Phylogenetic analysis of Tubificidae (Annelida, Clitellata) based on 18S rDNA sequences

Tubificids are aquatic clitellate worms, but recent analyses of morphological characters suggested that this family, as currently recognized, is paraphyletic. Sequences of the 18S rDNA gene of 40 protostome worm species (including 13 representatives of the Tubificidae) and 2 mollusc species were cladistically analyzed to test the monophyly of the Tubificidae and that of some of its constituent subfamilies. Under all alignments tested, the same general phylogenetic pattern emerged. The data support the idea that the Naididae, another clitellate taxon, is associated with some "rhyacodriline" groups within the Tubificidae. The data also corroborate the idea that the Tubificinae and the Limnodriloidinae are monophyletic but indicate that the Rhyacodrilinae and the Phallodrilinae are not. Bathydrilus does not appear to be closely related to other "phallodriline" genera.     

Phylogeny of Passerida (Aves: Passeriformes) based on nuclear and mitochondrial sequence data

Passerida is a monophyletic group of oscine passerines that includes almost 3500 species (about 36%) of all bird species in the world. The current understanding of higher-level relationships within Passerida is based on DNA-DNA hybridizations [C.G. Sibley, J.E. Ahlquist, Phylogeny and Classification of Birds, 1990, Yale University Press, New Haven, CT]. Our results are based on analyses of 3130 aligned nucleotide sequence data obtained from 48 ingroup and 13 outgroup genera. Three nuclear genes were sequenced: c-myc (498-510 bp), RAG-1 (930 bp), and myoglobin (693-722 bp), as well one mitochondrial gene; cytochrome b (879 bp). The data were analysed by parsimony, maximum-likelihood, and Bayesian inference. The African rockfowl and rockjumper are found to constitute the deepest branch within Passerida, but relationships among the other taxa are poorly resolved--only four major clades receive statistical support. One clade corresponds to Passeroidea of [C.G. Sibley, B.L. Monroe, Distribution and Taxonomy of Birds of the World, 1990, Yale University Press, New Haven, CT] and includes, e.g., flowerpeckers, sunbirds, accentors, weavers, estrilds, wagtails, finches, and sparrows. Starlings, mockingbirds, thrushes, Old World flycatchers, and dippers also group together in a clade corresponding to Muscicapoidea of Sibley and Monroe [op. cit.]. Monophyly of their Sylvioidea could not be corroborated--these taxa falls either into a clade with wrens, gnatcatchers, and nuthatches, or one with, e.g., warblers, bulbuls, babblers, and white-eyes. The tits, penduline tits, and waxwings belong to Passerida but have no close relatives among the taxa studied herein.     

The double sperm line in Isochaetides (Annelida,Clitellata, Tubificidae)

Sperm morphology and spermatogenesis were examined in the oligochaete annelid Isochaetides arenarius, a species belonging to the subfamily Tubificinae inhabiting the sediments of Lake Baikal. As all tubificines, Isochaetides produces two types of spermatozoa, named eusperm and parasperm. The eusperm are the fertilizing male gametes and consist, in sequence, of an acrosome, a nucleus, a mitochondrial mid-piece, and a tail. The parasperm have the same general architecture, but differ in cytological details: the acrosome is shorter, devoid of a perforatorium, and the acrosome vesicle has a different, simpler, shape. The nucleus is much shorter and rectilinear (the eusperm nucleus is twisted). The mid-piece mitochondria are less numerous but their overall volume is larger. The flagellum has a plasma membrane largely separated from the axoneme, and is devoid of glycogen granules. After mating, the two sperm types gather in the spermathecae to form spermatozeugmata; in these structures the parasperm form an external sheath involving the centrally located eusperm and their tails are connected by conspicuous septate junctions. Parasperm nuclei are produced through a process of fragmentation of the 'spermatocytes', whereas the flagellar basal bodies are produced by a process similar to that giving rise to basal bodies in ciliated epithelia.     

Molecular evidence for the non-monophyletic status of Naidinae (Annelida, Clitellata, Tubificidae)

Naidinae (former Naididae) is a group of small aquatic clitellate annelids, common worldwide. In this study, we evaluated the phylogenetic status of Naidinae, and examined the phylogenetic relationships within the group. Sequence data from two mitochondrial genes (12S rDNA and 16S rDNA), and one nuclear gene (18S rDNA), were used. Sequences were obtained from 27 naidine species, 24 species from the other tubificid subfamilies, and five outgroup taxa. New sequences (in all 108) as well as GenBank data were used. The data were analysed by parsimony and Bayesian inference. The tree topologies emanating from the different analyses are congruent to a great extent. Naidinae is not found to be monophyletic. The naidine genus Pristina appears to be a derived group within a clade consisting of several genera (Ainudrilus, Epirodrilus, Monopylephorus, and Rhyacodrilus) from another tubificid subfamily, Rhyacodrilinae. These results demonstrate the need for a taxonomic revision: either Ainudrilus, Epirodrilus, Monopylephorus, and Rhyacodrilus should be included within Naidinae, or Pristina should be excluded from this subfamily. Monophyly of four out of six naidine genera represented by more than one species is supported: Chaetogaster, Dero, Paranais, and Pristina, respectively.     

Ultrastructural investigation of coelomocytes in representatives of Naidinae and Rhyacodrilinae (Annelida,Clitellata, Tubificidae)

Various types of free‐floating cells are found in the coelomic fluid of representatives of several annelid groups. The ultrastructure of these “coelomocytes,” however, has been studied to a limited degree. In this study, we used a transmission electron microscope to investigate the coelomocytes in specimens of five species of Naidinae and three species of Rhyacodrilinae (all oligochaetous clitellates within the family Tubificidae). These were compared with each other and with previously described coelomocytes of representatives of other oligochaete taxa. Only one distinguishable coelomocyte type was found in the studied specimens: a round to oblong cell without pseudopodia or other appendages, primarily containing membrane‐bound granules of varying electron density, a prominent network of rough endoplasmic reticulum, and free ribosomes. This type differs to a great extent from most of the previously described coelomocytes, but shows similarities to certain types found in members of Enchytraeidae and Megascolecidae. Although we noticed some variation, we did not find any ultrastructural characters in these cells obviously useful for phylogenetic studies within Tubificidae. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Phylogeny and evolution of Burmanniaceae (Dioscoreales) based on nuclear and mitochondrial data

The mycoheterotrophic Burmanniaceae are one of the three families currently recognized in the order Dioscoreales. Phylogenetic inference using nucleotide sequences of the nuclear 18S rDNA region and the mitochondrial nad1 b-c intron revealed two well-supported, major lineages within the family, corresponding to the two tribes recognized in the family: Burmannieae and Thismieae. All data supported a strong relationship between Thismieae and Tacca (Dioscoreaceae) making both Burmanniaceae and Dioscoreaceae polyphyletic. The three largest Burmanniaceae genera, Burmannia, Gymnosiphon, and Thismia, are paraphyletic. The splitting of Burmanniaceae into Burmannieae and Thismieae indicates two independent origins of mycoheterotrophy and correlated loss of chlorophyll in Dioscoreales. In the genus Burmannia, in which many species still contain chlorophyll, the achlorophyllous species are nested in between the autotrophic species, suggesting many independent changes from autotrophy to heterotrophy or vice versa. A Bayesian relative rates test on the 18S rDNA data showed considerable variation in substitution rates among Burmanniaceae. The substitution rates in all Thismieae and many Burmannieae are significantly faster than in Dioscoreaceae, but there seems to be no correlation between rate increases and the loss of photosynthesis.     

A phylogenetic analysis of Tubificinae and Limnodriloidinae (Annelida, Clitellata, Tubificidae) using sperm and somatic characters

Marotta, R., Ferraguti, M. & Erséus, C. (2003). A phylogenetic analysis of Tubificinae and Limnodriloidinae (Annelida, Clitellata, Tubificidae) using sperm and somatic characters. — Zoologica Scripta , 32 , 255–278.
The spermatozoa and the sperm aggregates of 13 species belonging to four genera ( Smithsonidrilus, Limnodriloides, Thalassodrilides, Doliodrilus ) in the tubificid subfamily Limnodriloidinae (Annelida, Oligochaeta) were studied and compared with the spermatozoal patterns already described in the subfamily Tubificinae. Two characters considered exclusive for the Tubificinae were found in the more spermatologically variable Limnodriloidinae: the production of two kinds of spermatozoa, eusperm and parasperm, and the presence, in the spermathecae, of sperm aggregates formed by a combination of the two sperm types. A parsimony analysis was performed on the spermatozoal data of the species examined and compared with that based on the somatic characters of the same species: a critical revision of the already codified eusperm characters was carried out and the ultrastructure of parasperm was used as a new subset of spermatozoal characters. In a total evidence approach, a further parsimony analysis was run using a matrix combining both sets of characters. This analysis suggested that the double sperm line and the sperm aggregates composed of both eusperm and parasperm may well be homologous in tubificines and limnodriloidines. It thus supported the previous notion that Tubificinae and Limnodriloidinae are closely related and indicated that these subfamilies may be sister taxa.     

Phylogeny of Thaumastodermatidae (Gastrotricha: Macrodasyida) inferred from nuclear and mitochondrial sequence data

Background

Phylogenetic relationships within Gastrotricha are poorly known. Attempts to shed light on this subject using morphological traits have led to hypotheses lacking satisfactory statistical support; it seemed therefore that a different approach was needed.

Methodology/Principal Findings

In this paper we attempt to elucidate the relationships within the taxonomically vast family Thaumastodermatidae (Macrodasyida) using molecular sequence data. The study includes representatives of all the extant genera of the family and for the first time uses a multi-gene approach to infer evolutionary liaisons within Gastrotricha. The final data set comprises sequences of three genes (18S, 28S rDNA and COI mtDNA) from 41 species, including 29 thaumastodermatids, 11 non-thaumastodermatid macrodasyidans and a single chaetonotidan. Molecular data was analyzed as a combined set of 3 genes and as individual genes, using Bayesian and maximum likelihood approaches. Two different outgroups were used: Xenotrichula intermedia (Chaetonotida) and members of the putative basal Dactylopodola (Macrodasyida). Thaumastodermatidae and all other sampled macrodasyidan families were found monophyletic except for Cephalodasyidae. Within Thaumastodermatidae Diplodasyinae and Thaumastodermatinae are monophyletic and so are most genera. Oregodasys turns out to be the most basal group within Thaumastodermatinae in analyses of the concatenated data set as well as in analyses of the nuclear genes. Thaumastoderma appears as the sister taxon to the remaining species. Surprisingly, Tetranchyroderma is non-monophyletic in our analyses as one group of species clusters with Ptychostomella while another appears as the sister group of Pseudostomella.

Conclusions/Significance

Results in general agree with the current classification; however, a revision of the more derived thaumastodermatid taxa seems necessary. We also found that the ostensible COI sequences from several species do not conform to the general invertebrate or any other published mitochondrial genetic code; they may be mitochondrially derived nuclear genes (numts), or one or more modifications of the mitochondrial genetic code within Gastrotricha.     

Phylogeny of mysticete whales based on mitochondrial and nuclear data

Mysticetes or baleen whales are comprised of four groups: Eschrichtiidae, Neobalaenidae, Balaenidae, and Balaenopteridae. Various phylogenetic hypotheses among these four groups have been proposed. Previous studies have not satisfactorily determined relationships among the four groups with a high degree of confidence. The objective of this study is to determine the relationships among the mysticete whales. Mitochondrial and nuclear DNA were sequenced for phylogenetic analysis. Most species relationships determined using these data were well resolved and congruent. Balaenidae is the most basal group and Neobalaenidae is the second most basal and sister group to the balaenopterid-eschrichtiid clade. In this phylogenetic study, the resolution of Eschrichtiidae with two main lineages of Balaenopteridae was problematic. Some data partitions placed this group within the balaenopterids, and other partitions placed it as a sister taxon to the balaenopterids. An additive likelihood approach was used to determine the most optimal trees. Although it was not found in the combined phylogenetic analyses, the "best" tree found under the additive likelihood approach was one with a monophyletic Balaenopteridae.     

A proposal to regard the former family Naididae as a subfamily within Tubificidae (Annelida, Clitellata)

A new species of Placobdelloides is described from crocodiles and river turtles in the Singapore Zoological Gardens. Placobdelloides stellapapillosa is three annulate, has one pair of eyes on somite I or II, six pairs of testisacs, two annuli between the gonopores, one post anal annulus, seven pairs of lobed crop caeca and 12–14 unique star-shaped papillae on the dorsal surface of each annulus. Star-shaped papillae on annulus a2 are typically larger and more pronounced than on annuli a1 and a3. Additional minute cone-shaped papillae may occur between the larger star-shaped papillae or may replace these papillae on annuli a1 and a3. Eggs and young (100–200) are attached by their posterior suckers directly to the ventral surface of the parent.     

Phylogeny of Saururaceae based on mitochondrial matR gene sequence data

DNA sequences of matR gene from three species of Saururaceae and the selected outgroups, Chloranthus holostegius and Zippelia begoniaefolia, are reported. All DNA sequences of six species in four genera of Saururaceae and the two outgroups are analyzed on PAUP 4.0 8b to reconstruct the phylogeny. A single matR gene tree is generated from parsimony, distance, and likelihood analyses, respectively. The three trees with the same topology are slightly different in bootstrapping support for some clades. The result indicates that Saururaceae is monophyletic. Anemopsis is sister to Houttuynia, and the two genera form the first diverging lineage of the family. The sister group relationship between Saururus and Gymnotheca is also supported by a relatively high bootstrap value. The result is different from all the former phylogenetic opinions on Saururaceae based on morphology, but it is supported by the evolution of flower-bract stalk in Saururaceae. In addition, some characteristics of the matR gene are analyzed. The MatR gene is a relatively better tool to reconstruct the molecular clock because the base substitution bias greatly decreases in the gene.     

Phylogeny of Cyttaria inferred from nuclear and mitochondrial sequence and morphological data

Cyttaria species (Leotiomycetes, Cyttariales) are obligate, biotrophic associates of Nothofagus (Hamamelididae, Nothofagaceae), the southern beech. As such Cyttaria species are restricted to the southern hemisphere, inhabiting southern South America (Argentina and Chile) and southeastern Australasia (southeastern Australia including Tasmania, and New Zealand). The relationship of Cyttaria to other Leotiomycetes and the relationships among species of Cyttaria were investigated with newly generated sequences of partial nucSSU, nucLSU and mitSSU rRNA, as well as TEF1 sequence data and morphological data. Results found Cyttaria to be defined as a strongly supported clade. There is evidence for a close relationship between Cyttaria and these members of the Helotiales: Cordierites, certain Encoelia spp., Ionomidotis and to a lesser extent Chlorociboria. Order Cyttariales is supported by molecular data, as well as by the unique endostromatic apothecia, lack of chitin and highly specific habit of Cyttaria species. Twelve Cyttaria species are hypothesized, including all 11 currently accepted species plus an undescribed species that accommodates specimens known in New Zealand by the misapplied name C. gunnii, as revealed by molecular data. Thus the name C. gunnii sensu stricto is reserved for specimens occurring on N. cunninghamii in Australia, including Tasmania. Morphological data now support the continued recognition of C. septentrionalis as a species separate from C. gunnii. Three major clades are identified within Cyttaria: one in South America hosted by subgenus Nothofagus, another in South America hosted by subgenera Nothofagus and Lophozonia, and a third in South America and Australasia hosted by subgenus Lophozonia, thus producing a non-monophyletic grade of South American species and a monophyletic clade of Australasian species, including monophyletic Australian and New Zealand clades. Cyttaria species do not sort into clades according to their associations with subgenera Lophozonia and Nothofagus.     

Phylogeny of Chloranthus (Chloranthaceae) based on nuclear ribosomal ITS and plastid TRNL-F sequence data

The internal transcribed spacers (ITS) of the nuclear ribosomal DNA and trnL-F region of the chloroplast DNA were sequenced for all ten species of Chloranthus and the outgroup Sarcandra (Chloranthaceae). Parsimony analyses of separate and combined data sets strongly suggest that Chloranthus is monophyletic and can be divided into two major clades: one containing C. erectus, C. spicatus, C. serratus, C. henryi, C. sessilifolius, and C. oldhamii (Clade A), and the other comprising C. angustifolius, C. fortunei, C. nervosus, and C. japonicus (Clade B). Taxonomically, these two clades correspond to Bentham and Hooker's sections Euchloranthus and Tricercandra. Within Clade A, two subclades, corresponding to Solms-Laubach's sections Triandri and Brachyuri, can be recognized. Solms-Laubach's subgenera Fruticosi and Herbacei, however, were resolved as paraphyletic, and thus the traditional division of Chloranthus on the basis of growth habit was not supported. Evidence from ITS and trnL-F sequences, in agreement with morphology, anatomy, and cytology, strongly suggest that Chloranthus consists of two groups that morphologically may be distinguished by their androecial characters. The present study also supports the hypothesis that the tripartite androecium of Chloranthus may have arisen by splitting of a single stamen with two marginal thecae.     

New alluroidids (Annelida,Clitellata) from Guyana

A total of 30 microbialites at two sites in Lake Clifton, Yalgorup National Park, Western Australia, were sampled by coring to quantify the associated fauna with these organo-sedimentary structures. Twenty five species of aquatic fauna were recorded from the cores, comprising 20 species of metazoan, predominantly Crustacea (including Melita kauerti (Amphipoda), Exosphaeroma cf. serventii (Isopoda); and Cyprideis australiensis(Ostracoda); Polychaeta (Capitella cf. capitata); nematodes; and five species of Foraminifera (Protista). Multivariate analysis of the five numerically most abundant taxa (amphipods, isopods, ostracods, polychaetes, nematodes) separated microbialites by season and submergence. Numbers of all taxa, particularly polychaetes and amphipods, were much higher in spring than in autumn, and in permanently-inundated than in seasonally-inundated microbialites. The exception was higher numbers of juvenile polychaetes in seasonally-inundated microbialites at the northern site in spring. This study showed that modern thrombolitic microbialites can co-exist with a diverse invertebrate fauna and serves as a baseline for future studies of interactions between microbialites and fauna.     

Phylogeny of Carpinus and subfamily Coryloideae (Betulaceae) based on chloroplast and nuclear ribosomal sequence data

Phylogenetic studies were conducted for Carpinus and the subfamily Coryloideae (Betulaceae) using sequences of the chloroplast matK gene, the trnL-trnF region (trnL intron, and trnL [UAA] 3' exon-trnF [GAA] intergenic spacer) and the psbA-trnH intergenic spacer, and the nuclear ribosomal ITS regions. The combined analyses of the three chloroplast regions suggest that Coryloideae is monophyletic; Ostryopsis is sister to the Carpinus - Ostrya clade; Corylus is monophyletic and sister to the Ostrya - Carpinus - Ostryopsis clade; Ostrya is paraphyletic; and within Carpinus, species of sect. Carpinus from eastern Asia form a monophyletic group, whereas the positions of C. betulus from Europe and C. caroliniana from eastern North America are unresolved within the Carpinus clade. The cpDNA tree generated in this study is largely congruent with the previously published ITS results, but the ITS tree places Carpinus sect. Distegocarpus as sister to the Ostrya - Carpinus sect. Carpinus clade. Future work is needed to examine the relationships within the Ostrya - Carpinus clade, evaluate the generic status of Ostrya, and test the phylogenetic position of Ostryopsis.