Fluvial range expansion, allopatry, and parallel evolution in a Danubian snail lineage (Neritidae: Theodoxus)

The snails Theodoxus danubialis and Theodoxus prevostianus form a single clade native to freshwaters of south-eastern Europe whose inter- and intraspecific relationships remain unresolved. The present study utilized a phylogeographical approach to clarify the relationship of these species as well as to reconstruct the evolutionary and demographic history of populations in the western portion of their range. Phylogenetic, population genetic, and nested clade analyses reveal a clade that has distributed itself upriver from a lower Danube River source population and become genetically distinct primarily through range expansion and localized allopatric divergence. Notably, this geographical pattern is replicated phylogenetically in the form of two cytochrome c oxidase subunit I (CO I) lineages that are present simultaneously in individual snails. Haplotypes from polymorphic individuals form two distinct clades, both of which show phylogenetic and nucleotide substitution patterns consistent with a mitochondrial origin, and whose common ancestor must have occurred in a lower Danube source population. Separated allopatrically from their Danubian relatives, populations of T. danubialis in northern Italy have also undergone substantial range expansion, much more recently than Danube watershed lineages. In addition to repeated patterns of range expansion, parallelism is found in T. prevostianus , which is shown to be a nonmonophyletic taxon of remarkable morphological and ecological similarity.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 603–617.     

A comprehensive phylogeny of the bumble bees (Bombus)

Bumble bees ( Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades ( short-faced and long-faced ) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and for evaluating character evolution in the bumble bees.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 161–188.     

Uncovering tropical diversity: six sympatric cryptic species of Blepharoneura (Diptera: Tephritidae) in flowers of Gurania spinulosa (Cucurbitaceae) in eastern Ecuador

Diversification of phytophagous insects is often associated with changes in the use of host taxa and host parts. We focus on a group of newly discovered Neotropical tephritids in the genus Blepharoneura , and report the discovery of an extraordinary number of sympatric, morphologically cryptic species, all feeding as larvae on calyces of flowers of a single functionally dioecious and highly sexually dimorphic host species ( Gurania spinulosa ) in eastern Ecuador. Molecular analyses of the mitochondrial cytochrome oxidase-I gene from flies reared from flowers of G. spinulosa reveal six distinct haplotype groups that differ by 7.2–10.1% bp (uncorrected pairwise distances; N  = 624 bp). Haplotype groups correspond to six distinct and well-supported clades. Members of five clades specialize on the calyces of flowers of a particular sex: three clades comprise male flower specialists; two clades comprise female flower specialists; the sixth clade comprises generalists reared from male and female flowers. The six clades occupy significantly different morphological spaces defined by wing pigmentation patterns; however, diagnostic morphological characters were not discovered. Behavioural observations suggest specific courtship behaviours may play a role in maintaining reproductive isolation among sympatric species. Journal compilation  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 779–797. No claim to original US government works.     

Genetic differentiation amongst fragrant orchids (Gymnadenia conopsea s.l.) in the British Isles

Genetic variation was examined in five microsatellite loci to seek evidence of genetic differentiation and restricted gene flow that would support the taxonomic division of Gymnadenia into three species ( G. borealis , G. conopsea , and G. densiflora ). A total of 107 alleles was detected in 17 populations from England, Scotland, and Ireland. The mean expected heterozygosities ranged from 0.48 to 0.81. The differentiation in allele frequencies amongst populations that had been assigned to each taxon on the basis of morphology was sufficiently large to support the taxa as distinct species. Phylogenetic trees based on microsatellite allele frequencies, as well as assignment tests, supported the existence of three distinct groups with at least partial restriction of gene flow between them. There was substantial homozygote excess, leading to high F _IS estimates, for most loci in most populations. This is unlikely to have been a result of widespread null alleles, and more probably reflects a high level of inbreeding in G. conopsea . This inference requires further investigation. The implications of the results of this and other taxonomic studies for the conservation of Gymnadenia in Britain are discussed.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 349–360.     

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 among Acestrorhynchus species (Ostariophysi: Characiformes: Acestrorhynchidae)

A total of 104 osteological and external morphological features were examined in 13 species of Acestrorhynchus and 15 outgroup taxa to advance a hypothesis of relationships within the genus. Two most parsimonious hypotheses corroborate the monophyly of Acestrorhynchus but differ in the hypothesized relationships of Acestrorhynchus heterolepis . Three proposed supraspecific assemblages are at least partially correlated with groups of species previously diagnosed on the basis of colour pattern: (1) Acestrorhynchus britskii , Acestrorhynchus grandoculis , Acestrorhynchus microlepis , and Acestrorhynchus minimus ; (2) Acestrorhynchus falcirostris , Acestrorhynchus isalineae , and A. Acestrorhynchus nasutus ; and (3) Acestrorhynchus abbreviatus , Acestrorhynchus altus , Acestrorhynchus falcatus , Acestrorhynchus lacustris , and Acestrorhynchus pantaneiro . In one hypothesis A. heterolepis is proposed as the closest relative of the clade formed by A. falcirostris , A. isalineae , and A. nasutus , and in the alternative hypothesis it is proposed as a sister species of the clade formed by A. abbreviatus , A. altus , A. falcatus , A. lacustris , and A. pantaneiro . Relationships among species of the latter clade remain unresolved. Two independent episodes of reduction of body size are hypothesized to have occurred within the genus: one associated with the clade formed by A. grandoculis and A. minimus , and the other with the clade formed by A. isalineae and A. nasutus . © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 691–757.     

Weintrauboa, a new genus of pimoid spiders from Japan and adjacent islands, with comments on the monophyly and diagnosis of the family Pimoidae and the genus Pimoa (Araneoidea, Araneae)

The spider genus Weintrauboa new genus (Araneae, Pimoidae) is described to place two species of pimoids from Japan and adjacent islands that were formerly classified in the linyphiid genus Labulla . Weintrauboa contortipes (Karsch) new comb., the type species, and W. chikunii (Oi) new comb. are redescribed. Parsimony analysis of morphological characters provides robust support for the monophyly of the genus Weintrauboa and corroborates the monophyly of Pimoa , Pimoidae, and the clade Linyphiidae plus Pimoidae. New diagnoses for Pimoa and Pimoidae are provided.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 261–281     

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.     

High genetic variability in Neotropical myophilous orchids

The patterns of genetic variability of seven Bulbophyllum Thouars species were investigated using 14 enzymatic loci, and compared with those of other Neotropical myophilous orchid species. The genetic variability estimated was very high ( H _e = 0.39–0.61, P  = 86–100%, A  = 2.6–3.8), despite the small population sizes, which may be explained by the vegetative reproduction. Of the three species with multiple populations, only B. epiphytum showed moderate values of genetic structuring. Environmental characteristics and seed dispersal mechanisms are important in understanding the differences in the population structure observed. Almost all pairs of species showed low genetic similarity, indicating a long period of divergence. Our results suggest that B. adiamantinum and B. insectiferum , species currently classified in different sections of the genus, are more closely related than previously thought.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 33–40.     

Phylogenetic relationships based on morphology among the Diospyros (Ebenaceae) species endemic to the Mascarene Islands

The Diospyros (Ebenaceae) species which colonized the Mascarene Islands, namely Mauritius, Reunion and Rodrigues, have been decimated over the years by human settlements. Of the 14 endemic species that have been described and collected for herbaria, Diospyros angulata is now believed to be extinct in Mauritius. The phylogenetic relationships of the 14 Diospyros species were determined using maximum parsimony analysis of 35 morphological characters. This analysis separated the Mascarene Diospyros into two major clades, with D. revaughanii , D. egrettarum and D. leucomelas grouped in the same strongly supported most basal clade while the rest of the species formed the other major clade. High bootstrap values were obtained for the sister species D. angulata and D. boutonania , and the clade clustering the upland species D. neraudii , D. nodosa and D. pterocalyx . There was also relatively strong support for the clade comprising D. hemiteles and D. melanida , which are located in mid altitude regions. These results indicate that Diospyros species most probably colonized the coastal areas of Mauritius and then moved to mid altitude habitats before finally reaching the upland regions. There are also strong indications that D. borbonica and D. diversifolia , endemic to Reunion and Rodrigues, respectively, resulted from migrations from Mauritius.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 307–313.     

Molecular evidence for phylogenetic relationships of Jacquemontia reclinata House (Convolvulaceae) – a critically endangered species from south Florida

Jacquemontia reclinata House (Convolvulaceae) is a federally listed endangered species endemic to coastal strand habitat of south-eastern Florida. In order to identify the closest relatives of J. reclinata , a phylogenetic reconstruction was performed based on nucleotide sequence data from the internal transcribed spacer region of nuclear ribosomal DNA and from the trnH-psbA intergenic spacer of chloroplast DNA. The three putative relatives of J. reclinata and the majority of Jacquemontia Choicy species known to occur in the Caribbean Islands were included. The strict consensus tree concurs with previous morphological studies, indicating that J. reclinata is closely related to the Caribbean species J. cayensis Britton, J. curtisii Hallier f., and J. havanensis (Jacq.) Urb. These three species and J. reclinata form an unresolved clade. Nucleotide divergence within this clade is low, suggesting that the group is recent, perhaps diversifying after the latest glacial period. These taxa need to be studied further using population-level DNA markers. Most species endemic to the Caribbean Basin form a strongly supported clade. Our phylogeny supports Convolvulus nodiflorus Desr. as part of Jacquemontia . In addition, J. ovalifolia (West) Hallier f. ssp. sandwicensis (A. Gray) K.R. Robertson is sister to the Antillean endemic J. solanifolia (L.) Hallier f. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 443–454.     

Taxonomic notes on the genus Crataegus (Rosaceae) in Turkey

The Turkish Crataegus taxa were investigated using morphological, palynological, and anatomical characters. A new series ( Crataegus Section Crataegus Series Peshmenia ), two new species ( Crataegus peshmenii and Crataegus christensenii ), and one variety ( Crataegus rhipidophylla var. kutahyaensis ) are described. Furthermore, Crataegus  ×  browicziana has been assigned to Crataegus rhipidophylla with a new status. Illustrations of the described taxa and their distribution map are also given. The lectotype for Crataegus yaltirikii is designated here.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 231–240.     

Intraspecific variation in Anolis sagrei mirrors the adaptive radiation of Greater Antillean anoles

Anolis lizards of the Greater Antilles represent one of the classic examples of vertebrate adaptive radiation. The same morphological types ('ecomorphs') have evolved repeatedly in response to similar ecological pressures on different islands. We tested whether patterns of within species diversification were congruent with between species patterns and the processes leading to the adaptive radiation of Greater Antillean anoles by measuring variation in performance-related morphological characters in the brown anole, Anolis sagrei . We measured morphological and genetic variation in two different habitat types on each of five islands in the Bahamas. We estimated population structure and rates of gene flow within and among islands using eight microsatellite markers. Intraspecific variation in performance-related morphological characters was similar to the pattern of interspecific variation that characterizes the adaptive radiation of this group in the Greater Antilles. For example, limb length was correlated with perch diameter within A. sagrei as has also been shown among species of anole. Morphological divergence in traits has occurred despite relatively high levels of gene flow both within and among islands. These results are discussed in the context of the divergence-with-gene-flow model of speciation. The results provide important intraspecific evidence that the diversification of anoles has been shaped by natural selection and show how ecologically-based selection pressures explain diversification at both the population and species levels.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 189–199.     

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.     

Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

With multiple well-preserved specimens, tyrannosaurid dinosaurs are ideal subjects for exploring the role of the neck for feeding in an extinct amniote clade. Detailed exploration of the morphology of avian and crocodilian neck muscles grounds phylogenetic inference of musculature of tyrannosaurids. Introduced methods of kinematic and physiological inference explicate action and function of tyrannosaurid neck muscles, and the explicit method of extant behavioural interpolation (EBI) allows inference of generalized behaviours. Anteriorly originating craniocervical muscles of tyrannosaurids, and insertions of the neck dorsiflexor m. transversospinalis cervicis, were similar to those of birds, while the major respective head dorsiflexor and lateroflexor, m. transversospinalis capitis and m. longissimus capitis superficialis, resembled their homologues in crocodilians. Because the proposed inferential methods are cumulative and emphasize mutual falsification, behavioural effects of muscle function become immediately testable rather than assumed. Kinematic inference of muscle moment topology, and bracketed and unipolar inference of muscle activity, facilitate EBI of rapid gaze shifts, lateral and sagittal strikes, inertial feeding, and tearing of flesh by specific permutational actions of tyrannosaurid neck muscles. Several examples of EBI are tested further through biomechanical considerations.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 759–808.     

A new species of Lavatera Sect. Olbia (Medik.) DC. (Malvaceae) from north-east Morocco

As a result of recent fieldwork in north-east Morocco, a new woody Malvaceae species, Lavatera valdesii Molero & J.M. Monts., is described here. The morphological characters that link it to other species in Sect. Olbia (Medik.) DC. are discussed, as are those traditionally used in defining the species that make up this section. A recently described species of this group, Lavatera plazzae Atzei, is established here as a synonym of L. stenopetala Coss. & Durieu ex Batt., a species previously considered as an endemic of east Algeria. On the basis of morphological characters, we propose the exclusion of both L. thuringiaca and L. cashmiriana Cambess. from Sect. Olbia.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 445–454.     

Species-richness of the Anopheles annulipes complex (Diptera: Culicidae) revealed by tree and model-based allozyme clustering analyses

The Australasian Anopheles annulipes complex contains at least ten sibling species, some of which are important vectors of myxomatosis in rabbits. We aimed to establish how many species occurred among specimens from 61 sites throughout Australia, scored for 32 putative allozyme loci. We compared the number of species predicted from tree-based clustering of operational taxonomic units (OTUs) with that from a novel model-based Bayesian clustering approach for individual genotypes. We rejected the hypothesis of conspecificity of OTUs if they differed by at least 20% fixed differences and 0.300 Nei's standard genetic distance D . According to these criteria, 18–25 species occur, making this the most species-rich anopheline complex known to date. A conservative estimate from the Bayesian analysis was 15–20 species. There was large overlap in the assignment of individuals to clusters inferred from the Bayesian and tree-based analyses. The genetic clustering of northern and southern distributed species and an apparent cline in alleles of the enzyme glucose phosphate isomerase suggest that a latitude-dependent factor, such as temperature, may have played a role in speciation and the subsequent distribution of species. Ecological niche modelling of clusters predicted that none occur in New Guinea, emphasizing that additional, as yet unsampled, species may occur.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 523–539.