A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata)

The taxonomic placement of an enigmatic species of wasp known from two specimens in Late Cretaceous New Jersey amber is investigated through cladistic analyses of 90 morphological characters for 33 terminals ranging across non-Aculeata, non-Chrysidoidea, most subfamilies of Chrysidoidea and all genera of Plumariidae (the family to which the fossils were initially assigned), based on use of exemplars. The fossil taxon is apparently basal in Chrysidoidea, most likely sister to Plumariidae, but perhaps sister to the remaining chrysidoids, or even sister to Chrysidoidea as a whole. It is described as representing a new family, Plumalexiidaefam. n., containing a single species, Plumalexius rasnitsynigen. et sp. n. Previous estimates of relationships for the genera of Plumariidae and for the higher taxa of Chrysidoidea are mostly confirmed. The importance of outgroup choice, and additivity and weighting of characters are demonstrated.     

Progress in erigonine spider phylogeny—the <Emphasis Type="Italic">Savignia</Emphasis>-group is not monophyletic (Araneae: Linyphiidae)

We present the most inclusive study on the higher-level phylogeny of erigonine spiders, including about 30% of all erigonine genera. By expanding the previously most comprehensive analysis (Miller and Hormiga Cladistics 20:385–442, 2004) we tested the robustness of its results to the addition of closely related taxa, and also the monophyly of the Savignia-group defined by Millidge (Bulletin of the British Arachnological Society 4:1–60, 1977). The character matrix was expanded by adding 18 newly scored species in 15 genera, and also includes all species scored by other authors. This adds up to 98 species in 91 erigonine genera plus 13 outgroup taxa. The parsimony analysis led to eight fully resolved most parsimonious trees (L=1084). The topology concerning the taxa basal to the ‘distal erigonines’ remained unchanged, and the latter clade still shares 67% of all nodes with the original analysis. The Savignia-group is not monophyletic at genus level with respect to Saloca diceros and Alioranus pastoralis, and the same applies at species level in Diplocephalus and Erigonella. From the Savignia-group, Glyphesis servulus, Diplocephalus cristatus, Savignia frontata, and two representatives each of Erigonella, Dicymbium and Araeoncus combine to form a monophyletic clade.     

A phylogenetic analysis of subtribe Pleurothallidinae (Orchidaceae)

A cladistic analysis of subtribe, Pleurothallidinae (Orchidaceae) is based on 45 anatomical/ morphological characters. The ingroup members comprise 24 genera; the large genus Pleurothallis consists of two subgenera and ten species complexes. Three taxa representing subtribes Laeliinae and Arpophyllinae are designated as outgroup. Eight most parsimonious trees were discovered using computer assisted software (length = 230; CI = 0.27). The hypothesis that subtribe Pleurothallidinae has undergone a unilinear reduction in the number of pollinia is not supported by this study. Although the eight-pollinia state as represented by Octomeria apparently is plesiomorphic, the two-pollinia and four-pollinia states arose early in the phylogeny of the subtribe. Both two-and four-pollinia states subsequently reappeared as parallelisms. The six-pollinia state exhibited in Brachionidium is autapomorphic. This cladistic analysis suggests that Pleurothallis is not a natural genus and, perhaps may be divided into several discrete genera.     

A phylogenetic analysis of the nursery-web spider family Pisauridae, with emphasis on the genera Architis and Staberius (Araneae: Lycosoidea)

This study presents a general cladistic analysis of pisaurid genera, with emphasis on the Neotropical genera Architis Simon and Staberius Simon. The analysis was based on a matrix with 21 terminal taxa: eight species of Architis , Staberius spinipes (Taczanowski), seven exemplars of other pisaurid genera and five outgroup taxa. These terminals were scored for 59 morphological and three behavioural characters. An analysis with all characters equally weighted resulted in two most parsimonious trees, differing only in the position of Architis colombo Santos. In both trees Pisauridae arouse as a monophyletic group, and the exemplars of Architis composed a clade with S. spinipes . Contrary to hypotheses from the literature, Thalassius Simon emerged as sister-group of all remaining pisaurids, not as a close relative of Dolomedes Latreille. The genera Tinus F.O.P. -Cambridge and Thaumasia Perty appeared in the trees as the closest relatives of Architis and Staberius Simon. The analysis strongly supported S. spinipes as the sister-group of A. helveola (Simon), indicating that Architis as currently delimited is paraphyletic. Based on these results, Staberius is considered a subjective junior synonym of Architis . Additionally, the genus Mimicosa Petrunkevitch, originally described in Tetragnathidae, is transferred to Pisauridae and considered a junior synonym of Architis . Two species are proposed as junior synonyms of A. spinipes comb.n. Mimicosa spinosa Petrunkevitch and Staberius lemoulti Caporiacco.     

Cladistics and the comparative morphology of linyphiid spiders and their relatives (Araneae, Araneoidea, Linyphiidae)

This paper provides the first quantitative cladistic analysis of linyphiid morphology. Classical and novel homology hypotheses for a variety of character systems (male and female genitalia, somatic morphology, spinneret silk spigot morphology, etc.) are critically examined and studied within a phylogenetic context. Critical characters have been illustrated. A sample of linyphiid taxa (nine genera in four subfamilies), five species of Pimoa (Pimoidae), and two other araneoid families (Tetragnathidae and Araneidae, represented by Tetragnatha and Zygiella , respectively) were used to study the implications of the phylogeny of Pimoidae for the systematics of linyphiids. The phylogenetic relationships of these 16 exemplar taxa, as coded for the 47 characters studied, were analysed using numerical cladistic methods. In the preferred cladogram Pimoidae and Linyphiidae are sister groups, Stemonyphantinae are sister group to the remaining linyphiids, and Mynogleninae are sister group to the clade composed of Erigoninae plus Linyphiinae. These results agree with the relationships recently proposed by Wunderlich, except by finding erigonines as the sister group to linyphiines rather than to mynoglenines.     

Phylogeny of Neotropical Protoneuridae (Odonata: Zygoptera) and a preliminary study of their relationship with related families

Abstract.  A cladistic analysis of Neotropical Protoneuridae was performed on a data matrix of 48 morphological characters and 43 terminal taxa. Representatives of Paleotropical Protoneuridae, Platycnemididae and Isostictidae were included to test their relationships with Neotropical Protoneuridae. Coenagrionidae was chosen as the outgroup, but alternative analyses with Platycnemididae as the outgroup were also performed. Protoneuridae appears as a polyphyletic clade, with its Paleotropical component being more closely related to Platycnemididae and Isostictidae. Neotropical Protoneuridae appear as a monophyletic clade; included genera considered monophyletic or valid monotypic taxa are Epipleoneura Williamson, 1915 ; Idioneura Selys, 1860; Junix Rácenis, 1968; Neoneura Selys, 1860; Peristicta Hagen in Selys, 1860; Roppaneura Santos, 1966; and Lamproneura De Marmels, 2003. A key to the Neotropical Protoneuridae genera is included.     

Revision of the Agathidinae (Hymenoptera: Braconidae) with comparisons of static and dynamic alignments

The phylogeny of the Agathidinae (Insecta: Hymenoptera: Braconidae) is investigated based on morphological and sequence data from the D2–3 regions of 28S rDNA. Morphology and molecular data were run simultaneously and separately and the molecular and combined data sets were analyzed using both static, Clustal W, and dynamic, POY, alignments. Both alignments were conducted under a variety of gap costs and results are compared. Sixty‐two ingroup exemplars representing 22 genera and six outgroup taxa representing two subfamilies and five genera were included. Numerous taxa at the generic and tribal levels were tested for monophyly and the evolutionary history of several characters is discussed. The tribe Agathidini s.s. is found to be a derived member of the Microdini and the two are synonymized under the older name, Agathidini s.l. Support is substantial for the tribes Cremnoptini and Disophrini and Earinini but equivocal for the Agathidini s.l. At the generic level, Bassus is found to be polyphyletic. Numerous new synonymies and combinations are proposed. © The Willi Hennig Society 2006.     

Rapid evolutionary divergences in reef fishes of the family Acanthuridae (Perciformes: Teleostei)

A phylogenetic analysis of the sugeonfish family Acanthuridae was conducted to investigate: (a) the pattern of divergences among outgroup and basal ingroup taxa, (b) the pattern of species divergences within acanthurid genera, (c) monophyly in the genus Acanthurus, and (d) the evolution of thick-walled stomach morphology in the genera Acanthurus and Ctenochaetus. Fragments of the 12S, 16S, t-Pro, and control region mitochondrial genes were sequenced for 21 acanthurid taxa (representing all extant genera) and four outgroup taxa. Unweighted parsimony analysis produced two optimal trees. Both of these were highly incongruent with a previous morphological phylogeny, especially with regard to the placement of the monotypic outgroups Zanclus and Luvarus. The maximum likelihood tree and the morphological phylogeny were not significantly different and the conflicting branches were very short. Split decomposition analysis identified conflict in the placement of long basal branches separated by short internodes, providing further evidence that long branch attraction is an important cause of disagreement between molecular and morphological trees. Parametric bootstrapping rejected hypotheses of monophyly of: (a) the genus Acanthurus and (b) a group containing representatives of Acanthurus/Ctenochaetus with thick-walled stomachs. The branching pattern of the likelihood and split decomposition trees indicates that evolution in the acanthurid clade has involved at least three periods of intense speciation.     

Head capsule characters in the Hymenoptera and their phylogenetic implications

The head capsule of a taxon sample of three outgroup and 86 ingroup taxa is examined for characters of possible phylogenetic significance within Hymenoptera. 21 morphological characters are illustrated and scored, and their character evolution explored by mapping them onto a phylogeny recently produced from a large morphological data set. Many of the characters are informative and display unambiguous changes. Most of the character support demonstrated is supportive at the superfamily or family level. In contrast, only few characters corroborate deeper nodes in the phylogeny of Hymenoptera.     

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     

Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: further evidence from a nuclear gene of the systematic chaos in the family

The family Cyprinidae is the largest freshwater fish group in the world, including over 200 genera and 2100 species. The phylogenetic relationships of major clades within this family are simply poorly understood, largely because of the overwhelming diversity of the group; however, several investigators have advanced different hypotheses of relationships that pre- and post-date the use of shared-derived characters as advocated through phylogenetic systematics. As expected, most previous investigations used morphological characters. Recently, mitochondrial DNA (mtDNA) sequences and combined morphological and mtDNA investigations have been used to explore and advance our understanding of species relationships and test monophyletic groupings. Limitations of these studies include limited taxon sampling and a strict reliance upon maternally inherited mtDNA variation. The present study is the first endeavor to recover the phylogenetic relationships of the 12 previously recognized monophyletic subfamilies within the Cyprinidae using newly sequenced nuclear DNA (nDNA) for over 50 species representing members of the different previously hypothesized subfamily and family groupings within the Cyprinidae and from other cypriniform families as outgroup taxa. Hypothesized phylogenetic relationships are constructed using maximum parsimony and Basyesian analyses of 1042 sites, of which 971 sites were variable and 790 were phylogenetically informative. Using other appropriate cypriniform taxa of the families Catostomidae (Myxocyprinus asiaticus), Gyrinocheilidae (Gyrinocheilus aymonieri), and Balitoridae (Nemacheilus sp. and Beaufortia kweichowensis) as outgroups, the Cyprinidae is resolved as a monophyletic group. Within the family the genera Raiamas, Barilius, Danio, and Rasbora, representing many of the tropical cyprinids, represent basal members of the family. All other species can be classified into variably supported and resolved monophyletic lineages, depending upon analysis, that are consistent with or correspond to Barbini and Leuciscini. The Barbini includes taxa traditionally aligned with the subfamily Cyprininae sensu previous morphological revisionary studies by Howes (Barbinae, Labeoninae, Cyprininae and Schizothoracinae). The Leuciscini includes six other subfamilies that are mainly divided into three separate lineages. The relationships among genera and subfamilies are discussed as well as the possible origins of major lineages.     

Direct optimization and multiple alignment of 28S D2–D3 rDNA sequences: problems with indels on the way to a molecular phylogeny of the cryptine ichneumon waSPS (Insecta: Hymenoptera)

A molecular phylogeny of the Cryptinae based on the D2–D3 region of the 28S rDNA gene, including 176 cryptines representing 139 genera and an additional 77 outgroup genera, is presented. Direct optimization and multiple alignment each with a range of gap/substitution parameters were employed. The informal higher grouping of the ichneumonids were supported, but the relationship of the Ichneumoninae and Cryptinae was not resolved. Cryptinae tribes were supported but many genera formed constant groups that did not follow earlier subtribal divisions. We investigated artifactual placements of, e.g., Agriotypinae and Brachycyrtinae, and showed that they do not possess molecular synapomophies with the Ichenumoninae where direct optimization places them. The sequences of anomalous placed taxa were markedly longer (or in some cases shorter) than those of most other taxa, and this is explained by homoplastic indels at “hot spots” in the absence of large numbers of informative substitutions. We show that multiple alignment generally resulted in taxa with large insertions or deletions being placed in more plausible positions. The relationship of the genus Alomya is also explored. © The Willi Hennig Society 2006.     

Evolutionary relationships among the protostrongylidae (Nematoda: Metastrongyloidea) as inferred from morphological characters, with consideration of parasite-host coevolution.

The phylogeny of nematodes in the family Protostrongylidae (Nematoda: Metastrongyloidea) was reconstructed by cladistic analysis of 28 binary and multistate characters derived from comparative morphology. Analyses were hierarchical, and examined (1) relationships among genera, including 13 ingroup taxa and Metastrongylidae as an outgroup (single tree, 78 steps, consistency index [CI] = 0.705); and (2) relationships among genera and species groups, including 21 ingroup taxa and Metastrongylus apri as an outgroup (single tree, 76 steps, CI = 0.582). In the species-level tree, Protostrongylidae was divided into 2 major clades, 1 containing the subfamilies Muelleriinae (including the recently described Umingmtakstrangylus pallikuukensis), Elaphostrongylinae, and the Varestrongylinae (excluding Pneumocaulus kadenazii). Varestrongylus was paraphyletic as it included Pneumostrongylus calcaratus. The second major clade consisted of a paraphyletic group containing Protostrongylus spp. and Spiculocaulus leuckarti and, basal to this subclade, several other individual protostrongylid lineages. The various subclades generally correspond to the subfamilial divisions of the Protostrongylidae. The Neostrongylinae, however, is not supported as Neostrongylus and Orthostrongylus are not sister groups. Based on a large number of hypothesized synapomorphies, the elaphostrongylines appear to be a highly derived group of protostrongylids, a feature potentially correlated with their habitat localization in muscular and nervous tissues. The generic-level tree retained most of the primary structure revealed among the species but excluded the varestrongylines from the Muelleriinae + Elaphostrongylinae subclade. Artiodactyles of the family Cervidae are considered basal hosts for protostrongylids; secondary colonization in Caprini, Rupicaprini, and among lagomorphs is postulated.     

Phantoms of Gondwana?—phylogeny of the spider subfamily Mynogleninae (Araneae: Linyphiidae)

This is the first genus‐level phylogeny of the subfamily Mynogleninae. It is based on 190 morphological characters scored for 44 taxa: 37 mynoglenine taxa (ingroup) representing 15 of the 17 known genera and seven outgroup taxa representing the subfamilies Stemonyphantinae, Linyphiinae (Linyphiini and Micronetini), and Erigoninae, and a representative of the family Pimoidae, the sister‐group to Linyphiidae. No fewer than 147 of the morphological characters used in this study are new and defined for this study, and come mainly from male and female genitalia. Parsimony analysis with equal weights resulted in three most parsimonious trees of length 871. The monophyly of the subfamily Mynogleninae and the genera Novafroneta, Parafroneta, Laminafroneta, Afroneta, Promynoglenes, Metamynoglenes, and Haplinis are supported, whereas Pseudafroneta is paraphyletic. The remaining seven mynoglenine genera are either monotypic or represented by only one taxon. Diagnoses are given for all genera included in the analysis. The evolution of morphological traits is discussed and we summarize the diversity and distribution patterns of the 124 known species of mynoglenines. The preferred topology suggests a single origin of mynoglenines in New Zealand with two dispersal events to Africa, and does not support Gondwana origin.     

A phylogenetic analysis of subtribe Pleurothallidinae (Orchidaceae)

A cladistic analysis of subtribe, Pleurothallidinae (Orchidaceae) is based on 45 anatomical/ morphological characters. The ingroup members comprise 24 genera; the large genus Pkurothallis consists of two subgenera and ten species complexes. Three taxa representing subtribes Laeliinae and ArpophyUinae are designated as outgroup. Eight most parsimonious trees were discovered using computer assisted software (length = 230; CI = 0.27). The hypothesis that subtribe Pleurothallidinae has undergone a unilinear reduction in the number of pollinia is not supported by this study. Although the eight-pollinia state as represented by Octomeria apparently is plesiomorphic, the two-pollinia and four-pollinia states arose early in the phytogeny of the subtribe. Both two-and four-pollinia states subsequently reappeared as parallelisms. The six-pollinia state exhibited in Brachionidium is autapomorphic. This cladistic analysis suggests that Pkurothallis is not a natural genus and, perhaps may be divided into several discrete genera.     

Phyletic patterns of early development in gastropod molluscs

Cell lineage data for 30 exemplar gastropod taxa representing all major subclades and the outgroup Polyplacophora were examined for phylogenetic signal using cladistic analysis. Most cell lineages show phyletic trends of acceleration or retardation relative to the outgroup and more basal ingroup taxa, and when coded this variation is phylo-genetically informative. PAUP analyses of a cell lineage data set under three sets of character ordering assumptions produced similar tree topologies. The topologies of the strict consensus trees for both ordered and Dollo (near irreversibility of character transformations) character assumptions were similar, whereas the unordered character assumption recovers the least phyletic information. The cell lineage cladograms are also in agreement with the fossil record of the timing and sequence of gastropod subclade origination. A long branch lies between the Patellogastropoda+Vetigastropoda grade and the Neritopsina+Apogastropoda clade. The geological timing of this long branch is correlated with the first large-scale terrestrially derived eutrophication of the near-shore marine habitat, and one possible explanation for this branch may be a developmental shift associated with the evolution of feeding larvae in response to the more productive conditions in the near-shore water column. Although character transformations are highly ordered in this data set, developmental rate characters (like all other morphological and molecular characters) are also subject to homoplasy. Finally, this study further supports the hypothesis that early development of gastropod molluscs has conserved a strong phyletic signal for about half a billion years.     

Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera

Epyrini (Epyrinae) are the largest and morphologically most diverse group within Bethylidae. Evans' conception of Epyrini induces misclassification of several taxa and their diagnostic features, such as median lobe of clypeus projected, and fovea in lateroposterior angle of propodeal disc, are insufficient to place species in the tribe. No previous comprehensive cladistic study has recovered Epyrini as a clade, and no synapomorphies are reported for the tribe, suggesting that the group is artificial. We infer a phylogeny based on 391 informative morphological characters for 42 terminal taxa (31 ingroup, 11 outgroup). Monophyly of Epyrini is supported by a petiole with fused root and body as an autapomorphy. Anisepyris Kieffer, Laelius Ashmead, Holepyris Kieffer, Formosiepyris Terayama, Disepyris Kieffer, Bakeriella Kieffer and Trachepyris Kieffer were retrieved as monophyletic. Epyris Westwood and Chlorepyris Kieffer are paraphyletic. The addition of new characters was essential to define Epyrini, and the ventral region of the mesosoma, petiole and male genitalia may help to better understand genera and species limits. All resulting trees indicate that Epyrinae are polyphyletic. Tribes Epyrini and Sclerodermini are revalidated to subfamily status.     

Phylogeny of the Nymphaeaceae inferred from

The Nymphaeaceae are one of the most important taxa for understanding the origin and evolution of primitive angiosperms due to its basal position in the cladograms of the angiosperms recently given based both on several gene sequences and on morphological characters, but phylogenetic relationships within the family have not as yet been resolved. The sequences of nrDNA ITS region of 11 species representing seven genera of the Nymphaeaceae and one outgroup, Ceratophyllum demersum, were used to reconstruct the phylogeny of the family using PAUP4.0b4A. Three most parsimonious trees (Length=1125, CI=0.7618 and RI=0.7214) were obtained. In the consensus tree: (1) Nelumbo was basal to the other genera with a bootstrap value of 100% and occupied an isolated position, so it could be separated from the Nymphaeaceae and placed in its own family, Nelumbonaceae, and its own order, Nelumbonales; (2) Nuphar was monophyletic, nested in the basal position of clade II and was strongly supported to be the sister group of the other genera (excluding Nelumbo) of the Nymphaeaceae, suggesting that its traditional inclusion in the Nymphaeaceae should be maintained; (3) Cabomba and Brasenia were sister group and formed a subclade with a bootstrap value of 99%, indicating their close affinity; (4) Nymphaea was strongly supported to be the sister group of a subclade comprising Euryale and Victoria with a bootstrap value of 94%, indicating that they should be placed in the Nymphaeaceae.