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PubMed1996

Phylogenetic analysis of apiaceae subfamily apioideae using nucleotide sequences from the chloroplast rpoC1 intron.

Phylogenetic relationships among 25 members of Apiaceae (Umbelliferae) subfamily Apioideae, representing 7 of the 8 tribes and 8 of the 10 subtribes traditionally recognized in the subfamily, and 5 outgroups from Apiaceae subfamilies Hydrocotyloideae and Saniculoideae and allied families Araliaceae and Pittosporaceae have been inferred from nucleotide sequence variation in the intron of the chloroplast gene RNA polymerase C1 (rpoC1). Sequence divergence values in pairwise comparisons of unambiguous positions among all taxa ranged from 0 to 11.3% of nucleotides and averaged 3.8%. Trees derived from rpoC1 intron sequences estimated using maximum parsimony or maximum likelihood methods are of essentially similar topology, and indicate that (1) subfamily Apioideae, with Heteromorpha as its most basal element, is monophyletic and is a sister-group to Eryngium, the only representative examined of Apiaceae subfamily Saniculoideae, (2) Aralia (Araliaceae) arises from within a paraphyletic Apiaceae subfamily Hydrocotyloideae (represented by Centella and Hydrocotyle) and this clade is a sister-group to Apioideae+Saniculoideae, (3) there is a major phylogenetic division within Apioideae (excluding the basal Heteromorpha), with one clade comprising the genus Smyrnium and those taxa traditionally grouped in tribes Dauceae, Scandiceae, and Laserpitieae, and the other clade comprising all other examined taxa, and (4) relationships within each of these 2 major clades of Apioideae are largely equivocal owing to the low levels of nucleotide sequence divergence observed. Although rpoC1 intron sequences can provide valuable characters for addressing phylogenetic relationships among the outgroups and distantly related members of Apioideae, they have little power for resolving relationships among closely related taxa.

Base Sequence↗

PubMed1996

Phylogenetic origins of the asexual mycorrhizal symbiont Cenococcum geophilum Fr. and other mycorrhizal fungi among the ascomycetes.

The phylogenetic relationship of the asexual mycorrhizal fungus Cenococcum geophilam Fr. among sexual ascomycetes was examined by phylogenetic analysis of nucleotide sequence data from the nuclear small subunit (18S) ribosomal RNA genie region. A specific focus of this study was to test the hypothesis that the genus Elaphomyces is the closest sexual relative of C. geophilum. Thus nucleotide sequence data of five C. geophilum isolates, three Elaphomyces species, and 44 additional genera of ascomycetes were included in the phylogenetic analyses. The percentage of similarity among the 18S rDNA sequences of the C. geophilum isolates examined was 99.8 to 100%, indicating that C. geophilum is monophyletic. Percent similarity of nucleotide sequence among the three Elaphomyces species was also high and ranged from 99.4 to 99.5%. DNA parsimony and distance analysis of the sequence data separated these 2 genera on distant clades when sequence from 44 additional genera of ascomycetes was included. Parsimony and distance analyses positioned C. geophilum as a basal, intermediate lineage between the two Loculoascomycete orders, the Pleosporales and the Dothidiales, and strongly supported Elaphomyces to be of Plectomycete origin. Among the sexual Ascomycetes examined, which included representative taxa from four classes of filamentous Ascomycetes (Plectomycetes, Pyrenomycetes, Discomycetes, and Loculoascomycetes), no close sexual relative to C. geophilum was identified. At least four independent lineages of mycorrhizal fungi were identified among the ascomycetes examined.

Ascomycota↗

PubMed1997

Molecular phylogenetic study of Tephritidae (Insecta: Diptera) using partial sequences of the mitochondrial 16S ribosomal DNA.

We analyzed sequence data from 925 base pairs within the mitochondrial 16S ribosomal DNA from 34 tephritid species and two outgroup taxa. DNA sequences were amplified by the polymerase chain reaction and sequenced by the dideoxy chain-termination method. Patterns of nucleotide composition and substitution were assessed based on sequence comparison. Phylogenetic relationships among the included taxa were inferred using neighbor-joining and maximum parsimony methods. The trees were congruent with the morphological classification and suggested the following phylogenetic relationships: (1) monophyly of the tribe Trypetini, s. str., excluding Parastenopa; (2) a possible sister group relationship between Platyparea and the subfamily Tephritinae; (3) a sister group relationship between Hexachaeta and the tribe Toxotrypanini; (4) monophyly of the Dacinae and possible relationship of Oedicarena to this subfamily; (5) monophyly of the tribe Adramini; and (6) the nonmonophyletic nature of the genus Rhagoletis. We propose that the 16S rDNA shows great promise for helping to create a phylogenetically based higher classification of the Tephritidae.

Animals↗

PubMed1997

Re: Congruence and phylogenetic reanalysis of perching bird cytochrome b sequences.

In a study of the phylogenetic relationships among perching bird mtDNA sequences, Edwards et al. (1991; Proc. R. Soc. London Ser B. 243: 99-107) sequenced part of the mitochondrial cytochrome b gene from 13 perching birds (Passeriformes) and a woodpecker outgroup. However, recently the validity of part of the sequence of the hermit thrush (Catharus guttatus) in that study has been questioned. To determine the effect of inclusion of this apparently chimeric sequence on the conclusions of the original analysis, we reanalyzed these sequences using a different published hermit thrush sequence. In addition, we applied tests of congruence to examine the possibility that the aberrant phylogenetic behavior of chimeric mtDNA sequences might be detected. The reanalysis confirms the ability of slow evolving first and second codon positions of cytochrome b sequences to resolve deep branches in the avian tree. The fact that the new data set does not reject the DNA hybridization tree of these species probably indicates poor ability of the cytochrome b sequences to discriminate among alternative trees, rather than consistency among data sets. Statistical testing of trees based on individual amplified segments of mtDNA indicates that congruence tests may be one useful way of identifying chimeric mtDNA sequences when they have not been detected in the laboratory or during standard phylogenetic analysis.

Animals↗

PubMed1997

Phylogenetic relationships of cottontails (Sylvilagus, Lagomorpha): congruence of 12S rDNA and cytogenetic data.

The genus Sylvilagus, which comprises the New World cottontail rabbits, contains several commercially important as well as endangered (or threatened) species. Understanding the evolution of this group is pertinent to their management and conservation. The purpose of this study was to examine the evolutionary history of the cottontails using sequence data from the mitochondrial 12S rRNA gene. The 12S data provide a robust phylogeny which was supported under a variety of phylogenetic approaches and transition/transversion (Ti/Tv) weighting schemes. Stem and loop regions of the gene were analyzed separately and two different methods of estimating Ti/Tv ratios were employed. The phylogeny obtained was consistent with available cytogenetic information. The 12S data indicate that separate generic status for the pygmy rabbit, Brachylagus idahoensis, is warranted based on its phylogenetic position and sequence divergence values. Additionally, the taxa which are geographically adjacent are also phylogenetically closely related; for example, the marsh rabbit, S. palustris, and the swamp rabbit, S. aquaticus, are sister taxa, as are the mountain cottontail, S. nuttallii, and desert cottontail, S. audubonii. This finding suggests that recent vicariance events might explain the diversification of several cottontail lineages.

Animals↗

PubMed1997

Consistency, characters, and the likelihood of correct phylogenetic inference.

Computer simulations of character-state evolution in 8, 16, 32, and 64 ingroup taxa with a known set of relationships demonstrate that the maximum probability of correct phylogenetic inference increases with the number of variable (or informative) characters and their consistency index and decreases with the number of taxa, when the consistency index has been standardized to eliminate its dependence on the number of taxa. Equations for the probability of correct phylogenetic inference and for the standardized consistency indices (including or excluding autapomorphies) are derived. Given that actual studies based on DNA restriction sites and sequences generate more characters with a higher level of consistency than comparable studies based on morphology, calculations suggest that such molecular studies may often provide a more precise guide to phylogenetic relationships.

Computer Simulation↗

PubMed1998

Phylogenetic relationships of toads in the Bufo bufo species group from the eastern escarpment of the Tibetan Plateau: a case of vicariance and dispersal.

Phylogenetic relationships among Tibetan populations of the Bufo bufo species group are investigated using 1063 bases of mitochondrial DNA sequence from the genes encoding ND1 (subunit one of NADH dehydrogenase), tRNA(Ile), tRNA(Gln), tRNA(Met), and ND2. The aligned sequences contain 181 phylogenetically informative characters across all taxa sampled. Two hypotheses for colonization of the Tibetan Plateau are tested. A vicariant hypothesis predicts monophyly of populations from high elevations. A dispersalist hypothesis predicts monophyly of populations in each of two river drainages (Yangtze and Yellow rivers), which requires nonmonophyly of populations from high elevations. Both hypotheses are rejected in favor of a third hypothesis that combines elements of vicariance and dispersal. The most parsimonious phylogenetic tree places the high-elevation species, B. andrewsi, as the sister taxon to the other Asian Bufo populations; these high-elevation populations are postulated to have had a vicariant origin approximately 5 million years before present. The high-elevation population recognized as B. minshanicus is nested within low-elevation populations of B. gargarizans and is suggested to have dispersed onto the Tibetan Plateau more recently.

Animals↗

PubMed1998

Cryptic species in a "living fossil" lineage: taxonomic and phylogenetic relationships within the genus Lepidurus (Crustacea: Notostraca) in North America.

Lineages which exhibit little morphological change over geologic time are evolutionarily and ecologically interesting, but often taxonomically difficult. For some morphologically conservative groups, not only is it almost impossible to identify significant changes in fossil forms over time, but the relationships among extant populations are often poorly understood due to lack of known characters which clearly delimit species. Notostracan crustaceans are a classic example of such "living fossils." The paradoxical characteristics of long-term stasis in gross morphology and hypervariability of many individual morphological characters make notostracans an especially taxonomically challenging group. We used molecular and biochemical techniques to investigate the taxonomic and phylogenetic relationships of four nominal species within the genus Lepidurus in North America, three of which had been alternately abandoned, resurrected, or synonymized under a single, globally distributed morphospecies since their original descriptions in the 1800s. Data from a 330-bp sequence of the mitochondrial 12S rDNA gene and from nine allozyme loci consistently indicate five highly genetically divergent clades among the populations we used to represent the four nominal species. Diagnostic molecular characters, magnitudes of genetic divergence among clades, and the fact that these genetically distinct clades have broadly overlapping geographical ranges strongly suggest that the five clades are reproductively isolated species. One of the nominal species (L. couesii) is not monophyletic, but rather consists of two species which are not sister taxa. The other three nominal species (L. lemmoni, L. packardi, and L. bilobatus) are supported as valid phylogenetic species. The best current hypothesis for phylogenetic relationships among the five species is provided by a simultaneous analysis of both 12S rDNA and allozyme data, which places L. bilobatus and L. "couesii"-1 as sister taxa and L. "couesii"-2 as the most basal of all the Lepidurus species included in this study. These results point to the existence of cryptic species within the current classification scheme for Lepidurus, the need for further taxonomic work within the Notostraca in general, and the role that genetic techniques can play in clarifying the systematics of morphologically conservative groups.

Animals↗

PubMed1998

Phylogenetic relationships of terrestrial Australo-Papuan elapid snakes (subfamily Hydrophiinae) based on cytochrome b and 16S rRNA sequences.

Phylogenetic relationships among the venomous Australo-Papuan elapid snake radiation remain poorly resolved, despite the application of diverse data sets. To examine phylogenetic relationships among this enigmatic group, portions of the cytochrome b and 16S rRNA mitochondrial DNA genes were sequenced from 19 of the 20 terrestrial Australian genera and 6 of the 7 terrestrial Melanesian genera, plus a sea krait (Laticauda) and a true sea snake (Hydrelaps). These data clarify several significant issues in elapid phylogeny. First, Melanesian elapids form sister groups to Australian species, indicating that the ancestors of the Australian radiation came via Asia, rather than representing a relict Gondwanan radiation. Second, the two major groups of sea snakes (sea kraits and true sea snakes) represent independent invasions of the marine environment. Third, the radiation of viviparous Australian elapids is much older than has been suggested from immunological data. Parsimony analyses were unable to resolve relationships among the Australian radiation, a problem previously encountered with analyses of other (morphological, electrophoretic, karyotypic, immunological) data sets on these species. These data suggest that the reason for this continued difficulty lies in the timing of speciation events: the elapids apparently underwent a spectacular adaptive radiation soon after reaching Australia, such that divergences are ancient even within genera. Indeed, intrageneric divergences are almost as large as intergeneric divergences. Although this timing means that our sequence data cannot fully resolve phylogenetic relationships among the Australian elapids, the data suggest a close relationship of the following clades: Pseudonaja with Oxyuranus; Ogmodon with Toxicocalamus; Demansia with Aspidomorphus; Echiopsis with Denisonia; the "Notechis" lineage with Drysdalia coronoides; and Rhinoplocephalus and Suta with Drysdalia coronata. At least two of the Australian genera (Drysdalia and Simoselaps) appear to be paraphyletic. These sequence data support many of the conclusions reached by earlier studies using other types of data, but additional information will be needed before the phylogeny of the Australian elapids can be fully resolved.

Animals↗

PubMed1998

Molecular phylogenetics at the Felsenstein zone: approaching the Strepsiptera problem using 5.8S and 28S rDNA sequences.

Recent efforts to reconstruct the phylogenetic position of the insect order Strepsiptera have elicited a major controversy in molecular phylogenetics. We sequenced the 5.8S rDNA and major parts of the 28S rDNA 5' region of the strepsipteran species Stylops melittae. Their evolutionary dynamics were analyzed together with previously published insect rDNA sequences to identify tree estimation bias risks and to explore additional sources of phylogenetic information. Several major secondary structure changes were found as being autapomorphic for the Diptera, the Strepsiptera, or the Archaeognatha. Besides elevated substitution rates a significant AT bias was present in dipteran and strepsipteran 28S rDNA which, however, was restricted to stem sites in the Diptera while also affecting single-stranded sites in the Strepsiptera. When dipteran taxa were excluded from tree estimation all methods consistently supported the placement of Strepsiptera to within the Holometabola. When dipteran taxa were included maximum likelihood continued to favor a sister-group relationship of Strepsiptera with Mecoptera while remaining methods strongly supported a sister-group relationship with Diptera. Parametric bootstrap analysis revealed maximum likelihood as a consistent estimator if rate heterogeneity across sites was taken into account. Though the position of Strepsiptera within Holometabola remains elusive, we conclude that the evolution of dipteran and strepsipteran rDNA involved similar yet independent changes of substitution parameters.

Animals↗

PubMed1998

Base compositional bias and phylogenetic analyses: a test of the "flying DNA" hypothesis.

Phylogenetic methods can produce biased estimates of phylogeny when base composition varies along different lineages. Pettigrew (1994, Curr. Biol. 4:277-280) has suggested that base composition bias is responsible for the apparent support for the monophyly of bats (Chiroptera: megabats and microbats) from several different nuclear and mitochondrial genes. Pettigrew's "flying DNA" hypothesis makes several predictions: (1) that metabolic constraints associated with flying result in elevated levels of adenine and thymine throughout the genome of both megabats and microbats, (2) that the resulting base compositional bias in bats is sufficient to mislead phylogenetic methods and account for the support for bat monophyly from several nuclear and mitochondrial genes, and (3) that phylogenetic analysis using pairwise distances corrected for compositional bias should eliminate the support for bat monophyly. We tested these predictions by analyzing DNA sequences from two nuclear and three mitochondrial genes. The predicted base compositional bias does not appear to exist in some of the genes, and in other genes the differences in AT content are very small. Analyses under a wide diversity of criteria and models of evolution, including analyses that take base composition into account (using log-determinant distances), all strongly support bat monophyly. Moreover, simulation analyses indicate that even extreme bias toward AT-base composition in bats would be insufficient to explain the observed levels of support for bat monophyly. These analyses provide no support for the "flying DNA" hypothesis, whereas the monophyly of bats appears to be well supported by the DNA sequence data.

Animals↗

PubMed1998

Phylogenetic relations among percid fishes as inferred from mitochondrial cytochrome b DNA sequence data.

Hypotheses of relationship among genera of Percidae have been conflicting. Based on different phylogenetic premises, the evolution of small benthic forms in Percidae has been interpreted as resulting from either convergence or common ancestry. In order to assess various phylogenetic hypotheses of Percidae we collected complete sequences (1140 bp) of mitochondrially encoded cytochrome b for 21 species of percids. Seven species representing four additional families of Perciformes were used as outgroups. Maximum parsimony and minimum evolution analyses both recovered single shortest trees, and the results of these analyses were generally congruent with one another. All analyses consistently recovered three monophyletic groups in Percidae: Etheostomatinae (Ammocrypta, Crystallaria, Etheostoma, and Percina), Percinae (Perca and Gymnocephalus), and Luciopercinae (Stizostedion, Zingel, and Romanichthys). As a result of this analysis we present a revised classification of Percidae and discuss the phylogenetic evidence for the independent evolution of small benthic species within Etheostomatinae and Luciopercinae.

Animals↗

PubMed1999

The major opsin in bees (Insecta: Hymenoptera): A promising nuclear gene for higher level phylogenetics.

We report the phylogenetic utility of the nuclear gene encoding the long-wavelength opsin (LW Rh) for tribes of bees. Aligned nucleotide sequences were examined in multiple taxa from the four tribes comprising the corbiculate bees within the subfamily Apinae. Phylogenetic analyses of sequence variation in a 502-bp fragment (approx 40% of the coding region) strongly supported the monophyly of each of the four tribes, which are well established from previous studies of morphology and DNA. Trees estimated from parsimony and maximum likelihood analyses of LW Rh sequences show a strongly supported relationship between the tribes Meliponini and Bombini, a relationship that has been found uniformly in studies of other genes (28S, 16S, and cytochrome b). All of the tribal clades as well as relationships among the tribes are supported by high bootstrap values, suggesting the utility of LW Rh in estimating tribal and subfamily rank for these bees. The sequences exhibit minimal base composition bias. Both 1st + 2nd and 3rd position sites provide information for estimating a reliable tree topology. These results suggest that LW Rh, which has not been reported previously in studies of organismal phylogenetics, could provide important new data from the nuclear genome for phylogeny reconstruction.

Animals↗

PubMed1999

Chloroplast-expressed glutamine synthetase (ncpGS): potential utility for phylogenetic studies with an example from Oxalis (Oxalidaceae).

Chloroplast-expressed glutamine synthetase (ncpGS), a nuclear-encoded gene containing several introns, is introduced as a tool for phylogenetic studies at lower taxonomic levels. This gene is a member of a multigene family, but it diverged long ago from the cytosolic-expressed members of the family and appears to be single copy in the majority of taxa examined to date. The conservation of both coding sequence and position of introns has allowed the design of primers for use in a broad range of dicot taxa to amplify and sequence a region of ncpGS that contains four introns. The utility of this region in phylogenetic studies of congeneric species is illustrated by an example using eight Oxalis species. The four introns in these taxa are typical in size (76 to 136 bp), base composition (high T content), and structure (e.g., sequence of splice sites and putative branch points) for plant internal introns. Levels of variation among these ncpGS sequences compare favorably with those of the internal transcribed spacer of nuclear ribosomal DNA (ITS) from the same taxa, and results of phylogenetic analysis of ncpGS data are generally congruent with previous results using ITS.

Base Sequence↗

PubMed1999

Molecular phylogenetics and the evolution of labral spines among eastern Pacific ocenebrine gastropods.

Labral spines are sharp projections of the apertural lip found in some marine gastropods that are used to penetrate hard-shelled prey. The majority of gastropod genera that contain labral spine-bearing species are found in the subfamily Ocenebrinae (Gastropoda: Muricidae). To reconstruct the evolutionary history of labral spine-bearing and labral spine-lacking gastropods in the eastern Pacific (EP) Ocean, partial sequences of two mitochondrial genes (cytochrome oxidase I and 12S rRNA) were obtained from representative taxa. Despite high nucleotide bias, a variety of phylogenetic reconstruction methods produced the same tree topology. The traditional taxonomic view that all "Nucella-like" spine-bearing taxa in the EP belong to a monophyletic "Acanthina" is rejected due to nonmonophyly of this group. The more recently recognized "Acanthinucella" is also not monophyletic, and we therefore propose the new genus Mexacanthina for two Mexican species formerly assigned to Acanthinucella. The genus Ocinebrina, which first appears in the middle Eocene, is not a stem EP ocenebrine lineage and may also not be a monophyletic clade. Tracing the evolutionary history of labral spines among extant lineages indicates that the absence of a labral spine is ancestral for all EP ocenebrines. Ancestral conditions could not be resolved unambiguously for all nodes of the phylogeny based on extant taxa. However, by jointly considering both molecular phylogenetic relationships and the phylogenetic affinities of several extinct taxa, all remaining character state transformation can be inferred unambiguously. Based on this analysis, a labral spine likely evolved independently in at least four lineages of EP ocenebrines. Although homoplasy appears to characterize labral spine evolution among ocenebrine gastropods, the structural position of a labral spine was evolutionarily altered in one lineage, indicating that different types of labral spines do not necessarily reflect convergent evolution.

Animals↗

PubMed2000

Phylogenetic analysis of molluscan mitochondrial LSU rDNA sequences and secondary structures.

Mollusks are an extraordinarily diverse group of animals with an estimated 200,000 species, second only to the phylum Arthropoda. We conducted a comparative analysis of complete mitochondrial ribosomal large subunit sequences (LSU) of a chiton, two bivalves, six gastropods, and a cephalopod. In addition, we determined secondary structure models for each of them. Comparative analyses of nucleotide variation revealed substantial length variation among the taxa, with stylommatophoran gastropods possessing the shortest lengths. Phylogenetic analyses of the nucleotide sequence data supported the monophyly of Albinaria, Euhadra herklotsi + Cepaea nemoralis, Stylommatophora, Cerithioidea, and when only transversions are included, the Bivalvia. The phylogenetic limits of the mitochondrial LSU rRNA gene within mollusks appear to be up to 400 million years, although this estimate will have to be tested further with additional taxa. Our most novel finding was the discovery of phylogenetic signal in the secondary structure of rRNA of mollusks. The absence of entire stem/loop structures in Domains II, III, and V can be viewed as three shared derived characters uniting the stylommatophoran gastropods. The absence of the aforementioned stem/loop structure explains much of the observed length variation of the mitochondrial LSU rRNA found within mollusks. The distribution of these unique secondary structure characters within mollusks should be examined.

Animals↗

PubMed2000

Phylogenetic relationships and climatic adaptations in the Drosophila takahashii and montium species subgroups.

We analyze phylogenetic relationships among temperate, subtropical highland, and subtropical lowland species of the Drosophila takahashii and montium species subgroups based on sequence data of COI and Gpdh genes and discuss the evolution of temperate species in these subgroups with reference to their climatic adaptations. In the takahashii subgroup, D. lutescens (the temperate species) branched off first in the tree based on the combined data set, but D. prostipennis (the subtropical highland species) branched off first in the trees based on single genes. Thus, phylogenetic relationships in this subgroup are still ambiguous. In the montium subgroup, the cool-temperate species are phylogenetically close to the warm-temperate species, and these cool- and warm-temperate species form a cluster with the subtropical highland species. This suggests that perhaps the cool-temperate species derived from the warm-temperate species and the warm-temperate species derived from the subtropical highland species. In comparison with the subtropical lowland species, the subtropical highland species may be better able to colonize temperate areas since, as in the temperate species, they have an ability to develop their ovaries at moderately low temperature. However, the subtropical highland species, as well as the subtropical lowland species, were much less cold tolerant than the temperate species. Therefore, considerable genetic reformation would be required for both the subtropical highland and the subtropical lowland species to adapt to temperate climates.

Acclimatization↗

PubMed2000

Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida).

The phylogenetic relationships of the Clitellata were investigated using a data set with published and new complete or partial 18S rRNA and mtCOI gene sequences of 13 and 49 taxa representing 8 and 14 families, respectively. Three different alignments were considered for 18S, and the possible influence of departures from rate constancy among sites was evaluated by analyses using a Gamma model of rate heterogeneity. Maximum-likelihood estimates of the shape parameter alpha of the Gamma distribution were very low, whatever the alignment or the gene considered, suggesting that phylogenetic reconstructions taking into account the rate heterogeneity among sites are likely to be the most reliable. Analyzed separately, the two genes did not resolve the relationships among the Clitellata, but the consensus tree was congruent with the morphology-based relationships. Our data suggest the inclusion of the Euhirudinea, Acanthobdellida, and Branchiobdellida in the Oligochaeta and suggest the Lumbriculidae as the link between both assemblages. Although separate analyses of both genes, as well as different alignments for the 18S rRNA sequences, yielded conflicting results concerning the phylogenetic position of leeches and leech-like worms vis-à-vis the Oligochaeta, subsequent analyses using the Gamma model greatly reduced the observed inconsistencies. Our analyses show that among the Clitellata, the leeches and the leech-like and gutless worms represent significantly faster evolving lineages. It is suggested that the observed higher mutation rates may be explained by the fact that these lineages contain almost exclusively commensal and/or parasitic taxa.

Animals↗