Systematic revision of Nearctic species of Achalcus Loew (Diptera: Dolichopodidae) with comments on their phylogeny, ecology and zoogeography

The species of the genus Achalcus Loew from the Nearctic Region are revised and a key to males and females is provided. Achalcus utahensis (Harmston & Miller) and A. oregonensis (Harmston & Miller) are redescribed, and A. californicus sp.n., A. dytei sp.n. and A. similis sp.n. are described. Together with eight of the nine known Palaearctic species of the genus, they represent a monophyletic species group, which is characterized by the presence of five pairs of dorsocentral bristles and the incorporation of the epandrial setae onto the shaft of the epandrial lobe. Like their Palaearctic congeners, most Nearctic species occur in damp habitats, such as coastal forests and riparian vegetation. Most specimens were collected in the Pacific coastal states from British Columbia to California, but one species occurs in Utah, and one of six species which were not described because males are lacking, was discovered in Ontario.     

The history of a nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo)

Previous hypotheses of phylogenetic relationships among Nearctic toads (Bufonidae) and their congeners suggest contradictory biogeographic histories. These hypotheses argue that the Nearctic Bufo are: (1) a polyphyletic assemblage resulting from multiple colonizations from Africa; (2) a paraphyletic assemblage resulting from a single colonization event from South America with subsequent dispersal into Eurasia; or (3) a monophyletic group derived from the Neotropics. We obtained approximately 2.5 kb of mitochondrial DNA sequence data for the 12S, 16S, and intervening valine tRNA gene from 82 individuals representing 56 species and used parametric bootstrapping to test hypotheses of the biogeographic history of the Nearctic Bufo. We find that the Nearctic species of Bufo are monophyletic and nested within a large clade of New World Bufo to the exclusion of Eurasian and African taxa. This suggests that Nearctic Bufo result from a single colonization from the Neotropics. More generally, we demonstrate the utility of parametric bootstrapping for testing alternative biogeographic hypotheses. Through parametric bootstrapping, we refute several previously published biogeographic hypotheses regarding Bufo. These previous studies may have been influenced by homoplasy in osteological characters. Given the Neotropical origin for Nearctic Bufo, we examine current distributional patterns to assess whether the Nearctic-Neotropical boundary is a broad transition zone or a narrow boundary. We also survey fossil and paleogeographic evidence to examine potential Tertiary and Cretaceous dispersal routes, including the Paleocene Isthmian Link, the Antillean and Aves Ridges, and the current Central American Land Bridge, that may have allowed colonization of the Nearctic.     

Parasites of grass flies (Diptera, Chloropidae) from the order hymenoptera in the Holarctic region

Hymenopterous parasitoids of grass flies of the family Chloropidae from the Palaearctic and Nearctic regions are reviewed. These parasitoids belong to four superfamilies and 16 families of Hymenoptera and were reared from 39 species of Chloropidae in the Palaearctic (less than 6% of the fauna) and only from 10 species in the Nearctic Region. The majority of parasitoids are oligo-or polyphagous species. To a certain degree, the parasitoids are specialized on one of the three host groups: (1) species developing in shoots of cereal and meadow grasses; (2) forest species developing in cones of coniferous trees; and (3) species associated with the common reed, Phragmites australis. In the Palaearctic Region, the majority of parasitoids (91 species) were reared from Oscinella frit L. s. 1.; a significantly smaller number of these parasites is known from this host in the Nearctic Region, nearly half of the parasitoids being common for both regions. The next large group of parasitoids is associated with gall-inducing species of the genus Lipara Meigen (59 species) developing in the common reed. By contrast with two other groups of parasitoids, this one includes many species of the family Ichneumonidae. It should be noted that taxonomic diversity at the third trophic level is markedly greater than at the second level.     

A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)

Abstract The campoplegine genus Benjuminia is herein revised, with ten species being described as new, in addition to the five previously known species. Phylogenetic relationships of Benjuminia to other genera are discussed, and a cladogram presented for the species. Two monophyletic species groups are recognized, one being confined to the Palearctic and the other to the Nearctic. Examination of larval morphology not only fails to reveal larval characters for distinguishing Benjuminia but leads to the conclusion that use of larval morphology to identify campoplegine species is a dubious practice.     

The Identity of Simulium murmanum Enderlein, 1935 (Diptera: Simuliidae)

The identity of Simulium murmanum is re-established by designation of lectotype and paralectotype. The species is common and widespread in the northern Holarctic Region and has up to now been reported from the Nearctic Region as S. corbis and from the Palaearctic Region as S. relictum (and S. rostratum, auct., nec Lundstrüm).     

DNA barcoding and morphology reveal three cryptic species of Anania (Lepidoptera: Crambidae: Pyraustinae) in North America,all distinct from their European counterpart

Anania coronata (Hufnagel), a Holarctic species of pyraustine crambid moth, has long been treated as having two geographically separated subspecies – the nominotypical Anania coronata in the Palaearctic Region and Anania coronata tertialis (Guenée) in the Nearctic Region. Maximum likelihood and Bayesian inference analysis of mitochondrial DNA barcodes both recover four well‐supported, reciprocally monophyletic groups within Anania coronata. Qualitative and quantitative analyses of genital structures reveal diagnostic differences that correspond to the four barcode lineages. On the basis of both molecular and morphological evidence, we conclude that Anania coronata is actually a complex of four species. Anania coronata (Hufnagel) is restricted to Europe, whereas three species occur in North America: Anania tertialis (Guenée), Anania plectilis (Grote & Robinson) and Anania tennesseensis sp.n. Yang.     

Hispines (chrysomelidae, cassidinae) of la selva biological station, costa rica

Survey work from 1992-2001 identified 139 species of hispines at the lowland part of La Selva Biological Station, Costa Rica. The tribe Cephaloleiini was the most speciose with 58 species (41.7%) followed by the Chalepini with 55 (39.5%). The fauna is most closely related to that in South America but with some genera which are more speciose in the Nearctic Region. Plant associations are known for 88 (63.3%) of the species but many of these are merely collecting records, not host plant associations. The first plant associations are reported for Alurnus ornatus, Alurnus salvini, and Acentroptera nevermanni.     

Phylogeny and historical biogeography of Agabinae diving beetles (Coleoptera) inferred from mitochondrial DNA sequences

The Agabinae, with more than 350 species, is one of the most diverse lineages of diving beetles (Dytiscidae). Using the mitochondrial genes 16S rRNA and cytochrome oxidase I we present a phylogenetic analysis based on 107 species drawn mostly from the four main Holarctic genera. Two of these genera (Ilybius and Ilybiosoma) are consistently recovered as monophyletic with strong support, Platambus is never recovered as monophyletic, and Agabus is found paraphyletic with respect to several of the species groups of Platambus. Basal relationships among the main lineages are poorly defined, although within each of them relationships are in general robust and very consistent across the parameter space, and in agreement with previous morphological analyses. In each of the two most diverse lineages (Ilybius and Agabus including part of Platambus) there is a basal split between Palearctic and Nearctic clades, estimated to have occurred in the late Eocene. The Palearctic clade in turn splits into a Western Palearctic clade and a clade containing mostly Eastern Palearctic species, and assumed to be ancestrally Eastern Palearctic but with numerous transitions to a Holarctic or Nearctic distribution. These results suggest an asymmetry in the colonization routes, as there are very few cases of transcontinental range expansions originating from the Nearctic or the Western Palearctic. According to standard clock estimates, we do not find any transcontinental shift during the Pliocene, but numerous speciation events within each of the continental or subcontinental regions.     

Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences.

Based on extensive phenetic analyses, bats of the genus Myotis have been classically subdivided into four major subgenera each of which comprise many species with similar morphological and ecological adaptations. Each subgenus thus corresponds to a distinct "ecomorph" encompassing bat species exploiting their environment in a similar fashion. As three of these subgenera are cosmopolitan, regional species assemblages of Myotis usually include sympatric representatives of each ecomorph. If species within these ecomorphs are monophyletic, such assemblages would suggest extensive secondary dispersal across geographic areas. Conversely, these ecomorphological adaptations may have evolved independently through deterministic processes, such as adaptive radiation. In this case, phylogenetic reconstructions are not expected to sort species of the same ecomorph into monophyletic clades. To test these predictions, we reconstructed the phylogenetic history of 13 American, 11 Palaearctic, and 6 other Myotis species, using sequence data obtained from nearly 2 kb of mitochondrial genes (cytochrome b and nd1). Separate or combined analyses of these sequences clearly demonstrate the existence of several pairs of morphologically very similar species (i.e., sibling species) which are phylogenetically not closely related. None of the three tested subgenera constitute monophyletic units. For instance, Nearctic and Neotropical species currently classified into the three subgenera were clustered in a single, well-supported monophyletic clade. These species thus evolved independently of their ecological equivalents from the Palaearctic region. Independent adaptive radiations among species of the genus Myotis therefore produced strikingly similar evolutionary solutions in different parts of the world. Furthermore, all phylogenetic reconstructions based on mtDNA strongly supported the existence of an unsuspected monophyletic clade which included all assayed New World species plus M. brandtii (from the Palaearctic Region). This "American" clade thus radiated into a morphologically diverse species assemblage which evolved after the first Myotis species colonized the Americas. Molecular reconstructions support paleontological evidence that species of the genus Myotis had a burst of diversification during the late Miocene-early Pliocene epoch.     

The chromosomes of Lepturinae (Coleoptera: Cerambycidae) II. A study of eight more species,with focus on Desmocerus palliatus

Following the study of 28 species of Lepturinae (Coleoptera: Cerambycidae) the karyotypes of seven additional Palaearctic and one Nearctic species are established. The 19,X male karyotypes found in genera Stictoleptura (four species), Vadonia and Judolia (one species each) confirm the loss of Y chromosome in Lepturini. The 22,XY male karyotype of Cortodera humeralis is very close to that of some species of Rhagiini (genera Gaurotes, Acmaeops, Dinoptera, all 22,XY) and Grammoptera ruficornis (24,XY) recently reported. We propose that these taxa could form a monophyletic group within Rhagiini. The karyotype of the Nearctic species Desmocerus palliates (23,neoXneoXneoY) is quite different and characterized by the presence of many acrocentric chromosomes and a complex autosome–gonosome translocation. Its particular karyotype is compatible with its present classification within a separate tribe, the Desmocerini.     

New Wolbachia endosymbionts from Nearctic and Neotropical fleas (Siphonaptera)

Several species of fleas (Siphonaptera), ectoparasites of mammals and birds, have recently been shown to harbor species of Wolbachia. Here, we extend this data set to 20 more species of Siphonaptera (Rhopalopsyllidae, Stephanocircidae, Pulicidae, Ceratophyllidae, Ctenophthalmidae, Ischnopsyllidae, Leptopsyllidae, and Malacopsyllidae) from sylvatic populations throughout the Nearctic and Neotropical regions. Using polymerase chain reaction, we targeted the Wolbachia 16S ribosomal DNA (rDNA) gene. Applying maximum parsimony- and maximum likelihood-based algorithms, as well as statistical parsimony, we conducted a phylogenetic analysis of Wolbachia 16S rDNA to evaluate its position within the known Wolbachia spp. The analysis recovered the siphonapteran Wolbachia 16S rDNA sequences as a monophyletic group and shows multiple haplotype connections between the Neotropical and Nearctic Wolbachia strains of fleas.     

Molecular phylogeny of the speciose vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial DNA sequences

Voles of the genus Microtus represent one of the most speciose mammalian genera in the Holarctic. We established a molecular phylogeny for Microtus to resolve contentious issues of systematic relationships and evolutionary history in this genus. A total of 81 specimens representing ten Microtus species endemic to Europe as well as eight Eurasian, six Asian and one Holarctic species were sequenced for the entire cytochrome b gene (1140 bp). A further 25 sequences were retrieved from GenBank, providing data on an additional 23, mainly Nearctic, Microtus species. Phylogenetic analysis of these 48 species generated four well-supported monophyletic lineages. The genus Chionomys, snow voles, formed a distinct and well-supported lineage separate from the genus Microtus. The subgenus Microtus formed the strongest supported lineage with two sublineages displaying a close relationship between the arvalis species group (common voles) and the socialis species group (social voles). Monophyly of the Palearctic pitymyid voles, subgenus Terricola, was supported, and this subgenus was also subdivided into two monophyletic species groups. Together, these groupings clarify long-standing taxonomic uncertainties in Microtus. In addition, the "Asian" and the Nearctic lineages reported previously were identified although the latter group was not supported. However, relationships among the main Microtus branches were not resolved, suggesting a rapid and potentially simultaneous radiation of a widespread ancestor early in the history of the genus. This and subsequent radiations discernible in the cytochrome b phylogeny, show the considerable potential of Microtus for analysis of historical and ecological determinants of speciation in small mammals. It is evident that speciation is an ongoing process in the genus and that the molecular data provides a vital insight into current species limits as well as cladogenic events of the past.     

Evolutionary history of tree squirrels (Rodentia,Sciurini) based on multilocus phylogeny reconstruction

Pe?nerová, P. & Martínková, N. (2012). Evolutionary history of tree squirrels (Rodentia, Sciurini) based on multilocus phylogeny reconstruction. —Zoologica Scripta, 41, 211–219. Tree squirrels of the tribe Sciurini represent a group with unresolved phylogenetic relationships in gene trees. We used partial sequences of mitochondrial genes for 12S rRNA, 16S rRNA, cytochrome b and d‐loop, and nuclear irbp, c‐myc exon 2 and 3 and rag1 genes to reconstruct phylogenetic relationships within the tribe, maximizing the number of analysed species. Bayesian inference analysis of the concatenated sequences revealed common trends that were similar to those retrieved with supertree reconstruction. We confirmed congruence between phylogeny and zoogeography. The first group that diverged from a common ancestor was genus Tamiasciurus, followed by Palaearctic Sciurus and Indomalayan Rheithrosciurus macrotis. Nearctic and Neotropical Sciurus species formed a monophyletic group that included Microsciurus and Syntheosciurus. Neotropical Sciurini were monophyletic with a putative exception of Syntheosciurus brochus that was included in a polychotomy with Nearctic Sciurus in supertree analyses. Our data indicate that Sciurini tree squirrels originated in the northern hemisphere and ancestors of contemporary taxa attained their current distribution through overland colonization from the nearest continent rather than through trans‐Pacific dispersal.     

Global diversity of non-biting midges (Chironomidae; Insecta-Diptera) in freshwater

Chironomidae are common inhabitants of most aquatic habitats, and often dominate aquatic insect communities in both abundance and species richness. Species occur in all continents, including Antarctica, and most major oceanic islands that have been investigated. The family is divided into 11 subfamilies and 22 nominal tribes. Although individual species occur in a wide range of habitats from terrestrial to fully aquatic, a total of 339 genera and 4,147 species are unambiguously aquatic in their immature stages. Greatest species and generic richnesses occur in the Palaearctic Region and Nearctic Region, respectively, but this pattern may largely reflect historical patterns of past taxonomic research efforts. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Pugs of the tribe Eupitheciini (Lepidoptera,Geometridae): the history of study and distribution over biogeographic regions

The World fauna of the tribe Eupitheciini is the most species-rich in the family Geometridae. This tribe includes about 1900 species (almost 3000 species-group names) from 47 genera; about one third of the genera (15) are monotypic. The generic diversity of Eupitheciini is the highest in the Australian (38 genera, 11 of them endemic) and Oriental regions (32 genera, 4 endemic) and the lowest in the Neotropical Region (possibly one genus only). The faunas of different biogeographic regions can be arranged in following order by their species richness: the Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic Regions (166 species). Eupithecia is the most species-rich genus in the family Geometridae and the entire order Lepidoptera, and one of the largest genera in the whole World fauna of insects. The greatest number of species of this genus is recorded in the Palaearctic Region (466 species), where Eupithecia accounts for about 95% of the tribe Eupitheciini. The mainland of the Oriental Region (especially the Himalayas) is also very species-rich; however the proportion of the Eupithecia representatives decreases towards Malaysia, Sundaland, and the Australian Region (about 2% of the tribe). The Eupitheciini faunas have the greatest similarity at the generic level between the Oriental and Australian Regions (the Jaccard and Sørensen coefficient values being 0.62 and 0.77, respectively). The Palaearctic fauna is more similar to the Afrotropical and Oriental faunas at the genus-group level. On the whole, the fauna of the Nearctic Region is similar to that the West Palaearctic, with the exception of the fact that representatives of the genera Gymnoscelis and Chloroclystis are absent in North America, although two endemic genera Nasusina and Prorella are present. At the genus-group level, the Nearctic fauna of Eupitheciini is more similar to the Neotropical (the Jaccard and Sørensen coefficients 0.20 and 0.33, respectively) than to the Palaearctic fauna (0.17 and 0.29). The number of synonymies is very high in the tribe Eupitheciini because of the homogeneity of this group, whose species are difficult to identify without the use of elaborate anatomical techniques. Modern revisions, catalogues, surveys, and atlases on Eupitheciini are absent for many countries and large geographic regions. Revisions of pugs of the tribe Eupitheciini for some biogeographic regions are extremely difficult because of fragmentation of entomological collections including the type specimens of many species-group taxa. A large fraction of synonyms is characteristic of parts of the World with the best known faunas: Europe (64% of synonyms) and North America (39%). On the contrary, the lowest levels of synonymy are typical of the less known faunas of the regions situated at the equatorial latitudes, namely the Neotropical (9%) and Afrotropical (8%) ones.     

Morfologia y taxonomia de los Mesocyclops (Crustacea: Copepoda: Cyclopoida) de Mexico

A fundamental review of all species and subspecies of Mesocyclopsencountered in the territory of Mexico, a biogeographical transitionzone where the Neotropical Region bounds the Nearctic Regionin the south. The book begins with a sketch of the history of     

Molecular Systematics of Bats of the Genus Myotis (Vespertilionidae) Suggests Deterministic Ecomorphological Convergences

Based on extensive phenetic analyses, bats of the genus Myotis have been classically subdivided into four major subgenera each of which comprise many species with similar morphological and ecological adaptations. Each subgenus thus corresponds to a distinct “ecomorph” encompassing bat species exploiting their environment in a similar fashion. As three of these subgenera are cosmopolitan, regional species assemblages of Myotis usually include sympatric representatives of each ecomorph. If species within these ecomorphs are monophyletic, such assemblages would suggest extensive secondary dispersal across geographic areas. Conversely, these ecomorphological adaptations may have evolved independently through deterministic processes, such as adaptive radiation. In this case, phylogenetic reconstructions are not expected to sort species of the same ecomorph into monophyletic clades. To test these predictions, we reconstructed the phylogenetic history of 13 American, 11 Palaearctic, and 6 other Myotis species, using sequence data obtained from nearly 2 kb of mitochondrial genes (cytochrome b and nd1). Separate or combined analyses of these sequences clearly demonstrate the existence of several pairs of morphologically very similar species (i.e., sibling species) which are phylogenetically not closely related. None of the three tested subgenera constitute monophyletic units. For instance, Nearctic and Neotropical species currently classified into the three subgenera were clustered in a single, well-supported monophyletic clade. These species thus evolved independently of their ecological equivalents from the Palaearctic region. Independent adaptive radiations among species of the genus Myotis therefore produced strikingly similar evolutionary solutions in different parts of the world. Furthermore, all phylogenetic reconstructions based on mtDNA strongly supported the existence of an unsuspected monophyletic clade which included all assayed New World species plus M. brandtii (from the Palaearctic Region). This “American” clade thus radiated into a morphologically diverse species assemblage which evolved after the first Myotis species colonized the Americas. Molecular reconstructions support paleontological evidence that species of the genus Myotis had a burst of diversification during the late Miocene–early Pliocene epoch.     

A fossil nymphomyiid (Diptera) from the Baltic and Bitterfeld amber

Nymphomyia succina sp.n., the first fossil nymphomyiid fly (Diptera) from the Eocene Baltic and Bitterfeld amber is described and figured, based on four males. A probably conspecific female from the Baltic amber is also described and figured. With the discovery of this species, a gap in the distribution pattern of extant Nymphomyiidae is closed. It is probable that nymphomyiids colonized the eastern Nearctic Region via the Thule landbridge approximately 25–30 million years before present.     

Phylogeny and historical biogeography of the cynipoid wasp family Ibaliidae (Hymenoptera)

The Ibaliidae are a small family of cynipoid wasps, the members of which parasitize woodboring siricid larvae in hardwoods and conifers. The 19 currently recognized extant species occur mainly in the Northern Hemisphere. No fossils are known despite the presumed old age of the family. We present a cladistic analysis of ibaliid relationships at the species-level, mainly based on external skeletal characters of adults. The results indicate that the three genera ( Eileenella , Heteribalia , Ibalia ) and two subgenera of Ibalia ( Ibalia s. str. and Tremibalia ) recognized in the current classification are monophyletic. Three different categories of characters were compared for their phylogenetic usefulness. Homoplasy was found to be lowest for main structures, higher for sculptural characters, and still higher for colour differences. The historical biogeography of the family was reconstructed using dispersal–vicariance analysis in combination with palaeogeographical data. The results suggest that the family primarily diversified within the eastern Palaearctic–northern Oriental region. The nominate subgenus of Ibalia dispersed early to the western Nearctic, where it radiated; two species later spread throughout the Holarctic. The other subgenus of Ibalia shows an early eastern Palaearctic–eastern Nearctic disjunction which presumably dates back to the Eocene–Oligocene transition.     

Taxonomy and preliminary phylogeny of the parasitic genus Apocephalus, subgenus Mesophora (Diptera: Phoridae)

Abstract. The species of Apocephalus , subgenus Mesophora , are revised and twenty-eight species are recognized, including the following twenty-two new to science: from the Nearctic Region A. brunnipes, A. gemursus, A. pristinus, A. setialvus and A. unitarsus , and from the Neotropical Region A. absentis, A. adustus, A. anfractus, A. angustistylus, A. bisetus, A. brevicercus, A. curtus, A. gracilis, A. hansoni, A. leptotarsus, A. longistylus, A. micrepelis, A. moraviensis, A. prolatus, A. trisetus, A. tritarsus and A. truncaticercus. Additionally, three species known only from female specimens are described but not formally named. A lectotype is designated for A. mortifer Borgmeier, and immature stages of A. borealis, A. antennatus and A. mortifer are described. Unlike larvae of other Apocephalus species, all of which are parasites of ants (Hymenoptera: Formicidae), those of Mesophora species are parasites of various other hosts.