Evidence from mitochondrial genomics supports the lower Mesozoic of South Asia as the time and place of basal divergence of cypriniform fishes (Actinopterygii: Ostariophysi)

We analysed mitochondrial genomic sequences under maximum likelihood (ML) criteria to explore phylogenetic relationships, and performed historical biogeography analysis with divergence time estimation for fishes of Order Cypriniformes (Actinopterygii: Ostariophysi). We added mitogenomes for eight new cypriniforms and one outgroup to a data set comprising 53 and six outgroup mitogenomes from a previous study to make our taxon sampling geographically representative. The ML tree reconfirmed monophyly of four basal cypriniform clades (cyprinids, catostomids, gyrinocheilids, and loaches including balitorids and cobitids). It also recovered 18 monophyletic groups largely equivalent to the subfamilial rank, and resolved interrelationships among most of these subfamilial clades. However, lower bootstrap support for the ML tree and higher approximately unbiased (au) probabilities for alternative topologies around some branches indicated problems that still need to be resolved. Historical taxon biogeography by dispersal‐vicariance analysis, a parsimonious reconstruction of past ranges, and gain‐loss ratio analysis at the subfamilial level, identified the geographical region of basal cypriniform divergence as southern Asia. Bayesian divergence time analysis dated the basal otophysan split, which gave birth to Order Cypriniformes, to the late Triassic around 219.5 Mya. The basal cypriniform divergence took place during the late Jurassic around 155.9 Mya. These dates coincide with the onset and completion, respectively, of the Pangaean breakup. Taking biogeographical analysis and node dating into account, we consider the most likely candidate for the initial geographical range of Order Cypriniformes to be the south‐eastern area of Mesozoic Laurasia (present‐day southern Asia, excluding the Indian subcontinent). We also briefly discuss ecological implications of the group's divergence. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 633–662.     

Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae)

We infer for the first time the phylogenetic relationships of genera and tribes in the ecologically and evolutionarily well‐studied subfamily Nymphalinae using DNA sequence data from three genes: 1450 bp of cytochrome oxidase subunit I (COI) (in the mitochondrial genome), 1077 bp of elongation factor 1‐alpha (EF1‐α) and 400–403 bp of wingless (both in the nuclear genome). We explore the influence of each gene region on the support given to each node of the most parsimonious tree derived from a combined analysis of all three genes using Partitioned Bremer Support. We also explore the influence of assuming equal weights for all characters in the combined analysis by investigating the stability of clades to different transition/transversion weighting schemes. We find many strongly supported and stable clades in the Nymphalinae. We are also able to identify ‘rogue’ taxa whose positions are weakly supported (the different gene regions are in conflict with each other) and unstable. Our main conclusions are: (1) the tribe Coeini as currently constituted is untenable, and Smyrna, Colobura and Tigridia are part of Nymphalini; (2) ‘Kallimini’ is paraphyletic with regard to Melitaeini and should be split into three tribes: Kallimini s.s., Junoniini and Victorinini; (3) Junoniini, Victorinini, Melitaeini and the newly circumscribed Nymphalini are strongly supported monophyletic groups, and (4) Precis and Junonia are not synonymous or even sister groups. The species Junonia coenia, a model system in developmental biology, clearly belongs in the genus Junonia. A dispersal‐vicariance analysis suggests that dispersal has had a major effect on the distributions of extant species, and three biotic regions are identified as being centres of diversification of three major clades: the Palaearctic for the Nymphalis‐group, the Afrotropics for Junoniini and the Nearctic for Melitaeini. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 227–251.     

Molecular phylogeny and historical biogeography of the large carpenter bees, genus Xylocopa (Hymenoptera: Apidae)

The biogeographical history of major groups of bees with worldwide distributions have often been explained through hypotheses based on Gondwanan vicariance or long distance dispersal events, but until recently these hypotheses have been very difficult, if not impossible, to distinguish. New fossil data, comprehensive information on Mesozoic and Cenozoic coastline positions and the availability of phylogenetically informative DNA markers now makes it feasible to test these hypotheses for some groups of bees. This paper presents historical biogeographical analyses of the genus Xylocopa Latreille, based on phylogenetic analyses of species belonging to 22 subgenera using molecular data from two nuclear genes, elongation factor‐1α (EF‐1α) and phosphoenolpyruvate carboxykinase (PEPCK), combined with previously published morphological and mitochondrial data sets. Phylogenetic analyses based on parsimony and likelihood approaches resulted in several groups of subgenera supported by high bootstrap values (>85%): an American group with the Oriental/Palaearctic subgenera Nyctomelitta and Proxylocopa as sister taxa; a geographically diverse group (Xylocopa s.l); and a group consisting of African and Oriental subgenera. The relationships among these three clades and the subgenus Perixylocopa remained unresolved. The Oriental subgenus Biluna was found to be the sister group of all other carpenter bee subgenera included in this study. Using a relaxed molecular clock calibrated using fossil carpenter bees, we show that the major splits in the carpenter bee phylogeny occurred well after the final breakup of Gondwanaland (the separation of South America and Africa, 100 Mya), but before important Miocene fusion events. Ancestral area analysis showed that the genus Xylocopa most likely had an Oriental‐Palaearctic origin and that the present world distribution of Xylocopa subgenera resulted mainly from independent dispersal events. The influence of Pleistocene glaciations on carpenter bee distributions is also discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 249–266.     

A maximum likelihood approach to generate hypotheses on the evolution and historical biogeography in the Lower Volga Valley regions (southwest Russia)

The evolution of the diverse flora in the Lower Volga Valley (LVV) (southwest Russia) is complex due to the composite geomorphology and tectonic history of the Caspian Sea and adjacent areas. In the absence of phylogenetic studies and temporal information, we implemented a maximum likelihood (ML) approach and stochastic character mapping reconstruction aiming at recovering historical signals from species occurrence data. A taxon-area matrix of 13 floristic areas and 1018 extant species was constructed and analyzed with RAxML and Mesquite. Additionally, we simulated scenarios with numbers of hypothetical extinct taxa from an unknown palaeoflora that occupied the areas before the dramatic transgression and regression events that have occurred from the Pleistocene to the present day. The flora occurring strictly along the river valley and delta appear to be younger than that of adjacent steppes and desert-like regions, regardless of the chronology of transgression and regression events that led to the geomorphological formation of the LVV. This result is also supported when hypothetical extinct taxa are included in the analyses. The history of each species was inferred by using a stochastic character mapping reconstruction method as implemented in Mesquite. Individual histories appear to be independent from one another and have been shaped by repeated dispersal and extinction events. These reconstructions provide testable hypotheses for more in-depth investigations of their population structure and dynamics.     

Phylogenetic position of the adeleorinid coccidia (Myzozoa, Apicomplexa, Coccidia, Eucoccidiorida, Adeleorina) inferred using 18S rDNA sequences

Investigating the evolutionary relationships of the major groups of Apicomplexa remains an important area of study. Morphological features and host-parasite relationships continue to be important in the systematics of the adeleorinid coccidia (suborder Adeleorina), but the systematics of these parasites have not been well-supported or have been constrained by data that were lacking or difficult to interpret. Previous phylogenetic studies of the Adeleorina have been based on morphological and developmental characters of several well-described species or based on nuclear 18S ribosomal DNA (rDNA) sequences from taxa of limited taxonomic diversity. Twelve new 18S rDNA sequences from adeleorinid coccidia were combined with published sequences to study the molecular phylogeny of taxa within the Adeleorina and to investigate the evolutionary relationships of adeleorinid parasites within the Apicomplexa. Three phylogenetic methods supported strongly that the suborder Adeleorina formed a monophyletic clade within the Apicomplexa. Most widely recognized families within the Adeleorina were hypothesized to be monophyletic in all analyses, although the single Hemolivia species included in the analyses was the sister taxon to a Hepatozoon sp. within a larger clade that contained all other Hepatozoon spp. making the family Hepatozoidae paraphyletic. There was an apparent relationship between the various clades generated by the analyses and the definitive (invertebrate) host parasitized and, to lesser extent, the type of intermediate (vertebrate) host exploited by the adeleorinid parasites. We conclude that additional taxon sampling and use of other genetic markers apart from 18S rDNA will be required to better resolve relationships among these parasites.     

A mitogenomic phylogeny and genetic history of sable (Martes zibellina)

We assessed phylogeny of sable (Martes zibellina, Linnaeus, 1758) by sequence analysis of nearly complete, new mitochondrial genomes in 36 specimens from different localities in northern Eurasia (Primorye, Khabarovsk and Krasnoyarsk regions, the Kamchatka Peninsula, the Kuril Islands and the Urals). Phylogenetic analysis of mtDNA sequences demonstrates that two clades, A and BC, radiated about 200–300 thousand years ago (kya) according to results of Bayesian molecular clock and RelTime analyses of different mitogenome alignments (nearly complete mtDNA sequences, protein-coding region, and synonymous sites), while the age estimates of clades A, B and C fall within the Late Pleistocene (~ 50–140 kya). Bayesian skyline plots (BSPs) of sable population size change based on analysis of nearly complete mtDNAs show an expansion around 40 kya in the warm Karganian time, without a decline of population size around the Last Glacial Maximum (21 kya). The BSPs based on synonymous clock rate indicate that M. zibellina experienced demographic expansions later, approximately 22 kya. The A2a clade that colonized Kamchatka ~ 23–50 kya (depending on the mutation rate used) survived the last glaciation there as demonstrated by the BSP analysis. In addition, we have found evidence of positive selection acting at ND4 and cytochrome b genes, thereby suggesting adaptive evolution of the A2a clade in Kamchatka.     

Diversification process of stag beetles belonging to the genus Platycerus Geoffroy (Coleoptera: Lucanidae) in Japan based on nuclear and mitochondrial genes

Molecular phylogenies of the genus Platycerus in Japan were characterized based on the nuclear 28S ribosomal RNA and mitochondrial cytochrome oxidase subunit I (COI) genes. These analyses, combined with our previous morphological information, revealed a detailed diversification process of Platycerus in Japan, as well as estimates of their divergence times. Japanese species were monophyletic and were inferred to have diverged to the acuticollis species group and all other species ca 1.69 Mya. The acuticollis species group then appeared to split into P. viridicuprus and a group including P. takakuwai, P. albisomni and P. acuticollis ca 1.26 Mya. Other specific divergences have occurred primarily since ca 0.33 Mya. Comparing the molecular trees and the morphological tree, we also found introgression of the COI gene in some species. Genetic divergence of Platycerus has occurred intensely in southwestern Japan.     

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.     

A mitogenomic perspective on the phylogeny and biogeography of living caecilians (Amphibia: Gymnophiona)

The caecilians, members of the amphibian Order Gymnophiona, are the least known Order of tetrapods, and their intra-relationships, especially within its largest group, the Family Caeciliidae (57% of all caecilian species), remain controversial. We sequenced thirteen complete caecilian mitochondrial genomes, including twelve species of caeciliids, using a universal primer set strategy. These new sequences, together with eight published caecilian mitochondrial genomes, were analyzed by maximum parsimony, partitioned maximum-likelihood and partitioned Bayesian approaches at both nucleotide and amino acid levels, to study the intra-relationships of caecilians. An additional multiple gene dataset including most of the caecilian nucleotide sequences currently available in GenBank produced phylogenetic results that are fully compatible with those based on the mitogenomic data. Our phylogenetic results are summarized as follow. The caecilian family Rhinatrematidae is the sister taxon to all other caecilians. Beyond Rhinatrematidae, a clade comprising the Ichthyophlidae and Uraeotyphlidae is separated from a clade containing all remaining caecilians (Scolecomorphidae, Typhlonectidae and Caeciliidae). Within this large clade, Scolecomorphidae is the sister taxon of Typhlonectidae and Caeciliidae but this placement did not receive strong support in all analyses. Caeciliidae is paraphyletic with regard to Typhlonectidae, and can be divided into three well-supported groups: Caeciliidae group 1 contains the African caeciliids Boulengerula and Herpele; Caeciliidae group 2 contains Caecilia and Oscaecilia and it is the sister taxon of Typhlonectidae; Caeciliidae group 3 comprises the remaining species of caeciliids. The mitochondrial genome data were also used to calculate divergence times for caecilian evolution using the penalized likelihood method implemented in the program R8S. The newly obtained dating results are compatible with (but a little older than) previous time estimates mainly based on nuclear gene data. The mitogenomic time tree of caecilians suggests that the initial diversification of extant caecilians most probably took place in Late Triassic about 228 (195–260) Ma. Caeciliids currently distributed in India and the Seychelles diverged from their African and American relatives most probably in Late Jurassic about 138 (112–165) Ma, fairly close to the time (130 Ma) when Madagascar–India–Seychelles separated from Africa and South America. The split between the Indian caeciliid Gegeneophis and Seychellean caeciliids occurred about 103 (78–125) Ma, predated the rifting of India and the Seychelles (65 Ma).     

Phylogenetics, zoogeography, and the role of dispersal and vicariance in the evolution of the Rana catesbeiana (Anura: Ranidae) species group

The Rana catesbeiana species group consists of seven species, each variously distributed across eastern North America. We estimated the evolutionary relationships among 31 exemplars and used the phylogenetic hypothesis to examine the potential modes of speciation and relative role of dispersal in the evolution and zoogeography of this species group. Phylogenetic relationships based on 1554 combined base pairs of the cytochrome b and ND2 mitochondrial genes suggest that the species are closely related, having undergone rapid radiation from a common ancestor during the late Miocene or Pliocene. A Pleistocene origin for the rare R. okaloosae is suggested by its pattern of paraphyly with R. clamitans and by its geographically restricted distribution, although hybridization as the explanation for paraphyly cannot be ruled out. Dispersal–vicariance analysis suggested a Coastal Plain biogeographical region origin of the species group, supporting the notion that the region was an important centre of anuran diversification, with post-speciation dispersal playing a major role in explaining the distribution of the widespread species, R. catesbeiana , R. clamitans, and R. septentrionalis . High sea levels during the late Tertiary, greatly reducing and insularizing parts of the southern Coastal Plain region may have played a major role in the diversification of this group.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 601–624.     

Lineage diversification on an evolving landscape: phylogeography of the California newt, Taricha torosa (Caudata: Salamandridae)

We used mitochondrial cytochrome  b sequences (up to 778 bp) and starch gel electrophoresis (45 loci) to examine the phylogeographical history of 39 populations of the California newt, Taricha torosa . Phylogenetic and population genetic methods were integrated to infer history at deep and shallow time depths. Using a molecular clock, the subspecies T. t. torosa and T. t. sierrae were estimated to have diverged 7–13 Mya. Within T. t. torosa, genetically differentiated groups were identified along coastal California, in southern California, and in the southern Sierra Nevada. The coastal group exhibited isolation by distance, but a lack of genetic variation north of present-day Monterey was indicative of a recent range expansion. In southern California, a disjunct population in central San Diego County was genetically diverged from coastal populations to the north (Nei's genetic distance of 0.113). However, mtDNA and protein data were geographically discordant regarding the boundary between the coastal and southern Californian groups, and a biogeographical scenario was developed to account for this discordance. The southern Sierran clade of T. t. torosa was weakly diverged from coastal populations for mtDNA sequence variation, yet was strongly differentiated for allozyme variation (Nei's genetic distance of 0.17–0.20). Populations of T. t. sierrae exhibited substantial population structure, and showed a steeper pattern of isolation by distance than did coastal populations of T. t. torosa . These results are interpreted in consideration of the known geomorphological history of California.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 213–239.     

Phylogeography of the tailed frog (Ascaphus truei): implications for the biogeography of the Pacific Northwest

Tailed frogs are distributed in high-gradient streams within the disjunct mesic forests of the Pacific Northwest and represent the basal lineage of the anurans. We sequenced 1,530 nucleotides of the mitochondrial cytochrome b and NADH dehydrogenase subunit two genes from 23 populations and used parsimony, maximum-likelihood, and nested-clade analyses to estimate relationships among populations and infer evolutionary processes. We found two divergent haplotype clades corresponding with inland Rocky Mountain populations and coastal populations and separated by up to 0.133 substitutions per site. Within the coastal assemblage, haplotypes formed clades by mountain range with 0.010-0.024 substitutions per site divergence among populations. Inland haplotypes exhibited minimal genetic structure, with the exception of 0.021 substitutions per site distance between populations from the East Fork of the South Fork of the Salmon River and all other inland haplotypes. The magnitude of divergence between inland and coastal populations, as well as the paleobotanical record, suggest isolation of these lineages occurred during the late Miocene to early Pliocene, probably in response to the rise of the Cascade Mountains. Genetic structure within coastal and inland populations is consistent with isolation in refugia during the late Pliocene and early Pleistocene. Closely related inland haplotypes reflect range expansion following glaciation. The depth of divergence between inland and coastal populations supports the persistence of mesic forests within the inland Pacific Northwest throughout the Pleistocene and is congruent with patterns found in several other mesic forest species. Based on mitochondrial divergence and previous allozyme and morphological data, we recommend recognition of inland populations as a distinct species, Ascaphus montanus.     

Molecular systematics of the Asian torrent minnows (Ostariophysi: Psilorhynchidae) inferred from nuclear and mitochondrial DNA sequence data

The phylogenetic relationships between members of the South Asian family Psilorhynchidae are investigated using both mitochondrial and nuclear DNA sequence data. Phylogenetic hypotheses were derived from maximum likelihood and maximum parsimony analyses of a three gene concatenated data set, including cytochrome c oxidase subunit 1 (640 bp), cytochrome b (1140 bp) and exon 3 of the recombination‐activating gene 1 (~1500 bp). Our investigation provides strong support for the monophyly of two species groups of Psilorhynchus (the P. balitora and P. nudithoracicus species groups) and corroborates previous hypotheses on the phylogenetic position of the Western Ghats species Psilorhynchus tenura based on morphology. Basal relationships within Psilorhynchus were poorly supported and are worthy of further investigation. A fossil calibrated relaxed molecular clock estimates the split between Psilorhynchus and its sister group to have occurred within the Eocene/Oligocene, with subsequent diversification in the Miocene.     

Phylogeny and historical biogeography of the subtribe Aporiina (Lepidoptera: Pieridae): implications for the origin of Australian butterflies

The Australian fauna is composed of several major biogeographical elements reflecting different spatial and temporal histories. Two groups of particular interest are the Gondwanan Element, reflecting an ancient origin in Gondwana or southern Gondwana (southern vicariance hypothesis), and the Asian Element, reflecting a more recent origin in Asia, Eurasia or Laurasia (northern dispersal hypothesis). Theories regarding the origin and evolution of butterflies (Hesperioidea, Papilionoidea) in Australia are controversial, with no clear consensus. Here, we investigate the phylogenetic and historical biogeographical relationships of the subtribe Aporiina, a widespread taxon with disjunct distributions in each of the major zoogeographical regions. Attention is paid to origins of the subtribe in the Australian Region for which several conflicting hypotheses have been proposed for the Old World genus Delias Hübner. Our phylogenetic reconstruction was based on analysis of fragments of two nuclear genes (elongation factor‐1α, wingless) and one mitochondrial gene (cytochrome oxidase subunit I) for 30 taxa. Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined data set (2729 bp; 917 parsimony informative characters) recovered six major lineages within the monophyletic Aporiina, with the following topology: (Cepora + Prioneris + (Mylothris + (Aporia + Delias group + Catasticta group))). Given a probable age of origin of the stem‐group near the Cretaceous/Tertiary boundary (69–54 Mya), followed by diversification of the crown‐group in the early to mid Tertiary (57–45 Mya), we show that an origin of the Aporiina in either southern Gondwana or Laurasia is equally parsimonious, and that dispersal has played a major role in shaping the underlying phylogenetic pattern. We tentatively conclude that an origin in southern Gondwanan is more likely; however, neither hypothesis satisfactorily explains the present‐day distribution, and additional lower‐level phylogenies are needed to determine the directionality of dispersal events of several taxa and to reject one hypothesis over the other. Dispersal is inferred to have occurred primarily during cooler periods when land bridges or stepping‐stones were available between many of the zoogeographical regions. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 413–440.     

Eroding abodes and vanished bridges: historical biogeography of the substrate specialist pebble-mound mice (Pseudomys)

Aim  To determine whether the pronounced ecological importance of pebble mounds to pebble-mound mice ( Pseudomys ) is manifest in their continental biogeography.
Location  Northern Australia.
Methods  A GIS-based comparison was made between the habitats contained within the potential climatic distributions of mice, representing a null hypothesis of no habitat selection, and their actual distributions based on all known location records.
Results  All species had a clear preference for hilly, rocky landscapes with a surficial cover dominated by bedrock. Simple vegetation communities with relatively open eucalypt overstorey and grassy understorey were preferred. Highly degraded rocks and aggradational surfaces and plains were avoided. The extent of the summer monsoon may be important in determining the southern limits of the group's distribution. Major disjunctions between species were attributable to the presence of clay plains and sand sheets.
The behavioural requirement of pebble-mound mice for mounds determines their population distribution pattern and the distribution of the different species within the genus.
Main conclusions  The behavioural need for pebble mounds drives the distributional pattern of populations and species of pebble-mound mice. The initial spread of pebble-mound mice probably occurred during the late Pliocene or earliest Pleistocene. There has predominantly been degradation of the potential distribution of the group since that time due to the stability of Australian landscapes and Pleistocene planation and sand sheet development over large areas of northern Australia. This process is ongoing, and past regions of rocky contact between current distributions have disappeared, while the distributional limits of several species are steadily being reduced by erosion of hills and the spread of dune fields.     

The history of a nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo)

Previous hypotheses of phylogenetic relationships among Nearctic toads (Bufonidae) and their congeners suggest contradictory biogeographic histories. These hypotheses argue that the Nearctic Bufo are: (1) a polyphyletic assemblage resulting from multiple colonizations from Africa; (2) a paraphyletic assemblage resulting from a single colonization event from South America with subsequent dispersal into Eurasia; or (3) a monophyletic group derived from the Neotropics. We obtained approximately 2.5 kb of mitochondrial DNA sequence data for the 12S, 16S, and intervening valine tRNA gene from 82 individuals representing 56 species and used parametric bootstrapping to test hypotheses of the biogeographic history of the Nearctic Bufo. We find that the Nearctic species of Bufo are monophyletic and nested within a large clade of New World Bufo to the exclusion of Eurasian and African taxa. This suggests that Nearctic Bufo result from a single colonization from the Neotropics. More generally, we demonstrate the utility of parametric bootstrapping for testing alternative biogeographic hypotheses. Through parametric bootstrapping, we refute several previously published biogeographic hypotheses regarding Bufo. These previous studies may have been influenced by homoplasy in osteological characters. Given the Neotropical origin for Nearctic Bufo, we examine current distributional patterns to assess whether the Nearctic-Neotropical boundary is a broad transition zone or a narrow boundary. We also survey fossil and paleogeographic evidence to examine potential Tertiary and Cretaceous dispersal routes, including the Paleocene Isthmian Link, the Antillean and Aves Ridges, and the current Central American Land Bridge, that may have allowed colonization of the Nearctic.     

The complete mitochondrial genome of the Chinese hook snout carp Opsariichthys bidens (Actinopterygii: Cypriniformes) and an alternative pattern of mitogenomic evolution in vertebrate

The complete mitochondrial genome sequence of the Chinese hook snout carp, Opsariichthys bidens, was newly determined using the long and accurate polymerase chain reaction method. The 16,611-nucleotide mitogenome contains 13 protein-coding genes, two rRNA genes (12S, 16S), 22 tRNA genes, and a noncoding control region. We use these data and homologous sequence data from multiple other ostariophysan fishes in a phylogenetic evaluation to test hypothesis pertaining to codon usage pattern of O. bidens mitochondrial protein genes as well as to re-examine the ostariophysan phylogeny. The mitochondrial genome of O. bidens reveals an alternative pattern of vertebrate mitochondrial evolution. For the mitochondrial protein genes of O. bidens, the most frequently used codon generally ends with either A or C, with C preferred over A for most fourfold degenerate codon families; the relative synonymous codon usage of G-ending codons is greatly elevated in all categories. The codon usage pattern of O. bidens mitochondrial protein genes is remarkably different from the general pattern found previously in the relatively closely related zebrafish and most other vertebrate mitochondria. Nucleotide bias at third codon positions is the main cause of codon bias in the mitochondrial protein genes of O. bidens, as it is biased particularly in favor of C over A. Bayesian analysis of 12 concatenated mitochondrial protein sequences for O. bidens and 46 other teleostean taxa supports the monophyly of Cypriniformes and Otophysi and results in a robust estimate of the otophysan phylogeny.     

Comparative historical biogeography of Plateumaris leaf beetles (Coleoptera: Chrysomelidae) in Japan: interplay between fossil and molecular data

Aim In an attempt to use molecular and fossil data interactively in historical biogeography, we studied the phylogeography of five Plateumaris leaf beetles in Japan using mitochondrial cytochrome oxidase subunit I (COI) sequence data to explore interspecific differences in phylogeographical patterns and estimate the timings of colonization and geographical differentiation. Location A total of 461 beetles from five species on Hokkaido, Honshu and Kyushu islands of Japan were analysed with 117 beetles from three conspecies and two congeners from the mainland (Russia, including Sakhalin; Korea; Mongolia; Belgium; France; hereafter, the continent). Methods Using the sequence data from a 750‐bp portion of the COI gene, we studied the phylogeny of COI haplotypes, intraspecific population differentiation using analysis of molecular variance and the Mantel test, and intraspecific phylogeography using nested clade analysis. In addition, divergence times between the continental and Japanese lineages, as well as among the various Japanese lineages, were estimated using a Bayesian approach with node constraints based on fossil records of extant species. Results Three widely distributed species showed different degrees of geographical differentiation corresponding to their different colonization history in Japan. Bayesian estimates of divergence time revealed that one of two endemic species, which originated before the late Pliocene, attained intraspecific differentiation through the Pliocene and Pleistocene, whereas another endemic species has been confined in one locality, and three non‐endemic species colonized Japan after the mid‐Pleistocene. Main conclusions Molecular analyses of an insect group with relatively abundant fossil data can contribute greatly to the understanding of diverse biogeographical histories of related species in a region. Bayesian estimates of divergence time could be used to assess the variable evolutionary rates of the COI gene, and may be applied to other related insect species.     

The biogeography and historical ecology of Lobelia urens L.(r)(the heath lobelia) in southern England

This paper examines the past and present ecology and biogeography of Lobelia urens L. (the heath lobelia) which is threatened with extinction in southern England. A survey of the historical and documentary evidence of the distribution of the species is presented. Historical records exist for nineteen sites in the south but nowhere else in England. The status of L. urens is in decline in Britain with ten populations having been lost this century. The six remaining sites have been surveyed to describe the phytosociology of L. urens communities and the variability of the environmental factors controlling its distribution. A total of ninety-five quadrats containing 122 species were recorded and analysed using two-way indicator species analysis. Canonical correspondence analysis was used to assess the correlations between twenty-eight environmental variables and the floristic variation in the ninety-five quadrats. These were reduced to sixteen variables for the final analysis. L. urens is shown to be a member of rough, grass-heath communities (NVC M25 and W23) dominated by Molinia caerulea and situated on seasonally waterlogged, moderately acidic, nutrient-poor soils. The species is suited to both cyclical and sporadic soil disturbance regimes. The final discussion argues that existing populations will only be maintained and expansion of the species distribution occur, if conservationists prevent woodland succession in L. urens sites and periodically disturb the ground in locations where the species survives at present and also where it has been observed in the recent past. This link between historical ecology and present-day conservation management would appear very important. Further demographic and experimental work is needed to explore these predictions and their implications for the future conservation management of L. urens .