Intraerythrocytic development of species of Hepatozoon infecting ranid frogs: evidence for convergence of life cycle characteristics among apicomplexans

Intraerythrocytic development of the adeleorin apicomplexans Hepatozoon clamatae and Hepatozoon catesbianae were investigated in the bullfrog, Rana catesbeiana, the green frog, Rana clamitans melanota, and the Northern leopard frog, Rana pipiens. Merozoites emerging from hepatic meronts penetrated erythrocytes and underwent 1-3 rounds of binary fission to produce 2-8 merozoites. Following their release from infected erythrocytes, individual merozoites entered new cells and transformed into gamonts. Although this is the first report of intraerythrocytic development for a fully described species of Hepatozoon, a phylogenetic reanalysis of 11 species of Hepatozoon, 6 species representative of the 5 other hemogregarine taxa, 2 species of dactylosomatids, and 2 species of piroplasms, indicates that asexual reproduction of parasites within blood cells of vertebrates has arisen at least 3 times in the apicomplexan lineage that includes adeleorins and piroplasms. This method of asexual development, which is also observed in species of hemospororin genera such as Plasmodium, is discussed in the context of the evolution of apicomplexan life cycles. In addition to supporting the paraphyly of the genus Hepatozoon determined in an earlier study, this phylogenetic analysis featured a monophyletic group, consisting of the sister taxa Hemolivia and Karyolysus, that was the sister group to a clade consisting of the more derived hemogregarines, the dactylosomatids, and the piroplasms.     

Phylogeny and classification of Marantaceae

Relationships of Marantaceae were estimated from nucleotide sequence variation in the rps16 intron (plastid DNA) and from morphological characters. Fifty-nine species (21 genera) formed the ingroup, and 12 species (12 genera) of other Zingiberales formed the outgroup. There is no support for the traditional subdivision of Marantaceae into a triovulate and a uniovulate tribe or the informal groups previously proposed. The so-called Donax group forms a paraphyletic grade that is basal within Marantaceae. Thalia appears as the distal branch of this grade, but its position is not supported in jackknife analysis. The so-called Calathea group is monophyletic in all shortest trees but not supported with greater than 50% jackknife. The genus Calathea appears to be paraphyletic. The Maranta and Phrynium groups are clearly polyphyletic. Maranta, Koernickanthe , and genera of the Mymsma group, all neotropical, form a strongly supported monophyletic group. The sister of this group is the palaeotropical genus Halopegia. Koernickanthe is nested within Maranta , as this genus is traditionally circumscribed. The African genera Ataenidia and Marantochloa form a strongly supported clade in which Ataenidia is the sister group to Marantochloa . Based on phylogeny it is concluded that Africa, in spite of being much poorer in species, is the most likely ancestral area of Marantaceae     

A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations

Siphonaptera (fleas) is a highly specialized order of holometabolous insects comprising ～2500 species placed in 16 families. Despite a long history of extensive work on flea classification and biology, phylogenetic relationships among fleas are virtually unknown. We present the first formal analysis of flea relationships based on a molecular matrix of four loci (18S ribosomal DNA, 28S ribosomal DNA, Cytochrome Oxidase II, and Elongation Factor 1‐alpha) for 128 flea taxa from around the world representing 16 families, 25 subfamilies, 26 tribes, and 83 flea genera with eight outgroups. Trees were reconstructed using direct optimization and maximum likelihood techniques. Our analysis supports Tungidae as the most basal flea lineage, sister group to the remainder of the extant fleas. Pygiopsyllomorpha is monophyletic, as are the constituent families Lycopsyllidae, Pygiopsyllidae, and Stivaliidae, with a sister group relationship between the latter two families. Macropsyllidae is resolved as sister group to Coptopsyllidae with moderate nodal support. Stephanociricidae is monophyletic, as are the two constituent subfamilies Stephanocircinae and Craneopsyllinae. Vermipsyllidae is placed as sister group to Jordanopsylla. Rhopalopsyllidae is monophyletic as are the two constituent subfamilies Rhopalopsyllinae and Parapsyllinae. Hystrichopsyllidae is paraphyletic with Hystrichopsyllini placed as sister to some species of Anomiopsyllini and Ctenopariini placed as sister to Carterettini. Ctenophthalmidae is grossly paraphyletic with the family broken into seven lineages dispersed on the tree. Most notably, Anomiopsyllini is paraphyletic. Pulicidae and Chimaeropsyllidae are both monophyletic and these families are sister groups. Ceratophyllomorpha is monophyletic and includes Ischnopsyllidae, Ceratophyllidae, and Leptopsyllidae. Leptopsyllidae is paraphyletic as are its constituent subfamilies Amphipsyllinae and Leptopsyllinae and the tribes Amphipsyllini and Leptopsyllini. Ischnopsyllidae is monophyletic. Ceratophyllidae is monophyletic, with a monophyletic Dactypsyllinae nested within Ceratophyllinae, rendering the latter group paraphyletic. Mapping of general host associations on our topology reveals an early association with mammals with four independent shifts to birds. © The Willi Hennig Society 2008.     

Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei)

Morphological character analyses indicate that Rhyacichthyidae, Odontobutidae, Eleotridae, and Xenisthmidae are the basal families within the perciform suborder Gobioidei. This study uses DNA sequence data to infer the relationships of genera within these families, as well as determine the placement of more derived gobioids (family Gobiidae) and the identity of the outgroup to Gobioidei. Complete sequences of the mitochondrial ND1, ND2, COI, and cyt b genes (4397 base pairs) are analyzed for representatives of 27 gobioid genera and a variety of perciform and scorpaeniform outgroup candidates; the phylogeny is rooted with a beryciform as a distal outgroup. The single most parsimonious tree that results indicates that, of the outgroups sampled, the perciform family Apogonidae is most closely related to Gobioidei. Gobioidei is monophyletic, and Rhyacichthys aspro is the most basal taxon. The remainder of Gobioidei is resolved into clades corresponding to the families Odontobutidae (plus Milyeringa) and Eleotridae+Xenisthmidae+Gobiidae. Within Eleotridae, the subfamily Butinae (minus Milyeringa) is paraphyletic with respect to Gobiidae, and Eleotrinae is paraphyletic with respect to Xenisthmidae. Other than these groupings, the primary disagreement with the current morphology-based classification is that the molecular data indicate that the troglodytic Milyeringa should be placed in Odontobutidae, not Butinae, although support for this placement is weak. The most basal lineage of Gobioidei is known from the freshwaters of the Indo-Pacific, with marine-dwelling lineages arising several times independently in the group. The phylogeny also indicates that different gobioid lineages are distributed in Asia, Africa, Madagascar and the Neotropics. Five sister pairs of basal gobioid species inhabit Atlantic and Pacific drainages of Panama, with widely varying divergences.     

Phylogenetic analysis of the class Sporozoea (phylum Apicomplexa Levine, 1970): evidence for the independent evolution of heteroxenous life cycles

A phylogenetic analysis of representative genera in the class Sporozoea was undertaken using biological and morphological features to infer evolutionary relationships among the widely recognized groups in the class. Gregarines were used as a functional outgroup to the remaining sporozoa (adeleids, eimeriorins, haemosporinids, and piroplasms). The piroplasms were shown to be closely related to the adeleid parasites. Species of Babesia and Theileria were shown not to form a sister group to the haemosporinids as has been frequently suggested. The data indicate that the biologically diverse family Haemogregarinidae should be divided into at least 3 families (Haemogregarinidae Neveu-Lemaire, 1901, containing the genera Haemogregarina and Cyrilia; Karyolysidae Wenyon, 1926, containing the genus Karyolysus; Hepatozoidae Wenyon, 1926, containing the genus Hepatozoon) because the 4 genera currently within the family do not form a monophyletic group. Correlation between parasite phylogeny and taxonomic affinities of their definitive hosts suggests that the definitive hosts of heteroxenous sporozoa are their ancestral hosts. Heteroxenous sporozoan life cycles apparently have evolved independently to adapt to changes in the feeding behaviors of their definitive hosts.     

An ITS phylogeny of Balsamorhiza and Wyethia (Asteraceae: Heliantheae)

The relationships among the species of Balsamorhiza and Wyethia (Asteraceae: Heliantheae) were examined using data from the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA. The ITS sequences were obtained from nine species of Balsamorhiza and 14 species of Wyethia as well as seven outgroup genera. Five of the outgroup genera were members of the subtribe Engelmanniinae of the tribe Heliantheae, the subtribe that includes Balsamorhiza and Wyethia. The resulting trees show that Balsamorhiza and Wyethia together form a monophyletic group. Balsamorhiza alone is monophyletic, but neither of its two sections is monophyletic. Wyethia is paraphyletic. One group of Wyethia species, including all members of sections Alarconia and Wyethia as well as W. bolanderi from section Agnorhiza, is monophyletic and sister to Balsamorhiza. The other species of Wyethia (all placed in section Agnorhiza) are part of a polytomy along with the clade composed of Balsamorhiza plus the rest of Wyethia.     

Phylogenetic relationships among genera of Aphidiinae (Hymenoptera: Braconidae) based on DNA sequence of the mitochondrial 16S rRNA gene

Phylogenetic relationships among forty‐nine taxa representing twenty‐four genera of Aphidiinae (Hymenoptera: Braconidae) were investigated using DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony analysis. Seven species in six other subfamilies of Braconidae were used as outgroup. The results suggested that members of Aphidiinae are monophyletic. The basal lineage of Aphidiinae was Aclitus in weighted and unweighted parsimony analyses and Praini was basal relative to Ephedrini. With the exception of Pauesia and Aphidius, all genera were monophyletic. The results support generic status for Euaphidius, but not for Lysaphidus. Diaeretus leucopterus was internal to a clade composed of three Pauesia species, suggesting that the latter genus may be paraphyletic. A combined analysis that included DNA sequence of 16S rRNA, NADH1 dehydrogenase and 28S rRNA resulted in more robust cladograms with topologies similar to those inferred from the 16S rRNA gene sequence alone. The results are compared to previously proposed phylogenies of Aphidiinae based on morphological and molecular characters.     

Phylogenetic position of the adeleorinid coccidia (Myzozoa, Apicomplexa, Coccidia, Eucoccidiorida, Adeleorina) inferred using 18S rDNA sequences

Investigating the evolutionary relationships of the major groups of Apicomplexa remains an important area of study. Morphological features and host-parasite relationships continue to be important in the systematics of the adeleorinid coccidia (suborder Adeleorina), but the systematics of these parasites have not been well-supported or have been constrained by data that were lacking or difficult to interpret. Previous phylogenetic studies of the Adeleorina have been based on morphological and developmental characters of several well-described species or based on nuclear 18S ribosomal DNA (rDNA) sequences from taxa of limited taxonomic diversity. Twelve new 18S rDNA sequences from adeleorinid coccidia were combined with published sequences to study the molecular phylogeny of taxa within the Adeleorina and to investigate the evolutionary relationships of adeleorinid parasites within the Apicomplexa. Three phylogenetic methods supported strongly that the suborder Adeleorina formed a monophyletic clade within the Apicomplexa. Most widely recognized families within the Adeleorina were hypothesized to be monophyletic in all analyses, although the single Hemolivia species included in the analyses was the sister taxon to a Hepatozoon sp. within a larger clade that contained all other Hepatozoon spp. making the family Hepatozoidae paraphyletic. There was an apparent relationship between the various clades generated by the analyses and the definitive (invertebrate) host parasitized and, to lesser extent, the type of intermediate (vertebrate) host exploited by the adeleorinid parasites. We conclude that additional taxon sampling and use of other genetic markers apart from 18S rDNA will be required to better resolve relationships among these parasites.     

Phylogeny of the Anomala (Crustacea, Decapoda, Reptantia) based on the ossicles of the foregut

We present a cladistic analysis of the Anomala based on 66 ingroup species and 5 outgroup representatives. Based on a comparative analysis of the morphology of the foregut we scored 124 characters related to size, shape, and fusion of foregut ossicles and other foregut structures. Our parsimony analysis resulted in 30 equally parsimonious trees which differ mainly at the lower hierarchical level. Our study reveals two large clades within Anomala. One large clade consists of Galatheoidea and Chirostyloidea. The internal relationships show a monophyletic Porcellanidae nested within a group comprising paraphyletic Galatheidae, and Munididae as well as Munidopsidae. The other large clade contains Aegla as sister group to a monophyletic group consisting of the Hippoidea and a clade formed by Lomis and the Paguroidea. Coenobitidae are nested within paraphyletic Diogenidae and Lithodidae are nested within paraphyletic Paguridae. The results are discussed in the context of other morphological and molecular analyses. Furthermore, some aspects of carcinization are touched upon; in particular, an anomalan stem species with a, at least to some extent, ventrally folded pleon is suggested.     

Polemoniaceae phylogeny and classification: implications of sequence data from the chloroplast gene ndhF

The chloroplast gene ndhF was used to study phylogenetic relationships of the Polemoniaceae at two levels: among members of the Ericales and among genera of the family. Sequence data for interfamilial analyses consisted of 2266 bp for 14 members of the Ericales, including four species of the Polemoniaceae, plus three outgroup taxa. The Polemoniaceae were found to be related to Diospyros, Fouquieria, the Primulales, Rhododendron, and Impatiens, but relationships among taxa were generally not well supported. The precise position of the Polemoniaceae within the Ericales remains obscure. Data for intrafamilial analyses consisted of 1031 bp for 27 species of the Polemoniaceae, including at least one species from most genera of the family, plus five outgroup taxa. A single most parsimonious tree was identified. The analyses suggested that subfamily Cobaeoideae, excluding Loeselia, is monophyletic and that Huthia is sister to Cantua. Acanthogilia was sister to the remainder of subfamily Cobaeoideae. Subfamily Polemonioideae plus Loeselia formed four subclades that were strongly supported as monophyletic and represent the major lineages of the subfamily.     

A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae)

We present the most comprehensive analysis of higher-level relationships in gall wasps conducted thus far. The analysis was based on detailed study of the skeletal morphology of adults, resulting in 164 phylogenetically informative characters, complemented with a few biological characters. Thirty-seven cynipid species from thirty-one genera, including four genera of the apparently monophyletic Cynipini and almost all of the genera in the other tribes, were examined. The outgroup included exemplar species from three successively more distant cynipoid families: Figitidae (the sister group of the Cynipidae), Liopteridae and Ibaliidae. There was considerable homoplasy in the data, but many groupings in the shortest tree were nonetheless well supported, as indicated by bootstrap proportions and decay indices. Partitioning of the data suggested that the high level of homoplasy is characteristic of the Cynipidae and not the result of the amount of available phylogenetically conservative characters being exhausted. The analysis supported the monophyly of the Cynipini (oak gall wasps) which, together with the Rhoditini (the rose gall wasps), Eschatocerini and Pediaspidini formed a larger monophyletic group of gall inducers restricted to woody representatives of the eudicot subclass Rosidae. The inquilines (Synergini) were indicated to be monophyletic, whereas the Aylacini, primarily herb gall inducers, appeared as a paraphyletic assemblage of basal cynipid groups. The shortest tree suggests that the Cynipidae can be divided into three major lineages: one including the inquilines, the Aylacini genera associated with Rosaceae, and Liposthenes ; one consisting entirely of Aylacini genera, among them Aulacidea , Isocolus and Neaylax ; and one comprising the woody rosid gallers (the oak and rose gall wasps and allies), the Phanacis-Timaspis complex and the Aylacini genera associated with Papaveraceae.     

Molecular characterization of Hepatozoon species in reptiles from the Seychelles

Hepatozoon parasites were examined for the first time in reptiles from the Seychelles Islands. Although both prevalence and intensity were low, Hepatozoon species were detected in individuals from 2 endemic species, the lizard Mabuya wrightii and the snake Lycognathophis seychellensis. This was confirmed using visual identification and through sequencing part of the 18s rRNA gene. Phylogenetic analysis indicates that the Hepatozoon on the Seychelles form a monophyletic lineage, although more data are clearly needed to stabilize estimates of relationships based on this marker.     

Molecular evidence for the phylogeny of Australian gekkonoid lizards

Partial sequences of the mitochondrial 12S rRNA and nuclear c-mos genes were determined for 12 species of gekkonoid lizards representing the four major taxa of the Australian region, the Diplodactyhni and Carphodactylini (forming the subfamily Diplodactylinae), the Pygopodidae and the Gekkoninae. One further species represented a non-Australian gekkonoid lineage, the Eublepharinae. The combined sequence data were used to reconstruct the underlying molecular phylogeny. We used the molecular phylogeny to test the monophyly of the diplodactyline tribes and conflicting hypotheses of relationships of die pygopods and of the genus Oedura. Monophyly of the Diplodactylinae is supported, while pygopods form a monophyletic sister lineage to all Diplodactylinae. The molecular data support the monophyly of the Diplodactyhni, with Oedura firmly placed as a diplodactylin. Monophyly of the Carphodactylini is not supported. The four carphodactylin genera form a paraphyletic cluster at the base of the Diplodactyhni. Pygopods are nested within the traditional Gekkonidae and pygopods plus diplodactylines form a well-supported monophyletic group with respect to the remaining gekkonoids, the gekkonines and eublepharines.     

Phylogeny of the fish leeches (Oligochaeta, Hirudinida, Piscicolidae) based on nuclear and mitochondrial genes and morphology

The phylogeny of the Piscicolidae was analysed from combined 18S rDNA, cytochrome c oxidase subunit I (CO-I), nicotinamide adenine dinucleotide dehydrogenase subunit I (ND-I) and morphological data using parsimony. A worldwide distribution of Piscicolidae was represented for the first time. While the family Piscicolidae was supported as a monophyletic group, the traditional subfamilies based on morphology were not supported. The Platybdellinae was polyphyletic and formed four distinct clades, and Bathybdella sawyeri did not group with any other platybdellins. The Piscicolinae was also polyphyletic, also forming four distinct clades. The pontobdellin genus Stibarobdella was shown to be the basal taxon within the Piscicolidae; however, the Pontobdellinae was found to be paraphyletic if Oxytonostoma was included. The genera Aestabdella , Austrobdella, and Malmiana were found to be paraphyletic; the genera Calliobdella, Cystobranchus, and Platybdella were found to be polyphyletic. The species Myzobdella lugubris was not found to be monophyletic. It is proposed that Oxytonostoma be transferred out of the Pontobdellinae, that Aestabdella be synonymized with Pterobdella , that Calliobdella vivida be returned to Cystobranchus , that Gonimosobdella be synonymized with Cystobranchus , and that Piscicolaria be synonymized with Myzobdella . The synonymy of Malmiana and Heptacyclus is confirmed, with Heptacyclus having priority. Piscicola milneri is confirmed to be a separate species from Piscicola geometra .     

Genera and species of Baetidae in Japan: <Emphasis Type="Italic">Nigrobaetis,Alainites, Labiobaetis</Emphasis>, and <Emphasis Type="Italic">Tenuibaetis</Emphasis> n. stat. (Ephemeroptera)

We describe morphological characters of the genera Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis n. stat. and provide generic situations of six Japanese species: Nigrobaetis chocoratus n. comb., N. sacishimensis n. comb., Alainites atagonis, A. florens, A. yoshinensis, and Tenuibaetis pseudofrequentus. To evaluate the polarities of the morphological characters and the monophyly of Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis, character states of these four genera were compared with the genus Cloeon as an outgroup. Labiobaetis is considered to be a monophyletic group supported by a wide paraglossa. Tenuibaetis is a monophyletic group that is distinguishable from the related genera by robust setae with a medial ridge on the dorsomedian surface of the nymphal femur. We did not find any synapomorphic characters of Nigrobaetis or Alainites. Although we tentatively treat Nigrobaetis and Alainites as distinct genera, they are considered to be paraphyletic taxa.     

螟黄足盘绒茧蜂复合群支序系统学研究（膜翅目：茧蜂科：小腹茧蜂亚科）

对螟黄足盘绒茧蜂复合群Cotesia flavipes complex (膜翅目：茧蜂科：小腹茧蜂亚科)分布全世界的5个种和外群侧沟茧蜂Microplitis 及荻茧蜂Diolcogaster 的25个性状,以及复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂Cotesia kariyai、粉蝶盘绒茧蜂Cotesia glomerata 的24个性状,分别进行比较研究,并运用支序分析的方法探讨该复合群内5个种种间的系统发育关系。支序分析表明螟黄足盘绒茧蜂复合群是一单系群,二化螟盘绒茧蜂C. chilonis 和大螟盘绒茧蜂C. sesamiae 近缘,芦螟盘绒茧蜂C. chiloluteelli 和汉寿盘绒茧蜂C. hanshouensis 近缘,螟黄足盘绒茧蜂C. flavipes相对独立。 以上研究表明无论是以近缘属作外群还是以同属其它种作外群,所得结果基本上都能反映螟黄足盘绒茧蜂复合群各种之间的分类地位。     

Molecular phylogenetics of the Pectinidae (Mollusca: Bivalvia) and effect of increased taxon sampling and outgroup selection on tree topology

Evolutionary relationships of the Pectinidae were examined using two mitochondrial genes (12S rRNA, 16S rRNA) and one nuclear gene (Histone H3) for 46 species. Outgroup taxa from Propeamussidae, Spondylidae and Limidae were also sequenced to examine the impact of outgroup choice on pectinid topologies. Our phylogenetic analyses resolved the Pectinidae as monophyletic, but many of the subfamilies and tribes within the family do not form monophyletic clades. The paraphyletic Aequipectinini group is the most basal member of the Pectinidae, with the Chlamydinae and Palliolinae representing the most recently derived pectinid groups. These results are in contrast with the current morphological hypothesis of Pectinidae evolution, which suggests the Chlamydinae and Pallioline are basal groups within the Pectinidae. Ingroup topology was found to be sensitive to outgroup choice and increasing taxon sampling within the Pectinidae resulted in more robust phylogenies.