Phylogenetic position of Nerillidae and Aberranta (Polychaeta, Annelida), analysed by direct optimization of combined molecular and morphological data

The phylogenetic position of the most speciose meiofaunal polychaete family, Nerillidae, has remained contentious. Recent hypotheses have generally focused on the fact that Nerillidae shares with Aciculata (a major polychaete subgroup) features such as compound chaetae, ventral buccal organ and short ventrolateral palps. Here we present the first phylogenetic analysis of Aciculata, together with Nerillidae, combining morphological and molecular data. We also include Aberrantidae, previously referred to or placed near to spiomorph polychaetes, but recently referred to Aciculata, possibly close to Nerillidae. The data sets of 24 terminals contain 53 morphological characters and nearly complete sequences of 18S rRNA. The sequences were analysed simultaneously with the morphological data by direct optimization in the program POY with a variety of parameter settings (costs of gaps: transversions: transitions). The various settings resulted in markedly different phylogenetic hypotheses, but on the basis of congruence (ILD) the results of two parameter settings were chosen. In all analyses, the three included nerillid species constituted a monophyletic group. Only two analyses provided fully resolved cladograms. The morphological analysis gave poor resolution and the position of the nerillids was equivocal. The two molecular‐based cladograms (minimizing ILD) were also poorly resolved, but one provided a position for nerillids next to Eunice pennata and Nothria conchilega, from the subgroup Eunicida within Aciculata. The two cladograms of the combined analyses (minimizing ILD) were fully resolved and placed nerillids in a terminal position next to Aberranta sp., within a clade of eunicidan species. The study showed that the analytical conditions for the homology assignment of 18S rRNA strongly influenced the phylogenetic results. The various previous proposals on the phylogenetic position of the Nerillidae are reviewed, some of which are in accordance with the results of the present study.     

Phylogeny,biogeography and the stepwise evolutionary colonization of intertidal habitat in the Liparocephalini based on morphological and molecular characters (Coleoptera: Staphylinidae: Aleocharinae)

A phylogenetic analysis of the tribe Liparocephalini Fenyes is presented based on morphological and molecular characters. The data set comprised 50 adult morphological characters, partial COI (907 bp), COII (366 bp) and 12S rDNA (325–355 bp), and nearly complete sequences of 18S rDNA (1768–1902 bp) for 21 species. Eighteen species of liparocephaline beetles from all eight genera and three outgroups, are included. The sequences were analysed separately and simultaneously with morphological characters by direct optimization in the program POY4 and by partitioned Bayesian analysis for the combined data. The direct optimization (DO) tree for the combined data under equal weighting, which also shows a minimum incongruence length difference value, resulted in a monophyletic Liparocephalini with the following patterns of phylogenetic relationships (outgroup ((Baeostethus, Ianmoorea) (Paramblopusa ((Amblopusa, Halorhadinus) (Liparocephalus, Diaulota))))). A sensitivity analysis using 16 different parameter sets for the combined data shows the monophyly of the liparocephalines and all its genera under all parameter sets. Bayesian analysis resulted in topological differences in comparison with the DO tree under equal weighting only in the position of the genus Paramblopusa and clade (Amblopusa + Halorhadinus), which were reversed. Historical biogeography and the stepwise evolutionary colonization of intertidal habitat in the Liparocephalini are discussed. Based on the biogeographical analyses, we hypothesize that the ancestor of the Liparocephalini occurred along the Panthallassan Ocean, the direct antecedent of the Pacific Ocean, followed by repeated dispersals to the Nearctic from the Palearctic. We also hypothesize that ancestors of the Liparocephalini appear to have arisen in the littoral zone of beaches and then colonized rocky reef areas in the low tidal zone later through high‐ to mid‐tide zones. © The Willi Hennig Society 2009.     

Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae,review of austral genera,and new larval descriptions

Abstract We develop a morphological dataset for the rove beetle subfamily Euaesthetinae comprising 167 morphological characters (135 adult and 32 larval) scored from 30 terminal taxa including 25 ingroup terminals (from subfamilies Euaesthetinae and Steninae) and five outgroups. Four maximum parsimony analyses using different sets of terminals and character sets were run to test the monophyly of (1) Euaesthetinae, (2) Steninae, (3) Euaesthetinae + Steninae, (4) euaesthetine tribes Austroesthetini, Alzadaesthetini, Euaesthetini, Fenderiini and Stenaesthetini, and (5) the ten currently known austral endemic genera together. Analyses of adult and larval character sets separately and in combination recovered the monophyly of Euaesthetinae, Steninae, and both subfamilies together, with strong support. Analysis of 13 ingroup terminals for which complete data were available suggests that monophyly of Euaesthetinae is supported by 19 synapomorphies (13 adult, six larval), of Steninae by 23 synapomorphies (14 adult, nine larval), and of both subfamilies together by 24 synapomorphies (21 adult, three larval). Within Euaesthetinae, only the tribe Stenaesthetini was recovered as monophyletic based on adult characters, and in no analyses were the ten austral endemic genera recovered as a monophyletic group. Phylogenetic relationships among euaesthetine genera were weakly supported, although analyses including adult characters supported monophyly of Octavius and Protopristus separately, and of Octavius + Protopristus, Austroesthetus + Chilioesthetus and Edaphus + Euaesthetus. Steninae may include a third genus comprising two undescribed species probably possessing a ‘stick–capture’ method of prey capture, similar to that in Stenus. These two species formed a strongly supported clade recovered as the sister group of Stenus based on adult characters. Diagnoses and a key to adults are provided for the 15 euaesthetine genera currently known from the austral region (Australia, New Zealand, South Africa and southern South America). Euaesthetine larvae previously were known only for Euaesthetus, and we describe the larvae of nine more genera and provide the first larval identification key for genera of Euaesthetinae.     

Phylogeny of the North American catfish family Ictaluridae (Teleostei: Siluriformes) combining morphology,genes and fossils

We performed the first combined‐data phylogenetic analysis of ictalurids including most living and fossil species. We sampled 56 extant species and 16 fossil species representing outgroups, the seven living genera, and the extinct genus ?Astephus long thought to be an ictalurid. In total, 209 morphological characters were curated and illustrated in MorphoBank from published and original work, and standardized using reductive coding. Molecular sequences harvested from GenBank for one nuclear and four mitochondrial genes were combined with the morphological data for total evidence analysis. Parsimony analysis recovers a crown clade Ictaluridae composed of seven living genera and numerous extinct species. The oldest ictalurid fossils are the Late Eocene members of Ameiurus and Ictalurus. The fossil clade ?Astephus placed outside of Ictaluridae and not as its sister taxon. Previous morphological phylogenetic studies of Ictaluridae hypothesized convergent evolution of troglobitic features among the subterranean species. In contrast, we found morphological evidence to support a single clade of the four troglobitic species, the sister taxon of all ictalurids. This result holds whether fossils are included or not. Some previously published clock‐based age estimates closely approximate our minimum ages of clades.     

Reconstructing the phylogeny of the Sipuncula

Sipunculans are marine spiralian worms with possible close affinities to the Mollusca or Annelida. Currently 147 species, 17 genera, 6 families, 4 orders and 2 classes are recognized. In this paper we review sipunculan morphology, anatomy, paleontological data and historical affiliations. We have conducted cladistic analyses for two data sets to elucidate the phylogenetic relationships among sipunculan species. We first analyzed the relationships among the 45 species of Phascolosomatidea with representatives of the Sipunculidea as outgroups, using 35 morphological characters. The resulting consensus tree has low resolution and branch support is low for most branches. The second analysis was based on DNA sequence data from two nuclear ribosomal genes (18S rRNA and 28S rRNA) and one nuclear protein-coding gene, histone H3. Outgroups were chosen among representative spiralians. In a third analysis, the molecular data were combined with the morphological data. Data were analyzed using parsimony as the optimality criterion and branch support evaluated with jackknifing and Bremer support values. Branch support for outgroup relationships is low but the monophyly of the Sipuncula is well supported. Within Sipuncula, the monophyly of the two major groups, Phascolosomatidea and Sipunculidea is not confirmed. Of the currently recognized families, only Themistidae appears monophyletic. The Aspidosiphonidae, Phascolosomatidae and Golfingiidae would be monophyletic with some adjustments in their definition. The Sipunculidae is clearly polyphyletic, with Sipunculus nudus as the sister group to the remaining Sipuncula, Siphonosoma cumanense the sister group to a clade containing Siphonosoma vastumand the Phascolosomatidea, and Phascolopsis gouldi grouping within the Golfingiiformes, as suggested previously by some authors. Of the genera with multiple representatives, only Phascolosoma and Themiste are monophyletic as currently defined. We are aiming to expand our current dataset with more species in our molecular database and more detailed morphological studies.     

Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae)

Euptychiina is the most species‐rich subtribe of Neotropical Satyrinae, with over 450 known species in 47 genera (14 monotypic). Here, we use morphological characters to examine the phylogenetic relationships within Euptychiina. Taxonomic sampling included 105 species representing the majority of the genera, as well as five outgroups. A total of 103 characters were obtained: 45 from wing pattern, 48 from genitalia and 10 from wing venation. The data matrix was analysed using maximum parsimony under both equal and extended implied weights. Euptychiina was recovered as monophyletic with ten monophyletic genera, contrasting previous DNA sequence‐based phylogenies that did not recover the monophyly of the group. In agreement with sequence‐based hypotheses, however, three main clades were recognized: the ‘Megisto clade’ with six monophyletic and three polyphyletic genera, the ‘Taygetis clade’ with nine genera of which three were monophyletic, and the ‘Pareuptyhia clade’ with four monophyletic and two polyphyletic genera. This is the first morphology‐based phylogenetic hypothesis for Euptychiina and the results will be used to complement molecular data in a combined analysis and to provide critical synapomorphies for clades and genera in this taxonomically confused group.     

Phylogeny of feather mite subfamily Avenzoariinae (Acari: Analgoidea: Avenzoariidae) inferred from combined analyses of molecular and morphological data.

Phylogenetic relationships among feather mites of the subfamily Avenzoariinae (Acari: Analgoidea: Avenzoariidae) were reconstructed by parsimony analysis of a combined data matrix. We analyzed 41 morphological characters and 246 molecular characters from a fragment of the 16S rDNA. Morphological trees were well supported at deep branches (genera and above), but showed much less support and resolution within genera. Molecular analyses produced trees with better resolution and support on terminal branches and worse support on basal branches. I(MF) index for the combined matrix pointed to the significant congruence of both data subsets with the whole of the data. The topology of the combined tree was close to the morphological tree in the deep branches and had well-resolved terminal branches as in the molecular tree. This suggests a considerable level of complimentarity between the two data sets. An analysis of association patterns of the mites and their hosts was conducted based on the results of the combined analyses for the Avenzoariinae and a phylogeny of their charadriiform hosts (compiled from various bird phylogeny hypotheses). The trees could be reconciled by the invoking of 12-13 cospeciation events, 6-7 duplications, 2 host shifts, and 26-29 sorting events. This suggests a high degree of cospeciation.     

Toward a Better Understanding of the Phylogeny of the Asplanchnidae (Rotifera)

We investigated the phylogenetic relationships of Family Asplanchnidae using both morphological and molecular data. The morphological database, comprising 23 characters from 19 taxa (15 Asplanchnidae and 4 outgroups), was compiled from a survey of the literature and our own observations; the molecular data (ITS and V4 region nuclear regions and mitochondrial cox1) was sequenced from specimens that we collected. Our analysis of the morphological data set (maximum parsimony) yielded 12 most-parsimonious trees with a tree length of 27 steps. From this analysis we conclude (1) Asplanchnidae is a monophyletic group as are the three genera comprising it, (2) there is no compelling support for the argument that Asplanchna should be separated into two discrete genera, and (3) there is some support for the proposal that Asplanchnidae and Synchaetidae are sister groups. Our analysis of the molecular data set supports the first two of these conclusions while the sister group of the family varied depending on the gene region analyzed and families and genera included. Current understanding of the phylogeny of Asplanchnidae is hampered by the need for additional informative morphological characters and a lack of molecular data for the genus Harringia and several other members of the Asplanchnidae.     

A molecular and morphological phylogeny of the extant Augochlorini (Hymenoptera,Apoidea) with comments on implications for biogeography

The Augochlorini Beebe is a New World tribe of bees comprising 663 described species. Relationships among the genera of this monophyletic tribe remain uncertain. Here I provide a comprehensive phylogeny using morphological and molecular information. In all, 54 Augochlorini species plus 16 outgroups and 3017 molecular and 105 morphological characters were analysed. Sequences for four genes were analysed using Bayesian inference, maximum likelihood and parsimony. Morphological characters were taken from a literature review and analysed alone and in combination with molecular data using parsimony. The monophyly of Augochlorini and most genera is confirmed, with divergence of the main lineages of the tribe around 55–20 Ma. Seven clades were supported by most analyses and are here treated as genus‐level groups, as follows (combined analysis topology): (Corynura group, (Chlerogella group, (Rhinocorynura group, (Augochloropsis, (Megaloptidia group, (Neocorynura group, (Augochlora group, Megalopta group))))))). According to this topology, dim‐light foraging and cleptoparasitism arose three times in the tribe. According to my hypothesis, the diversification of Augochlorini may have begun as a response to vicariant events, including the split of the Neotropical/Andean regions and marine transgressions in the Amazon region.     

Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution

Magnoliales, consisting of six families of tropical to warm-temperate woody angiosperms, were long considered the most archaic order of flowering plants, but molecular analyses nest them among other eumagnoliids. Based on separate and combined analyses of a morphological matrix (115 characters) and multiple molecular data sets (seven variable chloroplast loci and five more conserved genes; 14 536 aligned nucleotides), phylogenetic relationships were investigated simultaneously within Magnoliales and Myristicaceae, using Laurales, Winterales, and Piperales as outgroups. Despite apparent conflicts among data sets, parsimony and maximum likelihood analyses of combined data converged towards a fully resolved and well-supported topology, consistent with higher-level molecular analyses except for the position of Magnoliaceae: Myristicaceae + (Magnoliaceae + (( Degeneria + Galbulimima ) + ( Eupomatia + Annonaceae))). Based on these results, we discuss morphological evolution in Magnoliales and show that several supposedly plesiomorphic traits are synapomorphies of Magnoliineae, the sister group of Myristicaceae (e.g. laminar stamens). Relationships within Annonaceae are also resolved with strong support ( Anaxagorea basal, then ambavioids). In contrast, resolution of relationships within Myristicaceae is difficult and still incomplete, due to a very low level of molecular divergence within the family and a long stem lineage. However, our data provide good evidence that Mauloutchia is nested among other Afro-Malagasy genera, contradicting the view that its androecium and pollen are plesiomorphic  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 125–186.     

Phylogenetic analysis of the Amphitritinae (Polycnaeta: Terebellidae)

The phylogenetic relationships of the Amphitritinae (Polychaeta: Terebellidae) were studied using parsimony analysis of 22 external morphological characters. To choose outgroups to polarize the characters, I carried out a preliminary analysis of the relationships of the four terebellid subfamilies and the Trichobranchidae. The single most parsimonious tree from the analysis supports monophyly of the Terebellidae by the presence of ventral glandular shields. However, this character is homoplasious within the Terebellomorpha, and further evaluation of the Terebellidae is recommended. Artacama and Thelepus were chosen as outgroups for the analysis of amphitritine genera. The generic level analysis yielded seven equally parsimonious trees, which are consistent in their topologies except for the relationships among seven genera in one large clade. In all trees, Artacama is the sister taxon to a large clade within the Amphitritinae; the Artacaminae is therefore synonymized with the Amphitritinae, which is diagnosed by the presence of double rows of uncini. Within the Amphitritinae, the status of several monotypic genera is questioned; plesiomorphic character states indicated by the analysis are discussed. The results presented are offered as working hypotheses of the relationships among amphitritine genera. The large number of homoplasies indicated by the analysis emphasizes the need to further evaluate these hypotheses using additional characters. With a robust phylogenetic hypothesis of amphitritine relationships, a re-classification of the group based on apomorphic character states can be undertaken, and questions regarding the evolution of morphological characters, reproductive modes, or biogeographical patterns can be properly addressed.     

Phylogenetic relationships in Dichroplus Stål (Orthoptera: Acrididae: Melanoplinae) inferred from molecular and morphological data: testing karyotype diversification

The neotropical genus Dichroplus and related genera are characterized by a relatively uniform external morphology and a remarkably divergent male genitalia and hence its taxonomy is controversial. It also shows an extreme karyotypic diversification. In this study we used molecular and morphological characters to test the monophyly of the genus and to evaluate chromosome evolution. Twenty‐seven species from Dichroplus and related genera were included in the analysis. Morphological characters refer to the general morphology, male genitalia and female structures. Molecular studies were performed, sequencing part of two mitochondrial genes, cytochrome oxidase I and II. Independent and combined phylogenetic analyses of the data were performed under maximum parsimony. The karyotypic characters (rearrangements) were either mapped onto the combined topology or combined with the other data sets. While the molecular analysis confirms some results attained with morphology, some others do not. All point towards the paraphyly of the genus. Our results show the relevance of morphological data in phylogenetic studies because morphology and molecules supply complementary evidence. The mapping of chromosome characters on the combined tree shows that the most extreme karyotype, in D. silveiraguidoi, is a derived condition, probably reached through several centric fusions, and that X‐autosome centric fusions were recurrently fixed during the evolution of the group. © The Willi Hennig Society 2005.     

Phylogeny of West AfricanCaryedon(Coleoptera: Bruchidae): Congruence between Molecular and Morphological Data

Seed beetles belonging to the Old World genusCaryedonfeed in the seeds of various Caesalpinioideae, Mimosoideae, and Combretaceae. In an attempt to resolve broad phylogenetic relationships within the genus, we obtained 332 base pair sequences of mitochondrial 12S ribosomal DNA and morphological data for the 16 West AfricanCaryedonspecies. Morphological characters were analyzed under maximum parsimony and sequences were compared under maximum parsimony, maximum likelihood, and neighbor joining. Using a partition homogeneity test, we determined that morphological and molecular data sets were combinable. Combined data were analyzed under maximum parsimony. Morphological and molecular trees were congruent at the species group level and total evidence analyses yielded the same topologies as molecular data with each of the three outgroups used. Four main terminal clades are recognized, each corresponding with a group of species generally feeding on the same host plant family, subfamily, genus, or species. The monophyly of legume feedingCaryedonis supported by both data sets, and Combretaceae feeders split in two monophyletic assemblages.     

Analysis of the systematic relationships among ticks of the genera Rhipicephalus and Boophilus (Acari: Ixodidae) based on mitochondrial 12S ribosomal DNA gene sequences and morphological characters

A portion of mitochondrial 12S rDNA sequences (337-355 base pairs) and 63 morphological characters of 36 hard-tick species belonging to 7 genera were analyzed to determine the phylogenetic relationships among groups and species of Rhipicephalus and between the genera Rhipicephalus and Boophilus. Molecular and morphological data sets were first examined separately. The molecular data were analyzed by maximum parsimony (MP), maximum likelihood, and neighbor-joining distance methods; the morphological data were analyzed by MP After their level of congruence was evaluated by a partition homogeneity test, all characters were combined and analyzed by MP. The branches of the tree obtained by combining the data sets were better resolved than those of the trees inferred from the separate analyses. Boophilus is monophyletic and arose within Rhipicephalus. Boophilus species clustered with species of the Rhipicephalus evertsi group. Most of the clustering within Rhipicephalus was, however, consistent with previous classifications based on morphological data. Morphological characters were traced on the molecular reconstruction in order to identify characters diagnostic for monophyletic clades. Within the Rhipicephalus sanguineus complex, the sequences of specimens morphologically identified as Rhipicephalus turanicus were characterized by a high level of variability, indicating that R. turanicus-like morphology may cover a spectrum of distinct species.     

PHYLOGENYAND HISTORICAL ECOLOGY OF THE DESMARESTIACEAE (PHAEOPHYCEAE) SUPPORT A SOUTHERN HEMISPHERE ORIGIN1

Phylogenetic relationships in the Desmarestiales (Phaeophyceae) were inferred among the monotypic Arthrocladia (Arthrocladiaceae) and 27 isolates from Desmarestiaceae, representing 17 taxa of Desmarestia and the monotypic Antarctic genera Himantothallus and Phaeurus. Phaeurus and Arthrocladia were used as outgroups. Parsimony analyses of nuclear ribosomal DNA internal transcribed spacer (ITS1 and ITS2) sequences, in which gaps were both included and excluded, yielded well-resolved trees with a consistent general branching pattern. A parallel analysis of nine morphological and life-history characters and three ecological characters yielded a similar tree but provided little resolution in the terminal clades. The position of the monotypic Arthrocladia villosa within the Desmarestiales is consistent with monophyly for the order, but its position as the most primitive desmarestialean is not resolvable from the molecular data set. The basal position of Phaeurus, the Antarctic Desmarestia species, and Himantothallus is consistent with the hypothesis of a Southern Hemisphere origin for the family Desmarestiaceae. The more recent Northern Hemisphere “aculeata” clade evolved from an Antarctic ancestor. A “D. aculeata-like” species was ancestral to a lineage characterized by annual sporophytes with high sulfuric acid content, which radiated into many species, widely distributed in both hemispheres. Mapping of morphological and ecological characters onto the molecular tree confirm the informativeness of sulfuric acid-containing vacuoles and unilocular sporangial types. There is good congruence between phylogenetic tree topology and temperature impints in relation to biogeographic distribution, supporting the theory that temperature tolerance is a conservative trait.     

Problems in polychaete systematics

Some of the intriguing issues in current polychaete systematics are reviewed. (1) The root of the `polychaete' tree. Currently there are two major hypotheses concerning the root position among polychaetes. One is based on rooting cladograms with outgroups such as Mollusca and result in simple-bodied taxa such as Opheliidae and Questidae forming a basal annelid grade along with Clitellata. Other hypotheses do not use outgroup rooting but involve scenarios on the evolution of the group and would place taxa in Aciculata as basal annelids, thus making Aciculata and Phyllodocida paraphyletic. Molecular sequence data has been of little help in resolving this issue thus far, largely due to limited taxon sampling. (2) Paraphyly. Owing, in part, to a tradition involving the emphasis on differences among taxa, and the application of Linnean ranks (e.g., family), paraphyly is undoubtedly a widespread phenomenon in polychaete systematics. An example of this has been proposed already for Spionidae. If the tree topology and rooting used by Blake & Arnofsky (1999) is correct, Spionidae is made paraphyletic by the recognition of the following four family-ranked taxa; Trochochaetidae, Poecilochaetidae, Longosomatidae and Uncispionidae. Another possible example is seen with Cirratulidae. A preliminary cladistic analysis shows that it is entirely possible that seven other taxa recognised as families may be nested within Cirratulidae. These include Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae, Flotidae, Poeobiidae and Sternaspidae. (3) Problematic taxa. Apart from the problems exposed by the analysis of Cirratuliformia, the position of some of these groups, such as Aberranta, Alciopidae, Hesionides,Lopadorhynchidae, Microphthalmus,Nerillidae, Spinther,Tomopteridae and Sabellariidae, is discussed.     

基于28S rDNA D2基因片段与形态特征的矛茧蜂亚科系统发育研究（膜翅目：茧蜂科）

本研究选取矛茧蜂亚科Doryctinae（昆虫纲Insecta：膜翅目Hymenoptera：茧蜂科Braconidae）的6族15属18种做内群,茧蜂科其它7亚科11属11种做外群,首次结合同源核糖体28S rDNA D2基因序列片段和100个形态学和解剖学特征对该亚科进行了系统发育学研究。利用“非圆口类"的小腹茧蜂亚科Microgastrinae为根,以PAUP*4.0和MrBayes 3.0B4软件分别应用最大简约法（MP）和贝叶斯法对矛茧蜂亚科的分子数据和分子数据与非分子数据的结合体进行了运算分析;并以PAUP*4.0对矛茧蜂亚科的28S rDNA D2基因序列片段的碱基组成与碱基替代情况进行了分析。结果表明：矛茧蜂亚科的28S rDNA D2基因序列片段的GC含量在39.33%～48.28%之间变动,而对于碱基替代情况来讲,矛茧蜂亚科各成员间序列变异位点上颠换（transversion）大于转换（transition）。不同的分析算法所产生的系统发育树都表明矛茧蜂亚科是一个界限分明的单系群;在矛茧蜂亚科内,除了吉丁茧蜂族Siragrini为单系群外,其他族（矛茧蜂族Doryctini和方头茧蜂族Hecabolini）都是并系群。对于矛茧蜂亚科内各属之间的相互亲缘关系,不同算法所得的系统发育树的拓扑结构不完全一致,表明矛茧蜂亚科内（属及族）的系统发育关系还有待于进一步研究。     

Phylogeny of Branchiopoda (Crustacea) based on a combined analysis of morphological data and six molecular loci

The phylogenetic relationships of branchiopod crustaceans have been in the focus of a number of recent morphological and molecular systematic studies. Although agreeing in some respects, major differences remain. We analyzed molecular sequences and morphological characters for 43 branchiopods and two outgroups. The branchiopod terminals comprise all eight “orders”. The molecular data include six loci: two nuclear ribosomal genes (18S rRNA, 28S rRNA), two mitochondrial ribosomal genes (12S rRNA, 16S rRNA), one nuclear protein coding gene (elongation factor 1α), and one mitochondrial protein coding gene (cytochrome c oxidase subunit I). A total of 65 morphological characters were analyzed dealing with different aspects of branchiopod morphology, including internal anatomy and larval characters. The morphological analysis resulted in a monophyletic Phyllopoda, with Notostraca as the sister group to the remaining taxa supporting the Diplostraca concept (“Conchostraca” + Cladocera). “Conchostraca” is not supported but Cyclestheria hislopi is the sister group to Cladocera (constituting together Cladoceromorpha) and Spinicaudata is closer to Cladoceromorpha than to Laevicaudata. Cladocera is supported as monophyletic. The combined analysis under equal weighting gave results in some respects similar to the morphological analysis. Within Phyllopoda, Cladocera, Cladoceromorpha and Spinicaudata + Cladoceromorpha are monophyletic. The combined analysis is different from the morphological analysis with respect to the position of Notostraca and Laevicaudata. Here, Laevicaudata is the sister group to the remaining Phyllopoda and Notostraca is sister group to Spinicaudata and Cladoceromorpha. A sensitivity analysis using 20 different parameter sets (different insertion–deletion [indel]/substitution and transversion/transition ratios) show the monophyly of Anostraca, Notostraca, Laevicaudata, Spinicaudata, Cladoceromorpha, Cladocera, and within Cladocera, of Onychopoda and Gymnomera under all or almost all (i.e., 19 of 20) parameter sets. Analyses with an indel‐to‐transversion ratio up to 2 result in monophyletic Phyllopoda, with Laevicaudata as sister group to the remaining Phyllopoda and with Spinicaudata and Cladoceromorpha as sister groups. Almost all analyses (including those with higher indel weights) result in the same topology when only ingroup taxa are considered. © The Willi Hennig Society 2007.