Phylogeny and classification of the Phyllodocidae (Polychaeta)

The phylogenetic relationships of the benthic phyllodocid genera are estimated by a parsimony analysis based on 26 morphological characters. It is concluded that the cephalization within Eteone has led to a reduction of segment 1 and loss of the first pair of tentacular cirri. Biramous parapodia (like in Austrophyllum and Notophyllum ) constitute an ancestral state, which has evolved independently also within Phyllodoce . A new subfamily, Notophyllinae, is erected, and the two additional subfamilies, Phyllodocinae and Eteoninae, are redefined. Pareteoninae is treated as a junior synonym to Phyllodocinae, and Lugiinae as a nomen dubium. A number of genera are redefined and new combinations introduced. Based on type material Zverlinum and Phyllouschakovius are synonymized with Phyllodoce, Nipponophyllum with Notophyllym, Vitiaziphyllum with Eumida, Pareteone, Eulalia (Euphylla) and Compsanaitis with Paranaitis. More tentatively proposed synonymizations include Bergstroemia with Austrophyllum, Prophyllodoce with Phyllodoce , and Pseudeulalia with Protomystides. Eulalia lapsus is proposed as a replacement name for E. anoculata Fauchald, 1972, junior homonym to E. anoculata Hartman & Fauchald, 1971, and a number of species are synonymized and new combinations introduced. A checklist of phyllodocids together with information on the location of their type specimens is included.     

On the Anatomy of the Central Nervous System of Phyllodocidae (Polychaeta) and the Phylogeny of Phyllodocid Genera: a New Alternative

A reinvestigation of the central nervous system of the Phyllodocidae appeared necessary as the existing literature proved insufficient for detailed comparisons with other polychaete families. Many earlier opinions turned out to be quite contradictory, especially as concerns the morphological value of the anterior end appendages. In the present paper the microanatomy of the brain and the anterior ventral cord, as well as the innervation of, inter alia, the cephalic appendages, the nuchal organs and the alimentary canal of Eulalia viridis (main object of study), Eumida sanguinea, Pterocirrus macroceros, Sige fusigera, Eteone picta, E. foliosa, Phyllodoce groenlandica, P. longipes, P. mucosa, Paranaitis wahlbergi, Notophyllum foliosum and Nereiphylla lutea are described. The results are summarized in schematic diagrams and compared with corresponding observations on other polychaete families. Thereby, some earlier opinions about the nature of the cephalic appendages of the Phyllodocidae are strengthened while others are rejected. Most significantly, it is concluded that the so-called ventrolateral antennae of phyllodocids are homologous with the palps of other polychaetes and the median antenna found in certain genera is homologous with the nuchal papilla found in most other genera of the family. These partly new or revised characters and character states formed the basis for a reconsideration of earlier ideas about the phylogeny of the Phyllodocidae: a new alternative.     

On the phylogeny of the Phyllodocidae (Polychaeta Annelida): an alternative

The phylogeny of the Phyllodocidae (Polychaeta Annelida) is examined in an analysis which includes previously unnoticed characters of the nervous system. Concentrating on four problems of importance in understanding the phylogeny of the family, a study of the nervous system demonstrated that: 1. A median antenna, such as that present in species of Eulalia, is not homologous to the nuchal papilla of, for example, Eteone and Phyllodoce; 2. Nuchal organs, presumably found in all phyllodocids, are basically homologous, but four separate types may be recognized; 3. The anterior concentration of the first segments (particularly their tentacular cirri), which is evident in most phyllodocids, is accompanied by different alterations of the ventral nervous system. The polarities of these changes are not self-evident; 4. A study of the ventral nervous system of Eteone reveals that the first apparent segment, bearing two pairs of tentacular cirri, must be regarded as homologous to the second segment of all other phyllodocids. Furthermore, this genus descended from forms in which a segment was present anterior to the first apparent segment, the first two segments possibly bearing tightly packed tentacular cirri. In connection with other characters, this information was used to produce a cladogram, which has some surprising features. The most unexpected result is the prediction that Eteone is the sister group of a taxon containing the genera Paranaitis, Chaetoparia, and Phyllodoce. The results are discussed and compared to those of other investigators.     

Three new phyllodocid species (Polychaeta) from the Trondheimsfjord in Norway, including some notes concerning the validity of the genus Steggoa

Three new phyllodocid species are described from the deep coral reefs in the Trondheimsfjord in Norway: Eulalia hanssoni sp.n., Eulalia microoculata sp.n. and Pterocirrus nidarosiensis sp.n. The genus Steggoa Bergstrom, 1914 is considered a junior synonym of Eulalia Savigny, 1817.     

A phylogenetic analysis of Coelopidae (Diptera) based on morphological and DNA sequence data

The phylogenetic relationships of 22 species of Coelopidae are reconstructed based on a data matrix consisting of morphological and DNA sequence characters (16S rDNA, EF-1alpha). Optimal gap and transversion costs are determined via a sensitivity analysis and both equal weighting and a transversion cost of 2 are found to perform best based on taxonomic congruence, character incongruence, and tree support. The preferred phylogenetic hypothesis is fully resolved and well-supported by jackknife, bootstrap, and Bremer support values, but it is in conflict with the cladogram based on morphological characters alone. Most notably, the Coelopidae and the genus Coelopa are not monophyletic. However, partitioned Bremer Support and an analysis of node stability under different gap and transversion costs reveal that the critical clades rendering these taxa non-monophyletic are poorly supported. Furthermore, the monophyly of Coelopidae and Coelopa is not rejected in analyses using 16S rDNA that was manually aligned. The resolution of the tree based on this reduced data sets is, however, lower than for the tree based on the full data sets. Partitioned Bremer support values reveal that 16S rDNA characters provide the largest amount of tree support, but the support values are heavily dependent on analysis conditions. Problems with direct comparison of branch support values for trees derived using fixed alignments with those obtained under optimization alignment are discussed. Biogeographic history and available behavioral and genetic data are also discussed in light of this first cladogram for Coelopidae based on a quantitative phylogenetic analysis.     

蝗科高级阶元的分子系统发育(英文）

迄今,蝗科内各分类阶元之间的系统发生关系大部分是未知的。本文用来自中国24种蝗科昆虫的12SrDNA和16SrDNA2个基因的联合序列(共795bp)数据,以锥头蝗科的锥头蝗(Pyrgomorpha conica)为外群,重建了分子系统树。研究结果表明,在12SrDNA与16SrDNA组成的联合数据中,转换的替代速率明显比颠换的替代速率高得多,核酸的替代已经发生了饱和。分子系统树表明:斑翅蝗亚科是一单系群,该亚科是一个合法的亚科,但斑腿蝗亚科和蝗亚科都不是单系群;斑翅蝗亚科在蝗科内是一个相对原始的类群,而稻蝗亚科比斑翅蝗亚科相对进化,比蝗科的其他亚科的种类相对原始。     

A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea)

An hypothesis of phylogenetic relationships of Asilidae and its constituent taxa is presented, combining morphological and DNA sequence data in a total evidence framework. It is based on 77 robber fly species, 11 Asiloidea outgroup species, 211 morphological characters of the adult fly, and approximately 7300 bp of nuclear DNA from five genes (18S and 28S rDNA, AATS, CAD, and EF-1α protein-encoding DNA). The equally weighted, simultaneous parsimony analysis under dynamic homology in POY resulted in a single most parsimonious cladogram with a cost of 27,582 (iterative pass optimization; 27,703 under regular direct optimization). Six of the 12 included subfamily taxa are recovered as monophyletic. Trigonomiminae, previously always considered as monophyletic based on morphology, is shown to be non-monophyletic. Two of the three Trigonomiminae genera, Holcocephala Jaennicke, 1867 and Rhipidocephala Hermann, 1926, group unexpectedly as the sister taxon to all other Asilidae. Laphriinae, previously seen in the latter position, is the sister group of the remaining Asilidae. Five other subfamily taxa, i.e. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Tillobromatinae, and Willistonininae, are also shown to be non-monophyletic. The phylogenetic relationships among the higher-level taxa are partly at odds with findings of a recently published morphological study based on more extensive taxon sampling. The total evidence hypothesis is considered as the most informative one, but the respective topologies from the total-evidence, morphology-only, and molecular-only analyses are compared and contrasted in order to discuss the signals from morphological versus molecular data, and to analyze whether the molecular data outcompete the fewer morphological characters. A clade Apioceridae+Mydidae is corroborated as the sister taxon to Asilidae.     

DIVERSITY AMONG TETRASPORALEAN TAXA: EVIDENCE FROM ULTRASTRUCTURE AND MOLECULES

Recent phylogenetic studies of tetrasporalean exemplars using 18S rDNA revealed the extent of diversity among taxa in this non-monophyletic group (Booton et al. 1998). In particular, two distinct tetrasporalean lineages were identified; one group allied with chlamydomonadalean taxa and the other group comprising taxa now placed in a separate order, the Chaetopeltidales. Using these observations as a framework, a new investigation of diversity among a broader sampling of putative tetrasporalean genera (e.g. Asterococcus , Chloranomala , Chlorophysema , Gloeocystis , Gloeodendron , Palmella , Paulschulzia , Physocytium , Schizochlamys and Tetraspora ) was undertaken. Phylogenetic studies of both 18S and 26S rDNA were compared with ultrastructural investigations of vegetative cells. Molecular phylogenetic analyses corroborate the earlier 18S rDNA results, but also reveal additional diversity. The new data raise doubts regarding the monophyly of two genera, Palmella and Tetraspora. The new data also link two enigmatic green algal genera, Physocytium and Heterochlamydomonas , in a long-branch lineage within the Chlamydomonadales. Another enigmatic genus, Schizochlamys , is allied with Bracteacoccus in the Sphaeropleales. Lastly, Chloranomala is resolved as an ally of Paulschulzia , Tetraspora sp., and the green flagellate, Lobomonas. Comparison of pyrenoid ultrastructure generally supports the molecular phylogenetic analyses, suggesting that this non-molecular character will be a useful marker for broad phylogenetic studies of chlamydomonadalean taxa. (Supported by NSF grant, DEB 9726588)     

ZYGOTE APPENDAGES (CIRRI), A NEW STRUCTURAL FEATURE IN THE SPHAEROPLEACEAE(CHLOROPHYCEAE)1

Cirri are curled appendages borne on the surface of mature zygotes in some members of the green algal family Sphaeropleaceae. Structures equivalent to cirri have not been reported previously in any green algae. We found cirri in four taxa: Sphaeroplea fragilis Buchheim et Hoffman, Atractomorpha porcata Hoffman, and two new undescribed species of Atractomorpha from Africa. Zygotes of seven other sphaeropleacean taxa were examined, but they did not produce cirri. Cirri may be either simple or compound. Simple cirri are less than 0.5 μm in diameter and show no evidence of longitudinal striations or subunits. Compound cirri are more massive (commonly 2.0–3.5 μm, and up to 4 μm diameter in S. fragilis) and possess longitudinal striations suggesting that each is formed of longitudinally arranged subunits ca. 0.5 μm in diameter. Simple and compound cirri are formed by zygotes of all four taxa that produce cirri. Both types may occur on the same zygote. Nonetheless, the patterns of variation in length, shape, and distribution of the two types of cirri are sufficiently distinct to characterize each taxon. Energy dispersive X-ray microanalysis (EDX) shows that cirri lack appreciable mineralization and are thus primarily organic in nature. The potential value of cirri as a taxonomic character in the Sphaeropleaceae is discussed.     

Phylogeny of Hesionidae (Aciculata, Polychaeta), assessed from morphology, 18S rDNA, 28S rDNA, 16S rDNA and COI

We assess phylogenetic relationships within the polychaete family Hesionidae from morphological data combined with nucleotide data from 18S rDNA, 28S rDNA, 16S rDNA and COI. Parsimony and Bayesian analyses were performed on two data sets; the first was based on a more restricted set of terminals with both morphological and molecular data (17 ingroup terminals), while the second included additional taxa with morphological data only (25 ingroup terminals). The different sets of terminals yielded fully congruent results, as did the parsimony and the Bayesian analyses. Our results indicate high levels of homoplasy in traditionally used morphological characters in the group, and that Hesioninae, Gyptini and Gyptis are nonmonophyletic. Hesionini (mainly Hesione and Leocrates ), Psamathini (mainly Hesiospina , Micropodarke , Nereimyra , Psamathe and Syllidia ), Ophiodrominae (Gyptini and Ophiodromini) and Ophiodromini (mainly Heteropodarke , Ophiodromus and Podarkeopsis ) are monophyletic and agree with previous classifications, and Hesionini is probably the sister to all other hesionids. The placements of the small hesionids capricornia and Lizardia , the hydrothermal vent taxa Hesiodeira and Hesiolyra , and the newly described Hesiobranchia , remain uncertain.     

中国黄金茅属与楔颖草属的新分类等级

本文在黄金茅属Eulalia Kunth下发表2新种,即E.splendensKeng et S.L.Chen与E.yunnanensis Keng et S.L.Chen,在楔颖草属Apocopis Nees下发表一新变种A.wrightii var.macrantha S.L.Chen。     

Basal teleosts and the question of elopomorph monophyly. Morphological and molecular approaches

The methodology used by previous authors to resolve the relationships of the elopomorphan taxa is criticized. The morphological characters that have been proposed to support the monophyly of the Elopomorpha are reviewed and it is shown that most of them are weak. A new hypothesis of relationships is proposed on the basis of nucleotidic sequences of ribosomal RNA 18S, 16S and 12S. In order to really test all the possible relationships, the monophyly of the Elopomorpha was not considered a priori. The tree was rooted on Amia calva and Lepisosteus osseus and the ingroup taxa sampling was subsequently increased. The obtained topology shows that the Elopomorpha is a non-monophyletic taxon. Elopiforms, anguilliforms, albuliforms and notacanthiforms are considered here as four monophyletic, incertae sedis taxa among basal teleosts.     

Vrijenhoekia balaenophila, a new hesionid polychaete from a whale fall off California

Vrijenhoekia balaenophila gen. nov., sp. nov. (Polychaeta, Hesionidae) is described from a whale carcass at near 3000 m depth in Monterey Canyon off the coast of California. The phylogenetic relationships of V. balaenophila are assessed in a parsimony analysis of morphological data together with nucleotide data from 28S rDNA, 16S rDNA and cytochrome oxidase I genes. Within the hesionids V. balaenophila belongs to Psamathini, where it is the sister group to Sirsoe . Among psamathins it is morphologically distinguished by having six glandular lip pads around the mouth opening, papilla-shaped neuropodial lobes on segment 3, extreme length of the dorsal cirri, and by a characteristic growth pattern in which the maximum number of segments is already formed in subadults, and further growth takes place through size increase of the segments.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 625–634.     

Ecological parallelism and cryptic species in the genus Ophiothrix derived from mitochondrial DNA sequences

We addressed the long-standing problem of species assignment of two nominal species of the genus Ophiothrix (Echinodermata, Ophiuroidea) by phylogenetic analysis of a segment of the mitochondrial 16S rDNA. Our phylogeny identified two distinct mitochondrial lineages that do not correspond to the present species assignments. Individuals of the endemic Mediterranean species O. quinquemaculata were clustered with individuals of O. fragilis in both mitochondrial lineages. We thus suggest that these taxa are not biological species but ecotypes. Differences between the two ecotypes in morphological and physiological characteristics may be explained by adaptation to environmental conditions at different water depths. Despite the observed ecomorphological variability within each of the two major mitochondrial lineages, the large genetic distance (9.0-12.0%) between them does suggest the existence of two distinct biological species. Their reproductive isolation could result from differences in reproductive strategy rather than by ecological and/or morphological differentiation.     

Chromosomal localization of 45S and 5S rDNA in 14 species and the implications for genome evolution of genus Epimedium

Studying the genome structure of Epimedium has been hindered by the large genomes and uniform karyotypes. Consequently our understanding of the genome organization and evolutionary changes of Epimedium is extremely limited. In the present study, the 45S and 5S rDNA loci of 14 Epimedium species were physically mapped by double-probe FISH for the first time. Results showed the following: (1) Chromosomes I and II of all 14 species examined, except for E. shuichengense, hosted one pair of 45S rDNA sites, respectively. Most of the 45S rDNA sites gave clear signals and were positioned in the distal regions of the short arms. (2) All species studied of section Diphyllon were found to have one pair of 5S rDNA sites localized in the interstitial regions of the long arm of chromosome IV, and the two species of section Epimedium, E. alpinum and E. pubigerum, had two pairs of 5S rDNA sites localized in the interstitial regions of the long arm of chromosomes IV and V, respectively. (3) In section Diphyllon, all species of small flower taxa, except E. shuichengense, had three pairs of 45S rDNA sites, clearly more than species of big flower taxa, except E. davidii, with two pairs of 45S rDNA sites. Based on the 45S and 5S rDNA distribution patterns and other chromosomal morphological characteristics, six pairs of chromosomes can be unambiguously identified in all 14 Epimedium species. The stable differentiation in 45S and 5S rDNA FISH patterns between the two sections suggests that chromosomal rearrangements and transpositional events played a role in the splitting of the two sections, and section Diphyllon may be more primitive than section Epimedium. In the same way, big flower taxa may be more primitive than small flower taxa in section Diphyllon.     

The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology

Phylogenetic relationships among the holometabolous insect orders were inferred from cladistic analysis of nucleotide sequences of 18S ribosomal DNA (rDNA) (85 exemplars) and 28S rDNA (52 exemplars) and morphological characters. Exemplar outgroup taxa were Collembola (1 sequence), Archaeognatha (1), Ephemerida (1), Odonata (2), Plecoptera (2), Blattodea (1), Mantodea (1), Dermaptera (1), Orthoptera (1), Phasmatodea (1), Embioptera (1), Psocoptera (1), Phthiraptera (1), Hemiptera (4), and Thysanoptera (1). Exemplar ingroup taxa were Coleoptera: Archostemata (1), Adephaga (2), and Polyphaga (7); Megaloptera (1); Raphidioptera (1); Neuroptera (sensu stricto = Planipennia): Mantispoidea (2), Hemerobioidea (2), and Myrmeleontoidea (2); Hymenoptera: Symphyta (4) and Apocrita (19); Trichoptera: Hydropsychoidea (1) and Limnephiloidea (2); Lepidoptera: Ditrysia (3); Siphonaptera: Pulicoidea (1) and Ceratophylloidea (2); Mecoptera: Meropeidae (1), Boreidae (1), Panorpidae (1), and Bittacidae (2); Diptera: Nematocera (1), Brachycera (2), and Cyclorrhapha (1); and Strepsiptera: Corioxenidae (1), Myrmecolacidae (1), Elenchidae (1), and Stylopidae (3). We analyzed approximately 1 kilobase of 18S rDNA, starting 398 nucleotides downstream of the 5' end, and approximately 400 bp of 28S rDNA in expansion segment D3. Multiple alignment of the 18S and 28S sequences resulted in 1,116 nucleotide positions with 24 insert regions and 398 positions with 14 insert regions, respectively. All Strepsiptera and Neuroptera have large insert regions in 18S and 28S. The secondary structure of 18S insert 23 is composed of long stems that are GC rich in the basal Strepsiptera and AT rich in the more derived Strepsiptera. A matrix of 176 morphological characters was analyzed for holometabolous orders. Incongruence length difference tests indicate that the 28S + morphological data sets are incongruent but that 28S + 18S, 18S + morphology, and 28S + 18S + morphology fail to reject the hypothesis of congruence. Phylogenetic trees were generated by parsimony analysis, and clade robustness was evaluated by branch length, Bremer support, percentage of extra steps required to force paraphyly, and sensitivity analysis using the following parameters: gap weights, morphological character weights, methods of data set combination, removal of key taxa, and alignment region. The following are monophyletic under most or all combinations of parameter values: Holometabola, Polyphaga, Megaloptera + Raphidioptera, Neuroptera, Hymenoptera, Trichoptera, Lepidoptera, Amphiesmenoptera (Trichoptera + Lepidoptera), Siphonaptera, Siphonaptera + Mecoptera, Strepsiptera, Diptera, and Strepsiptera + Diptera (Halteria). Antliophora (Mecoptera + Diptera + Siphonaptera + Strepsiptera), Mecopterida (Antliophora + Amphiesmenoptera), and Hymenoptera + Mecopterida are supported in the majority of total evidence analyses. Mecoptera may be paraphyletic because Boreus is often placed as sister group to the fleas; hence, Siphonaptera may be subordinate within Mecoptera. The 18S sequences for Priacma (Coleoptera: Archostemata), Colpocaccus (Coleoptera: Adephaga), Agulla (Raphidioptera), and Corydalus (Megaloptera) are nearly identical, and Neuropterida are monophyletic only when those two beetle sequences are removed from the analysis. Coleoptera are therefore paraphyletic under almost all combinations of parameter values. Halteria and Amphiesmenoptera have high Bremer support values and long branch lengths. The data do not support placement of Strepsiptera outside of Holometabola nor as sister group to Coleoptera. We reject the notion that the monophyly of Halteria is due to long branch attraction because Strepsiptera and Diptera do not have the longest branches and there is phylogenetic congruence between molecules, across the entire parameter space, and between morphological and molecular data.     

Morphology and Small Subunit rDNA Phylogeny of Two New Marine Urostylid Ciliates,Caudiholosticha marina sp. nov. and Nothoholosticha flava sp. nov. (Ciliophora,Hypotrichia)

Two marine urostylid ciliates, Caudiholosticha marina sp. nov. and Nothoholosticha flava sp. nov., isolated from intertidal sediment in the Yellow Sea, are investigated using morphological and small subunit rDNA phylogenetic analyses. Caudiholosticha marina is 210?310 μm × 40?55 μm in vivo, and has 10?20 macronuclear nodules, 23?37 midventral cirral pairs extending to 5?8 transverse cirri, and two caudal cirri. It differs from congeners by its marine habitat, larger size, macronuclear arrangement pattern and high number of midventral pairs. Molecular phylogenetic analyses indicate a polyphyly of Caudiholosticha. Nothoholosticha flava is yellow to brownish and 240?320 μm × 40?60 μm sized, and has a bipartite adoral zone, six frontal cirri in atypical bicorona, usually four frontoterminal, one buccal and 5?7 transverse cirri and 28?54 midventral pairs. Phylogenetic analyses allocate N. flava as sister of N. fasciola, type of the genus. The two Nothoholosticha species differ distinctly by the presence/absence of frontoterminal cirri, a feature often used to define genera in the Hypotrichia. However, the SSU rDNA sequence similarity between these two species is 99.3%, which weakens the justification for separating the new isolate at genus level. The taxonomic significance of frontoterminal cirri is discussed based on morphological and molecular data.     

Morphology,morphogenesis, and molecular phylogeny of a new freshwater ciliate,Gonostomum jangbogoensis n. sp. (Ciliophora,Hypotricha), from Victoria Land,Antarctica

In a study on ciliate diversity, we discovered the new hypotrich species, Gonostomum jangbogoensis n. sp., in freshwater from Terra Nova Bay, Victoria Land, southeast Antarctica. We describe its morphology and morphogenesis using standard methods, and the SSU rRNA gene phylogeny is provided as well. Morphology of Gonostomum jangbogoensis n. sp. is characterized as follows: slender to elongated body shape; grayish under low magnification; cortical granules present; 32–41 adoral membranelles; 3 enlarged frontal cirri; 1 buccal cirrus; 2 frontoterminal cirri; 3 or 4 frontoventral cirral pairs, 2 pretransverse cirri, 6–7 transverse cirri; 13–19 left and 18–26 right marginal cirri; 17–23 paroral kinetids; 3 dorsal kineties; 3 caudal cirri; 2 macronuclear nodules with 1–3 micronuclei. The morphogenesis of the new species confirms that it has at least seven frontal-ventral-transverse cirral anlagen, which is also reported in Gonostomum sp. 1 sensu Shin from Korea. Even though these two populations occur very far from each other, the morphometric data prove that this character state, the seven cirral anlagen, is a stable feature across these populations and might be an apomorphy. The phylogenetic analyses show that the genus Gonostomum is non-monophyletic and that the new species is a sister to G. bromelicola.