Shared nucleotide composition biases among species and their impact on phylogenetic reconstructions of the Drosophilidae

Compositional changes are a major feature of genome evolution. Overlooking nucleotide composition differences among sequences can seriously mislead phylogenetic reconstructions. Large compositional variation exists among the members of the family Drosophilidae. Until now, however, base composition differences have been largely neglected in the formulations of the nucleotide substitution process used to reconstruct the phylogeny of this important group of species. The present study adopts a maximum-likelihood framework of phylogenetic inference in order to analyze five nuclear gene regions and shows that (1) the pattern of compositional variation in the Drosophilidae does not match the phylogeny of the species; (2) accounting for the heterogeneous GC content with Galtier and Gouy's nucleotide substitution model leads to a tree that differs in significant aspects from the tree inferred when the nucleotide composition differences are ignored, even though both phylogenetic hypotheses attain strong nodal support in the bootstrap analyses; and (3) the LogDet distance correction cannot completely overcome the distorting effects of the compositional variation that exists among the species of the Drosophilidae. Our analyses confidently place the Chymomyza genus as an outgroup closer than the genus Scaptodrosophila to the Drosophila genus and conclusively support the monophyly of the Sophophora subgenus.     

Phylogeny of the Saprininae reveals interesting ecological shifts in the history of the subfamily (Coleoptera: Histeridae)

A morphological data set for the histerid beetle subfamily Saprininae comprising 95 adult morphological characters scored (multistate coding) from 72 terminal taxa and four outgroups was developed in order to analyse and determine the relationships amongst the genera and subgenera of the Saprininae subfamily. Cladograms were rooted with exemplars of Dendrophilinae (genus Dendrophilus), Bacaniini (genus Bacanius), Abraeinae (genus Chaetabraeus), and Anapleini (genus Anapleus). Parsimony‐based phylogenetic analyses were performed based on the type species of each genus and subgenus of the Saprininae occurring around the world, with the exception of three taxa: Paramyrmetes foveipennis (type species of the genus Paramyrmetes), Satrapister nitens (type species of the genus Satrapister) and Xerosaprinus (Auchmosaprinus) laciniatus (type species of the subgenus Auchmosaprinus) that were not available. In addition, in order to test the monophyly of several questionable genera, multiple exemplars were added in a few cases. The analysis also included an exemplar of an apparently undescribed genus. The results of the analysis confirm the monophyly of the subfamily supported by two unique synapomorphies: (1) presence of sensory structures of the antenna; and (2) presence of the antennal cavity, as well as several other weaker synapomorphies. However, the phylogeny inferred here shows mostly low support for the deeper branches and consequently no major changes in the Saprininae classification are proposed. The presented cladogram is discussed together with its implications for the evolution of the subfamily. The most informative characters and their respective states are outlined. Multiple shifts in lifestyles have evolved during the evolutionary history of the group. Taxa found near the root of the cladogram are mostly nidicolous or myrmecophilous, and inquiliny is presumed to be the plesiomorphic lifestyle of the subfamily. The nidicolous lifestyle has undergone several transformations to other lifestyles and myrmecophily has evolved three times independently during the evolution of the subfamily. Termitoxeny has evolved two times independently in the group whereas ecological adaptation for life in caves has likewise evolved two times independently. The analyses yielded a large clade of predominantly psammophilous taxa; psammophily is thought to have evolved once and has been subsequently lost several times. © 2014 The Linnean Society of London     

基于线粒体COI基因的大蚊属Tipula系统发育及长角大蚊亚属Tipula (Sivatipula)分类地位初步研究(双翅目:大蚊总科)

大蚊属Tipula Linnaeus,1758是大蚊科中种类最多的属,目前其单系性尚未得到全面验证.此外,长角大蚊亚属Tipula (Sivatipula) Alexander,1964因其极长的触角以及独有的精子泵结构,明显不同于大蚊属其他亚属,使其亚属的分类地位存在争议.本研究基于COI序列对19个大蚊属物种及5个其他属物种进行了系统发育分析,并计算了物种间的遗传距离.研究结果表明:(1)邻接树(NJ)和最大似然树(ML)均显示长角大蚊亚属与大蚊属其他亚属未形成单系,大蚊属的单系性没有得到支持;(2)基于遗传距离和系统发育分析并结合形态信息,结果显示长角大蚊亚属独立于大蚊属内其他亚属,应将其提升为属级分类单元.     

Phylogenetic relationships in the genus mus,based on paternally,maternally, and biparentally inherited characters

Several species in the rodent genus Mus are used as model research organisms, but comparative studies of these mice have been hampered by the lack of a well-supported phylogeny. We used DNA sequences from six genes representing paternally, maternally, and biparentally inherited regions of the genome to infer phylogenetic relationships among 10 species of Mus commonly used in laboratory research. Our sample included seven species from the subgenus Mus; one species each from the subgenera Pyromys, Coelomys, and Nannomys; and representatives from three additional murine genera, which served as outgroups in the phylogenetic analyses. Although each of the six genes yielded a unique phylogeny, several clades were supported by four or more gene trees. Nodes that conflicted between trees were generally characterized by weak support for one or both of the alternative topologies, thus providing no compelling evidence that any individual gene, or part of the genome, was misleading with respect to the evolutionary history of these mice. Analysis of the combined data resulted in a fully resolved tree that strongly supports monophyly of the genus Mus, monophyly of the subgenus Mus, division of the subgenus Mus into Palearctic (M. musculus, M. macedonicus, M. spicilegus, and M. spretus) and Asian (M. cervicolor, M. cookii, and M. caroli) clades, monophyly of the house mice (M. m. musculus, "M. m. molossinus," M. m. castaneus, and M. m. domesticus), and a sister-group relationship between M. macedonicus and M. spicilegus. Other clades that were strongly supported by one or more gene partitions were not strongly supported by the combined data. This appears to reflect a localized homoplasy in one partition obscuring the phylogenetic signal from another, rather than differences in gene or genome histories.     

Molecular phylogeny of the subgenus Drosophila (Diptera, Drosophilidae) with an emphasis on Neotropical species and groups: a nuclear versus mitochondrial gene approach

The genus Drosophila has played an essential role in many biological studies during the last 100 years but much controversy and many incompletely addressed issues still remain to be elucidated regarding the phylogeny of this genus. Because information on the Neotropical species contained in the subgenus Drosophila is particularly incomplete, with this taxonomic group being underrepresented in many studies, we designed a study to answer some evolutionary questions related to these species. We subjected at least 41 Drosophilidae taxa to a phylogenetic analysis using a 516-base pair (bp) fragment of the alpha-methyldopa (Amd) nuclear gene and a 672 bp fragment of the mitochondrial cytochrome oxidase subunit II (COII) gene both individually and in combination. We found that the subgenus Drosophila is paraphyletic and subdivided into two main clusters: the first containing species traditionally placed in the virilis-repleta radiation and the second assembling species of the immigrans-Hirtodrosophila radiation. Inside the first of these clusters we could detect the monophyly of both the flavopilosa (the sister-clade of the annulimana group) and the mesophragmatica (closely related to the repleta group) species groups. Concerning the immigrans-Hirtodrosophila lineage, Zaprionus, Liodrosophila, Samoaia, and Hirtodrosophila were the early offshoots, followed by the immigrans, quinaria, testacea, and funebris species groups. The tripunctata radiation appears to be a derived clade, composed of a paraphyletic tripunctata group, intimately interposed with members of the cardini, guarani, and guaramunu species groups. Overall, the COII gene yielded a poor phylogenetic performance when compared to the Amd gene, the evolutionary hypothesis of which agreed with the total evidence tree. This phenomenon can be explained by the fast saturation of transitional substitutions in COII, due to strong biases in both base composition and substitution patterns, as also by its great among-site rate variation heterogeneity.     

Phylogenetic classification of the Drosophilidae Rondani (Diptera): the role of morphology in the postgenomic era

Molecular sequences now overwhelm morphology in phylogenetic inference. Nonetheless, most molecular studies are conducted on a limited number of taxa, as DNA rarely can be analysed from old museum types or fossils. During the last 20 years, more than 150 molecular studies have challenged the current phylogenetic classification of the family Drosophilidae Rondani based on morphological characters. Most studies concerned a single genus, Drosophila Fallén, and included only few representative species from 17 out of the 78 genera of the family. Therefore, these molecular studies were unable to provide an alternative classification scheme. A supermatrix analysis of seven nuclear and one mitochondrial genes (8248 bp) for 33 genera was conducted using outgroups from one calyptrate and four ephydroid families. The Bayesian phylogeny was consistent with previous molecular studies including whole genome sequences and divided the Drosophilidae into four monophyletic clades. Morphological characters, mostly male genitalia, then were compared thoroughly between the four clades and homologous character states were identified. These states were then checked for 70 genera and a revised phylogenetic, family‐group classification for the Drosophilidae is proposed. Two genera –Cladochaeta Coquillett and Diathoneura Duda – of the tribe Cladochaetini Grimaldi are transferred to the family Ephydridae. The Drosophilidae is divided into two subfamilies: Steganinae Hendel (30 genera) and Drosophilinae Rondani (43 genera). A further two genera, Apacrochaeta Duda and Sphyrnoceps de Meijere, are incertae sedis, and Palmophila Grimaldi, is synonymized with Drosophila syn.n. The Drosophilinae is subdivided into two tribes: the re‐elevated Colocasiomyini Okada (nine genera) and Drosophilini Okada. The paraphyly of the genus Drosophila was not resolved to avoid affecting the binomina of important laboratory model species; however, its subgeneric classification was revised in light of molecular and morphological data. Three subgenera, namely Chusqueophila Brncic, Phloridosa Sturtevant and Psilodorha Okada, were synonymized with the subgenus Drosophila (Drosophila) Fallén syns.n. Among the 45 species groups and 5 species complexes of Drosophila (Drosophila), 22 groups and 1 complex were transferred to the subgenus Drosophila (Siphlodora) Patterson & Mainland and 6 groups, 2 species subgroups and 3 complexes are considered incertae sedis within the genus Drosophila. Different morphological characters provide different signals at different phylogenetic scales: thoracic characters (wing venation and presternal shape) discriminate families; grasping and erection‐related characters discriminate subfamilies to tribes; whereas phallic paraphyses, i.e. auxiliary intromittent organs, discriminate genera and Drosophila subgenera. The study shows the necessity of analysing morphological characters within a molecular phylogenetic framework to translate molecular phylogenies into taxonomically‐comprehensive classifications.     

A cladistic analysis of Geranium subgen. Erodioidea (Picard) Yeo (Geraniaceae)

Subgenus Erodioidea , comprising 19 species, is diagnosed exclusively by the type of fruit discharge and associated morphological characters, all of which are of widespread occurrence within the Geraniaceae. We have no evidence to suggest that the three included sections ( Erodioidea, Aculeolata and Subacaulia) form a monophyletic group. Cladistic analysis of a data set containing 30 morphological characters has confirmed our doubts of monophyly of the subgenus by placing part of the outgroup within the ingroup. Subsequent analyses of the two included non-monotypic sections, Erodioidea and Subacaulia , give different results. For both sections, the monophyly assumption is reasonably well-supported by characters. However, while analysis of sect. Erodioidea results in an explicit hypothesis of relationships and a fully resolved cladogram, resolution within sect. Subacaulia is low. For analysis of the latter, seven different outgroups were used and, in each one, the successive approximations weighting procedure (available in the program Hennig 86) was followed to improve resolution. Despite this, lower nodes in the cladogram remained unresolved, and strict consensus trees of all the analyses resulted in only a few replicated subterminal clades. These results are discussed in connection with biogeographical data and with the hypothesis that representatives of sect. Subacaulia are schizoendemics.     

Phylogenetic analysis of Phlebotomus species belonging to the subgenus Larroussius (Diptera, psychodidae) by ITS2 rDNA sequences

In the genealogy of Phlebotomus (Diptera: Psychodidae), morphological analyses have indicated that the subgenus Larroussius is a monophyletic group which is most closely related to the subgenera Transphlebotomus and Adlerius. We conducted a phylogenetic analysis of the relationships among six representative species of the subgenus Larroussius and one species representatitive of the Phlebotomus subgenus, assessing sequences of the Second Internal Transcribed Spacer (ITS2) of the ribosomal RNA (rRNA). Three of the species (P. perniciosus, P. ariasi and P. perfiliewi perfiliewi) were collected in different parts of the Mediterranean area. The trees estimated from parsimony and neighbour-joining analyses supported the monophyly of the Larroussius subgenus inferred from the morphological analysis. According to our data, P. ariasi may be a sister group to the rest of the Larroussius subgenus, although additional sequence data are needed to confirm this observation. Our results suggest that P. perniciosus and P. longicuspis are distinct species, in spite of the fact that there are only slight morphological differences. The strict congruence between the phylogeny of the Larroussius subgenus inferred from the ITS2 sequences and that based on morphological studies further confirmed the ability of the spacer sequence to identify recently-derived affiliations.     

Molecular phylogeny of the Korean Rivellia species (Diptera: Tephritoidea: Platystomatidae) based on 16S rDNA sequences: Testing morphological hypotheses using molecular data

The phylogeny of the genus Rivellia Robineau‐Desvoidy was inferred from mitochondrial 16S ribosomal (r)DNA gene sequences of 13 Korean Rivellia species and six species representing other platystomatid genera and the family Tephritidae. We compared the inferred molecular phylogeny with the previously published morphological cladogram. As a result, the following phylogenetic relationships were recognized: (i) monophyly of the genus Rivellia; (ii) monophyly of the R. syngenesiae species group; (iii) R. depicta and R. apicalis (which were not previously placed in any species group) were recognized as a sister group of the R. syngenesiae species group; and (iv) monophyly of the R. basilaris species group was recognized to a limited extent. These results, even though geographically limited, provide a new insight into the phylogeny of the genus Rivellia. They clearly show the utility of 16S rDNA for phylogenetic analysis of the genus Rivellia. Additional study involving samples from different geographical areas will be needed to gain a better understanding of the adaptive radiation of this species‐rich genus.     

Molecular and morphological phylogenetic analysis of an insular radiation in Pacific black flies (Simulium)

Ecological adaptation within islands may have figured prominently in the insular radiation of black flies (subgenus Inseliellum) in the Society Islands, French Polynesia. To aid in understanding the sequence of ecological shifts in this group, we have constructed a phylogeny by using morphology, the cytochrome oxidase I (COI) gene, and the small ribosomal subunit (12S) gene. The strong influence of COI on the combined analysis tree was evident from its contribution to the partitioned Bremer support (62%). The net effect of including 12S was to reduce overall tree support. Different character sets resolved different portions of the combined analysis tree, with COI resolving recent lineages, 12S resolving basal relationships, and morphology supporting the monophyly of taxa having smaller larval feeding fans (oviceps group). The Partition Homogeneity and Kashino-Hasegawa tests indicated significant incongruence between morphological and mitochondrial data. The Templeton test revealed that morphology and the combined (COI + 12S) mitochondrial data were incongruent. This conflict stems primarily from disagreement over the monophyly of taxa having much smaller larval feeding fans. Either convergence in a subset of morphological characters, low phylogenetic signal among mitochondrial sequences, or lineage-sorting causing the mitochondrial data to track an incorrect evolutionary history may be responsible for these results.     

Molecular phylogenetics of the genus Ceratitis (Diptera: Tephritidae)

The Afrotropical fruit fly genus Ceratitis MacLeay is an economically important group that comprises over 89 species, subdivided into six subgenera. Cladistic analyses of morphological and host use characters have produced several phylogenetic hypotheses for the genus. Only monophyly of the subgenera Pardalaspis and Ceratitis (sensu stricto) and polyphyly of the subgenus Ceratalaspis are common to all of these phylogenies. In this study, the hypotheses developed from morphological and host use characters are tested using gene trees produced from DNA sequence data of two mitochondrial genes (cytochrome oxidase I and NADH-dehydrogenase subunit 6) and a nuclear gene (period). Comparison of gene trees indicates the following relationships: the subgenus Pardalaspis is monophyletic, subsection A of the subgenus Pterandrus is monophyletic, the subgenus Pterandrus may be either paraphyletic or polyphyletic, the subgenus Ceratalaspis is polyphyletic, and the subgenus Ceratitis s. s. might not be monophyletic. In addition, the genera Ceratitis and Trirhithrum do not form reciprocally monophyletic clades in the gene trees. Although the data statistically reject monophyly for Trirhithrum under the Shimodaira-Hasegawa test, they do not reject monophyly of Ceratitis.     

Aphis (Hemiptera: Aphididae) species groups found in the Midwestern United States and their contribution to the phylogenetic knowledge of the genus

A phylogeny of the genus Aphis Linnaeus, 1 758 was built primarily from specimens collected in the Midwest of the United States. A data matrix was constructed with 68 species and 41 morphological characters with respective character states of alate and apterous viviparous females. Dendrogram topologies of analyses performed using UPGMA (Unweighted Pair Group Method with Arithmetic Mean), Maximum Parsimony and Bayesian analysis of Cytochrome Oxidase I, Elongation Factor 1‐α and primary endosymbiont Buchnera aphidicola 16S sequences were not congruent. Bayesian analysis strongly supported most terminal nodes of the phylogenetic trees. The phylogeny was strongly supported by EF1‐α, and analysis of COI and EF1‐α molecular data combined with morphological characters. It was not supported by single analysis of COI or Buchnera aphidicola 16S. Results from the Bayesian phylogeny show 4 main species groups: asclepiadis, fabae, gossypii, and middletonii. Results place Aphis and species of the genera Protaphis Börner, 1952, Toxoptera Koch, 1856 and Xerobion Nevsky, 1928 in a monophyletic clade. Morphological characters support this monophyly as well. The phylogeny shows that the monophyletic clade of the North American middletonii species group belong to the genus Protaphis: P. debilicornis (Gillette & Palmer, 1929 ), comb. nov., P. echinaceae (Lagos and Voegtlin, 2009 ), comb. nov., and P. middletonii (Thomas, 1879 ). The genus Toxoptera should be considered a subgenus of Aphis (stat. nov.). The analysis also indicates that the current genus Iowana Frison, 1954 should be considered a subgenus of Aphis (stat. nov.).     

Phylogeny of the Baldratiina (Diptera: Cecidomyiidae) inferred from morphological, ecological and molecular data sources, and evolutionary patterns in plant-galler relationships

The phylogeny of the gall-midge subtribe Baldratiina (Diptera: Cecidomyiidae) was reconstructed from molecular (partial sequence of the mitochondrial 12S rDNA), morphological and ecological data sets, using 16 representative species of most of the genera. The morphological and ecological data were combined in a single character matrix and analyzed separately from the molecular data, resulting in an eco-morphological cladogram and a molecular cladogram. Attributes of galls and host associations were superimposed on the molecular cladogram in order to detect possible trends in the evolution of these traits. The cladograms resulting from the two independent analyses were statistically incongruent, although both provide evidence for the monophyly of the genera Baldratia and Careopalpis and the paraphyly of the genera Stefaniola and Izeniola. The results suggest a minor impact of the morphological characters traditionally used in the classification of the Baldratiina, whereas ecological data had a major impact on the phylogenetic inference. Mapping of gall and host attributes on the molecular cladogram suggests that multi-chambered stem galls constitute the ancestral state in the subtribe, with several subsequent shifts to leaf galls. It is concluded that in contrast to other studied groups of gall insects, related baldratiine species induce different types of galls, attesting to speciation driven by gall-type shifts at least as often as host shifts.     

Phylogenetic relationships within the genus Jesogammarus (Crustacea, Amphipoda, Anisogammaridae) deduced from mitochondrial COI and 12S sequences

The genus Jesogammarus contains 16 species in two subgenera, Jesogammarus and Annanogammarus. To examine relationships among species in the genus, a molecular phylogenetic study including eight species of the former subgenus and four of the latter was conducted using partial DNA sequences of the mitochondrial COI and 12S rRNA genes. MP, NJ, and ML trees based on the combined COI and 12S data indicated monophyly of the subgenus Annanogammarus, though the monophyly of Jesogammarus was left unresolved. Consistent with few morphological differences, Jesogammarus (A.) naritai and J. (A.) suwaensis showed low genetic differentiation and did not show reciprocal monophyly, which suggests a close affinity of these taxa.     

Phylogeny of Geomydoecus and Thomomydoecus pocket gopher lice (Phthiraptera: Trichodectidae) inferred from cladistic analysis of adult and first instar morphology

The phylogeny for all 122 species and subspecies of chewing lice of the genera Geomydoecus and Thomomydoecus (Phthiraptera: Trichodectidae) hosted by pocket gophers (Rodentia: Geomyidae) is estimated by a cladistic analysis of fifty-eight morphological characters obtained from adults and first instars. The data set has considerable homoplasy, but still contains phylogenetic information. The phylogeny obtained is moderately resolved and, with some notable exceptions, supports the species complexes proposed by Hellenthal and Price over the the last two decades. The subgenera G. (Thaelerius) and T. (Thomomydoecus) are both shown to be monophyletic, but the monophly of subgenus T. (Jamespattonius) could not be confirmed, perhaps due to the lack of first-instar data for one of its component species. The nominate subgenus of Geomydoecus may be monophyletic, but our cladogram was insufficiently resolved to corroborate this. Mapping the pocket gopher hosts onto the phylogeny reveals a consistent pattern of louse clades being restricted to particular genera or subgenera of gophers, but the history of the host-parasite association appears complex and will require considerable effort to resolve.     

Ultrastructural features of spermatozoa and their phylogenetic application in Zaprionus (Diptera,Drosophilidae)

The genus Zaprionus consists of approximately 60 species of drosophilids that are native to the Afrotropical region. The phylogenetic position of Zaprionus within the Drosophilidae family is still unresolved. In the present study, ultrastructural features of spermatozoa of 6 species of Zaprionus as well as the species Drosophila willistoni and Scaptodrosophila latifasciaeformis were analyzed. The ultrastructure revealed that the species have the same flagellar ultrastructure. Two mitochondrial derivatives, one larger than the other, close to the axoneme were present, primarily in D. willistoni (subgenus Sophophora). Except for Z. davidi and Z. tuberculatus, the analyzed species had paracrystalline material in both mitochondrial derivatives. Moreover, the testes showed 64 spermatozoa per bundle in all of the species. In the cluster analysis, 6 Zaprionus species were grouped closely, but there were some incongruent positions in the cladogram. The results indicated that sperm ultrastructure is an important tool for elucidating the phylogeny and taxonomy of insects.     

The phylogenetic relationships of flies in the family Drosophilidae deduced from mtDNA sequences.

The phylogenetic relationships of several taxa from representative genera, subgenera, groups, and subgroups in the Drosophilidae were examined using sequences from a 905-bp mtDNA fragment. Conventional cloning and sequencing techniques were used to obtain nucleotide sequences. In addition, polymerase chain reaction primers were designed for the rapid amplification and sequencing of this region for the species examined in the Drosophilidae. Phylogenetic analysis was done by cladistic techniques. Because of the coding nature of the 905-bp mtDNA fragment, several separate analyses of these sequences were performed. The genera Scaptomyza and Hirtodrosophila occupy ancestral branching positions in the molecular phylogeny. The genera Chymomyza and Zaprionus have intermediate branching positions, while the subgenera Drosophila and Sophophora are in the most derived position in the molecular phylogeny. Within the subgenus Sophophora, there is little resolution using these sequences, while within the subgenus Drosophila, D. melanica, D. funebris, and D. pinicola form a clade in a derived part of the phylogeny, with D. robusta and D. immigrans branching in an intermediate position in the phylogeny. D. mercatorum, a member of the repleta species group, occupies an ancestral position in the molecular phylogeny.     

The origin and possible time of divergence of the Hawaiian Drosophilidae: evidence from DNA sequences.

The origins and times of divergence of the speciose Hawaiian Drosophilidae are examined using mtDNA sequences. The Hawaiian Drosophilidae are resolved as the sister group to the subgenus Drosophila. No one member of the subgenus Drosophila could be determined to be more closely related to the Hawaiian Drosophilidae than could any other. This result suggests that the Hawaiian Drosophilidae arose before the diversification of the subgenus Drosophila and after the divergence of the subgenus Sophophora. In light of fossil evidence, this phylogenetic scenario suggests that the Hawaiian Drosophilidae lineages are no younger than 30 Myr.     

Molecular phylogeny of horsetails (<Emphasis Type="Italic">Equisetum</Emphasis>) including chloroplast <Emphasis Type="Italic">atpB</Emphasis> sequences

Equisetum is a genus of 15 extant species that are the sole surviving representatives of the class Sphenopsida. The generally accepted taxonomy of Equisetum recognizes two subgenera: Equisetum and Hippochaete. Two recent phylogenetical studies have independently questioned the monophyly of subgenus Equisetum. Here, I use original (atpB) and published (rbcL, trnL-trnF, rps4) sequence data to investigate the phylogeny of the genus. Analyses of atpB sequences give an unusual topology, with E. bogotense branching within Hippochaete. A Bayesian analysis based on all available sequences yields a tree with increased resolution, favoring the sister relationships of E. bogotense with subgenus Hippochaete.     

Cladistic analyses of the animal kingdom

A recently published book on the phylogeny of the animal kingdom, written by the first author, provided a classification based on a 'manual' cladistic analysis at the phylum level. We have extracted a data matrix consisting of 61 characters for 32 phyla from this book and treated it in more formal analyses using three different parsimony programs. Following a posteriori weighting, one cladogram emerged as the most parsimonious explanation of the data. This cladogram is compared to those in recent publications. Congruence is greatest with the phylogeny published by the first author, as the monophyly of 18 of the 21 supraphyletic categories proposed therein are supported in our cladogram. The exceptions are Aschelminthes, Frotornaeozoa and Neorenalia, but the latter group does emerge as a monophyletic taxon in a number of equally parsimonious, equally weighted trees. Comparisons with other recent phytogenies show varying degrees of divergence, especially concerning the monophyly of Spiralia and Aiticulata, both of which are advocated in the present paper. Significant characters of most of the supraphyletic taxa proposed by the first author are discussed. C1996 The Linnean Society of London