Phylogenetic analysis and revision of the linyphiid spider genus Solenysa (Araneae: Linyphiidae: Erigoninae)

In this paper we carry out a taxonomic revision and phylogenetic analysis of the linyphiid spider genus Solenysa Simon, 1894. A total of 12 species is treated here, including five new species collected from China and Japan: Solenysa akihisai Tu sp. nov., Solenysa lanyuensis Tu sp. nov., Solenysa retractilis Tu sp. nov., Solenysa tianmushana Tu sp. nov. , and Solenysa yangmingshana Tu sp. nov. Solenysa circularis Gao, Zhu & Sha, 1993 is a junior synonym of Solenysa protrudens Gao, Zhu & Sha, 1993. We have assembled two different character matrices to reconstruct the phylogenetic relationships of Solenysa. In the first matrix (Matrix 1), five representative species of Solenysa were added to the morphological dataset of Miller & Hormiga to test the monophyly of the genus and its placement within Linyphiidae. The genitalic structures and somatic morphology of Solenysa were studied by means of scanning electron microscopy for the first time. To infer the species‐level phylogenetic relationships of Solenysa we produced a second matrix (Matrix 2) that includes all 12 Solenysa species and six outgroup species chosen from the results of the analysis of the first matrix. The two most parsimonious trees from the analysis of Matrix 1 support the monophyly of Solenysa and its placement within the ‘Distal Erigonines’ clade. The single most parsimonious tree resulting from the analysis of the second matrix suggests that the Solenysa clade includes four monophyletic groups, each group represented by a distinct genitalic pattern. The morphology of Solenysa, both somatic and genitalic, is highly autapomorphic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 484–530.     

New mammal remains from the Late Cretaceous Bostobe Formation (Northeast Aral Sea Region,Kazakhstan)

Four recently collected mammal specimens from the Upper Cretaceous (Santonian–?Campanian) Bostobe Formation in the northeastern Aral Sea Region, Kazakhstan are attributed to Asioryctitheria indet. (an edentulous dentary fragment) and the zhelestid Parazhelestes sp. cf. P. mynbulakensis (a maxillary fragment with a double-rooted canine, an M1, and a dentary fragment including m3). These new records double the known mammal fauna from this formation, which previously included the zhelestid Zhalmouzia bazhanovi and Zhelestidae indet. The taxonomic and ecological structure of the mammal assemblage from the Bostobe Formation can, on present evidence, be considered close to the other eutherian dominated Late Cretaceous mammal assemblages of Central Asia. This region is important in particular in the search for Late Cretaceous ancestors of crown-group eutherian mammal clades (Placentalia).     

Phylogeny of Early Cretaceous spatangoids (Echinodermata: Echinoidea) and taxonomic implications

A phylogenetic analysis of 36 species provides a test for the taxonomy and the history of Early Cretaceous spatangoids. Most taxonomic units from genera to suborders are consistent with the proposed phylogenetic framework. We retain Hemiasterina, Micrasterina, Hemiasteridae, Schizasteridae, Hemiaster , Heteraster , Mecaster , and Periaster as original monophyletic groups. However, all of these clades originate without the classical apomorphies normally ascribed to them. We suggest a revision of their diagnoses and of the generic attributions of basal species. Some ill-defined, 'primitive', and paraphyletic taxa are recognised: Toxaster , Epiaster , Palhemiaster , and Toxasteridae. Even if they do not have phylogenetic meaning, they are retained here, pending a more complete revision.     

Phylogenetic analysis and a time tree for a large drosophilid data set (Diptera: Drosophilidae)

Drosophila is the genus responsible for the birth of experimental genetics, but the taxonomy of drosophilids is difficult because of the overwhelming diversity of the group. In this study, we assembled sequences for 358 species (14 genera, eight subgenera, 57 species groups, and 65 subgroups) to generate a maximum‐likelihood topology and a Bayesian timescale. In addition to sampling an unprecedented diversity of Drosophila lineages, our analyses incorporated a geographical perspective because of the high levels of endemism. In our topology, Drosophila funebris (Fabricius, 1787) (the type species of Drosophila) is tightly clustered with the pinicola subgroup in a North American clade within subgenus Drosophila. The type species of other drosophilid genera fall within the Drosophila radiation, presenting interesting prospects for the phylogenetic taxonomy of the group. Our timescale suggests that a few drosophilid lineages survived the Cretaceous–Palaeogene (K‐Pg) extinction. The drosophilid diversification began during the Palaeocene in Eurasia, but peaked during the Miocene, an epoch of drastic climatic changes. The most recent common ancestor of the clades corresponding to subgenera Sophophora and Drosophila lived approximately 56 Mya. Additionally, Hawaiian drosophilids diverged from an East Asian lineage approximately 26 Mya, which is similar to the age of the oldest emerging atoll in the Hawaiian–Emperor Chain. Interestingly, the time estimates for major geographical splits (New World versus Asia and Africa versus Asia) were highly similar for independent lineages. These results suggest that vicariance played a significant role in the radiation of fruit flies. © 2013 The Linnean Society of London     

Reexamination of the morphological evidence for the cohort Epitheria (Mammalia,Eutheria)

Novacek and co-workers recognized a monophyletic clade Epitheria, comprising all eutherians except edentates and the extinct palaeoryctoids, on the basis of two synapomorphies: a stirrupshaped stapes and a foramen ovale enclosed within the alisphenoid. To evaluate this phylogenetic hypothesis, we reexamined the distributions of stapedial morphologies and positions of the foramen ovale across Recent and extinct mammals and nonmammalian cynodonts. The states and distributions of the stapes and forament ovale characters used by Novacek and coworkers were modified by recognizing two stapedial characters (one relating to shape of the crura, the other to the nature of the foramen) and a single, multistate foramen ovale character (within, behind, and lateral to the alisphenoid). The taxon-character matrix used by Novacek (1989, 1992b), substituting our amended stapedial and foramen ovale characters and adding several previously unscored extinct taxa and three new characters, was subjected to a series of PAUP manipulations. Identified among the most parsimonious trees were three major topologies for the base of Eutheria: (1) a polytomy including an Edentata/Ungulata clade, (2) a polytomy with Edentata and Ungulata as separate clades, and (3) Edentata and (when included) Palaeoryctoidea as the successive outgroups to a monophyletic Epitheria. We conclude that topology 2 best reflects the current state of knowledge. An edentate/ungulate clade is supported by three characters (from the mastoid region and subarcuate fossa); however, other morphological studies require modification of the distributions of these characters in xenarthrans and bassal ungulates, thereby eliminating support for this clade. In nearly all manipulations, obtaining a monophyletic Epitheria required that one or two steps be added to the most parsimonious trees. When a monophyletic Epitheria was obtained, it was supported by a triangular stapes and, in some trees, the reappearance of a stapedial artery (lost earlier at the level of Recent therians) and a transpromontorial internal carotid artery. In the most parsimonious trees, a foramen ovale within the alisphenoid was an equivocal synapomorphy of Recent therians or cutherians, and a stapes with strongly convex crura (our state closest to the stirrup-shaped state of Novacek and co-workers) appeared independently within various eutherian lineages. The reduction or loss of the stapedial foramen was identified as an independent event in monotremes and within marsupials and various eutherian lineages.To whom correspondence should be addressed.     

Phylogenetic analysis of the Mexican freshwater crabs of the tribe Pseudothelphusini (Decapoda: Brachyura: Pseudothelphusidae)

The Mexican pseudothelphusid crabs are classified in one subfamily, three tribes, and 13 genera. Up to now, 56 species have been recognized, distributed in a strictly Neotropical pattern, with some of them reaching the state of Sonora on the western slope of Mexico. The tribe Pseudothelphusini is the most diverse, with five genera and 35 species, all of them endemic to Mexico: the two most species‐rich genera are Pseudothelphusa, with 23 species, and Tehuana, with eight species; Epithelphusa includes two species, whereas Disparithelphusa and Smalleyus are monotypic. The Pseudothelphusini lack an updated systematic revision, which could serve as a framework to analyse the monophyletic origin of the group, to clarify the relationships among genera and species, as well as to resolve the taxonomic status of various species complexes. In the present study, an exhaustive morphological revision was conducted using somatic and sexual characters. A phylogenetic analysis was performed using 77 characters and 183 character states, taken from 41 species. Ten trees of the same length were obtained using PAUP 4.0 through a heuristic search. The results show that the tribe as it is actually known constitutes a paraphyletic group, in which the species of Epithelphusa and Pseudothelphusa puntarenas are excluded from the internal group. According to the obtained results, the tribe Pseudothelphusini s.s. includes five genera: Smalleyus, Pseudothelphusa, Tehuana, and two new ones to accommodate Pseudothelphusa galloi and Pseudothelphusa sulcifrons, respectively. This new arrangement considers the provisional suppression of the genus Disparithelphusa, which remained as another species of Pseudothelphusa throughout the cladistic analysis. The phylogenetic results show a strong congruence with the distribution of the species, in several cases grouping species that form morphological clines along a geographical gradient. The previously proposed southern origin of the tribe Pseudothelphusini gains support with the results obtained. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 457–481     

Cretaceous Therian Tarsals and the Metatherian-Eutherian Dichotomy

Diverse metatherian and eutherian tarsal remains from the Late Cretaceous (middle-late Turonian) Bissekty Formation, Kyzylkum Desert, Uzbekistan (ca 90 MYA) are described. Their functional and taxonomic properties, along with those of other tarsal evidence, led to a reassessment of polarity hypotheses of therian, metatherian, and eutherian cruropedal attributes, and the consequences of this for phylogeny of taxa. There are calcaneal remains of several types of marsupials, and a single astragalus that probably belongs to one of these. This represents greater taxonomic diversity than the dental record suggests. Exceptionally large and distally extending peroneal processes, and small and steeply angled calcaneocuboid articulations facing mediodistally, as seen in the Early Cretaceous Sinodelphys and other Cretaceous and Paleogene taxa, attest not only to the metatherian status of these specimens, but also to the retention of many ancestral therian features, even more so than in both the Tiupampa and Itaboraí marsupials of the South American Paleocene (both the calcanea and the astragalus suggest therian traits that are decidedly unlike those of symmetrodonts). Calcanea allocated to the deltatherioid species at Bissekty testify unequivocally to their metatherian affinity. The morphology of the best represented sample of eutherian calcanea from Bissekty, presumably of a number of zhelestid species, appears to be more derived than that of the Late Cretaceous/Paleocene Protungulatum in having a much more reduced peroneal process and a calcaneocuboid articulation that faces distally, oriented nearly at a 90° angle to the long axis of the calcaneus. In fact, this distally facing facet, common in later eutherians (except for lineages in the Paleogene record, and various Carnivora), may not be diagnostic of either the protoeutherian, or even of the protoplacentalian, in spite of its presence in Eomaia. Many putatively “basal” lineages have derived characters, hence such outgroups should not be considered the unequivocal repositories of only ancestral character states.     

晚白垩世精美临河盗龙(Linheraptor exquisitus)的分类位置及其对驰龙科系统学的意义（英文）

驰龙类的精美临河盗龙(Linheraptor exquisitus)于2010年基于一件近完整骨架命名,化石采自内蒙古西部巴彦满达呼大门地点上白垩统乌兰苏海组。然而,最近三项研究认为精美临河盗龙是蒙古乌哈托喀地点上白垩统捷达克赫塔组发现的白魔龙(Tsaagan mangas)的晚出异名。本文依据61个形态学特征区分精美临河盗龙和白魔龙,否认了上述观点。许多特征来自对精美临河盗龙正型标本先前未修理部分,特别是头骨左侧的观察。这些新观察支持和加强了我们先前认为的精美临河盗龙和白魔龙属于两个不同属种的观点。精美临河盗龙具有的一些特征之前被认为是白魔龙或者其他驰龙类的自近裔特征,这表明进步的骨骼特征在驰龙类中的分布异常复杂。认为精美临河盗龙和白魔龙为同物异名的观点实际上忽略了很多微小的形态变异。增加分类取样可以将那些看似显著的鉴定特征转化为更加细微的形态变异,而这些形态变异对于精细的系统发育分析具有潜在的重要意义。在未来的研究中,利用连续数据的严格定量方法,有助于处理类似信息。     

A revision and phylogenetic analysis of Stapeliopsis (Apocynaceae)

A survey of morphological characters is carried out for Stapeliopsis . The information obtained from this is combined with molecular data from the plastid trn L-F DNA region and ITS1 of the nuclear encoded 18S−26S rRNA cistron, to obtain a hypothesis of the evolutionary relationships among the species. It is shown that Stapeliopsis is monophyletic in a combined molecular and morphological analysis. Stapeliopsis is sister to a clade containing Huernia , Orbea and Tromotriche . The species of Stapeliopsis group into two clades. One contains S. khamiesbergensis , S. neronis and S. urniflora , and this is highly supported. The remaining species fall into an unsupported clade in which S. exasperata is sister to the others. The genera Hermanschwartzia Plowes and Neopectinaria Plowes are rejected. It is shown that a synapomorphy for Stapeliopsis is the laterally flattened inner corona-lobes, which touch the anthers only at their bases. Eight species of Stapeliopsis are recognized, with no subgeneric divisions.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 125–155.     

New choristoderes (Reptilia: Diapsida) from the Upper Cretaceous and Palaeocene, Alberta and Saskatchewan, Canada, and phylogenetic relationships of Choristodera

New choristoderan fossils from the Late Cretaceous and Palaeocene of Alberta and Saskatchewan, Canada, are described: incomplete maxillae and dentaries from the Upper Cretaceous Oldman Formation, Alberta, extend the range of the primitive Cteniogenys from the Jurassic in the North American Western Interior; an incomplete dentary from the Palaeocene Ravenscrag Formation, Saskatchewan, comprises the first occurrence of the crocodile-like Simoedosaurus in Canada and the earliest record of the genus; well-preserved skulls and mandibles from the Oldman and Horseshoe Canyon formations, Alberta, document a new species of Champsosaurus and clarify the status of previously known species of the genus. New information about Asian choristoderes supports a Tchoiria- ( Ikechosaurus + Simoedosaurus ) relationship, contrary to previous work. Choristoderes share no convincing synapomorphies with either Lepidosauromorpha or Archosauromorpha, but occupy a more basal position within Diapsida, possibly as a sister-taxon with Neodiapsida (Younginiformes + Sauria).     

A revision of the genus Dinofelis (Mammalia, Felidae)

This paper consists of a taxonomic and systematic revision of the extinct felid genus Dinofelis (Felidae, Ma-chairodontinae) and an analysis of its ecomorphology and evolution. Dinofelis has a broad distribution, with material from all northern continents and Africa, the latter of which was the apparent centre of evolution of the genus. We describe new material of Dinofelis from a number of sites in eastern Africa and reconsider all previously described material. We name two new species and identify several other distinct species-level taxa but refrain from naming these due to a paucity of well-preserved material. At the same time, we synonymize the two named Asian species, D. cristata and D. abeli , of which the former has priority. There are few characters useful in systematic analysis, but we can suggest at least one migration from eastern to southern Africa. Ecomorphological analysis of both craniodental and postcranial characters suggests that Dinofelis in many respects converged on modern pantherine cats in morphology and behaviour, a trend culminating in the South African D. barlowi and the Asian D. cristata , which are the most pantherine-like of all machairodont felids. This trend is reversed in the evolution of the youngest species, D. piveteaui , which is also the most machairodont in its ecomorphology. The timing of the extinction of Dinofelis is difficult to determine. Outside Africa material is scarce at all times, while in Africa the apparent extinction of Dinofelis at about 1.4 Mya coincides with the end of the good, semi-continuous fossil record present in eastern Africa from about 4 Mya onwards. Dating of Kanam East (with D. piveteaui) to the Jaramillo Subchron (1.070-0.990 Mya) suggests possible survival considerably later. Thus, the extinction datum for Dinofelis cannot at present be firmly established.     

Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence

This study is undertaken in order to evaluate specific hypotheses of relationship among extant and extinct sloths (Mammalia, Xenarthra, Tardigrada). Questions of particular interest include the relationship among the three traditional family groupings of extinct ground sloths and the monophyletic or diphyletic origin of the two genera of extant tree sloths. A computer‐based cladistic investigation of the phylogenetic relationships among 33 sloth genera is performed based upon 286 osteological characteristics of the skull, lower jaw, dentition and hyoid arch. Characters are polarized via comparisons with the following successive outgroups, all members of the supraordinal grouping Edentata: the Vermilingua, or anteaters; the Cingulata, or armadillos and glyptodonts; the Palaeanodonta; and the Pholidota, or pangolins. The results of the analysis strongly corroborate the diphyly of living tree sloths, with the three‐toed sloth Bradypus positioned as the sister‐taxon to all other sloths, and the two‐toed sloth Choloepus allied with extinct members of the family Megalonychidae. These results imply that the split between the two extant sloth genera is ancient, dating back perhaps as much as 40 Myr, and that the similarities between the two taxa, including their suspensory locomotor habits, present one of the most dramatic examples of convergent evolution known among mammals. The monophyly of the three traditional ground sloth families Megatheriidae, Megalonychidae and Mylodontidae is confirmed in the present study, and the late Miocene–Pleistocene nothrotheres are shown to form a clade. It is suggested that this latter clade merits recognition as a distinct family‐level grouping, the family Nothrotheriidae. The monophyly of the Megatherioidea, a clade including members of the families Megatheriidae, Megalonychidae and Nothrotheriidae, is also supported. Within Megatherioidea, the families Nothrotheriidae and Megatheriidae form a monophyletic group called the Megatheria. The relationships within the families Megatheriidae and Mylodontidae are fully and consistently resolved, although the hypothesized scheme of relationships among the late Miocene to Pleistocene members of the mylodontid subfamily Mylodontinae differ strongly from any proposed by previous authors. Within the family Megalonychidae, Choloepus is allied to a monophyletic grouping of West Indian sloths, although the relationships within this clade are not fully resolved. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 255–305.     

About some aporrhaid and strombid gastropods from the Late Cretaceous

Sixteen species of the Stromboidea (Gastropoda) from the Late Cretaceous are described in this study. One species belongs to the Strombidae, the others to eight genera and subgenera of the Aporrhaidae. Eight species are new. They have been found in the Cenomanian of Kassenberg in western Germany, the Coniacian of the Mungo River in Cameroon, the Santonian of the Eastern Cape Province of South Africa, the Santonian/Campanian of the Ariyalur area in southern India, the Campanian of northern Spain, and the Maastrichtian of the Western Desert in Egypt. Two species ofDrepanocheilus from South Africa are considered stem-group representatives of the aporrhaids which inhabited the Weddellian Province in the early Tertiary. One new species of the strombid genusHippocrenes is regarded as an early species of the evolutionary lineage leading to the Seraphsidae. The new species areLatiala? ponsi, Drepanocheilus herberti, Drepanocheilus triliratus, Drepanocheilus (Tulochilus) jouberti, Graciliala quaasi, Kaunhowenia catalanica, Kaunhowenia punctata andHippocrenes kussi.     

Molecular phylogenetics and taxonomic revision of Habenaria section Pentadactylae (Orchidaceae,Orchidinae)

As a step towards a revision of the sectional classification of Neotropical species of Habenaria, we focus here on section Pentadactylae. In its current delimitation, this is the largest of the 14 New World sections and embraces a group of 34 morphologically heterogeneous species. We expanded the sampling of Neotropical species currently placed in this section and performed Bayesian, maximum likelihood and parsimony analyses using nucleotide sequences from one nuclear (internal transcribed spacer, ITS) and three plastid (matK, trnK intron, rps16–trnK) DNA regions. In addition, morphological features of these species were reassessed. Based on our analyses, we propose that Habenaria section Pentadactylae should be recircumscribed to include only seven species: H. pentadactyla (the type species of the section), H. dutrae, H. ekmaniana, H. exaltata, H. henscheniana, H. megapotamensis and H. montevidensis. Thirty‐two species previously assigned to the section grouped within unrelated clades and are therefore excluded from the section. There are no unambiguous morphological synapomorphies for the section, but the group can be confidently recircumscribed and identified on the basis of a combination of diagnostic morphological vegetative and floral characters. Morphological floral features in Habenaria montevidensis are distinct from those of other species in the section, probably as a result of a shift to diurnal pollinators. Following a taxonomic revision of the group, H. crassipes is placed under the synonymy of H. exaltata and neotypes are designated for H. crassipes, H. montevidensis and H. recta (= H. ekmaniana). All species in the section live in marshes or wet grasslands from northern Argentina to central Brazil; most species are concentrated in southern Brazil. Most species are probably rare, and five may be threatened according to the World Conservation Union (IUCN) criteria. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 47–73.     

A preliminary phylogeny of the Pterasteridae (Echinodermata,Asteroidea) and the first fossil record: Late Cretaceous of Germany and Belgium

The Pterasteridae comprises a diversified group of extant largely deep-sea starfishes. Generic diagnoses have been based classically on soft tissue characters and skeletal architecture. A preliminary phylogeny of sixteen extant species is here worked out by cladistic analysis. The resulting tree suggests monophyly of extant genera and the validity of dissociated plates for identification of genera. Fossil remains of Pterasteridae are here described for the first time. By comparison with extant species, all the skeletal remains from the lower Upper Campanian of Belgium and from the lower Maastrichtian of Germany are tentatively assigned to the genusPteraster. The fossil record of starfishes is poor, but the present Late Cretaceous pterasterids provide one more piece of evidence of the high diversity of starfishes during the Mesozoic. Known Late Cretaceous and Paleogene fossils are broadly similar, which suggests the end-Cretaceous extinction event did not cause major turnover in asteroid faunal composition. As suggested for other starfish groups, both the fossil record of deep-sea Pterasteridae in shelf settings and tree topology imply an onshore-offshore evolutionary trend.      

Phylogenetic analysis of eleven species of Biomphalaria Preston, 1910 (Gastropoda: Planorbidae) based on comparisons of allozymes

Freshwater snails in the genus Biomphalaria transmit Schistosoma mansoni in Africa, South America and the Caribbean region. Although considerable attention has been given to the identification of species, little is known of evolutionary relationships among the species. A phylogenetic analysis of 25 populations representing 11 species was performed on 25 enzyme loci examined using starch gel electrophoresis. A phylogenetic analysis of the individual populations produced 60 trees of equal length. The 60 trees have a consistency index value of 75.9% and a retention index value of 76.5%. The phylogenetic analysis provided strong support for the monophyly of Biomphalaria with either 14 or 15 synapomorphies uniting all of the species included and separating them from the outgroup, two species of Helisoma. Four nominal species represented by multiple populations formed monophyletic groups. Populations of B. sudanica, B. choanomphala, and B. alexandrina were interspersed. Ten arrangements were obtained for the populations of these three species. A variety of ingroup taxa were used to root the trees, and all provided support for the use of Helisoma species as an outgroup. In all of the trees obtained, the African species together formed a monophyletic group. In none of the trees obtained did the neotropical species form a monophyletic group. A constrained analysis requiring the monophyly of the neotropical species as well as the African species resulted in 90 trees just two steps longer than the shortest trees. Analysis of the species from either hemisphere alone resulted in decreased resolution, as measured by an increase in the number of trees obtained. This finding suggests that further comparisons of species from the two hemispheres will be of considerable value. Finally, two species which are resistant to infection with S. mansoni were included among the eleven studied. Neither of these species formed the sister group to all of the other species included, indicating that susceptibility is the plesiomorphic state, and that resistance is derived. Similarly, in none of the trees obtained did the two resistant species fall out as sister taxa, indicating that resistance arose independently twice.