The relationship of Sophora sect. Edwardsia (Fabaceae) to Sophora tomentosa, the type species of the genus Sophora, observed from DNA sequence data and morphological characters

Sophora tomentosa , the type species of the genus Sophora , is shown by phylogenetic analyses of rbc L and ITS sequence data to be sister to Sophora sect. Edwardsia . S. tomentosa and most of the species from sect. Edwardsia share hypogeal germination, exstipulate leaves, and terete filaments. These species have buoyant seeds, and are distributed by ocean currents throughout the pantropics ( S. tomentosa ) and around southern temperate oceanic islands (sect. Edwardsia ). S. tomentosa differs from the species of sect. Edwardsia by its frutescent growth habit, terminal elongate inflorescence and smooth-walled legume. S. macrocarpa is unusual in sect. Edwardsia as its leaves have stipules, the filaments are winged, and the legume is smooth-walled.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 439–446.     

Disentangling causes of disjunction on the South Island of New Zealand: the Alpine fault hypothesis of vicariance revisited

Many elements of the flora and fauna of New Zealand's South Island show disjunct distributions with conspecific populations or closely-related species that occur in the north-west and south separated by a central gap. Three events have been implicated to account for this pattern: Pleistocene glaciations, Pliocene mountain building, or displacement along the Alpine fault, the border of the Pacific and Australian plates stretching diagonally across the South Island from south-west to north-east that formed during the Miocene. Disjunct distributions of species level taxa are probably too young to be due to Alpine fault vicariance. It has therefore been suggested that the biogeographical impact of the Alpine fault, if any, should be apparent on deeper phylogenetic levels. We tested this hypothesis by reconstructing the phylogenetic relationships of the hydrobiid gastropods of New Zealand based on mitochondrial DNA fragments of cytochrome oxidase subunit I (CO I ) and 16S rDNA. The creno- and stygobiont species of this family are typically poor dispersers. Therefore, ancient patterns of distribution may be conserved. The phylogenetic reconstructions were in accordance with the Alpine fault hypothesis uniting genera occurring on either side of the fault. Divergence estimates based on a molecular clock of CO I indicated splits predating the Pliocene uplift of the Alps.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 361–374.     

Mitochondrial DNA phylogeny of the woodpecker genus Veniliornis (Picidae, Picinae) and related genera implies convergent evolution of plumage patterns

The woodpecker genus Veniliornis comprises 12 species, all restricted to the New World tropics. The seemingly distantly related genus Picoides is broadly distributed in Eurasia and North America with two putative species, P. lignarius and P. mixtus , occurring in South America. The two genera are clearly distinct with respect to general plumage colouration and patterning as well as habitat utilization and thus traditionally have been placed in different tribes. Phylogenetic analyses of mtDNA sequences from the COI and cyt b genes indicated that both genera are reciprocally paraphyletic. The two South American species of Picoides belong to a clade comprising most species of Veniliornis , but V. fumigatus of Central and north-western South America belongs to a clade comprising species of Picoides . The mtDNA tree also indicated that Veniliornis is not closely related to the genus Piculus, as is implicit in current classifications. Misclassifications involving Veniliornis at both the generic and tribal levels appear to result from convergent evolution of plumage traits in specific forest types. We infer that the common ancestor of Veniliornis entered South America approximately at the time the Isthmus of Panama was formed, and diversification within South America was rapid.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 611–624.     

Species diversity and endemism in the Daphnia of Argentina: a genetic investigation

Although the temperate regions of South America are known to have a diverse daphniid fauna, there has been no genetic evaluation of the existing taxonomic system or of the affinities between the North and South American faunas. The present study analyses mitochondrial DNA sequences and allozyme variation to investigate species diversity in 176 Daphnia populations from Argentina. This work established the presence of at least 15 species in Argentina, six of which are either undescribed or are currently misidentified and two of which represent range extensions of North American taxa. Eleven of the Argentine species appear endemic to South America, while the remaining four also occur in North America. In the latter cases, the close genetic similarity between populations from North and South America indicates the recent exchange of propagules between the continents. While biological interactions and habitat availability have undoubtedly contributed to the observed species distributions, chance dispersal has apparently played a dominant role in structuring large-scale biogeographical patterns in this genus and probably in other passively-dispersed organisms.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 171−205.     

A new species of Selaginella from Hainan Island, China, with a key to the Hainan species

Selaginella hainanensis X.C.Zhang & Noot. sp. nov. , a new species from Hainan Island, South China, is described and illustrated. Diagnostic notes and a key to the species distributed in Hainan are also provided.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 323–327.     

Development, phylogeny, and taxonomy of Bostrycapulus (Caenogastropoda: Calyptraeidae), an ancient cryptic radiation

Calyptraeid gastropods are well know for the taxonomic difficulties caused by their simple, phenotypically variable shells. In this paper I demonstrate that what was previously considered to be a single species, Crepidula aculeata , is an ancient (3–15 Myr) cryptic species complex made up of at least eight species, and that this group should be placed in the genus Bostrycapulus . Despite the difficulty in finding diagnostic adult shell and anatomical features upon which species can be unambiguously identified, DNA sequences, protoconch morphology, embryonic morphology and developmental characters clearly differentiate these eight species. A single species with direct development and nurse eggs is present in the South Atlantic, and a species with planktotrophic development occurs in the equatorial Pacific. The species from Japan, Australia, Florida, the Pacific coasts of Mexico and Central America, and the Cape Verde Islands all have direct development. Most of these species are separated by > 15% divergence in COI sequence data. The fossil record of Bostrycapulus goes back to the Miocene, which agrees with genetic estimates of divergences within the genus ranging from 3 to 15 Mya. Surprisingly, these ancient species differ only slightly in morphology from each other and genetic differentiation does not correlate with geographical distance. I revise the genus Bostrycapulus on the basis of differences in adult morphology, embryonic morphology, mode of development, protoconch morphology, and DNA sequence data. I also describe four new species ( B. pritzkeri sp. nov., B. odites sp. nov., B. latebrus sp. nov. and B. urraca sp. nov. ) and remove three others ( B. gravispinosus , B. calyptraeformis , and B.  cf. tegulicius ) from synonymy with B. aculeatus .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 75−101.     

New fossil record of Lactoridaceae in southern South America: a palaeobiogeographical approach

Lactoridaceae are a monotypic family confined to Masatierra Island, Juan Fernández Archipelago, in the Pacific Ocean. It grows in the understorey of a subtropical montane rain forest. Lactoridaceae most probably originated in southern South Africa in the Cretaceous, with the oldest records in the Turonian–Campanian, and reached its widest palaeogeographical distribution by the Maastrichtian, extending into Australia, India, Antarctica, and North and South America. In this paper, we report a new fossil find of lactoridaceous tetrads from the early Miocene of eastern Patagonia, southern South America. This record is the youngest and geographically one of the closest to the extant Lactoris distribution area. Patagonian fossil material shows greater similarities to extant L. fernandeziana Phil. than to any other described morphotaxon. The family may have migrated into South America, either via Africa (through the Atlantic Ocean) or Antarctica, by the Maastrichtian, growing in eastern Patagonia up to the early Miocene. Arid conditions established in this region by the middle–late Miocene onwards would have determined the restriction of forests to the western lands. Lactoridaceae may have followed a similar migration pattern towards the Pacific coast of South America. The shifting of Lactoridaceae towards Masatierra Island would have occurred in the last 4 Myr by long‐distance dispersal events (perhaps by birds). © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 41–50.     

Historical biogeography of Hexabathynella , a cosmopolitan genus of groundwater Syncarida (Crustacea, Bathynellacea, Parabathynellidae)

Hexabathynella is the only cosmopolitan genus of the order Bathynellacea (Crustacea). The known species number 18, found in Europe (9), Africa (1), South America (2), North America (3) and Australia and New Zealand (3). Phylogenetic analysis suggests that the least derived species are those from South America and the most derived those from the Iberian Peninsula, North America and Australia. The five species with the most plesiomorphic characters occur in salt or brackish water, which supports a marine origin for the genus. Phylogenetic and biogeographical analyses suggest that the distribution of the genus can be explained by dispersion and a double vicariant biogeographical model based on plate tectonics and the evolution of the Tethys during the Mesozoic and Cenozoic.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 457–466.     

Evolution of St Helena arborescent Astereae (Asteraceae): relationships of the genera Commidendrum and Melanodendron

The cladistic relationships of endemic Commidendrum (four species) and Melanodendron (one species) from St Helena were inferred from sequences of ITS1 and ITS2 of nuclear ribosomal DNA. Despite showing a range of morphological and ecological variation, the four species of Commidendrum , C. spurium , C. robustum , C. rotundifolium , and C. rugosum , form a closely related monophyletic group with percentage sequence divergence between 0.2 and 0.9%. Melanodendron integrifolium is sister to Commidendrum indicating that the two genera may have evolved from a common ancestor that arrived in St Helena via a single dispersal event. The closest relatives of Commidendrum and Melanodendron appear to be South African, in the predominantly shrubby genus Felicia , although further sampling of South African Astereae is required. We discuss the evolution and adaptive radiation of these rare and threatened species with particular reference to the possible role of heterochrony.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 69–83.     

New carnivorous sponges (Porifera, Poecilosclerida) collected from manned submersibles in the deep Pacific

Nine new species of carnivorous sponges, namely Cladorhiza segonzaci , Chondrocladia koltuni , Chondrocladia lampadiglobus , Asbestopluma agglutinans , Asbestopluma ( Helophloeina ) formosa , Abyssocladia huitzilopochtli , Abyssocladia inflata , Abyssocladia dominalba and Abyssocladia naudur , are described. The subgenus Helophloeina Topsent, 1929 and the genus Abyssocladia Lévi, 1964 are revived and redefined, with an identification key for Abyssocladia which at present includes seven species. Eight of the sponges were collected from the French IFREMER manned submersible Nautile near active hydrothermal sites of the East Pacific Rise and of the North Fiji and Lau Basins, one from the Russian submersible Mir 2 in the Northwest Pacific near Bering Island, and one from the US submersible Alvin south of Easter Island. Their life conditions are described from direct observations from the submersibles. Some remarks are presented on the taxonomy of Cladorhizidae and more generally of carnivorous Poecilosclerida. The study suggests a very high degree of diversity in the deep Pacific carnivorous sponges.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 553–584.     

Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence

Despite the considerable research that has focused on the evolutionary relationships and biogeography of the genus Bufo, an evolutionary synthesis of the entire group has not yet emerged. In the present study, almost 4 kb of DNA sequence data from mitochondrial (12S, tRNA^Val, and 16S) and nuclear (POMC; Rag-1) genes, and 83 characters from morphology were analysed to infer a phylogeny of South American toads. Phylogenies were reconstructed with parsimony and maximum likelihood and Bayesian model-based methods. The results of the analysis of morphological data support the hypothesis that within Bufo , some skull characters (e.g. frontoparietal width), correlated with the amount of cranial ossification, are prone to homoplasy. Unique and unreversed morphological synapomorphies are presented that can be used to diagnose recognized species groups of South American toads. The results of all phylogenetic analyses support the monophyly of most species groups of South American Bufo . In most DNA-only and combined analyses, the South American (minus the B. guttatus and part of the ' B. spinulosus ' groups), North American, Central American, and African lineages form generally well-supported clades: ((((((((South America) (North America + Central America)) Eurasia) Africa) Eurasia) South America) West Indies) South America). This result confirms and extends prior studies recovering South American Bufo as polyphyletic. The biogeographical results indicate that: (1) The origin of Bufo predates the fragmentation of Gondwana; (2) Central and North American species compose the sister group to a large, 'derived' clade of South American Bufo ; and (3) Eurasian species form the sister group to the New World clade.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 146 , 407–452.     

Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodontidae) and its evolutionary and biogeographical implications

The largest genus of salamanders, Bolitoglossa (Plethodontidae), is widespread in tropical America, where it occurs in diverse habitats and elevations, from high elevation grasslands to lowland rain forest . It has the most extensive geographical range of any salamander genus. While most species occur in Middle America, it ranges throughout most of tropical South America as well. Phylogenetic analysis of 1196 bp of two mitochondrial genes (cytochrome b , 16S RNA) from 55 species offers strong support for the monophyly of the genus and sorts the species into a number of clades. Taking into account morphology, distribution, general ecology, and prior systematic and taxonomic studies, we recognize seven subgenera, four of them new: Bolitoglossa Duméril, Bibron et Duméril, 1854, Eladinea Miranda Ribeiro, 1937, Magnadigita Taylor, 1944, Mayamandra , Nanotriton , Oaxakia and Pachymandra . All South American and some lower Middle American species are included in a single well -supported clade, Eladinea . At the species level our analyses uncover the existence of large genetic diversity within morphologically homogeneous taxa. We propose the new combination: B. (Eladinea) paraensis (Unterstein, 1930) stat. nov. , for Brazilian salamanders previously included under B. altamazonica . We evaluate evidence for the multiple colonization of the tropical lowlands by morphologically derived species groups. South America was invaded by members of one clade, Eladinea , which we infer to have dispersed to South America prior to closure of the Panamanian Portal. Despite the relatively long history of salamanders in South America, that continent now accounts for a relatively small proportion of the lineages and species of neotropical salamanders.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 325–346.     

The geography of diversification in the mormoopids (Chiroptera: Mormoopidae)

The traditional explanation of the distribution of the Mormoopidae is that this family originated in southern Central America or northern South America, later expanding its range north to Mexico and the West Indies, and differentiating into eight species. An alternative fossil-based hypothesis argues that the family originated in the northern Neotropics, reached the Caribbean early in its history, and dispersed to South America after the completion of the Isthmus of Panama. The present study analyses new and previously published sequence data from the mitochondrial 12S, tRNA^val, 16S, and cytochrome b , and the nuclear Rag 2, to evaluate species boundaries and infer relationships among extant taxa. Fixed differences in cytochrome b often coincide with published morphological characters and show that the family contains at least 13 species. Two additional, morphologically indistinct, lineages are restricted to Suriname and French Guiana. Phylogeny-based inferences of ancestral area are equivocal on the geographical origin of mormoopids, in part because several internal nodes are not resolved with the available data. Divergences between Middle American and Antillean populations are greater than those between Mexico/Central America and South America. This suggests that mormoopids diversified in northern Neotropics before entering South America. A northern neotropical origin for mormoopids is congruent with both the Tertiary fossil record and recent phylogenetic hypotheses for the sister family to the Mormoopidae, the Phyllostomidae.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 101–118.     

A new diploid species of Hieracium sect. Pannosa (Asteraceae) from Bulgaria

Hieracium petrovae Vladimirov & Szeląg sp. nov. , a new diploid (2 n  = 18) species in H. sect. Pannosa Zahn, is described and illustrated from the Rhodope Mountains, South Bulgaria, and compared with related taxa. It grows in relict habitats in crevices of limestone rock together with many Balkan endemics. H. petrovae is morphologically similar to taxa from the H. pannosum , H. pilosissimum and H. heldreichii groups (collective species sensu Zahn) to some of which it is a presumed ancestral species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 261–265.     

Intercontinental distribution of Plagiochila corrugata (Plagiochilaceae, Hepaticae) inferred from nrDNA ITS sequences and morphology

Plagiochila sect. Vagae is a large pantropical clade that is characterized morphologically by frequent terminal branching, vegetative distribution by propagules on the ventral surface of the leaves and a capsule wall with thickenings in all layers. Plagiochila corrugata from Brazil is characterized by strongly undulate, toothed leaf margins and represents the only known neotropical species of sect. Vagae with unispiral elaters. Plagiochila cambuena from Madagascar is distinguished by the same features. Maximum likelihood and parsimony analyses of 38 nrDNA ITS sequences of Plagiochila reveal P. corrugata and P. cambuena in a weakly (ML) to well (MP) supported monophyletic lineage within P.  sect.  Vagae . As an outcome of the morphological and molecular investigation, P. cambuena is relegated to the synonymy of P. corrugata. Plagiochila corrugata is placed in a Vagae -subclade with 11 further American species. The range of P. corrugata can be ascribed to long-range dispersal from the Neotropics rather than a Gondwanan distribution. Species from tropical Asia and Africa are placed at the base of the Vagae clade. Branch length within P.  sect.  Vagae points to a sudden radiation.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 469–481.     

Phylogenetic relationships of thorny catfishes (Siluriformes: Doradidae) inferred from molecular and morphological data

Doradidae is a putatively monophyletic group of South American freshwater catfishes containing 30 extant genera and 72 valid species. Only one study to date has attempted to estimate phylogenetic relationships among doradids. This morphological analysis partitioned species into two basal genera ( Wertheimeria and Francisodoras ) and a crown group of three subfamilies (Platydoradinae, Astrodoradinae and Doradinae) whose relationships were unresolved. No subsequent work has been done to resolve the subfamilial trichotomy or to assess whether postulated intergeneric relationships are accurate. We address this problem with complete sequences (2.5 kilobases, kb) of mitochondrial 12S and 16S rRNA genes and partial (1.3 kb) sequences of the nuclear elongation factor-1 alpha (EF1α) gene from representatives of 23 doradid genera (43 species) and 13 outgroups from additional siluriform families. Phylogenetic analysis of these data yields strong support for the monophyly of Doradidae and Astrodoradinae (as well as other relationships), but otherwise shows significant conflict with morphological results. A partial re-examination of published morphological data indicates that many characters may have been incorrectly polarized and many taxa have incorrect state assignments. Our results provide a framework for ongoing efforts to describe the species-level diversity of this poorly understood neotropical family.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 551–575.     

Killickia (Lamiaceae): a new genus from KwaZulu-Natal, South Africa

The genus Killickia is described to accommodate the South African endemic species formerly placed in Micromeria sect. Hesperothymus . Morphological data, as well as results from unpublished phylogenetic studies support its separation from the genera Micromeria and Clinopodium . A new species Killickia lutea Bräuchler is described and three new combinations are made. Killickia is characterised as comprising solitary- or few-flowered cymes, a campanulate to subcampanulate (obconical) calyx with similar teeth, a corolla tube with two pubescent ridges and nutlets with scattered minute hairs. A thickened marginal vein in the leaves as typical for Micromeria is absent. As currently understood all species are restricted to the Drakensberg mountains and KwaZulu-Natal Midlands in eastern South Africa. A key to the species, brief notes on anatomy and ecology are provided.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 575–586.