Taxonomic studies in Chiloscyphus Corda (Jungermanniales: Lophocoleaceae) based on nrITS sequences and morphology

Maximum parsimony and likelihood analyses of 40 Lophocoleaceae nrITS sequences and 6 Plagiochilaceae sequences (outgroup) lead to a robust phylogeny of Chiloscyphus. Four main lineages are assigned to as Chiloscyphus subgenera Chiloscyphus, Lophocolea, Connati and Notholophocolea. Chiloscyphus subgen. Connati is resolved sister to the remainder of this genus. Chiloscyphus subgenus Lophocolea is subdivided into sections Heterophylli (incl. sect. Semiteretes, syn. nov.), Lophocolea, Microlophocolea, and Novae-Zeelandiae. Five accessions of Chiloscyphus pallescens with a chromosome number of n = 18 form a robust monophyletic lineage that is placed sister to a well supported clade with 4 accessions of C. polyanthos [n = 9]. Chiloscyphus mandonii is placed in the synonymy of C. latifolius.     

Acceptance of Liochlaena Nees and Solenostoma Mitt., the systematic position of Eremonotus Pearson and notes on Jungermannia L. s.l. (Jungermanniidae) based on chloroplast DNA sequence data

Nucleotide sequence variation of the chloroplast rbcL gene of 78 representatives of Jungermanniales suborders Jungermanniineae and Cephaloziineae and five outgroup species is analysed using maximum parsimony and maximum likelihood. Jungermannia s.l. is resolved in three independent lineages corresponding to Jungermannia subgen. Jungermannia, Liochlaena and Solenostoma (incl. subgen. Plectocolea). Based on the outcome of the phylogenetic analyses we adopt a narrow generic concept of Jungermannia and accept the genera Liochlaena and Solenostoma. Contrary to proposed hypotheses, the monospecific genus Eremonotus is not nested in Scapaniaceae, Gymnomitriaceae, Cephaloziellaceae or Cephaloziaceae but placed in a clade together with Jungermannia s.str., Liochlaena, Delavayella and Leiocolea. These genera possess female involucres consisting solely of leaf tissue (perianths). Related clades comprise genera with female involucres originating at least partly from stem tissue. We propose to include Delavayella, Eremonotus, Jungermannia s.str., Leiocolea and Liochlaena in Jungermanniaceae based on the phylogenetic reconstructions of cpDNA variation and the distribution of perianths in Jungermanniales suborder Jungermanniineae.     

The systematic identity of Chiloscyphus trichocoleoides,a new liverwort species from New Zealand,uncovered by morphological and molecular evidence

Abstract

A new liverwort species from Western Nelson Ecological Province of New Zealand known from a single sterile specimen is described. It is unusual in having well-developed stem paraphyllia, and the leaves and underleaves are divided with three levels of ciliate division in a pinnate manner. This morphology suggested membership of Trichocoleaceae or Trichotemnomaceae, but other features of the plant made placement in either family seem unnatural. We determined its position by conducting phylogenetic analyses of rbcL, rps4 and trnL–F sequence datasets from 35 exemplars, and further confirmation of the position was carried out by an extended analysis using rbcL sequence dataset of 59 exemplars. The surprising result is that it belongs within the genus Chiloscyphus, where it is unique in these features. We describe this new species as Chiloscyphus trichocoleoides. Placing the new species in Chiloscyphus has broadened the generic concept of Chiloscyphus, and its phylogenetic relationship within and to other genera such as Clasmatocolea and Heteroscyphus needs further investigation. The results also show that the same morphological features seen in Chiloscyphus and in Trichocoleaceae and Trichotemnomaceae have developed independently.     

Multi-locus phylogeny clarifies the systematics of the Australo-Papuan robins (Family Petroicidae, Passeriformes)

The Australo-Papuan family Petroicidae (Aves: Passeriformes) has been the focus of much systematic debate about its relationships with other passerine families, as well as relationships within the family. Mostly conservative morphology within the group limits the effectiveness of traditional taxonomic analyses and has contributed to ongoing systematic debate. To assess relationships within the family, we sampled 47 individuals from 26 species, representing the majority of genera and species, for four loci: 528 base pairs (bp) of C-myc, 501 bp of BA20454 and 336 bp of BA23989 from nuclear DNA and 1005 bp of the mitochondrial ND2 gene. There was consensus between individual loci and overall support for major lineages was strong. Partitioned Bayesian analyses of all four loci produced a fully resolved and very well-supported phylogeny that addresses many of the previous systematic debates in this group. The Eopsaltriinae as construed is monophyletic with the exception of Eopsaltria flaviventris, which is nested within Microeca as an unremarkable member of that genus. This relationship is corroborated by morphology and egg color and pattern. Petroicinae as currently construed was not monophyletic and comprised two lineages that are paraphyletic with respect to each other. The third subfamily, Drymodinae, remains incertae sedis. The mangrove robin, Peneonanthe pulverulenta, of tropical Australia and New Guinea is nested within a clade that also contained the sampled species of Peneothello and Melanodryas, a novel relationship. Preliminary biogeographic and divergence time estimates from these results are discussed and a new subfamily arrangement proposed.     

Phylogeny of the Anomala (Crustacea, Decapoda, Reptantia) based on the ossicles of the foregut

We present a cladistic analysis of the Anomala based on 66 ingroup species and 5 outgroup representatives. Based on a comparative analysis of the morphology of the foregut we scored 124 characters related to size, shape, and fusion of foregut ossicles and other foregut structures. Our parsimony analysis resulted in 30 equally parsimonious trees which differ mainly at the lower hierarchical level. Our study reveals two large clades within Anomala. One large clade consists of Galatheoidea and Chirostyloidea. The internal relationships show a monophyletic Porcellanidae nested within a group comprising paraphyletic Galatheidae, and Munididae as well as Munidopsidae. The other large clade contains Aegla as sister group to a monophyletic group consisting of the Hippoidea and a clade formed by Lomis and the Paguroidea. Coenobitidae are nested within paraphyletic Diogenidae and Lithodidae are nested within paraphyletic Paguridae. The results are discussed in the context of other morphological and molecular analyses. Furthermore, some aspects of carcinization are touched upon; in particular, an anomalan stem species with a, at least to some extent, ventrally folded pleon is suggested.     

Phylogeny of Chinese <Emphasis Type="Italic">Polystichum</Emphasis> (Dryopteridaceae) based on chloroplast DNA sequence data (<Emphasis Type="Italic">trnL-F</Emphasis> and <Emphasis Type="Italic">rps4-trnS</Emphasis>)

Polystichum is one of the largest and most taxonomically complex fern genera in China. The evolutionary relationships of Chinese Polystichum and related genera, and the relationship between our Polystichum phylogeny and ecogeographic distribution, were tested by the use of DNA sequence data. Fifty-one species of Polystichum and 21 species in allied genera were sequenced for the plastid intergenic spacers rps4-trnS and trnL-F. Maximum parsimony and Bayesian phylogenetic analyses of both individual and combined data sets showed that Chinese Polystichum as commonly recognized was paraphyletic: one clade (the CCPC clade) included Cyrtomidictyum lepidocaulon, two Cyrtogonellum species, three Cyrtomium species, and a small number of Polystichum species usually occurring on limestone. A second clade, Polystichum sensu stricto, included the remainder of the Polystichum species; these often occur on non-limestone substrates. The remaining Cyrtomium species formed the third clade. Three subclades resolved within Polystichum sensu stricto (s.s.) clade do not correspond with recent sectional classifications, and we outline the issues relevant to a new classification for the genus. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Phylogeny of the Celastreae (Celastraceae) and the relationships of Catha edulis (qat) inferred from morphological characters and nuclear and plastid genes

The phylogeny of Celastraceae tribe Celastreae, which includes about 350 species of trees and shrubs in 15 genera, was inferred in a simultaneous analysis of morphological characters together with nuclear (ITS and 26S rDNA) and plastid (matK, trnL-F) genes. A strong correlation was found between the geography of the species sampled and their inferred relationships. Species of Maytenus and Gymnosporia from different regions were resolved as polyphyletic groups. Maytenus was resolved in three lineages (New World, African, and Austral-Pacific), while Gymnosporia was resolved in two lineages (New World and Old World). Putterlickia was resolved as nested within the Old World Gymnosporia. Catha edulis (qat, khat) was resolved as sister to the clade of Allocassine, Cassine, Lauridia, and Maurocenia. Gymnosporia cassinoides, which is reportedly chewed as a stimulant in the Canary Islands, was resolved as a derived member of Gymnosporia and is more closely related to Lydenburgia and Putterlickia than it is to Catha. Therefore, all eight of these genera are candidates for containing cathinone- and/or cathine-related alkaloids.     

Morphological data indicates two major clades of the subtribe Gorteriinae (Asteraceae-Arctotideae)

The phylogeny of subtribe Gorteriinae (Asteraceae‐Arctotideae) is investigated by means of cladistic analysis of morphological characters. Two sister groups are formed, namely a Gorteria clade also containing Hirpicium and Gazania, and a Berkheya clade, which also contains Cullumia, Cuspidia, Didelta and Heterorhachis. The Gorteria clade has strong jackknife support and is diagnosed by four morphological characters (leaves with longitudinally striate hairs, fringed anther apical appendages, pollen of the “Gazania‐type”, and subulate‐ensiform, ascending style sweeping hairs) that are unique within the Asteraceae. The Berkheya clade is moderately supported and diagnosed by two characters without contradiction (spiny leaves, and mamillate, large style sweeping hairs). Hirpicium and Berkheya are paraphyletic, with the other, morphologically more homogeneous genera (Gorteria, and Gazania, Cullumia, Cuspidia, Didelta and Heterorhachis, respectively) nested within them. There is some evidence for a radiation of species of the summer rainfall area of South Africa and tropical Africa and the corresponding species are nested within a grade confined to the Cape Floristic Region. © The Willi Hennig Society 2006.     

Implications of morphological phylogenetics for the placement of the generaAdenocaulon andEriachaenium (Asteraceae)

Adenocaulon andEriachaenium are two problematic genera because their tribal and subfamilial placement in Asteraceae is uncertain. Previous cladistic analyses based on molecular data undertaken to analyze the relationships within Asteraceae, placeAdenocaulon in the tribe Mutisieae (Cichorioideae). This paper investigates cladistic relationships amongAdenocaulon andEriachaenium and tribes of subfamilies Cichorioideae and Asteroideae using morphological data. Thirty-eight characters were scored across 52 genera selected as exemplar taxa to represent the current classification system. In the analysis (one tree, length = 86, c.i. = 0.55, r.i. = 0.64)Adenocaulon andEriachaenium are sister taxa and appear as an isolated clade nested in Cichorioideae. A new, tentative position among the tribes of the paraphyletic Cichorioideae is proposed for these two isolated genera.     

Phylogenetic relationships among species and genera of Lycoperdaceae based on ITS and LSU sequence data from north European taxa

Phylogenetic relationships, limits of species, and genera within Lycoperdaceae, were inferred by use of ITS and LSU nu-rDNA sequence data. Lycoperdaceae was confirmed as monophyletic, and Mycenastrum corium as a sister taxon to the ingroup. Four major clades were identified and received weak to moderate support and correspond with the genera Lycoperdon, Bovista, Calvatia, and Disciseda. The Lycoperdon clade includes species from Lycoperdon, Vascellum, Morganella, Handkea, Bovistella, and Calvatia. The structure within the Lycoperdon clade is unresolved and several clades are more or less unsupported, which suggests treating the supported Lycoperdon clade as the genus Lycoperdon. L. nigrescens and L. caudatum occur on single branches and their phylogenetic positions could not be resolved. The phylogenetic analyses identified 31 species of Lycoperdon, 11 species of Bovista, six species of Calvatia, and two species of Disciseda. In Lycoperdon three new species were recognized. A new species closely related to B. limosa is identified and discussed. A classification of Lycoperdaceae is proposed based on the results of the phylogenetic analyses. Morphological characters of species within and among identified clades are discussed.     

A phylogeny of Lophocoleaceae-Plagiochilaceae-Brevianthaceae and a revised classification of Plagiochilaceae

The Lophocoleaceae-Plagiochilaceae-Brevianthaceae clade is a largely terrestrial, subcosmopolitan lineage of jungermannialean leafy liverworts that may include significantly more than 1000 species. Here we present the most comprehensively sampled phylogeny available to date based on the nuclear ribosomal internal transcribed spacer region and the chloroplast markers rbcL and rps4 of 372 accessions. Brevianthaceae (consisting of Brevianthus and Tetracymbaliella) form a sister relationship with Lophocoleaceae; this lineage is in turn sister to Plagiochilaceae. Plagiochila is resolved monophyletic subsequent to exclusion of Plagiochila radiculosa; this species is placed in a new genus Cryptoplagiochila. Chiastocaulon and a polyphyletic Acrochila nest in Plagiochilion; these three genera are united under Chiastocaulon to include the Plagiochilaceae species with dominating or exclusively ventral branching. The generic classification of the Lophocoleaceae is still unresolved. We discuss alternative approaches to obtain strictly monophyletic genera by visualizing their consistence with the obtained consensus topology. The presented phylogeny will serve as a basis for follow-up studies including several thousand accessions. These studies will enable revision of current hypotheses on species diversity and distribution of Lophocoleaceae-Plagiochilaceae-Brevianthaceae and allow for a reconstruction of their evolution in time and space.     

Paraphyly of Veronica (Veroniceae; Scrophulariaceae): Evidence from the Internal Transcribed Spacer (ITS) Sequences of Nuclear Ribosomal DNA

Veronica (Veroniceae; Scrophulariaceae) and segregated genera, such as Hebe from New Zealand has been debated intensively in the past. We conducted an analysis of sequence data from the internal transcribed spacer region of nuclear ribosomal DNA (ITS) to evaluate the validity of segregate genera and the monophyly of Veronica. According to the results presented here, Veronica is paraphyletic, with the Hebe complex, Synthyris, and Paederota nested within the larger Veronica clade. Pseudolysimachion is in a basal polytomy of the expanded Veronica clade in the strict consensus tree and might be nested within Veronica as well. Clades within Veronica do not correspond to sections traditionally recognized. This study provides a first estimation of the phylogeny of Veroniceae using molecular data and can serve as a starting point for future investigations of Veronica and relatives. Received 24 July 2000/ Accepted in revised form 19 October 2000     

Ulocladium systematics revisited: phylogeny and taxonomic status

The genus Ulocladium represents phaeodictyosporic Hyphomycetes that produce conidia that are essentially obovoid in shape. Previous molecular studies that included Ulocladium and related taxa in Alternaria, Embellisia, and Stemphylium revealed a conflict between morphology and phylogeny, and Ulocladium was supported as polyphyletic with a paraphyletic core group. Moreover, the genus consistently resolved within a larger Alternaria/Ulocladium clade, resulting in paraphyly of Alternaria and questions as to the taxonomic status of Ulocladium. In the present study, 13 Ulocladium species and three genetic loci were included for a more comprehensive systematic analysis of the genus than had previously been conducted. Total genomic DNA was extracted from representative taxa and sequences were determined for the nuclear internal transcribed spacer region, including the 5.8S rDNA gene, and the protein-coding genes glyceraldehyde-3-phosphate dehydrogenase and Alt a1. Subsequent phylogenetic analyses based on maximum parsimony and Bayesian methods included related Alternaria, Embellisia, and Stemphylium spp. Results supported previous findings of polyphyletic and paraphyletic relationships of Ulocladium among other taxa. Ten Ulocladium species clustered into a core Ulocladium clade and all taxa possessed the key diagnostic feature of Ulocladium, namely, conidia essentially obovoid in shape. However, A. cheiranthi and E. indefessa also clustered within this group with high bootstrap support but did not posses this diagnostic feature. This paraphyletic clade resolved basal to the core Alternaria clade with high bootstrap support, unlike previous studies in which its position was imbedded within the primary Alternaria clade. Thus, the status of the genus as an independent lineage and a unique taxon is strongly supported. As previously reported, U. alternariae and U. oudemansii, which posses the key conidium characteristics of Ulocladium, clustered as a separate clade sister to the core Ulocladium clade. Further studies are necessary to determine if these taxa represent an independent lineage or share a common ancestor with other Ulocladium species. Obovoid conidia were poorly represented in the isolate of U. lanuginosum that was included in these analyses (the only U. lanuginosum isolate currently available), and the isolate resolved as A. radicina based upon all three loci sequenced. Based upon these data and the origin of the isolate, which was originally deposited as A. malvae, a reassessment of its identity is supported.     

Genetic diversity and phylogenetic position of the subclass Astomatia (Ciliophora) based on a sampling of six genera from West African oligochaetes (Glossoscolecidae, Megascolecidae), including description of the new genus Paraclausilocola n. gen

To more confidently assess phylogenetic relationships among astome ciliates, we obtained small subunit (SSU) rRNA sequences from nine species distributed in six genera and three families: Almophrya bivacuolata, Eudrilophrya complanata, Metaracoelophrya sp. 1, Metaracoelophrya sp. 2, Metaracoelophrya intermedia, Metaradiophrya sp., Njinella prolifera, Paraclausilocola constricta n. gen., n. sp., and Paraclausilocola elongata n. sp. The two new species in the proposed new clausilocolid genus Paraclausilocola n. gen. are astomes with no attachment apparatus, two files of contractile vacuoles, and an arc-like anterior suture that has differentiations of thigmotactic ciliature on the anterior ends of the left kineties of the upper surface. Phylogenetic analyses were undertaken using neighbor-joining, Bayesian inference, maximum likelihood, and maximum parsimony. The nine species of astomes formed a strongly supported clade, showing the subclass Astomatia to be monophyletic and a weakly supported sister clade to the scuticociliates. There were two strongly supported clades within the astomes. However, genera assigned to the same family were found in different clades, and genera assigned to the same order were found in both clades. Thus, astome taxa appear to be paraphyletic when morphology is used to assign species to genera.     

Systematics and phylogeny of Vasseuromys (Mammalia,Rodentia, Gliridae) with a description of a new species from the late Miocene of eastern Europe

Vasseuromys is a species‐rich genus of small‐ to medium‐sized glirids spanning the latest Oligocene to late Miocene of Europe and western Asia. Despite extensive discoveries over the past 50 years, little phylogenetic work has been done on Vasseuromys. This study presents the first phylogenetic analysis of the genus that includes all the described species and a new taxon Vasseuromys tectus sp. nov. from the late Miocene of eastern Europe, providing the first insights into the evolutionary relationships within the clade. Results suggest that the genus is clearly paraphyletic. Two strongly supported genus‐level clades are recognized within ‘Vasseuromys’: a restricted Vasseuromys clade (containing the three species, V. pannonicus, V. rugosus and V. tectus) and the Glirulus clade that includes ‘Vasseuromys’ duplex. The remaining ‘Vasseuromys’ species are found to constitute a set of paraphyletic taxa, with the polyphyletic ‘Ramys’ nested within it. The genus Gliruloides is synonymized with Glirulus. Vasseuromys tectus sp. nov. is the most derived member of the genus in having a greater number of cheek teeth ridges including constantly present anterotrope, centrotrope, second prototrope on M1–2, third metatrope on M2, two to three posterotropids on p4 and strong ectolophids on lower molars. The results of the study confirm a European origin for Vasseuromys while suggesting that the late Miocene species of the genus dispersed from the east in the early Turolian.     

Phylogenetic relationships and biogeographic patterns in North American members of Carex section Acrocystis (Cyperaceae) using nrDNA ITS and ETS sequence data

Carex section Acrocystis currently includes 27 taxa in North America. Recent phylogenetic studies have suggested that the North American and some but not all of the Eurasian species form a clade. Relationships and biogeographic patterns among species in this core-Acrocystis group are explored here using nuclear ribosomal (nrDNA) internal transcribed spacer region (ITS) and nrDNA external transcribed spacer region (ETS) sequence data. While maximum parsimony analysis of the ITS and ETS data provides only a moderately resolved branching structure for species relationships within the core-Acrocystis clade, maximum likelihood analysis provides a more resolved hypothesis of relationships in the section. The core-Acrocystis clade consists of a grade of Eurasian and primarily western North American species, with a well-supported clade of only eastern North American species nested within this grade. ITS and ETS types do not coalesce within many species or species complexes. Possible explanations for the non-coalescent nature of ITS and ETS copies in Acrocystis are explored, including lineage sorting, hybridization, and cryptic species.     

A Total Evidence Phylogeny of the Arctoidea (Carnivora: Mammalia): Relationships Among Basal Taxa

A total evidence phylogenetic analysis was performed for 14 extant and 18 fossil caniform genera using a data matrix of 5.6 kbp of concatenated sequence data from six independent loci and 80 morphological characters from the cranium and dentition. Maximum parsimony analysis recovered a single most parsimonious cladogram (MPC). The topology of the extant taxa in the MPC agreed with previous molecular phylogenies. Phylogenetic positions for fossil taxa indicate that several taxa previously described as early members of extant families (e.g., Bathygale and Plesictis) are likely stem taxa at the base of the Arctoidea. Taxa in the “Paleomustelidae” were found to be paraphyletic, but a monophyletic Oligobuninae was recovered within this set of taxa. This clade was closely related to the extant genera Gulo and Martes, therefore, nested within the extant radiation of the family Mustelidae. This analysis provides a resolution to several discrepancies between phylogenies considering either fossil taxa or extant taxa separately, and provides a framework for incorporating fossil and extant taxa into comprehensive combined evidence analyses.     

Phylogenetic relationships in Blumea (Asteraceae: Inuleae) as evidenced by molecular and morphological data

The present study, based on sequences of cpDNA (trnL-F & psbA-trnH) and nrDNA (ITS) and morphology, examined the evolutionary relationships in Blumea and its position among related genera. The results confirmed that the closest relatives of Blumea are Caesulia, Duhaldea and Pentanema p.p., and showed that the monotypic genera Blumeopsis and Merrittia are nested within Blumea. In Blumea s.l., two major, well-supported clades were recognised and a single species, the widespread Blumea balsamifera, that could not be placed with certainty relative to the two main clades. The two main clades differ in habit, ecology and distribution. The Blumea densiflora clade contains shrubs and subshrubs of evergreen forests, distributed from continental Asia to New Guinea and Polynesia, whereas the Blumea lacera clade is a widespread paleotropical group that comprises mostly annual, weedy herbs of open forests and fields.