Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

Systematics of African lowland rainforest Praomys (Rodentia, Muridae) based on molecular and craniometrical data

Representatives of the genus Praomys occur throughout the African intertropical zone. It is unclear how many species this genus contains, nor do we know the exact distribution ranges and phylogenetic relationships of these taxa. Using molecular (16S rRNA gene sequencing) and morphological (multivariate craniometry) analyses we clarify the taxonomy and phylogenetic relationships among the Praomys occurring in Africa's lowland tropical rainforests. We studied most species known from this area, based on specimens collected in seven countries (Guinea, Ivory Coast, Cameroon, Central African Republic, Gabon, Congo Republic and Democratic Republic of Congo). In our study, Praomys appears to be monophyletic. Our results identify two species complexes: the jacksoni complex includes at least two species ( P. jacksoni and P. mutoni ) and the tullbergi complex contains at least four species ( P. tullbergi , P. rostratus , P. misonnei , P. petteri ). Although the 16S rRNA gene appears insufficient to resolve the phylogenetic relationships among all the members of the tullbergi species complex, it is suitable for the identification of most of the studied species, and its use has allowed us to redefine the geographical limits of several species.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 539–553.     

Delimiting species in the hoplonemertean genus Tetrastemma (phylum Nemertea): morphology is not concordant with phylogeny as evidenced from mtDNA sequences

The marine genus Tetrastemma contains monostiliferous hoplonemertean (phylum Nemertea) species which mostly undergo direct development with no free-swimming stages of larvae. The lack of a pelagic phase, and the fact that many benthic species lay eggs attached to the bottom substrate, are obvious restrictions on dispersal and gene flow. Nevertheless, some of the species, for example T. candidum and T. melanocephalum , are described as ubiquitous and are reported from waters all over the world. We studied genetic variation and evolutionary relationships in order to assess whether they are concordant with external characters in samples of nine morphologically distinct forms formally named as different Tetrastemma species, from different geographical localities. We estimated the phylogeny and species network based on 539 base pairs of the mitochondrial protein-coding gene cytochrome oxidase-1 (CO1) for 30 ingroup specimens. From this, we assessed the evolutionary history and phylogenetic relationships between these forms. We conclude that in most cases there was no correspondence between evolutionary lineage and morphotype. Our results thus indicate that morphological species delimitation in nemerteans may be questionable, and that this in turn may have a profound effect upon estimates of species diversity within the phylum.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 201–212.     

A revision of Cerdia (Caryophyllaceae)

Cerdia is a poorly understood genus of minute cushiony plants endemic to Mexico that is traditionally placed in subfamily Paronychioideae, tribe Polycarpeae (Caryophyllaceae). Morphological observations and phylogenetic analysis of ITS DNA sequences were carried out to determine the number of species, heretofore controversial, that constitute the genus as well as its position within Caryophyllaceae. Samples of Cerdia were collected through its entire range of distribution and together with previous specimens were used in morphological analyses. Nineteen terminal taxa were considered in a phylogenetic analysis with representatives of the different lineages in Caryophyllaceae. Results indicated that Cerdia is a monotypic genus, including only one variable species, C .  virescens , but its phylogenetic affinities remain doubtful.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 1–13.     

Phylogenetic relationships in the subgenus Mus (genus Mus, family Muridae, subfamily Murinae): examining gene trees and species trees

Over eight kilobases (kb) of sequence from eight genes including two mitochondrial loci, Cyt b and 12S, and six nuclear loci, B2m , Zp3 , Tcp1, Sry, Smcx and Smcy , were used to investigate phylogenetic relationships among 11 taxa representing eight species within the rodent genus Mus . Particular attention was given to discerning relationships among species within the subgenus Mus including members of a Palearctic clade ( M. musculus , M. spicilegus , M. macedonicus and M. spretus ) and members of an Asian clade ( M. caroli , M. cookii and M. cervicolor ), as previous studies using different datasets have produced different topologies for taxa within these two groups. While parsimony analyses of the combined eight-gene dataset yielded a single, fully resolved tree, support values were lower for nodes resolving relationships within the Palearctic and Asian clades than they were elsewhere in the tree. In addition, a maximum likelihood analysis of the same eight-gene dataset yielded different topologies for both the Palearctic and the Asian clades. Both observations are indicative of clade instability. The nature of this instability was explored through a comparison with our previous study in which we included the two mitochondrial loci and only four of the six nuclear genes, and through an analysis of partitioned data, specifically mitochondrial vs. nuclear genes. This study underscores the importance of considering among-site rate variation in phylogeny reconstruction. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 653–662.     

A molecular approach to the phylogenetic relationships of the western palaearctic Helicoidea (Gastropoda: Stylommatophora)

Molecular phylogenetic relationships among 45 members of the Helicoidea (Gastropoda: Stylommatophora) were examined using partial mitochondrial 16S rRNA sequences. Phylogenetic relationships were inferred using maximum parsimony, maximum likelihood and Bayesian methods. The reconstructed phylogenies showed a good degree of support for more recent branches, but gave little support to deeper nodes. Mitochondrial rDNA data further confirmed monophyletic status of helicids, recognized monachine hygromiid and bradybaenid clades and resolved a number of relationships in the helicelline hygromiids. With the respect to the latter assemblage, most of the anatomically based groups are confirmed, corroborating the diagnostic value of the dart-sac complex and a close affinity between Ichnusomunda sacchii and species of the genus Cernuella . Nevertheless, some well resolved branches challenge previous systematic arrangements, grouping species previously placed in different arrangements. In particular, support was not found for the monophyly of helicelline hygromiids with pedal penial innervation. Possible explanations for these incongruencies are suggested. 16S sequence data are appropriate for studies of relationships within the different species groups and less so for recovery of more ancient radiations in the Helicoidea. It will be valuable to combine the 16S data with other gene sequences to estimate basal relationships.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 501–512.     

Molecular phylogeny of the genus Mus (Rodentia: Murinae) based on mitochondrial and nuclear data

The genus Mus encompasses at least 38 species divided into four subgenera: Mus , Pyromys , Nannomys and Coelomys . The subgenus Mus , which comprises the house mouse and related species, is by far the most extensively studied, although the subgenus Nannomys is the most speciose. Although the relationships within the subgenus Mus are rather well characterized, those between subgenera are still unclear. In the present study, phylogenetic analyses of the whole genus were performed using a larger species sample of Nannomys than in previous studies, and a nuclear gene (IRBP) in addition to mitochondrial data (cytochrome b and 12S rRNA). Members of the Acomyinae and Murinae were used as outgroups. Separate and combined analyses were performed with maximum parsimony, maximum likelihood and Bayesian methods, and divergence times were estimated. The results showed that the monophyly of the genus Mus and of each subgenus was strongly supported by the three genes and the combined analysis. The phylogenies derived from the three genes were on the whole congruent; however, several conflicting topologies were observed such as the relationships between the three Asian species of the subgenus Mus ( caroli , cervicolor and cookii ). Increasing the taxonomic sampling of Nannomys did not satisfactorily improve the resolution of relationships between the four subgenera. In addition, molecular calibrations indicate that the Mus and Nannomys radiation coincided with major environmental changes.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 417–427.     

Phylogenetic relationships of a new genus and species of microteiid lizard from the Atlantic forest of north-eastern Brazil (Squamata, Gymnophthalmidae)

A new genus and species of a short limbed and slightly elongate gymnophthalmid lizard is described from the Atlantic rain forests of north-eastern Brazil. The new genus is also characterized by short and stout pentadactyl limbs, presence of prefrontals, absence of frontoparietals, distinctive ear opening and eyelid, two pairs of genials, a distinct collar, smooth, quadrangular, dorsal scales, quadrangular ventrals, fused postfrontal and postorbital bones, and two pairs of sternal ribs. The geographical distribution of the new taxon extends from the state of Rio Grande do Norte to the northern bank of the Rio São Francisco in the state of Alagoas. All specimens were obtained in leaf litter, an observation which agrees with the fossorial habits suggested by the body shape and massive aspect of the head of this species. A phylogenetic analysis based on external morphology, osteology, and molecular data recovered the new lizard as the sister genus to Anotosaura , and Colobosauroides as the sister group to these two.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 543−557.     

Morphological character evolution and molecular trees in sepiids (Mollusca: Cephalopoda): is the cuttlebone a robust phylogenetic marker?

The sepiids are characterized by their cuttlebone or sepion, an internal shell resulting from secondary mineralization of a chitinous 'gladius'. The various species are identified using a combination of shell criteria and 'soft-part' characters. Using mitochondrial genes, we established phylogenetic relationships of sepiids including the three accepted genera ( Sepia , Sepiella , Metasepia ) and a species complex of uncertain status ( Doratosepion ). We showed the Sepia genus to be paraphyletic and found no direct correlation between geographical distribution and systematics. We mapped, on the molecular tree, shell characteristics commonly used as reliable diagnostic criteria for taxonomy. Due to the plasticity of the shell, these characters did not appear phylogenetically informative. In an attempt to define systematic categories related to phylogenetic relationships, new clear synapomorphies need to be established for each genus, necessitating a revision of the genus Sepia .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 139–150.     

Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data

The phylogenetic relationships of orbiniid taxa were reconstructed based on sequence data of the mitochondrial 16S rRNA and nuclear 18S rRNA genes. Both genes were analysed separately and in combination using maximum likelihood, Bayesian inference and maximum parsimony. Regardless of the method used, a clade consisting of the investigated Orbiniidae, Methanoaricia dendrobranchiata and Questa was strongly supported by the 18S dataset. The analysis of the combined dataset suggests inclusion of M. dendrobranchiata within the Orbiniidae with close relationships to species of Orbinia and Phylo, rather than as a sister taxon to all other orbiniids. Evidence is given for the paraphyletic status of Leitoscoloplos , Naineris , Orbinia , Phylo and Scoloplos , which represent the most species-rich genera of the Orbiniidae. It is thus reasoned that the morphological characters presently used for genus diagnosis are not informative for cladistic analysis. No support is found for the hypothesis that taxa of the Protoariciinae represent juveniles of Orbiniinae. Instead, in the case of Protoaricia oerstedi , strong support for a progenetic origin is found.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 59−73.     

Description of a new Eocene osteoglossid fish and additional information on †Singida jacksonoides Greenwood and Patterson, 1967 (Osteoglossomorpha), with an assessment of their phylogenetic relationships

A new genus and species of osteoglossomorph fish is described from the Eocene Mahenge site in north central Tanzania. The new species is sympatric with the previously described osteoglossomorph † Singida jacksonoides , but is easily distinguished from it by the presence in the new species of teeth on the jaws and entopterygoid. The new species shares characters with the Heterotidinae, such as the two arms of the preopercle being of similar length and the posterior end of the maxilla lying on the dentary, and other characters with the Osteoglossinae (including † Phareodus ), for example the enlarged pectoral fin ray and the shape of the opercular bone. New specimens of † S. jacksonoides also have been recovered from the site, providing more information on the osteology of this fish. The relationships of these two Mahenge osteoglossomorphs are assessed, with both fishes placed within the Osteoglossidae, but of uncertain position in the family. Addition of the new species to previous cladistic data sets, and the revised information for † S. jacksonoides , causes the subfamily Osteoglossinae to be nonmonophyletic.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 213–228.     

Molecular phylogeny of the western Atlantic species of the genus Portunus (Crustacea, Brachyura, Portunidae)

The genus Portunus encompasses a comparatively large number of species distributed worldwide in temperate to tropical waters. Although much has been reported about the biology of selected species, taxonomic identification of several species is problematic on the basis of strictly adult morphology. Relationships among species of the genus are also poorly understood, and systematic review of the group is long overdue. Prior to the present study, there had been no comprehensive attempt to resolve taxonomic questions or determine evolutionary relationships within this genus on the basis of molecular genetics. Phylogenetic relationships among 14 putative species of Portunus from the Gulf of Mexico and other waters of the western Atlantic were examined using 16S sequences of the rRNA gene. The resultant molecularly based phylogeny disagrees in several respects with current morphologically based classification of Portunus from this geographical region. Of the 14 species generally recognized, only 12 appear to be valid. We recommend that P. vossi be hereafter regarded as a junior synonym of P. spinimanus and that P. bahamensis be regarded as a junior synonym of P. depressifrons . Our analysis suggests that western Atlantic members of the genus can be subdivided into at least three well-defined clades. Pending further molecular analyses with a large subset of species, it appears that the genus is not monophyletic and that it warrants further taxonomic revision.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 211–220.     

Phylogenetic relationships of the north-eastern Atlantic and Mediterranean blenniids

The phylogenetic relationships of 27 north-eastern Atlantic and Mediterranean blennioids are analysed based on a total of 1001 bp from a combined fragment of the 12S and 16S mitochondrial rDNA. The most relevant results with implications in current blenniid taxonomy are: (1) Lipophrys pholis and Lipophrys (=  Paralipophrys ) trigloides are included in a well-supported clade that by the rule of precedence must be named Lipophrys ; (2) the sister species of this clade are not the remaining species of the genus Lipophrys but instead a monotypic genus comprising Coryphoblennius galerita ; (3) the smaller species of Lipophrys were recovered in another well-supported and independent clade, which we propose to be recognized as Microlipophrys ; (4) although some authors included the genera Salaria and Lipophrys in a single group we have never recovered such a relationship. Instead, Salaria is more closely related to the genera Scartella and Parablennius ; (5) the genus Parablennius , which was never recovered as a monophyletic clade, is very diverse and may include several distinct lineages; (6) the relative position of Aidablennius sphynx casts some doubts on the currently recognized relationships between the different blenniid tribes. Meristic, morphological, behavioural and ecological characters support our results and are also discussed. The possible roles of the tropical West African coast and the Mediterranean in the diversification of blenniids are discussed.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 283–295.     

Systematics of the spider genus Metabus O. P.-Cambridge, 1899 (Araneoidea: Tetragnathidae) with additions to the tetragnathid fauna of Chile and comments on the phylogeny of Tetragnathidae

The spider genus Metabus (Tetragnathidae) previously included nine species: the type M. gravidus O. P.-Cambridge, 1899 – junior synonym of Leucauge ocellata (Keyserling) – from Central America and eight species from Chile. In this paper, the classification of the Metabus species-complex is revised, and two new genera, with three new species and five new combinations, are described. Allende gen. nov. is created for four Chilean species not congeneric with the type of Metabus : the type A. puyehuensis sp. nov. , A. patagiatus (Simon) comb. nov. , A. nigrohumeralis (F. O. P.-Cambridge) comb. nov. and A. longipes (Nicolet) comb. nov. Further additions to the Chilean fauna are under the new genus Mollemeta gen. nov. – created for M. edwardsi (Simon) comb. nov. – and three new species of Chrysometa : C. acinosa sp. nov. , C. levii sp. nov. and C. maitae sp. nov. Metabus now includes four species: M. ocellatus (Keyserling) comb. nov. , M. debilis (O. P.-Cambridge) comb. nov. , M. ebanoverde sp. nov. and M. conacyt sp. nov. All of these species were included in a phylogenetic analysis of 38 tetragnathid and 12 orbicularian outgroup terminals scored for 105 morphological and behavioural characters. The results suggest that Metabus as previously circumscribed is polyphyletic. The phylogenetic relationships within tetragnathids are briefly discussed. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 285–335.     

Khaosokia caricoides, a new genus and species of Cyperaceae from Thailand

Khaosokia caricoides , D.A. Simpson, Chayam. & J. Parn., a newly discovered genus and species of Cyperaceae is described and illustrated. The genus is characterized by a narrowly paniculate dioecious inflorescence with 2–4 nodes, each of the nodes having a leaf-like inflorescence bract that exceeds the inflorescence. Spikelets in both sexes are linear-cylindric and each flower has seven perianth bristles. The nutlet was immature in the specimens examined. The photosynthetic pathway is C₃. Khaosokia is endemic to limestone cliffs in peninsular Thailand; its conservation status is assessed as Vulnerable (VU B1a + 2a). It has affinities to tribes Cariceae, Dulichieae and some members of Scirpeae, but the exact nature of these relationships has yet to be determined. A revised key to the genera of Cyperaceae in Thailand is presented.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 357–364.     

A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae

The publication of the scientific name Monjurosuchus splendens in 1940 documented the first tetrapod fossil of the later world-renowned Jehol Biota. For more than half a century since this discovery, however, Monjurosuchus has remained as a monotypic genus of the family Monjurosuchidae, and the relationships of the family with choristoderes have not been correctly recognized until quite recently. In this paper, a new monjurosuchid is named and described based on a nearly complete skull and postcranial skeleton from the Early Cretaceous Chiufotang Formation exposed near Chaoyang, western Liaoning Province, China. This new material documents the first occurrence of monjurosuchid choristoderes outside the type Lingyuan area, and extends the geological range of the family from the Yixian Formation to the younger Chiufotang Formation. Cladistic analyses were conducted with inclusion of monjurosuchids, and the results support the placement of the family Monjurosuchidae as a primitive clade outside the Neochoristodera. A new classification scheme is proposed for choristoderes on the basis of the recovered phylogenetic framework of the group.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 427–444.