Evolutionary relationships among the Pulmonate land snails and slugs (Pulmonata, Stylommatophora)

We have incorporated an additional 56 species of land snails and slugs in our ribosomal (r) RNA molecular phylogeny. The new taxa include representatives of several important groups. The molecular tree now includes 160 species of stylommatophoran land snails and slugs in 144 genera in 61 families. In the rDNA tree, the Stylommatophora are principally divided into an 'achatinoid' and a 'non-achatinoid' clade. Within these clades, several major land snail groups, including the Orthurethra, Elasmognatha, Limacoidea, and Helicoidea, are supported. Overall, the rDNA molecular phylogeny has remained stable following the incorporation of the additional taxa, with these additions having little impact on the major evolutionary patterns in the tree. Taxonomic coverage of the Orthurethra, Orthalicidae, Camaenidae, and Bradybaenidae is increased significantly. The camaenids and bradybaenids form a complex, and both appear to be paraphyletic. Several families of uncertain affinity, such as the Sagdidae and Thyrophorellidae, are included for the first time. The Sagdidae are shown to belong to the Helicoidea, and the Thyrophorellidae to the Achatinoidea.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 593–610.     

Relationships among extant and fossil echimyids (Rodentia: Hystricognathi)

The echimyid rodents are the most diverse group of Neotropical hystricognaths, with approximately 40 extant and fossil genera. Craniodental characters are proposed in order to formulate hypotheses of phylogenetic relationships within the Echimyidae. A data matrix of 54 taxa and 50 characters is constructed and submitted to parsimony analyses using PAUP and WinClada programs. Analysis of the complete data set results in 47 448 most parsimonious trees 107 steps long. These trees are summarized in a strict consensus tree, which is taken as the main phylogenetic hypothesis resulting from this study. The monophyly of several currently recognized supraspecific taxa is not corroborated. These are: the subfamilies Eumysopinae, Echimyinae, Myocastorinae and Adelphomyinae; and the genera Proechimys , Echimys and Makalata . Conversely, the monophyly of Dactylomyinae and Trinomys is supported. New associations are proposed: (1) a clade comprising the extant Carterodon , Clyomys and Euryzygomatomys and the fossil Pampamys and Theridomysops placed at the base of the crown-group Echimyidae; (2) a clade uniting Proechimys , Hoplomys and Trinomys , which is the sister-taxon of (3) a clade including Mesomys , Lonchothrix , Myocastor and a clade with extant dactylomyines and echimyines and associated fossil taxa. Based on this phylogenetic hypothesis, patterns of tooth evolution in Echimyidae are discussed, and minimum ages for the divergence events within the family are estimated.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 445–477.     

Phylogeny and taxonomy of thegenera of south-western North American Euctenizinae trapdoor spidersand their relatives (Araneae: Mygalomorphae, Cyrtaucheniidae)

The primary focus of this paper is to evaluate the monophylyand intergeneric relationships of the cyrtaucheniid subfamily Euctenizinaeand to a lesser degree the monophyly of Cyrtaucheniidae. Using 71morphological characters scored for 29 mygalomorph taxa our cladisticanalysis shows that Cyrtaucheniidae is likely paraphyletic withrespect to the Domiothelina, the clade that comprises the Migidae,Actinopodidae, Ctenizidae, and Idiopidae. Together, the Domiothelinaand Cyrtaucheniidae have been treated as the Rastelloidina clade.A strict interpretation of rastelloid classification based on ourcladogram would require the establishment of four additional spiderfamilies. However, we choose to use informal names for these cladesso that these taxa can be validated by subsequent studies of mygalomorphphylogeny before formal names are introduced. The phylogenetic analysisalso serves as a vehicle for examining the patterns of homoplasyobserved in mygalomorphs. The secondary focus of this paper is ataxonomic revision of Euctenizinae genera from the south-westernUnited States that includes a key to its genera. The cyrtaucheniid genera Enrico and Astrosoga areconsidered junior synonyms of Eucteniza. Actinoxia and Nemesoides arejunior ­synonyms of Aptostichus . At present theNorth American Euctenizinae comprises these seven nominal genera: Eucteniza, Neoapachella gen.nov. ( Neoapachella rothi sp. nov. ), Myrmekiaphila,Entychides, Promyrmekiaphila, Apomastus gen. nov. ( Apomastusschlingeri sp. nov. ), and Aptostichus .  © 2002The Linnean Society of London . Zoological Journal of the LinneanSociety, 2002, 136 , 487−534     

A new species of Dyrosaurus (Crocodylomorpha, Dyrosauridae) from the early Eocene of Morocco: phylogenetic implications

A new dyrosaurid is described from the Ypresian of the phosphatic deposits of the Oulad Abdoun Basin of Morocco. It is based on numerous cranial and postcranial remains, allowing an almost complete reconstruction. This new Dyrosaurus species, Dyrosaurus maghribensis sp. nov. , is currently only known from Morocco. It differs from D. phosphaticus , present in contemporaneous levels of Algeria and Tunisia, by several autapomorpies, including a smooth dorsal margin of the parietal and widely opened choanae. A phylogenetic analysis, using 47 taxa and 234 morphological characters, shows the dyrosaurids as the sister taxon of pholidosaurids, which include Elosuchus , Sarcosuchus , Terminonaris and Pholidosaurus , and the thalattosuchians. Goniopholididae is a non-monophyletic group; however, if dyrosaurids are not included in the analysis, the result differs and the goniopholidids form a distinct clade. If Thalattosuchia is excluded, both Goniopholididae and Pholidosauridae become paraphyletic assemblages. Thus, phylogenetic problems remain with respect to longirostrine clade, and more attention should be paid to resolving their evolutionary relationships amongst the crocodyliforms.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 603–656.     

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.     

Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers

The Nymphaeales (water-lilies and relatives) represent one of the earliest branching lineages of angiosperms and comprise about 70 aquatic species. Here, we present a comprehensive study of phylogenetic relationships within the Nymphaeales from a dataset containing 24 representatives of the order, including all currently recognized genera and all subgenera of the genus Nymphaea , plus 5 outgroup taxa. Nine different regions of the chloroplast genome − comprising spacers, group II introns, a group I intron, and a protein coding gene − were analysed. This resulted in a character matrix of 6597 positions and an additional 369 characters obtained from coded length mutations. Maximum parsimony and Bayesian analyses of the complete dataset yielded congruent, fully resolved and well-supported trees. Our data confirm the monophyly of the Cabombaceae but do not provide convincing support for the monophyly of Nymphaeaceae with respect to Nuphar . Moreover, the genus Nymphaea is inferred to be paraphyletic with respect to Ondinea , Victoria and Euryale . In fact, the Australian endemic Ondinea forms a highly supported clade with members of the Australian Nymphaea subgenus Anecphya . In addition, Victoria and Euryale are inferred to be closely related to a clade comprising all night-blooming water-lilies ( Nymphaea subgenera Hydrocallis and Lotos ). An experimental approach showed taxon sampling to be of influence on the nodes reconstructed in core Nymphaeaceae. The results underscore that more diverse genera, if not clearly known to be monophyletic, should be represented by all major lineages.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 141–163.     

A phylogenetic analysis of the genera of Lejeuneaceae (Hepaticae)

The Lejeuneaceae are the largest family of the liverworts (Hepaticae), with almost a thousand species in 91 currently accepted genera. We analysed phylogenetic relationships of 69 genera, representing all major subfamilies and tribes recognized in the family, by using 49 informative morphological characters (31 gametophytic, 18 sporophytic), one chemical character, and applying equal and successive weighting of characters and parsimony analysis. In all trees recovered, the Lejeuneaceae were monophyletic with Nipponolejeunea (subfam. Nipponolejeuneoideae) forming the basalmost lineage. The remaining genera clustered in two major groups, the monophyletic Lejeuneoideae (52 genera) and the paraphyletic Ptychanthoideae (16 genera). Within each, several multigeneric lineages corresponding in part to previously described taxa were recovered: the Acrolejeuneinae and Ptychanthinae clades in the Ptychanthoideae, and the Brachiolejeuneinae, Lejeuneeae and Tuyamaella–Cololejeunea clades in the Lejeuneoideae. Bryopteris , a genus sometimes treated as a separate family, was nested in the Ptychanthinae clade. The Tuyamaella–Cololejeunea lineage corresponded with three previously recognized subfamilies (Cololejeuneoideae, Myriocoleoideae and Tuyamaelloideae) and contained genera with neotenic features, in two subclades. These features seemed to have originated by multiple heterochronic events: single origins were detected for 'protonemal neoteny' and 'primary neoteny', whereas 'secondary neoteny' probably evolved twice. Relationships within the large Lejeuneeae clade (43 genera) remained largely unresolved, although several putative lineages were detected in majority rule trees. Additional characters such as DNA sequences may provide better phylogenetic resolution in this group.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 391–410.     

The evolution of CAM in the subfamily Pitcairnioideae (Bromeliaceae)

A molecular phylogeny for the subfamily Pitcairnioideae was inferred to examine the distribution of crassulacean acid metabolism in the subfamily. For this purpose, a neighbour-joining tree with p-distances was built using a MatK chloroplast gene data set. The phylogenetic results of our analysis confirmed the monophyletic condition of most genera examined: Brocchinia , Dyckia , Encholirium , Fosterella , Hechtia and Puya . A paraphyletic basal sequence showed Hechtia branching off from the basal node, followed by Brocchinia , Cottendorfia  +  Navia phelpsiae, and Puya . The remaining taxa were divided into two groups: (a) Deuterocohnia meziana , Dyckia , Encholirium ; Fosterella ; Deuterocohnia spp. +  Pitcairnia heterophyla ; (b) Pepinia , Pitcairnia spp. and Navia igneosicola . The basal placement of the CAM genera Hecthia indicates that CAM may be a 'primitive condition' in Pitcairnioideae and that C₃ species may have lost the ability to induce CAM. In this molecular tree, CAM metabolism appeared scattered throughout the tree. Current knowledge, however, does not exclude the possibility that CAM arose only once and it has been switching on and off in various lineages. Further detailed studies on photosynthetic metabolisms and the phylogenetic distribution of characters will provide a better basis on which to evaluate photosynthetic origins.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 261–268.     

Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla)

The systematics and taxonomy of early Eocene equids are investigated. A paraphyletic sequence of equid taxa is recovered from a phylogenetic analysis of 40 taxa and 121 characters. This analysis supports the identification of Hyracotherium as a primitive equoid and its restriction to the genotype, Hyracotherium leporinum . Sifrhippus gen. nov. is erected for the sister taxon of all other equids, Hyracotherium sandrae Gingerich. Minippus gen. nov., the next more-derived equid clade, is erected for two small equids, M. index Cope and M. jicarillai nov. species. Arenahippus gen. nov. is erected for the next three sequentially more-derived equid taxa, A. grangeri Kitts, A. aemulor Gingerich, and A. pernix Marsh. The genus Xenicohippus , which is the next more-derived equid clade, is redefined to include X. craspedotum Cope. Eohippus Marsh, the next more-derived equid taxon, is resurrected for E. angustidens Cope. Pliolophus , the only early Eocene equid from Europe, is identified as the sister taxon to Protorohippus , a sequence of successively more-derived equid taxa consisting of P. montanum Wortman and P. venticolum Cope. Protorohippus venticolum is identified as the sister taxon of Orohippus . Systemodon Cope is resurrected for S. tapirinum Cope. This taxon has historically been placed within Hyracotherium yet this analysis allies it with Cymbalophus near the base of the perissodactyl radiation. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 134 , 141–256.     

Phylogeny and evolutionary history of the Aplodontoidea (Mammalia: Rodentia)

Athough over a hundred species of fossil aplodontoids have been described since the extant species, Aplodontia rufa (the mountain beaver), was first described by Rafinesque in 1817, a thorough survey of the relationships among all the species in this clade has not been undertaken since McGrew's study in 1941. Here, a complete phylogenetic analysis of all published species of aplodontoids is used to reconstruct the evolutionary relationships within the clade, and to present an updated classification of the Aplodontoidea. Several of the traditionally recognized subfamilies are found to be paraphyletic, namely the Prosciurinae, the Allomyinae, and the Meniscomyinae. Others, however, including the Aplodontinae and the Mylagaulidae, appear to be monophyletic. These latter two taxa, which include all of the hypsodont members of the aplodontoid clade, seem to be sister taxa. The history of the aplodontoid clade shows several episodes of rapid diversification in the Early Oligocene, the Late Oligocene, and the Early to Middle Miocene. The Ansomyinae and Aplodontinae show comparatively low speciation rates. The patterns of change in morphology and evolutionary rates suggest a need for a more detailed study of the causes of diversification, extinction, and ecological change in this lineage.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 769–838.     

Phylogeny of the Nudibranchia

More than 100 morphological, anatomical and histological characters pertaining to the Nudibranchia are discussed in the course of a phylogenetic analysis. Based on our own investigations on anatomy, histology and published literature, the polarity of each character is assessed by comparing its expression with outgroups, chiefly the Pleurobranchoidea, but also with other opisthobranch taxa (Cephalaspidea, Anaspidea, Sacoglossa), and ultimately with the Pulmonata, Vetigastropoda and Caenogastropoda. By extracting 46 applicable characters expressed in 30 taxa, a phylogenetic analysis of the major groups of the Nudibranchia is performed by using the computer program PAUP. Nudipleura new taxon (containing the Pleurobranchoidea plus Nudibranchia) is defined. The monophyly of the Nudibranchia, the Anthobranchia (containing the Bathydoridoidea plus Doridoidea), and the Cladobranchia (containing the Dendronotoidea plus Aeolidoidea and 'Arminoidea') are confirmed and autapomorphies are highlighted. The paraphyly of the 'Arminoidea' is demonstrated. Incongruities at lower taxonomic level are discussed.     

Revision of Pseudonereis (Polychaeta, Nereididae)

A taxonomic revision of Pseudonereis (Polychaeta, Nereididae) shows that some of the described taxa are very similar in most morphological characteristics. The revision includes all ten taxa considered valid, and are redescribed from type material. Lectotypes are designated for Pseudonereis anomala Gravier, 1901, Pseudonereis noodti (Hartmann-Schröder, 1962) and Pseudonereis trimaculata Horst, 1924. The widely geographically distributed and well-known P. gallapagensis Kinberg, 1865 and P. variegata ( Grube, 1857 ) show striking morphological resemblance to less well-known taxa with similar distribution. Paragnath variation in populations of P. anomala is discussed relating to its geographical distribution. Pseudonereis trimaculata is recorded from Australia for the first time. Taxa belonging to Pseudonereis are predominantly tropical and subtropical. A cladistic analysis using parsimony is included to test for monophyly of Pseudonereis . A monophyletic clade including all Pseudonereis taxa is given low bootstrap support. This clade is supported by the synapomorphies: presence of paragnaths in closely spaced comb-like rows on the maxillary ring on the pharynx, and presence of p-bar paragnaths in Areas II–IV and VII–VIII. Several of the included taxa share the shield-shaped paragnath in Area VI, which serves to distinguish Pseudonereis spp. from Perinereis spp. Paragnaths of the type p-bars and shield-shaped bar is described for the first time; the latter character is different from the smooth bar-shaped paragnaths in Area VI as has previously been described in these taxa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 145–176.     

Taxonomic relationships of the western Asian taxa of Gentiana sect. Pneumonanthe

The taxonomic position of western Asian members of Gentiana sect. Pneumonanthe has long been a matter of conflict. In this paper, the six western Asian species currently recognized as belonging to sect. Pneumonanthe ( G. boissieri , G. calycina , G. freyniana , G. gelida , G. paradoxa , and G. septemfida ) are compared and their relationships established using a morphological phylogenetic analysis. Seed testa and flower and leaf morphological characters were studied and 11 characters were selected for a cladistic analysis. Euro-Siberian and Far Eastern taxa of sect. Pneumonanthe ( G. pneumonanthe and G. scabra ) were used as outgroups. Our results suggest the presence of two morphologically distinct clades within the western Asian gentians: a Septemfida and a Gelida clade. G. calycina and G. freyniana show close affinities to G. boissieri and G. gelida , and are distinct from G. septemfida s.l. Biogeographical aspects of the two groups are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 197–208.     

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.     

New clades of euthyneuran gastropods (Mollusca) from 28S rRNA sequences

Recent morphological and molecular results on phylogeny of euthyneuran gastropods, which include opisthobranchs and pulmonates, have greatly diminished previous supposed resolution of their phylogenetic relationships. In addition to recent morphological results, sequences of the D1 and D2 domains of the 28S rRNA are here analyzed by parsimony for 31 euthyneuran species. The molecular and previous morphological data sets were not congruent according to an ILD test, and morphological and molecular data could not be analyzed simultaneously. Consequently Bremer's Combinable Component Consensus was used to obtain a new tree, with the following supported molecular results: monophyly of a new clade of opisthobranchs including actively swimming Euthyneura, i.e., pelagic Gymnosomata and Thecosomata plus benthic Anaspidea; first molecular confirmation of monophylies of Hygrophila, including Chilina, Acteonoidea, and Sacoglossa, which include both shell-bearing species and slugs; and new confirmation of the monophyly of Stylommatophora. Morphological characters which support the new clades obtained here are discussed.     

Numerical taxonomy study of Salvia sect. Salvia (Labiatae)

Multivariate analysis was carried out in order to elucidate the taxonomic relationships between Salvia officinalis L., the type species of the genus, and S. fruticosa Mill., both taxa included in section Salvia (Labiatae). Seventy-five different herbarium specimens from all over the Mediterranean Region, belonging to these two taxa, were analysed. Twenty-four specimens belonging to S. lavandulifolia Vahl. s.l. , were used as outgroups. Twenty-six morphological characters were measured and a data matrix was constructed prior to multivariate analysis by means of R software. Multiple correspondence analysis was used to obtain a single dendrogram, applying Ward's minimum variance algorithm. This tree was used as a basis to propose a key for the determination of the species and subspecies studied. S. lavandulifolia is separated in two groups, one belonging to S. officinalis and another one representing what was called S. blancoana . Salvia officinalis L. ssp. gallica (W. Lippert) Reales, D. Rivera & Obón and S. officinalis L. ssp. oxyodon (Webb & Heldr.) Reales, D. Rivera & Obón are discussed as new combinations. In addition, a hybrid between S. officinalis and S. fruticosa is identified and its importance in gardening and cultivation is discussed. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 353–371.     

Genets and 'genet-like' taxa (Carnivora, Viverrinae): phylogenetic analysis, systematics and biogeographic implications

The subfamily Viverrinae is a taxon of uncertain systematic status. This study consists of cladistic analyses based on morphological characters of specimens belonging to the genera Genetta , Osbornictis , Poiana and Prionodon . Two levels of analysis are carried out, one concerning generic relationships (intergeneric analysis) and one dealing with the interrelationships of species within the genus Genetta (intrageneric analysis). In the first analysis, different outgroups were used in order to test the ingroup topology.
With regard to the intergeneric analysis, Osbornictis , Poiana and Prionodon , together with Genetta johnstoni , constitute a monophyletic group (including Nandinia ), which is the sister-group of a clade formed by the other species of genets. Thus, the genus Genetta is regarded as paraphyletic. Prionodon appears to be a derived taxon. The Poiana – Prionodon clade is well supported, especially by ultrastructural hair characters. The cladogram topology in the intrageneric analysis indicates an ecological transition from the rain forest genets to the savanna genets. This supports a rain forest origin of the genus Genetta , a conclusion which may be generalized to the entire study group. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 317–334.