composing the family Rhynchocinetidae. The SATé‐II method is considered superior to the other phylogenetic analyses employed, which failed to recognize these two major clades. Studies using more genes and a more complete species data set are needed to test yet unresolved inter‐ and intrafamilial systematic and evolutionary questions about this remarkable clade of caridean shrimps. © 2014 The Linnean Society of London     

中国似动蜉属及一新种记述 （蜉蝣目：扁蜉科）

似动蜉属Cinygmina是我国常见扁蜉科类群,分布广泛。本文对该属我国已知4种（湖南似动蜉C. hunanensis,斜纹似动蜉C. obliquistrita,红斑似动蜉C. rubromaculata,宜兴似动蜉C. yixingensis)和一新种（叉似动蜉Cinygmina furcata sp. Nov.）的鉴别特征作了对比和图示,并确认海南似动蜉C. hainanensis是红斑似动蜉C. rubromaculata的同物异名(新异名)。     

Rhithrogena trispina sp. n., a New Species from China (Ephemeroptera: Heptageniidae)

A new species, Rhithrogena trispina sp. n. from Tian-Mu-Shan Mountain of Southeast China, is described and figured in detail at the imaginal stage. From the position of gonopore on the penis lobe, it belongs to the znojkoi-group.     

Two phoretic black-flies (Diptera: Simuliidae) and their associated mayfly host (Ephemeroptera: Heptageniidae) in Cameroon

Simulium afronuri and S.lumbwanum are recorded living in association with nymphs of Afronurus in the forest zone of West Cameroon. Observations on infestation rates, attachment sites, larval size distribution and dispersion are recorded. Evidence of niche separation between the two simuliid species is discussed.     

Molecular phylogeny and morphological evolution of the Acantharia (Radiolaria)

Acantharia are ubiquitous and abundant rhizarian protists in the world ocean. The skeleton made of strontium sulphate and the fact that certain harbour microalgal endosymbionts make them key planktonic players for the ecology of marine ecosystems. Based on morphological criteria, the current taxonomy of Acantharia was established by W.T. Schewiakoff in 1926, since when no major revision has been undertaken. Here, we established the first comprehensive molecular phylogeny from single morphologically-identified acantharian cells, isolated from various oceans. Our phylogenetic analyses based on 78 18S rDNA and 107 partial 28S rDNA revealed the existence of 6 main clades, sub-divided into 13 sub-clades. The polyphyletic nature of acantharian families and genera demonstrates the need for revision of the current taxonomy. This molecular phylogeny, which highlights the taxonomic relevance of specific morphological criteria, such as the presence of a shell and the organisation of the central junction, provides a robust phylogenetic framework for future taxonomic emendation. Finally, mapping all the existing environmental sequences available to date from different marine ecosystems onto our reference phylogeny unveiled another 3 clades and improved the understanding of the biogeography and ecology of Acantharia.     

Molecular and morphological phylogeny of Diplazontinae (Hymenoptera,Ichneumonidae)

Klopfstein, S., Quicke, D. L. J., Kropf, C. & Frick, H. (2011) Molecular and morphological phylogeny of Diplazontinae (Hymenoptera, Ichneumonidae). —Zoologica Scripta, 40, 379–402. Parasitoid wasps are among the most species rich and at the same time most understudied of all metazoan taxa. To understand their diversification and test hypotheses about their evolution, we need robust phylogenetic hypotheses. Here, we reconstruct the phylogeny of the subfamily Diplazontinae using four genes and 66 morphological characters both in separate analyses and in a total evidence approach. The resulting phylogeny is highly resolved, with most clades supported by multiple independent data partitions. It contains three highly supported genus groups, for which we suggest morphological and behavioural synapomorphies. The placement of some of the genera, especially Xestopelta Dasch, is unexpected, but also supported by morphology. Most of the genera are retrieved as monophyletic, with the exception of the morphologically diverse genus Syrphoctonus Förster. We split this genus into three genera, including Fossatyloides gen. n., to restore the phylogeny–classification link. Conflict between the morphological and the molecular topology was mostly resolved in favour of the molecular partition in the total evidence approach. We discuss reasons for this finding, and suggest strategies for future taxon and character sampling in Diplazontinae.     

Molecular phylogeny evidence of altitudinal distribution and habitat adaptation in Korean Ephemera species (Ephemeroptera: Ephemeridae)

Molecular phylogeny of the four Korean Ephemera species, Ephemera orientalis, E. sachalinensis, E. strigata, and E. separigata, was inferred from 630 bp sequences of the partial mitochondrial cytochrome c oxidase I (COI) gene. Results indicated that mean intraspecific sequence divergences were 0.70%, whereas mean interspecific divergences were 15.75%, and 17 samples were distinguished to four species correctly by COI sequences. The results also demonstrated that four species of Korean Ephemera assembled a monophyletic group with high support in maximum parsimony and maximum likelihood analyses. This Ephemera group was divided into two major clades of E. orientalis–E. sachalinensis and E. strigata–E. separigata. Furthermore, we demonstrated that this phylogeny explained altitudinal and habitat adaptations of Korean Ephemera species. The E. orientalis–E. sachalinensis clade, a widespread and lowland‐adapted mayfly group, retained plesiomorphic traits such as paired stripes on abdominal segments and was regarded as plesiotypic in terms of habitat adaptation, compared to the geographically more limited and upland‐adapted E. strigata–E. separigata clade.     

Phylogenetic analyses of the isonychiid mayflies (Ephemeroptera: Isonychiidae) in the northeast palearctic region

We investigated the molecular phylogeny of isonychiid mayflies inhabiting the East Palearctic region, Isonychia (Isonychia) japonica, Isonychia (Isonychia) ignota, Isonychia (Isonychia) ussurica and Isonychia (Prinoides) shima. We discuss their genetic structures, phylogeny and phylogeography. We collected a total of 100 specimens of isonychiid mayfly species from 47 localities of the Northeast Palearctic region (the Japanese archipelago, the Korean peninsula, the Russian Far East and Mongolia). We analyzed the DNA sequences at the mtDNA COI and 16S rRNA regions, and the nuDNA Histone H3 region. As a result of our genetic analyses of the four Northeast Palearctic isonychiid mayflies, their monophyly at the species level was supported by both the mtDNA (COI and 16S rRNA regions) and the nuDNA (Histone H3 region). In addition, it also became clear that significantly large genetic differentiation exists at the inter‐species level; thus, the relationship of “shima + (japonica + (ignota + ussurica))” is supported. Among the four isonychiid mayflies of the Northeast Palearctic area, I. (P.) shima was shown to be a basal‐most linage within the included species     

Molecular phylogeny of flat‐footed flies (Diptera: Platypezidae): main clades supported by new morphological evidence

The molecular phylogeny of flat‐footed flies is inferred from analysis of DNA sequence data from the five mitochondrial genes 12S, 16S, COI, COII and CytB, and the nuclear gene 28S and discussed with the recent systematics based on morphological features. The Bayesian inference, maximum likelihood and maximum parsimony analyses included 42 species of 18 genera, representing all four extant subfamilies (Microsaniinae, Melanderomyiinae, Callomyiinae and Platypezinae) and all known genera except one (Metaclythia). Representatives of the brachycerous taxa Lonchopteridae, Phoridae, Sciadocerinae (Phoridae) and Opetiidae are used as outgroups, and Lonchoptera was used to root the trees. Our results show Platypezidae consisting of two well‐supported clades, the first with the subfamilies Melanderomyiinae + Callomyiinae and the second formed by subfamily Platypezinae. Genus Microsania was resolved as a separate lineage distant from Platypezidae which clustered with Opetiidae as its sister group, both together forming a sister group to Platypezidae. At the generic level, the genus Agathomyia proved not to be monophyletic in any of the analyses. The species Chydaeopeza tibialis is sister to Agathomyia sexmaculata, and consequently, the genus Chydaeopeza Shatalkin, 1992 is a new junior synonym of Agathomyia Verrall, 1901. Bifurcated setae on legs of adult Platypezidae are documented as a new synapomorphy of the family, exclusive of Microsania. Outstretched wings and only a small overlap of their surfaces at resting position are considered a new synapomorphy for the subfamily Platypezinae. Other phylogenetically important characters defining main clades are documented, and their relevance/validity in phylogenetic studies is discussed. The current systematic concept of Platypezidae is discussed, and new phylogenetic hypotheses are proposed.