The butterfly Danaus chrysippus (L.) in East Africa comprises polyphyletic, sympatric lineages that are, despite behavioural isolation, driven to hybridization by female-biased sex ratios

Mitochondrial and nuclear DNA information was analysed among four subspecies of the African Queen butterfly, Danaus (Anosia) chrysippus sensu lato (s.l.), along with four other Danaus species drawn from all three subgenera (D. (Danaus) plexippus, D. (Salatura) genutia, D. (A.) gilippus, D. (A.) eresimus) and two outgroup species from the same tribe, Tirumala septentrionis and Amauris niavius. A mitochondrial phylogeny derived from the 12S rRNA (347 bp) and COI (537 bp) loci indicates two very distinct haplotypes for subspecies D. (A.) c. dorippus, dorippus‐1 and dorippus‐2. Interestingly, dorippus‐1, on the one hand, and all other D. (A.) chrysippus haplotypes, on the other, are the most distantly related clades within the genus and have different most recent ancestors from different subgenera, though sharing the common ancestor of the monophyletic genus. A phylogeny based on the EF1‐α nuclear locus (400 bp) shows that the two well‐separated mitochondrial lineages of dorippus are identical for this gene and reciprocally monophyletic to the other D. (A.) chrysippus lineages. Thus, nuclear and cytoplasmic phylogenies are not only discordant, but also suggest that both D. (A.) chrysippus s.l. and subspecies dorippus are polyphyletic. Paradoxically, four African subspecies, chrysippus‐orange, chrysippus‐brown, alcippus and dorippus, though substantially vicariant, hybridize extensively in East Africa wherever the ranges of two or more of them overlap. Linkage disequilibrium, and hence sexual isolation, in sympatry between colour (nuclear) genes and unlinked mitochondrial (cytoplasmic) loci is consistent across populations and therefore indicates the operation of positive natural selection. Together with data from previous experimental and field work, our results suggest that extensive hybridization occurs among once allopatric or parapatric lineages, that are now nascent species. We deduce that hybridism among lineages in sympatry is currently enforced, in the face of assortative mate choice, by a bacterial symbiont, Spiroplasma, a male‐killer that forces females in female‐biased populations to pair with heterotypic males. In discussion we emphasize that neither D. (A.) chrysippus s.l. as presently circumscribed, nor its component clades, conform to any established concept of species. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 117–131.     

Hologenomic speciation: synergy between a male‐killing bacterium and sex‐linkage creates a ‘magic trait’ in a butterfly hybrid zone

Danaus chrysippus (L.) in Africa comprises four substantially isolated semispecies that are migratory and hybridize on a seasonal basis throughout the eastern and central part of the continent. In the hybrid zone (but not elsewhere), the butterfly is commonly host to a male killing endosymbiotic bacterium, Spiroplasma sp., which principally infects one semispecies, Danaus chrysippus chrysippus in Kenya. A W‐autosome mutation, inherited strictly matrilinearly, links B and C colour gene loci, which have thus gained sex‐linkage in chrysippus. We have monitored variation in sex ratio and genotype at the A and C colour gene loci for two extended periods of 18 months (2004–5) and 12 months (2009–10) in adults reared from wild eggs laid on trap plants in Kasarani, near Nairobi, Kenya. Additionally, in 2009–10, all surviving adult butterflies were screened for Spiroplasma infection. The hybridizing Kasarani population is highly atypical in three respects, and has apparently been so for some 30 years: first, the sex ratio is permanently female‐biased (as expected), although subject to seasonal fluctuation, being lowest (male/female) when D. c. chrysippus (cc) peaks and highest when Danaus chrysippus dorippus (CC) predominates; second, the population is invariably dominated by Cc heterozygotes of both sexes but especially females; and third, cc males are always scarce because they are systematically eliminated by male killing, whereas the CC genotype is male‐biased. It is this imbalance of sex versus genotype that determines the massive departure from Hardy–Weinberg equilibrium in the population, in part because cc females have little choice but to pair with C‐ males. We suggest that: first, Cc hybrids of both sexes fail to disperse in the company of either parental semispecies; second, Spiroplasma positive females carrying the W‐autosome mutation have a selective advantage over females that lack the translocation; third, the endoparasite and the translocation create a ‘magic trait’ linkage group that underlies hologenomic reproductive isolation between two emerging species, D. c. chrysippus and D. c. dorippus; and, fourth, that the predominance of males in dorippus suggests that individuals must be protected by a male‐killing suppressor gene. By contrast to the C locus, Aa heterozygotes are in substantial and permanent deficit, suggesting either assortative mating between AA (chrysippus and dorippus) and aa (Danaus chrysippus alcippus), or heterozygote unfitness, or both. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 92–109.     

Patterns and processes in the evolutionary history of parrotfishes (Family Labridae)

Phylogenetic reconstruction of the evolutionary relationships among 61 of the 70 species of the parrotfish genera Chlorurus and Scarus (Family Labridae) based on mitochondrial and nuclear gene sequences retrieved 15 well‐supported clades with mid Pliocene/Pleistocene diversification. Twenty‐two reciprocally monophyletic sister‐species pairs were identified: 64% were allopatric, and the remainder were sympatric. Age of divergence was similar for allopatric and sympatric species pairs. Sympatric sister pairs displayed greater divergence in morphology, ecology, and sexually dimorphic colour patterns than did allopatric pairs, suggesting that both genetic drift in allopatric species pairs and ecologically adaptive divergence between members of sympatric pairs have played a role in diversification. Basal species typically have small geographical ranges and are restricted to geographically and ecologically peripheral reef habitats. We found little evidence that a single dominant process has driven diversification, nor did we detect a pattern of discrete, sequential stages of diversification in relation to habitat, ecology, and reproductive biology. The evolution of Chlorurus and Scarus has been complex, involving a number of speciation processes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 529–557.     

Speciation on the coasts of the new world: phylogeography and the evolution of bindin in the sea urchin genus Lytechinus

Abstract Beginning with E. Mayr's study in 1954, tropical sea urchins have played an important role in studies of speciation in the sea, but what are the processes of cladogenesis and divergence that give rise to new species in this group? We attempt to answer this question in the genus Lytechinus. Unlike the majority of other tropical sea urchin genera, which have circumtropical distributions, Lytechinus is mostly confined to the tropics and subtropics of the New World. We sequenced a region of mitochondrial cytochrome oxidase I and the entire molecule of nuclear bindin (a sperm gamete recognition protein) of nearly all species in the genus, and we assayed isozymes of three partially sympatric closely related species and subspecies. We found that in both mitochondrial DNA (mtDNA) and in bindin the genus Lytechinus is paraphyletic, encompassing Sphaerechinus granularis as the sister species of L. euerces. The rest of the species are arranged in an Atlantic clade composed of L. williamsi and L. variegatus, and a Pacific clade containing L. anamesus, L. pictus, L. semituberculatus, and L. panamensis. Divergence between these clades suggests that they were separated no later than the closure of the Isthmus of Panama, and possibly before this time. Our data confirm that L. anamesus and L. pictus from California are a single species, and provide no evidence of differentiation between L. variegatus variegatus from the Caribbean and L. variegatus atlanticus from Bermuda. Lytechinus variegatus variegatus mtDNA is distinct from that of L. variegatus carolinus from the North American seaboard and the Gulf of Mexico, whereas their bindins are very similar. However, there is clear evidence of introgression of mtDNA between the two subspecies and they share alleles in all sampled isozyme loci. Lytechinus williamsi from the Caribbean shares mtDNA haplotypes with L. variegatus variegatus, and they also share isozymes in all assayed loci. Their bindin, however, is distinct and coalesces within each morphospecies. A private clade of mtDNA in L. williamsi may be indicative of former differentiation in the process of being swamped by introgression, or of recent speciation. Recent sudden expansions in effective population size may explain the predominance of a few mitochondrial haplotypes common to the two species. Despite the high divergence of bindin (relative to differentiation of mtDNA) between L. variegatus and L. williamsi, comparison of amino acid replacement to silent substitutions by various methods uncovered no evidence for positive selection on the bindin of any clade of Lytechinus. With the possible exception of L. williamsi and L. variegatus, our results are consistent with a history of allopatric speciation in Lytechinus. The molecular results from Lytechinus, along with those of other similar studies of sea urchins, suggest that the general speciation patterns deduced in the middle of last century by Mayr from morphology and geography have held up, but also have uncovered peculiarities in the evolution of each genus.     

Intraspecific molecular phylogeny and phylogeography of the Meriones meridianus (Rodentia: Cricetidae) complex in northern China reflect the processes of desertification and the Tianshan Mountains uplift

The phylogeographical patterns and demographic history of mitochondrial DNA (cytochrome b, N = 327; D‐loop, N = 252) and nuclear DNA (IRBP gene, N = 235) haplotypes were studied for the Meriones meridianus complex in northern China, a desert‐dwelling gerbil species complex. The phylogenetic analyses, which were performed on the separate and combined (mitochondrial + nuclear) datasets, revealed two divergent clades (Clade A and Clade B) corresponding to distinct geographical regions. Clade A contained the haplotypes found mostly in individuals from the Tianshan Mountains area. Clade B contained haplotypes from populations located in other deserts in northern China. The divergence times indicated that the history of the M. meridianus complex was influenced by the uplift of the Tianshan Mountains and climate‐induced habitat fluctuations. In the Pleistocene, the expansion of forests and grasslands during interglacial period led to the isolation of the M. meridianus complex, which preferred to inhabit deserts. Hence, long geological isolation and the M. meridianus complex adaptation to local ecological conditions led to its genetic divergence. Clade A had long‐lasting demographic stability, most likely because the populations of this clade remained in a stable desert environment for a long time. However, the extension of other deserts and disappearance of palaeolakes during the last glacial period resulted in demographic expansion of Clade B. Furthermore, our genetic data indicated that two subspecies may exist within the M. meridianus complex. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 362–383.     

Phylogeography and phylogenetic relationships of Malagasy tree and ground boas

Three species of boid snakes are recognized in Madagascar, namely the genus Sanzinia (one species and two subspecies) and the genus Acrantophis (two species). In the present study, we studied the patterns of genetic variation of these species across Madagascar using a fragment of the mitochondrial 16S rRNA gene in 77 specimens. To support the phylogenetic relationships of the lineages identified, three further gene fragments (cytochrome b, 12S rRNA and c‐mos) were analyzed in a reduced but representative set of samples. The results obtained corroborate that the genus Sanzinia includes two highly divergent mitochondrial lineages that evolved independently from each other on the east versus the west side of Madagascar. Each of these lineages presents a further subdivision that separates northern from southern groups. The nuclear marker showed no variation among the Malagasy boas, indicating either very low substitution rates in this gene or relatively recent speciation events coupled with high mitochondrial substitution rates. Because the broad geographic sampling detected no admixture among haplotypic lineages within Sanzinia, it is hypothesized that these may represent distinct species. Deviant haplotypes of snakes morphologically similar to Acrantophis dumerili indicate that this taxon may be a complex of two species as well. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 640–652.     

Evolutionary history of bulldog bats (genus Noctilio): recent diversification and the role of the Caribbean in Neotropical biogeography

We present phylogenetic relationships and phylogeographic patterns of the two species of bulldog bats, genus Noctilio. Using a comprehensive sampling of 118 individuals throughout the species distribution, we investigated the distribution of molecular variation in one nuclear and two mitochondrial markers. Phylogenetic trees do not recover Noctilio albiventris as a monophyletic group and point to three similar‐age intraspecific genetic lineages, suggesting cryptic diversity in this taxon. These lineages correspond to the subspecies previously proposed, and are strongly associated with major river basins in South America. Analyses also suggest a very recent origin for the fishing bat Noctilio leporinus, which probably originates from N. albiventris, with a population expansion corresponding to its invasion in South America. Based on our analysis, the speciation event was dated in the Pleistocene epoch and seems to be associated with the variation of the sea level in the Caribbean islands. The present work indicates how phylogeographic studies support the identification of independent evolutionary lineages, driving new systematic/taxonomic investigations, while at the same time shed light on the role of the Caribbean in shaping Neotropical bat fauna diversity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Hybridization between genetically and morphologically divergent forms of Rhagada (Gastropoda: Camaenidae) snails at a zone of secondary contact

Congeneric sympatry is rare in Australian camaenid land snails, and the reasons are poorly understood. As the first example of contact between species of the Western Australian genus Rhagada, we examined the parapatric zone between the coastally distributed Rhagada convicta and an undescribed inland species, which differ in size, shape, and banding of their shells and in their reproductive systems. Phylogenetic analysis of sequences of the mitochondrial 16S RNA gene indicated secondary contact between two distinct clades, with an average sequence divergence of 6.5%. Despite these substantial differences, the two forms hybridize in the narrow contact zone, resulting in intermediate phenotypes for shells and reproductive anatomy, and incongruence of the mitochondrial DNA (mtDNA) lineage and morphology. Hybridization appears to be asymmetric, however, as all morphological intermediates possessed the R. convicta mtDNA. Interbreeding between these deeply divergent forms challenges the use of reproductive structures as indicators of reproductive isolation in snail taxonomy. This is the first contact zone detected between mainland species of Rhagada and indicates that the widespread pattern of geographical replacement is not simply a result of allopatric divergence without subsequent contact. Instead, the perspective for understanding the radiation and geographical distributions must include interactions between the species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 348–362.     

A revised classification of the genus Matrona Selys, 1853 using molecular and morphological methods (Odonata: Calopterygidae)

An extensive review of the genus Matrona is presented based on mitochondrial (COI) and nuclear (ITS) sequences from 150 samples which cover all the known taxa of this genus. The separation of two main clades (oreades group: M. oreades, M. corephaea and M. taoi; basilaris group: M. basilaris, M. nigripectus, M. cyanoptera, M. japonica and M. annina) is strongly supported. The classification of all traditional recognized species is confirmed. The Hainan population separates very well from mainland M. basilaris populations, which is also confirmed by geometric morphometric analysis of wing shape. Given the implications of the molecular analysis the genus Matrona is grouped into two subgenera: subgen. Matrona (type species M. basilaris) and D ivortia subgen. nov. (type species M. oreades). A new species, M . ( M .) mazu sp. nov. , from Hainan is described. Brief taxonomic notes on the nine recognized species of the genus are given. Lectotype designations of M. basilaris and M. nigripectus are published. © 2015 The Linnean Society of London     

Molecular phylogeny of the Western Palaearctic Cordulegaster taxa (Odonata: Anisoptera: Cordulegastridae)

Although Odonata are a key component of many freshwater ecosystems, their taxonomy and evolutionary history is still far from being well resolved. In the present study, we report the first molecular phylogeny for the Western Palaearctic Cordulegaster genus (Odonata: Anisoptera: Cordulegastridae). We sequenced fragments of both mitochondrial and nuclear genes [cytochrome c oxidase I (COI) and Internal Transcribed Spacer‐1 (ITS‐1)] from eight species and 13 subspecies, from western, southern and central Europe, Turkey, and Morocco. Our data support the existence of two major groups corresponding to the traditional boltonii‐ and bidentata‐groups. Both groups are monophyletic based on COI sequences and the distinctiveness of Cordulegaster princeps, Cordulegaster trinacriae, Cordulegaster picta and Cordulegaster heros relative to Cordulegaster boltonii, and Cordulegaster helladica and Cordulegaster insignis relative to Cordulegaster bidentata, is confirmed. All species are also monophyletic for ITS‐1, with the exception of Cordulegaster helladica buchholzi, which shares the haplotype with C. insignis. Although moderate levels of genetic diversity were found within C. boltonii, there was no clear separation among the four subspecies, with the exception of the populations of Cordulegaster boltonii algirica from North Africa. Similarly, no genetic differentiation was found between the two subspecies of C. bidentata, Cordulegaster bidentata bidentata and Cordulegaster bidenta sicilica. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 49–57.     

Genital lock-and-key system and premating isolation by mate preference in carabid beetles (Carabus subgenus Ohomopterus)

The shapes and lengths of copulatory pieces and vaginal appendices of the carabid beetle subgenus Ohomopterus (genus Carabus) vary among species. In Japan, the species in the group with a medium body size (C. yaconinus, C. iwawakianus, C. maiyasanus, C. uenoi, C. arrowianus, C. esakii, and C. insulicola) are usually allopatric or parapatric, except at Mt Kongosan, where C. uenoi, C. iwawakianus, and C. yaconinus are sympatrically distributed. The degree of premating isolation by mate preference was high between sympatric populations, irrespective of the genetic distance between them. However, premating isolation was absent between parapatric populations. The degree of premating isolation for allopatric populations spanned a wide range of isolation values. Thus, mate discrimination by males seems to have evolved mostly between sympatric pairs. These results suggest two hypotheses. First, premating isolation has evolved through reinforcement or through reproductive character displacement after sympatric contact. Second, premating isolation has evolved in allopatry, and as a result of premating isolation, the species can coexist in sympatry. We also examined the degree of mechanical isolation between C. uenoi and C. iwawakianus (a sympatric pair), which have a very large difference in the length of the copulatory piece. The insertion success was low and only one female produced viable offspring among 15 crosses; however, death in females due to copulation was rare. For sympatric matings between C. uenoi and C. iwawakianus, a large difference in the genital size might reduce the gene flow with small mating costs. Gene flow that was significantly reduced by genital difference might cause either the evolution of premating isolation through reinforcement/reproductive character displacement or through the maintenance of a high degree of premating isolation following sympatric contact. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 145–154.     

Linkage disequilibrium and natural selection for mimicry in the Batesian mimic Hypolimnas misippus (L.) (Lepidoptera: Nymphalidae) in the Afrotropics

On two occasions, on opposite sides of the African continent (Cape Coast, Ghana, and Dar es Salaam, Tanzania), high adult population densities in the polymorphic butterfly Hypolimnas misippus (a presumed mimic of Danaus chrysippus) were followed by linkage disequilibrium in combinations of fore‐ and hindwing colour patterns. On both occasions, disequilibrium was caused by significant changes in morph frequencies favouring rarer and more mimetic forms. Recaptures were too few for analysis at Dar, although the changes there took place within a single generation and must have been the result of differential survival. Recapture rate data and survival rate estimates at Cape Coast support the hypothesis that selective predation was responsible, as does the observation of synchronous linkage disequilibrium at Dar in the model D. chrysippus, indicating parasitic mimicry. There was clear selection for the perfection of mimicry for forewings at Dar and for hindwings at Cape Coast. Disequilibrium is also reported for two other sites, Legon (Ghana) and Boksburg (South Africa) and, in all four sites, it was associated with an increase in the most mimetic forms. New chemical evidence is presented to support the contention that D. chrysippus is a defended model. Although all the evidence leads to the conclusion that H. misippus is a Batesian mimic of D. chrysippus, many questions remain, particularly with regard to the identity of predators, the episodic nature of selective predation events, and their apparent lack of lasting and significant impact on overall gene frequencies. We conclude that H. misippus presents both challenges and opportunities for studies on mimicry, and we suggest that linkage disequilibrium can be a useful generic indicator for Gestalt predation on polymorphic prey. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 180–194.     

Phylogenetic and biogeographical relationships of the Sander pikeperches (Percidae: Perciformes): patterns across North America and Eurasia

North America and Eurasia share several closely related taxa that diverged either from the breakup of the Laurasian supercontinent or later closures of land bridges. Their modern population structures were shaped in Pleistocene glacial refugia and via later expansion patterns, which are continuing. The pikeperch genus Sander contains five species – two in North America (S. canadensis and S. vitreus) and three in Eurasia (S. lucioperca, S. marinus, and S. volgensis) – whose evolutionary relationships and relative genetic diversities were previously unresolved, despite their fishery importance. This is the first analysis to include the enigmatic and rare sea pikeperch S. marinus, nuclear DNA sequences, and multiple mitochondrial DNA regions. Bayesian and maximum‐likelihood trees from three mitochondrial and three nuclear gene regions support the hypothesis that Sander diverged from its sister group Romanichthys/Zingel ～24.6 Mya. North American and Eurasian Sander then differentiated ～20.8 Mya, with the former diverging ～15.4 Mya, congruent with North American fossils dating to ～16.3–13.6 Mya. Modern Eurasian species date to ～13.8 Mya, with S. volgensis being basal and comprising the sister group to S. lucioperca and S. marinus, which diverged ～9.1 Mya. Genetic diversities of the North American species are higher than those in Eurasia, suggesting fewer Pleistocene glaciation bottlenecks. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 156–179.     

Molecular systematics,species limits,and diversification of the genus Dendrocolaptes (Aves: Furnariidae): Insights on biotic exchanges between dry and humid forest types in the Neotropics

The true diversity and interspecific limits in the Neotropical endemic avian genus Dendrocolaptes (Furnariidae) remain a highly controversial subject, with previous genus‐wide assessments, based mostly on morphological characters, producing poorly resolved phylogenies. The lack of well‐resolved, robust, and taxonomically densely sampled phylogenies for Dendrocolaptes prevents reliable inferences on the genus’ actual species diversity and evolutionary history. Here, we analyzed 2,741 base pairs of mitochondrial and nuclear genes from 43 specimens belonging to all species and the majority of subspecies described for Dendrocolaptes to evaluate species limits and reconstruct its diversification through time. Our phylogenies recovered a monophyletic Dendrocolaptes, with two main highly supported internal clades corresponding to the D. certhia and D. picumnus species complexes. Also, our analyses supported the monophyly of most Dendrocolaptes species recognized today, except D. picumnus, which was consistently recovered as paraphyletic with respect to D. hoffmannsi. A coalescent‐based test supported a total of 15 different lineages in Dendrocolaptes and indicated that the number of currently accepted species within the genus may be greatly underestimated. Particularly relevant, when combined with previous analyses based on plumage characters, comparative high levels of genetic differentiation and coalescent analyses support the recognition of D. picumnus transfasciatus as a full species that is already under threat. Ancestral area reconstructions suggest that diversification in Dendrocolaptes was centered in lowland Amazonia, with several independent dispersal events leading to differentiation into different adjacent dry and high elevation forest types throughout the Neotropics, mainly during the Middle and Late Pleistocene.     

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.     

Hidden species diversity of Australian burrowing snakes (Ramphotyphlops)

The worm‐like snakes (Scolecophidia; approximately 400 nominal extant species) have a conservative morphology and are among the most poorly‐known terrestrial vertebrates. Although molecular evidence has helped determine their higher‐level relationships, such data have rarely been used to discriminate among species. We generated a molecular data set for the continental Australian blindsnakes (genus Ramphotyphlops) to determine the concordance of molecular and morphological information in the taxonomic recognition of species. Our dataset included 741 specimens morphologically attributed to 27 nominal Ramphotyphlops species. We proposed species hypotheses (SHs) after analysis of sequences from a variable mitochondrial gene (cytochrome b) and examined these SHs with additional evidence from a nuclear gene (prolactin receptor) and geographical data. Although the nuclear marker was not as fast‐evolving and discriminating as the mitochondrial marker, there was congruence among the mitochondrial, nuclear, and geographical data, suggesting that the actual number of species is at least two times the current number of recognized, nominal species. Several biogeographical barriers and complex phytogeographical and geological patterns appeared to be involved in the division of some burrowing snake populations and, by consequence, in their diversification and speciation through isolation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 427–441.     

The origin of two cryptic species of African desert jerboas (Dipodidae: Jaculus)

The desert biota is exposed to extreme and variable conditions that shape its evolution and diversification processes. In this respect, the Jaculus jerboas have gained the attention of researchers as a result of their broad Saharan–Arabian distribution and their high and unexplained, morphological, anatomical, and molecular variation. In the present study, mitochondrial and nuclear genealogies where used to confirm monophyly of two cryptic species: Jaculus jaculus and Jaculus deserti. The reconstructed demography showed that the evolutionary histories of the species are markedly different and that the expansion into North‐West Africa by J. deserti was more recent than that of J. jaculus. The weak ecological separation between species and the signs of recent population growth and expansion of J. deserti suggest that its sympatric occurrence with J. jaculus is recent and that these species evolved in isolated populations, after diverging around the Pliocene–Pleistocene boundary. The importance of climate changes on the Sahara Desert biota is discussed in the context of genetic diversification. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 435–445.     

Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses

Adults of the Euthalia phemius complex, which is composed of three South‐East Asian nymphalid species, Euthalia phemius, Euthalia ipona, and Euthalia euphemia, were genetically analysed by examining mitochondrial and nuclear genes. The E. phemius complex was also examined morphologically, with particular emphasis on wing markings and male genitalia. No significant differences amongst the three species in the complex were detected with respect to either genetic distance or genital morphology. We therefore conclude that the three currently recognized Euthalia species belong to a single species. Accordingly, E. ipona is synonymized with E. phemius. Euthalia euphemia is treated as a subspecies of E. phemius. Type specimens of all taxa and a synonymic list for the E. phemius complex are also given. In addition, we briefly discuss the evolution and biogeography of the species complex. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 304–327.     

Phylogenetics,character evolution,and distribution patterns of the greenbriers,Smilacaceae (Liliales), a near‐cosmopolitan family of monocots

Smilacaceae, composed of Smilax and Heterosmilax, are a cosmopolitan family of > 200 species of mostly climbing monocots with alternate leaves characterized by reticulate venation, a pair of petiolar tendrils and usually prickly stems. Although there has been a long history of studying Smilax since Linnaeus named the genus in 1753, the phylogenetic history of this dioecious family remains unclear. Here we present results based on nuclear ribosomal internal transcribed spacer (nrITS) and plastid matK and rpl16 intron DNA sequence data from 125 taxa of Smilacaceae. Our taxon sampling covers all sections of Smilax and Heterosmilax and major distribution zones of the family; species from Ripogonaceae and Philesiaceae are used as outgroups. Our molecular analysis indicates that phylogenetic relationships largely contradict the traditional morphological classification of the family, instead showing a conspicuous geographical pattern among the species clades. The previously recognized genus Heterosmilax was found to be embedded in Smilax. Species in the family are separated into primarily New World and Old World clades, except for a single species lineage, Smilax aspera, that is sister to the remaining species of the family, but with poor statistical support. Ancestral character state reconstructions and examination of distribution patterns among the clades provide important information for future taxonomic revisions and historical biogeography of the group. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 535–548.