Biogeography and molecular phylogeny of the genus Schizothorax (Teleostei: Cyprinidae) in China inferred from cytochrome b sequences

Aim To test a vicariant speciation hypothesis derived from geological evidence of large‐scale changes in drainage patterns in the late Miocene that affected the drainages in the south‐eastern Tibetan Plateau. Location The Tibetan Plateau and adjacent areas. Methods The cytochrome b DNA sequences of 30 species of the genus Schizothorax from nine different river systems were analysed. These DNA sequences were analysed using parsimony, maximum likelihood and Bayesian methods. The approximately unbiased and Shimodaira–Hasegawa tests were applied to evaluate the statistical significance of the shortest trees relative to alternative hypotheses. Dates of divergences between lineages were estimated using the nonparametric rate smoothing method, and confidence intervals of dates were obtained by parametric bootstrapping. Results The phylogenetic relationships recovered from molecular data were inconsistent with traditional taxonomy, but apparently reflected geographical associations with rivers. Within the genus Schizothorax, we observed a divergence between the lineages from the Irrawaddy–Lhuit and Tsangpo–Parlung rivers, and tentatively dated this vicariant event back to the late Miocene (7.3–6.8 Ma). We also observed approximately simultaneous geographical splits within drainages of the south‐eastern Tibetan Plateau, the Irrawaddy, the Yangtze and the Mekong–Salween rivers in the late Miocene (7.1–6.2 Ma). Main conclusions Our molecular evidence tentatively highlights the importance of palaeoriver connections and the uplift of the Tibetan Plateau in understanding the evolution of the genus Schizothorax. Molecular estimates of divergence times allowed us to date these vicariant scenarios back to the late Miocene, which agrees with geological suggestions for the separation of these drainages caused by tectonic uplift in south‐eastern Tibet. Our results indicated the substantial role of vicariant‐based speciation in shaping the current distribution pattern of the genus Schizothorax.     

Heavily exploited but poorly known: systematics and biogeography of commercially harvested pythons (Python curtus group) in Southeast Asia

More than 100000 blood pythons (brongersmai) and short-tailed pythons (curtus and breitensteini) are taken from Borneo and Sumatra each year for the commercial leather trade. Traditionally, all have been treated as a single polytypic species (Python curtus) , with three subspecies differing in colour, size and geographic distribution. Analyses of DNA sequences and morphological data clarify the phylogenetic relationships, taxonomy and biogeography of this group. The lineage is monophyletic, and each of the three subspecies differs from the other two both morphologically and genetically. Given the morphological and genetic distinctiveness of each taxon, we here elevate the three subspecies to full species status. Python brongersmai is the most distinctive in terms of colour (of the three, only brongersmai has colour-morphs that are red or orange), size (it grows to 2.6 m, vs. approx. 2.0 m for the other taxa), and scalation (e.g. brongersmai has <166 ventral scales, vs. <166 in the other taxa and has two supralabials over each orbit, vs. one supralabial for the other two taxa). In terms of cytochrome b mitochondrial DNA sequence data, brongersmai is almost as distant genetically from the short-tailed pythons (8.9% divergence) as is the reticulated python (P. reticulatus: 10.3% divergence). The other two taxa (P. breitensteini from Kalimantan and P. curtus from western and southern Sumatra) are closely related (3% divergence), despite their disjunct distribution (separated by P. brongersmai). Sea-level fluctuations provide a plausible biogeographic scenario to explain phylogenetic divergence within this lineage. Given the distinctiveness of the component taxa, and the ease with which even dried skins can be identified to species level (based on ventral counts), the managers of this important commercial resource should no longer treat the P. curtus group as a single biological taxon.     

Genetic diversity and historical population structure of Schizopygopsis pylzovi (Teleostei: Cyprinidae) in the Qinghai–Tibetan Plateau

1. Complete sequences of 1140 base pair of the cytochrome b gene from 133 specimens were obtained from nine localities including the inflow drainage system, isolated lakes and outflow drainage system in Qinghai–Tibetan Plateau to assess genetic diversity and to infer population histories of the freshwater fish Schizopygopsis pylzovi. 2. Nucleotide diversities (π) were moderate (0.0024–0.0045) in populations from the outflow drainage system and Tuosuo Lake, but low (0.0018–0.0021) in populations from Qiadam Basin. It is probable that the low intra‐population variability is related with the paleoenvironmental fluctuation in Qiadam Basin, suggesting that the populations from Qiadam Basin have experienced severe bottleneck events in history. 3. Phylogenetic tree topologies indicate that the individuals from different populations did not form reciprocal monophyly, but the populations from the adjacent drainages cluster geographically. Most population pairwise F_ST tests were significant, with non‐significant pairwise tests between Tuosu Lake and Tuosuo Lake in the north‐west of the Qinghai–Tibetan Plateau. Analysis of molecular variance (amova ) indicates that the significant genetic variation was explained at the levels of catchments within and among, not among specific boundaries or inflow and outflow drainage systems. 4. The nested clade phylogeographical analysis indicates that historical processes are very important in the observed geographical structuring of S. pylzovi, and the contemporary population structure and differentiation of S. pylzovi may be consistent with the historical tectonic events occurred in the course of uplifts of the Qinghai–Tibetan Plateau. Fluctuations of the ecogeographical environment and major hydrographic formation might have promoted contiguous range expansion of freshwater fish populations, whereas the geological barriers among drainages have resulted in the fragmentation of population and restricted the gene flow among populations. 5. The significantly large negative F_s‐value (−24.91, P < 0.01) of Fu's F_s‐test and the unimodal mismatch distribution indicate that the species S. pylzovi underwent a sudden population expansion after the historical tectonic event of the Gonghe Movement. 6. The results of this study indicate that each population from the Qinghai–Tibetan Plateau should be managed and conserved separately and that efforts should be directed towards preserving the genetic integrity of each group.     

High levels of mitochondrial cytochrome b divergence in annual killifishes of the genus Cynolebias (Cyprinodontiformes, Rivulidae)

Phylogenetic relationships based on 324 base pairs of the mitochondrial cytochrome b gene were examined in 14 species of the genus Cynolebias. The monophyly of the genus relative to three outgroup taxa belonging to the family Rivulidae was supported by the sequence data. Bootstrap values corroborated the existence of intrageneric monophyletic units in both parsimony and Neighbour-joining analyses. These include a Cynolebias bellottii-C. wolterstorffl clade, a C. adloJfi-2, C. duraznensis, C. viarius and C adloffi-1 group, and a C. gymnoventris, C. luteoflammulatus pair and the strongly supported assemblage that includes C. prognalhus and C. cheradophilus clade. Phylogenetic relationships remain poorly supported for C. nigripinnis, C. qffinis and C. alexandri, and unresolved among the previous ingroup clades. Cytochrome b sequences reveal an unexpectedly high level of divergence among species of the genus Cynolebias. Consequently, cytochrome b shows good resolution of recent cladogenetic events but limited phylogenetic information at deeper nodes. High levels of sequence divergence span a broad range within Cynolebias. The highest sequence divergence (c. 28%) occurred among C. antenori and the remaining species of the genus. The minimum divergence value (4.5%) is exhibited by sympatric species C. cheradophilus and C. prognathus.     

中国西南部小鲵属的遗传分化

运用同工酶电泳和DNA测序技术,我们研究了中国东南部两种小鲵的遗传多样性。同工酶数据显示义乌小鲵和安吉小鲵之间存在中等水平的种间差异。而线粒体DNA序列数据则显示两种之间存在较高水平的种间差异。安吉小鲵种群存在明显的近亲繁殖现象;义乌小鲵种群间存在明显的分化。舟山群岛上义乌小鲵种群明显有别于大陆种群。     

The molecular phylogenetics of the genus Oligoryzomys (Rodentia: Cricetidae) clarifies rodent host–hantavirus associations

Several species of the genus Oligoryzomys are natural hosts of different hantavirus genotypes affecting humans. The systematics of the genus is confusing, which complicates the identification of the rodent host and hence the potential endemic areas of hantavirus pulmonary syndrome. In this study, we analyse molecular data to infer phylogenetic relationships among Central and South American specimens of Oligoryzomys, and compare our results with previously published data on karyotypic, geographic distribution and host–virus associations to solve contradictory taxonomic reports. We identified 25 clades, each one corresponding to a different putative species. The phylogenetic trees show that Oligoryzomys longicaudatus is strongly related to the Oligoryzomys flavescens complex, which comprises four clades; Oligoryzomys nigripes is related to Oligoryzomys stramineus, Oligoryzomys vegetus is related to Oligoryzomys fulvescens from Central America, and Oligoryzomys brendae is the sister species of Oligoryzomys aff. destructor. We identified the following rodent host–hantavirus genotype relationships: O. longicaudatus–Andes; O. flavescens ‘West'–Bermejo; O. flavescens ‘East'–Lechiguanas; O. nigripes–Juquitiba; Oligoryzomys microtis–Rio Mamore and Rio Mamore‐3; Oligoryzomys chacoensis–Oran; Oligoryzomys costaricencis–Choclo; Oligoryzomys delicatus–Maporal; Oligoryzomys utiaritensis–Castelo dos Sonhos; Oligoryzomys sp. RT2012–Rio Mamore‐4; Oligoryzomys sp. (and not Oligoryzomys fornesi)–Anajatuba. This work, besides contributing to the development of prevention programmes for hantavirus epidemiology in Latin America, represents a comprehensive update of the systematics of the genus Oligoryzomys. © 2014 The Linnean Society of London     

Out of Australia and back again: the world‐wide historical biogeography of non‐pollinating fig wasps (Hymenoptera: Sycophaginae)

Aim Figs (Ficus, Moraceae) are exploited by rich communities of often host‐specific phytophagous wasps. Among them, gall‐inducing Sycophaginae (Hymenoptera, Chalcidoidea) may share a common history with Ficus and their mutualistic pollinators (Agaonidae). We investigate here, for the first time, the phylogeny and biogeographical history of Sycophaginae and compare the timing of radiation and dispersion of major clades with available data on Ficus and fig pollinators. Reconstructing the history of their host colonization and association over space and time is central to understanding how fig wasp communities were assembled. Location World‐wide. Methods Maximum likelihood and Bayesian analyses were conducted on 4267 bp of mitochondrial and nuclear DNA to produce a phylogeny of all genera of Sycophaginae. Two relaxed clock methods with or without rate autocorrelation were used for date estimation. Analyses of ancestral area were also conducted to investigate the geographical origin of the Sycophaginae. Results The phylogeny is well resolved and supported. Our data suggest a post‐Gondwanan origin for the Sycophaginae (50–40 Ma) and two independent out‐of‐Australia dispersal events to continental Asia. Given palaeoclimatic and palaeogeographic records, the following scenario appears the most likely. The ancestor of Idarnes+Apocryptophagus migrated to Greater India through the Ninetyeast Ridge (40–30 Ma). The ancestor of Anidarnes+Conidarnes dispersed later via Sundaland (25–20 Ma). Idarnes and Anidarnes subsequently reached the New World via the North Atlantic land bridges during the Late Oligocene Warming Event. Apocryptophagus reached Africa c. 20 Ma via the Arabic corridors and returned to Australasia following the expansion of Sundaland tropical forests (20–10 Ma). Main conclusions Sycophaginae probably invaded the fig microcosm in Australia c. 50–40 Ma after the origin of their host plant. Once associated with figs, they dispersed out of Australia and radiated together with their host fig and associated pollinator through the tropics. We recorded a good coincidence of timing between dispersal events of Sycophaginae and continental connections. Furthermore, fruit pigeons that disperse figs probably spread out of Australasia through the Indian Ocean via the Ninetyeast Ridge c. 38 Ma. Therefore, our study highlights the potential for combining molecular phylogenetics with multiple methods of dating of interacting groups to reconstruct the historical biogeography of plant–herbivore associations.     

从Cyt b基因序列探讨Grophosoma属和Dybowskyia属的分类地位(半翅目:蝽科)

对蝽科11种昆虫的线粒体细胞色素b（Cyt b）基因部分序列进行测定和分析,在获得的长度为432bp的序列片段中,碱基T、C、 A、G的平均含量分别为38.8%、18.0%、31.6%和11.6%, A＋T平均含量为70.4%,明显高于G＋C含量（29.6 %）.密码子第三位点的A＋T含量高达86.1%.属种间序列变异丰富,有179个核苷酸位点发生变异,变异率为41.4%,碱基替换多发生于第三位点.以筛豆龟蝽Megacopta cribraria为外群构建的系统发育树表明：滴蝽属与蝽亚科其它属关系较远,条蝽属与蝽亚科其它属关系较近,结合形态特征与序列变异情况,支持将滴蝽属从蝽亚科划出并入舌盾蝽亚科,但条蝽属的分类地位仍需要进一步探讨.     

A new and highly divergent mitochondrial lineage in the Small Five-toed Jerboa,Allactaga elater,from Iran (Mammalia: Rodentia)

The Small Five-toed Jerboa, Allactaga elater, is a small rodent adapted to desert and semi-arid habitats with a widespread distribution around the Caucasus. Previous studies have suggested the occurrence of subspecific variation within the species but, except for a recent phylogeny of the genus Allactaga, most of the work done on the taxonomy of the group relies on morphological data only. To contribute to the current understanding of patterns of genetic diversity of A. elater we analysed one mitochondrial locus, cytochrome b, from 13 Iranian specimens. Comparing to a recent phylogeny, our results suggest the existence of two additional mitochondrial lineages, one that clusters within previously described lineages and a new and highly divergent one. The two novel mitochondrial lineages occur in the north and form two highly divergent monophyletic groups (Dxy = 14%), which likely separated during the Pleistocene.     

Phylogeny and phylogeography of squirrel monkeys (genus Saimiri) based on cytochrome b genetic analysis

Squirrel monkeys (genus Saimiri) are distributed over a wide area encompassing the Amazon Basin: French Guiana, Suriname, and Guyana, together with Western Panama and Western Costa Rica. The genus Saimiri includes a complex of species and subspecies displaying considerable morphological variation. Taxonomic and systematic studies have identified, in this genus, one to seven species comprising up to 16 subspecies. The phylogenetic relationships between these taxa are poorly understood. Molecular markers have yielded a consistent framework for the systematics of Central and South American Saimiri, identifying four distinct clades: S. oerstedii, S. sciureus, S. boliviensis, and S. ustus. Here, we reconsider the phylogenetic and biogeographic history of Saimiri on the basis of mitochondrial (mtDNA) sequence data, focusing mostly on individuals originating from the Amazon Basin. We studied 32 monkeys with well‐defined geographic origins and inferred the phylogenetic relationships between them on the basis of full‐length cytochrome b gene nucleotide sequences. The high level of gene diversity observed (0.966) is consistent with the high level of behavioral and morphological variation observed across the geographic range of the genus: 20 mtDNA haplotypes were identified with a maximum divergence of 4.81% between S. b. boliviensis and S. ustus. In addition to confirming the existence of the four clades previously identified on the basis of molecular characters, we suggest several new lineages, including S. s. macrodon, S. s. albigena, S. s. cassiquiarensis, and S. s. collinsi. We also propose new patterns of dispersion and diversification for the genus Saimiri, and discuss the contribution of certain rivers and forest refuges to its structuring. Am. J. Primatol. 72:242–253, 2010. © 2009 Wiley‐Liss, Inc.     

Molecular phylogeny of the genus Apatrobus (Coleoptera: Carabidae: Patrobinae) in western Japan

We performed a molecular phylogenetic analysis of the ground beetles Apatrobus (Carabidae), endemic to Japan, using the mitochondrial cytochrome c oxidase subunit I (COI) and the nuclear 28S rRNA (28S) genes. We focused on the species divergence in Kyushu, Shikoku and western Honshu and used 15 of 19 species and three populations with undetermined species in the DNA analysis. The gene trees showed that, of the Apatrobus species studied, A. hayachinensis Nakane from northern Honshu was not included in the monophyletic group of the other Apatrobus species and likely to be of a different genus. Divergence time estimation suggested that Apatrobus species excluding A. hayachinensis diverged 5.2 million years ago and the subsequent divergence of species occurred during the Pliocene and Pleistocene. In each of the main islands, Kyushu, Shikoku and Honshu, two or more distinct lineages occurred and all species had restricted distribution areas, suggesting that ancient dispersal and vicariance among the three main islands resulted in the nested biogeographical pattern of species distribution.     

New data on the distribution and genetic structure of Greek moles of the genus Talpa (Mammalia,Talpidae)

In this work, we employed molecular markers and confirmed the occurrence of two mole species, Talpa stankovici and Talpa europaea in Greece. For the first species, all analyses revealed three major phylogroups, exhibiting great genetic divergence, possibly due to a vicariant event. This event was probably the Peloponnisos’ insulation during the Pliocene and the following sea level fluctuations. A scenario of stable, large‐sized populations for a long period rather than rapid growth from small‐sized populations or founder events is supported. Additionally, T. stankovici’s distribution area is extended to a southernmost limit. For T. europaea there is evidence of low genetic divergence between Greek and Central and North European populations.     

Species dilemma of musk deer (Moschus spp) in India: molecular data on cytochrome c oxidase I suggests distinct genetic lineage in Uttarakhand compared to other Moschus species

Musk deer are of high conservation priority owing to poaching pressure because of its musk pod. Representation of musk deer status using genetics is poorly documented in India, and it is not confirmed as to how many species of musk deer are present. We characterize for the first time, the genetic diversity of musk deer from Uttarakhand using Cytochrome Oxidase sub-unit (COI) gene (486?bp) and compared with the data available for other species. Results revealed the presence of six haplotypes in the Uttarakhand population amongst 17 sequences. Of these, 12 sequences shared the single haplotype. The intra-species sequences divergence was 0.003–0.017, whereas divergence with other species of musk deer was 0.071–0.081. Bayesian phylogenetic tree revealed that samples from Uttarakhand formed a separate clade with respect to other species of musk deer, whereas three species distributed in China clustered in the same clade and showed low sequences divergence, i.e., 0.002–0.061. Because of different ecomorph reported, we suggest using the barcoding based approach for inter and intra-species distinction and delineating species boundaries across the range for effective conservation. Besides, systematic classification, DNA barcoding would also help in dealing wildlife offence cases for disposal of the legal report in court.     

Genetic structure and differentiation of four populations of Afghan Pika (Ochotona rufescens) in Iran based on mitochondrial cytochrome b gene

The population genetics of the Afghan Pika (Ochotona rufescens) was studied in Northern Khorasan Province, Iran. For prediction of the genetic differentiation of four populations, the DNA of mitochondrial cytochrome b of 32 individuals from four areas was sequenced and a Bayesian analysis based on the HKY model was constructed. In total, 15 polymorphic sites, 1125 conserved sites (98.7%) and 14 different haplotypes were found. The phylogenetic tree resulting from the Bayesian analysis and network analysis showed that all samples were clustered in two major groups and the haplotypes of the four populations did not separate geographically. An analysis of molecular variance (AMOVA) indicated that a large majority of the genetic variance was due to the variance within populations. The results of fixation indices showed significant genetic structure among populations in both methods. The pairwise F_st revealed that two northern populations have a significant genetic differentiation from two southern populations, but no significance pairwise F_st value was demonstrated between the closed populations. Nei's genetic distances between closed populations were not significant, while significant values occurred between distant populations. It seems that there is not a major discontinuity between populations of Afghan Pika based on cyt-b mitochondrial gene. However, phylogenetic analysis did not separate populations and a large majority of the genetic variance was found within populations. However, AMOVA analysis showed a significant level of genetic structure among populations (p?<?0.001) and between groups (p?<?0.5). It seems that these results suggest shallow genetic differentiation between populations of different geographic groups.     

Phylogeny and historical demography of economically important rodents of the genus Arvicanthis (Mammalia: Muridae) from the Nile Valley: of mice and men

The present study clarified the taxonomy, phylogeny and historical demography of semicommensal Nile rats ( Arvicanthis ) from the Nile Valley in Sudan. Nile rats are important crop pests and zoonotic disease reservoirs and are closely associated with agricultural settlements in the Nile Valley. Phylogenetic analysis of the complete cytochrome b gene (1140 bp) of 23 individuals from six localities in Sudan (from two previously recognized species, Arvicanthis niloticus and Arvicanthis testicularis ), supported the existence of only a single species, A. niloticus , from the Nile Valley. Historical demography of the Sudanese Nile Valley population inferred from mismatch coefficients indicated that an exponential population expansion event occurred approximately 144 000–288 000 years ago, corresponding in time with early human expansion and colonization from Africa to the Middle East, Europe, and the world. The inferred high level of gene flow and large size of Sudanese Nile populations of A. niloticus is consistent with historically recent (300 years ago) exponential human population growth and intense agricultural activity inferred from archaeological and historical evidence. Two African Arvicanthis clades were well supported by the broader phylogenetic analysis: (1) A. niloticus , Arvicanthis abyssinicus and Arvicanthis neumanni and (2) Arvicanthis rufinus and Arvicanthis ansorgei from western Africa. Within the first clade, divergence between lineages of A. niloticus s.s. from west and north-east Africa (8.9%) suggests specific recognition, but sampling of geographically intermediate localities is required. Based on hypothesized palaeodrainage and palaeoclimatic patterns, we propose a simple model for speciation of Arvicanthis in Africa.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 641–655.     

Molecular systematics,taxonomy and biogeography of the genus Cavia (Rodentia: Caviidae)

Phylogenetic analyses were conducted on cytochrome b sequence data of the most geographically and taxonomically broad sampling of Cavia taxa to date. Primary objectives included providing the first extensive molecular phylogenetic framework for the genus, testing the taxonomic and systematic hypotheses of previous authors and providing insight into the evolutionary and biogeographic history of the genus. Support was found for the morphologically defined species C. aperea, C. tschudii, C. magna and C. fulgida and the taxonomic placement of taxa previously subject to conflicting taxonomic opinions (e.g. C. nana, C. anolaimae and C. guianae) was further resolved. Additionally, we elevate the Ecuadorian C. a. patzelti to species status, restrict the distributional limits and suggest taxonomic affiliations of some C. tschudii subspecies, and provide strong evidence for the geographic origin of guinea pig domestication. Finally, we provide an estimated evolutionary timeline for the genus Cavia, which appears to extend well into the late Miocene.     

A marine fish follows Wallace's Line: the phylogeography of the three-spot seahorse (Hippocampus trimaculatus, Syngnathidae, Teleostei) in Southeast Asia

Aim To test the potential of two contrasting biogeographical hypotheses (‘Indian/Pacific Ocean Basin’ vs. ‘Wallace's Line’) to explain the distribution of genetic diversity among populations of a marine fish in Southeast Asia. Location The marine waters of Asia and Southeast Asia: from India to Japan, and east to the Indonesian islands of Sulawesi and Flores. Methods We sequenced a 696 base pair fragment of cytochrome b DNA of 100 individuals of Hippocampus trimaculatus Leach 1814 (three‐spot seahorse), obtained from across its range. We tested our hypotheses using phylogenetic reconstructions and analyses of molecular variance. Results Significant genetic divergence was observed among the specimens. Two distinct lineages emerged that diverged by an average of 2.9%. The genetic split was geographically associated, but surprisingly it indicated a major east–west division similar to the terrestrial Wallace's Line (Φ_ST = 0.662, P < 0.001) rather than one consistent with an Indian‐Pacific ocean basin separation hypothesis (Φ_ST = 0.023, P = 0.153). Samples from east of Wallace's Line, when analysed separately, however, were consistent with an Indian/Pacific Ocean separation (Φ_ST = 0.461, P = 0.005). The degree of genetic and geographical structure within each lineage also varied. Lineage A, to the west, was evolutionarily shallow (star‐like), and the haplotypes it contained often occurred over a wide area. Lineage B to the east had greater genetic structure, and there was also some evidence of geographical localization of sublineages within B. Main conclusions Our results indicate that the genetic diversity of marine organisms in Southeast Asia may reflect a more complex history than the simple division between two major ocean basins that has been proposed by previous authors. In particular, the east–west genetic division observed here is novel among marine organisms examined to date. The high haplotype, but low nucleotide diversity to the west of Wallace's Line is consistent with post‐glacial colonization of the Sunda Shelf. Additional data are needed to test the generality of these patterns.     

Phylogenetic relationships of the pygmy rice rats of the genus Oligoryzomys Bangs, 1900 (Rodentia: Sigmodontinae)

Sequences from two mitochondrial genes (cytochrome b and NADH1) were used to produce a molecular phylogeny for 12 named and two undescribed species of the genus Oligoryzomys. All analyses placed Oligoryzomys microtis as the most basal taxon, a finding consistent with previous studies that suggested the west‐central Amazon as a centre of origin for the tribe Oryzomyini to which Oligoryzomys belongs. Biogeographically, this suggests that Oligoryzomys had a South American origin, and later advanced northwards, entering Central America and Mexico more recently. Different analyses have provided consistent support for several additional clades that did not necessarily agree with the species groups hypothesized by previous studies. A molecular clock derived for these data suggests an origin for the genus of 6.67 Mya, with most speciation within the genus occurring between 3.7 and 1.5 Mya. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 551–566.