Phylogeny of Agriotypinae (Hymenoptera: Ichneumonidae), with comments on the subfamily relationships of the basal Ichneumonidae

Abstract. Agriotypinae is a small clade of aquatic parasitoid wasps with uncertain affinities. Its phylogeny was examined in a cladistic analysis of larval and adult morphological characters. The analysis found that Agriotypinae belongs within Ichneumonidae as sister group of Labeninae and related to Xoridinae and Cryptinae. All sixteen species in the subfamily should be assigned to Agriotypus Curtis. Two species groups were found, the Palaearctic armatus species group and the Oriental himalensis species group. Agriotypus chaoi sp.n. and Agriotypus masneri sp.n. are described from Vietnam and Atopotypus Chao is synonymized with Agriotypus forming Agriotypus succinctus (Chao) comb.n. A key and revision of the world species is provided, and the evolution of aquatic behaviour in Ichneumonidae is discussed.     

Systematics and evolution of the genus Paraceroglyphus and related taxa (Acari: Acaridae) associated with fleas (Insecta: Siphonaptera)

Collections of fleas from terrestrial Sciuridae from New Mexico and Montana yielded 2 species of acarid mites: Acarus monopsyllus from Ceratophyllus ciliatus and Paraceroglyphus cynomydis n. sp. from 4 species of Oropsylla. The genera Acarus, Paraceroglyphus, and Trichopsyllopus form a clade distinct from other genera of flea-associated mites, with Paraceroglyphus the sister group of the other 2 genera. Paraceroglyphus cynomydis is the sister group of a clade comprising P. xenopsylla and possibly P. californicus, with P. meles as the nearest outgroup.     

A new species of Oryzomys (Rodentia, Sigmondontinae) from southeast Brazil, with comments on the classification of the O. capito species group

A new species of Oryzomys is described for the coastal Atlantic region of Brazil. Oryzomys seuanezi sp. nov. was collected in eight localities throughout the Atlantic Forest from the south of the State of Bahia to the State of the Rio de Janeiro, mostly in lowlands. Belonging to the O. capito species group, O. seuanezi differs from the other species by its karyotype equal 2n = 48, FN = 60. A morphometric analysis of populations of some species of the group indicates a split between the samples of the Atlantic Forest and those from O. capito from Cerrados and Amazon in relation to size, the former being larger. The population from Amazonia and from the Cerrados belongs to the same species, O. goeldii being a synonym of O. capito. O. seuanezi sp. nov. is the sister taxon of O. oniscus, forming an Atlantic Forest clade. The close relationship of this group and some Peruvian populations of O. capito, probably a new species, is postulated.     

Phylogeny of the Augochlora clade with the description of four new species (Hymenoptera,Apoidea)

The Augochlora clade includes four genera: Augochlora Smith, Augochlorella Sandhouse, Ceratalictus Moure, and Pereirapis Moure. This is one of the richest and most widespread groups of Augochlorini bees. There are about 150 species, which occur from Argentina to Canada. The species of Augochlora clade are considered solitary to facultatively social, except Ceratalictus for which nothing is known. Wood nesting behavior arose once in the clade, in Augochlora sensu strictu. The objective of this study is to describe four new species and to present a revised phylogenetic analysis of the Augochlora clade for the placement of these species. The morphological matrix comprised 77 characters and 42 terminals, and resulted in two most parsimonious trees. The monophyly of the Augochlora clade is corroborated. Ceratalictus and Pereirapis are considered as sister groups and Ceratalictus inflexus sp. nov. came as sister to other species of Ceratalictus. Augochlora and Augochlorella are monophyletic and sister groups. Both extant subgenera of Augochlora were corroborated as monophyletic. Augochlorella comis is considered as sister group to the rest of Augochlorella species. All Augochlorella new species described belong to the Augochlorella ephyra group. Augochlorella kelliae sp. nov. is phylogenetically related to Augochlorella una. Augochlorella procliva sp. nov. and Augochlorella mavricera sp. nov. constitute a clade with Augochlorella acarinata. Including the new species, Augochlorella has 19 species and Ceratalictus 11 species. A revised key for species of Augochlorella and Ceratalictus is also presented in the Supplementary Information.     

Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF-1alpha genes

A phylogeny of the Neotropical members of the Tribe Troidini (Lepidoptera: Papilionidae) was obtained with sequences of three protein-coding genes: two mitochondrial (COI and COII), and one nuclear (EF-1alpha). Parsimony and Bayesian analyses of 33 taxa resulted in very similar trees regardless of method used with the 27 troidines always forming a monophyletic clade. Within Troidini, the genus Battus is sister group to the remaining troidines, followed by a clade formed by the Paleotropical taxa (here represented by three exemplars). The genus Euryades is the next branch, and sister group of Parides. The genus Parides is monophyletic, and is divided into four main groups by Maximum Parsimony analysis, with the most basal group composed of tailed species restricted to SE Brazil. Character optimization of ecological and morphological traits over the phylogeny proposed for troidines indicated that the use of several species of Aristolochia is ancestral over the use of few or a single host-plant. For the other three characters, the ancestral states were the absence of long tails, forest as the primary habitat and oviposition solitary or in loose group of several eggs.     

Acheilognathus changtingensis sp. nov., a new species of the cyprinid genus Acheilognathus (Teleostei: Cyprinidae) from Southeastern China based on morphological and molecular evidence

A new species of the bitterling genus Acheilognathus, Acheilognathus changtingensis sp. nov., was recently discovered from Changting County in Hanjiang River, Fujian Province, Southeastern China. It can be diagnosed by the following combination of characters: dorsal fin with three simple and 15 (occasionally 14) branched fin rays, anal fin with three simple and 12 (occasionally 11) branched fin rays; dorsal fin pale and anal fin slightly pale; white spots on anal-fin rays forming a transverse band, and anal fin margined with white band in males. Its unique characters are the many dispersed black spots on the dorsal, anal, pelvic, caudal fins, and on the head. Using the mitochondrial cytochrome Jb gene as a molecular marker, we reconstructed phylogenetic trees of A. changtingensis sp. nov. and other species in Acheilognathus to confirm its taxonomic status and study its speciation. Analyses of both morphological and molecular data consistently indicated the taxonomic status of the present new species. The results also show that A. changtingensis sp. nov. and Acheilognathus macropterus are sister species that diverged about 14.50 MYA by geographical isolation.     

Cretaceous and Eocene fossils of the rare extant genus Synneuron Lundstrom (Diptera: Canthyloscledidae): evidence of a true Pangean clade

The first two fossil species of the canthyloscelid genus Synneuron are described based on compression wings. Synneuron eomontana sp. nov. is described from the Middle Eocene Coal Creek Member of the Kishenehn Formation, in the USA, and Synneuron jelli sp. nov. is described from the Lower Cretaceous Koonwarra Fossil Bed of the Korumburra Group, in Australia. The wings are illustrated and compared to the extant species of the genus, to species of the three other recent genera of Canthyloscelidae and to an anisopodid. A phylogenetic analysis of the relationships between the species of Synneuron was performed. The Eocene fossil S. eomontana appears as sister of the pair of recent Holarctic species of the genus, while the Australian Cretaceous species S. jelli is sister of the clade with the species of Synneuron of the northern hemisphere. The sister group of Synneuron is the canthyloscelid clade (Hyperoscelis + Canthyloscelis), for which a middle Jurassic fossil is known. At the early Cretaceous, Gondwana was already separated from Laurasia and the disjunction between the species of Synneuron in Australia and the northern hemisphere clade of the genus suggest a true pangeic origin for the genus. The biology of the canthyloscelid larvae is shaped by its trophic specialization—xylosaprophagous. This suggests that the transition from the Pangean Jurassic gymnosperm-dominated forests to the late Cretaceous angiosperm-dominated forests may be related to the low recent diversity of Synneuron or of the canthyloscelids in the world—and maybe to the extinction of the genus in the southern hemisphere. This major turnover of the vegetation type along the Cretaceous may be also somehow related to the complete extinction of other groups of flies strictly associated with gymnosperms, as may be the case of the lower brachyceran family Zhangsolvidae. This speculation needs additional corroboration from other groups, that will become available with the combination of systematics, paleontology and biogeographical information of different early Cretaceous clades.     

A cladistic study of Arillastrum, Angophora and Eucalyptus (Myrtaceae)

A cladistic study of Anllastrum, Angophora and Eucalyptus (Myrtaceae). Transformed cladistic; character compatibility; branch and bound, and Farris-Wagner methods gave similar solutions in a cladistic study of Arillastrum, Angophora and Eucalyptus. These analyses, based on morphological characters, indicate that Eucalyptus is a monophyletic group and that its sister taxon is Angophora.
Within Eucalyptus , subgenera Blakella and Corymbia are sister taxa to all other groups; subgenera Monocalyptus, Idiogenes and Gaubaea form a monophyletic group with subgenus Monocalyptus sister to subgenera Idiogenes and Gaubaea ; subgenera Symphyomyrtus and Telocalyptus together also form a monophyletic group and, with Eucalyptus similis (subgenus Eudesmia group 4), are sister to the Monocalyptus group. Eucalyptus subgenus Telocalyptus (4 species), Eucalyptus subgenus Idiogenes (1 species) and Eucalyptus subgenus Gaubaea (2 species) should not be recognized as subgenera and some individual species need further examination. Eucalyptus subgenus Eudesmia is a paraphyletic group.
Some characters are identified as parallelisms, e.g. axillary inflorescences, sepaline operculum, bristle glands, and clustered anthers. A more congruent interpretation of the single operculum of Eucalyptus subgenus Monocalyptus as at least partly petaline rather than solely sepaline in origin is suggested.
The area relationships for the taxa are concordant with those derived from geological and climatological information. New Caledonia is sister area to Australia, and within Australia southwestern Australia is sister area to south-eastern and north-eastern Australia.     

†Cormopsocidae: A new family of the suborder Trogiomorpha (Insecta: Psocodea) from Burmese amber

Cormopsocidae n. fam . of the psocid suborder Trogiomorpha was proposed for a fossil species from mid‐Cretaceous Burmese amber, Cormopsocus groehni n. gen . & n. sp . This family was estimated to be the sister group of all other trogiomorphan taxa, but the possibility of much deeper divergence (i.e. placement as a sister group of all Psocodea) could not be excluded. Cormopsocus groehni retains many plesiomorphic features, which will contribute greatly to elucidating the ancestral state of Psocodea.     

A new subfamily of Trichomycteridae (Teleostei, Siluriformes), lower loricarioid relationships and a discussion on the impact of additional taxa for phylogenetic analysis

The Copionodontinae is described as a new subfamily of the neotropical catfish family Trichomycteridae. It comprises two new genera and three new species from north-eastern Brazil: Copionodon gen. nov. (including C. pecten sp. nov. and C. orthiocarinatus sp. nov. ) and Glaphyropoma gen nov. (including G. rodriguesi sp. nov. ) The Copionodontinae can be diagnosed externally by the anterior position of the dorsal fin, the presence of a well-developed adipose fin, and the strongly spatulate shape of the jaw teeth. The subfamily is hypothesized as monophyletic on the basis of several synapomorphies in internal and external anatomy. Copionodontines have the plesiomorphic condition of several characters, relative to all other trichomycterids, including the presence of ductus pneumaticus; the possession of separate pterosphenoids, sphenotics and prootics; the presence of the intercalarium; the complete infraorbital latero-sensory canal; the presence of the interhyal; and the wide lateral opening of the swimbladder capsule. Elsewhere within trichomycterids, these primitive traits are found only in Trichogenes. These and other characters support the hypothesis that copionodontines are the plesiomorphic sister group of all other trichomycterids, and that Trichogenes is their next successive sister group. Contrary to the currently accepted hypothesis, the monotypic Nematogenyidae is proposed as the sister group of the Trichomycteridae, and not of all remaining loricarioids. This change is to a major extent induced by the inclusion of copionodontines and Trichogenes in the analysis of lower loricarioid relationships. The present case is given as an example of the impact that undiscovered taxa, fossil or Recent, may have upon hypotheses of phylogenetic relationships.     

分子鉴定背瘤丽蚌中弯弓蚌螨的隐藏种

Unionicola sp.与弯弓蚌螨U.arcuata在形态结构上非常相似,为了鉴别弯弓蚌螨U.arcuata 和Unionicola sp.,本研究扩增出Unionicola sp.12S rRNA的部分基因片段,与基因数据库中5种寄生蚌螨序列进行比较分析.结果显示,Unionicola sp.与弯弓蚌螨U.arcuata遗传差异为0.145,而分子系统树表明,Unionicola sp.与弯弓蚌螨U.arcuata聚为一支,具较近的亲缘关系.我们推断Unionicola sp.是弯弓蚌螨U.arcuata的1个隐藏种.这可能是由于不同寄主蚌造成了宿主隔离,从而引起了蚌螨遗传基因的分化.     

Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF-1α genes

A phylogeny of the Neotropical members of the Tribe Troidini (Lepidoptera: Papilionidae) was obtained with sequences of three protein-coding genes: two mitochondrial (COI and COII), and one nuclear (EF-1α). Parsimony and Bayesian analyses of 33 taxa resulted in very similar trees regardless of method used with the 27 troidines always forming a monophyletic clade. Within Troidini, the genus Battus is sister group to the remaining troidines, followed by a clade formed by the Paleotropical taxa (here represented by three exemplars). The genus Euryades is the next branch, and sister group of Parides. The genus Parides is monophyletic, and is divided into four main groups by Maximum Parsimony analysis, with the most basal group composed of tailed species restricted to SE Brazil. Character optimization of ecological and morphological traits over the phylogeny proposed for troidines indicated that the use of several species of Aristolochia is ancestral over the use of few or a single host-plant. For the other three characters, the ancestral states were the absence of long tails, forest as the primary habitat and oviposition solitary or in loose group of several eggs.     

Relationships of the Macaronesian and Mediterranean floras: molecular evidence for multiple colonizations into Macaronesia and back-colonization of the continent in Convolvulus (Convolvulaceae)

A molecular phylogenetic analysis of the Macaronesian endemic species of Convolvulus was undertaken using data from the nuclear ribosomal internal transcribed spacer (ITS) regions. The results of the analysis support two introductions into Macaronesia from distantly related clades within Convolvulus and a subsequent back-colonization to the continent from within one of the clades. Hypothesized relationships between Macaronesian species and New World taxa and between the Canarian endemic C. caput-medusae and the Moroccan C. trabutianus are refuted. Both Macaronesian clades are shown to have Mediterranean sister groups although one is predominantly western Mediterranean and the other predominantly eastern Mediterranean in distribution. The patterns of colonization into Macaronesia demonstrated by Convolvulus and also by other multiple colonizing genera conform to either a pattern of phylogenetic distinctiveness or a checkerboard distribution of island lineages. Both are consistent with the hypothesis that niche preemption is responsible for the limited number of colonizations into the region. A review of sister group relationships demonstrates that, in common with Convolvulus, most Macaronesian groups have sister groups distributed in the near-continent (i.e., western Mediterranean). Disjunct sister group relationships (including Eastern Mediterranean disjunctions) occur in only 18% of groups.     

Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants

Abstract Colocasiomyia, a moderate‐sized genus in the subfamily Drosophilinae, comprises seventy (twenty‐six described and forty‐four undescribed) species. Several Colocasiomyia species have evolved intimate mutualisms with specific host plants, especially of the family Araceae: the flies depend throughout the entire life cycle, oviposition, larval growth, pupation, and adult feeding and mating, on inflorescences of their host plants, and in turn act as species‐specific pollinators for their host plants. To understand the evolution of this mutualism between Colocasiomyia flies and their host plants, the phylogenetic relationships of this genus and some possibly related taxa are inferred from a cladistic analysis based on sixty‐two characters of adult morphology. We conclude that Colocasiomyia is polyphyletic, with the C. arenga species group clearly separate. Colocasiomyia without the arenga group (Colocasiomyia proper) is sister to all other studied drosophilines, whereas the arenga group is relatively derived within the Drosophilinae. Within Colocasiomyia proper, four clades are recognized, three of which correspond to previously proposed species groups: the cristata, toshiokai and baechlii groups. The other clade, C. sp.1 aff. nepalensis+C. sp.2 aff. nepalensis, is defined as a new species group. Relationships amongst the four clades and three independent species (C. micheliae, C. gigantea and C. sp.K1) remain almost unresolved, except for a sister group relationship between the toshiokai and baechlii groups. The classification of species groups in Colocasiomyia is revised by erecting two new species groups (crassipes and zeylanica groups) in addition to the three known (baechlii, cristata and toshiokai) groups. Revision of the arenga group, which should be removed from Colocasiomyia, is left for future studies. The evolution of host plant selection in Colocasiomyia is discussed by mapping host plant taxa (families, subfamilies and tribes) on the phylogenetic tree deduced from the cladistic analysis. Cohabitation in the same host inflorescence by a pair of species with microallopatric niche separation on the spadix is hypothesized to have evolved independently at least more than twice in Colocasiomyia.     

Are the Platyhelminthes a monophyletic primitive group? An assessment using 18S rDNA sequences

In most zoological textbooks, Platyhelminthes are depicted as an early- emerging clade forming the likely sister group of all the other Bilateria. Other phylogenetic proposals see them either as the sister group of most of the Protostomia or as a group derived from protostome coelomate ancestors by progenesis. The main difficulty in their correct phylogenetic placing is the lack of convincing synapomorphies for all Platyhelminthes, which may indicate that they are polyphyletic. Moreover, their internal phylogenetic relationships are still uncertain. To test these hypotheses, new complete 18S rDNA sequences from 13 species of "Turbellaria" have been obtained and compared to published sequences of 2 other "Turbellaria," 3 species of parasitic Platyhelminthes, and several diploblastic and deuterostome and protostome triploblastics. Maximum-parsimony, maximum-likelihood, and neighbor-joining methods were used to infer their phylogeny. The results show the order Catenulida to form an independent early- branching clade and emerge as a potential sister group of the rest of the Bilateria, while the rest of Platyhelminthes (Rhabditophora), which includes the parasites, form a clear monophyletic group closely related to the protostomes. The order Acoela, morphologically considered as candidates to be ancestral, are shown to be fast-clock organisms for the 18S rDNA gene. Hence, long-branching of acoels and insufficient sampling of catenulids and acoels leave their position still unresolved and call for further studies. Within the Rhabditophora, our analyses suggest (1) a close relationship between orders Macrostomida and Polycladida, forming a clear sister group to the rest of orders; (2) that parasitic platyhelminthes appeared early in the evolution of the group and form a sister group to a still-unresolved clade made by Nemertodermatida, Lecithoepitheliata, Prolecithophora, Proseriata, Tricladida, and Rhabdocoela; and (3) that Seriata is paraphyletic.      

PHYLOGENY OF THE GENUS ROSTANGA (NUDIBRANCHIA), WITH DESCRIPTIONS OF THREE NEW SPECIES FROM SOUTH AFRICA

Rostanga elandsia sp. nov., Rostanga aureamala sp. nov. andRostanga phepha sp. nov. are characterized by having the radulawith slender innermost lateral teeth, which lack denticles onthe inner side of the cusp and have a single denticle on theouter side. The outermost lateral teeth of these three speciesare elongate, but shorter than in other species of the genus.In addition, R. aureamala is the only species of the genus withrachidian teeth and R. phepha is unique within the genus Rostanga byvirtue of its white coloration with dark spots. A phylogenetic analysis shows that the three new species fromSouth Africa and Rostanga setidens (Odhner, 1939) are the sistergroup of the rest of the genus. The species from Japan and MarshallIslands (North Pacific Ocean) are basal in the sister cladecontaining the other species of Rostanga Bergh, 1879. The tropicalIndo-Pacific species of Rostanga are not monophyletic. The Atlanticand Eastern Pacific species form a monophyletic, derived clade,being the sister group of Rostanga australis Rudman & Avern,1989, which has a narrow range restricted to south eastern Australia.The widespread Indo-Pacific species Rostanga bifurcata Rudman& Avern, 1989, is the sister group of Rostanga dentacus Rudman& Avern, 1989, also widesprad in the tropical western Pacific. This phylogeny suggest s a viariant origin of the Sourth African, Atlantic-EasternPacific, and probably North Pacific species, whereas in thetropical Indo-Pacific most sister speceis are sympatric. (Received 16 May 1999; accepted 31 July 2000)     

The Nemertodermatida are basal bilaterians and not members of the Platyhelminthes

Recent hypotheses on metazoan phylogeny have recognized three main clades of bilaterian animals: Deuterostomia, Ecdysozoa and Lophotrochozoa. The acoelomate and 'pseudocoelomate' metazoans, including the Platyhelminthes, long considered basal bilaterians, have been referred to positions within these clades by many authors. However, a recent study based on ribosomal DNA placed the flatworm group Acoela as the sister group of all other extant bilaterian lineages. Unexpectedly, the nemertodermatid flatworms, usually considered the sister group of the Acoela together forming the Acoelomorpha, were grouped separately from the Acoela with the rest of the Platyhelminthes (the Rhabditophora) within the Lophotrochozoa. To re-evaluate and clarify the phylogenetic position of the Nemertodermatida, new sequence data from 18S ribosomal DNA and mitochondrial genes of nemertodermatid and other bilaterian species were analysed with parsimony and maximum likelihood methods. The analyses strongly support a basal position within the Bilateria for the Nemertodermatida as a sister group to all other bilaterian taxa except the Acoela. Despite the basal position of both Nemertodermatida and Acoela, the clade Acoelomorpha was not retrieved. These results imply that the last common ancestor of bilaterian metazoans was a small, benthic, direct developer without segments, coelomic cavities, nephrida or a true brain. The name Nephrozoa is proposed for the ancestor of all bilaterians excluding the Nemertodermatida and the Acoela, and its descendants.     

Affinities between Asian non-human Schistosoma species,the S. indicum group,and the African human schistosomes

Schistosoma species have traditionally been arranged in groups based on egg morphology, geographical origins, and the genus or family of snail intermediate host. One of these groups is the 'S. indicum group' comprising species from Asia that use pulmonate snails as intermediate hosts. DNA sequences were obtained from the four members of this group (S. indicum, S. spindale, S. nasale and S. incognitum) to provide information concerning their phylogenetic relationships with other Asian and African species and species groups. The sequences came from the second internal transcribed spacer (ITS2) of the ribosomal gene repeat, part of the 28S ribosomal RNA gene (28S), and part of the mitochondrial cytochrome c oxidase subunit 1 (CO1) gene. Tree analyses using both distance and parsimony methods showed the S. indicum group not to be monophyletic. Schistosoma indicum, S. spindale and S. nasale were clustered among African schistosomes, while S. incognitum was placed as sister to the African species (using ITS2 and 28S nucleotide sequences and CO1 amino acid sequences), or as sister to all other species of Schistosoma (CO1 nucleotide sequences). Based on the present molecular data, a scenario for the evolution of the S. indicum group is discussed.     

Tetrablemmidae (Arachnida, Araneae), a spider family newly recorded from China

The family Tetrablemmidae is reported for the first time from China, with five new species and one new genus. Lehtinenia gen. n., which is erected to accommodate Lehtinenia bicornis sp. n., is characterized by the modified embolus, special modifications on chelicerae, and a Tetrablemma-type vulva. The other four new species are: Ablemma prominens sp. n., Brignoliella caligiformis sp. n., Brignoliella maoganensis sp. n., and Tetrablemma brevidens sp. n., all collected from caves. A phylogenetic analysis of the subfamily Tetrablemminae based on 41 morphological characters shows that the tribe Brignoliellini is the most basal group in the subfamily, rather than the sister group to the tribe Fallablemmini. Lehtinenia gen. n. and the genera Ablemma, Sulaimania, and Maijana together form a monophyletic group.