Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions

Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera (Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.     

The fourth paradigm pattern of post-glacial range expansion of European terrestrial species: the phylogeography of the Marbled White butterfly (Satyrinae, Lepidoptera)

Aim Three paradigm patterns of post‐glacial dispersal are known for terrestrial species in Europe. However, the possibility of a fourth arises with the Italian and the Balkan lineages expanding to central Europe and the Iberian one being trapped by the Pyrenees. We test this hypothesis by analysing the molecular biogeography of the Marbled White butterfly. Location Twelve populations distributed over a major part of the European range of Melanargia galathea and M. lachesis. Methods We studied 18 allozyme loci of 403 individuals from 12 populations. Butterflies were sampled in the field, frozen in liquid nitrogen and stored under these conditions until analysis. We used cellulose acetate plates for allozyme electrophoresis. Results We detected three major genetic lineages within the M. galathea/lachesis complex. The M. lachesis sample from the southern Pyrenees was strongly genetically differentiated from M. galathea (F_CT: 0.312). Melanargia galathea splits into two major genetic lineages (F_CT: 0.115), which both were found in post‐glacially invaded regions. The further differentiation within these lineages was comparably low (F_SC: 0.028). The genetic diversity within populations was high compared with other butterfly species. Main conclusions Our findings support the existence of a fourth pattern with only the Iberian lineage not contributing considerably to the post‐glacial colonization of central Europe. Preliminary studies in other butterfly species of dry grasslands support the importance of this pattern possibly representing a fourth paradigm. The high genetic diversity within populations might be one reason for the recently observed expansions at the northern distribution limits.     

The genetic structure of natural and reintroduced roe deer (Capreolus capreolus) populations in the Alps and central Italy,with reference to the mitochondrial DNA phylogeography of Europe

The first hypervariable fragment (HVI) of the mitochondrial DNA control region was sequenced in 90 individuals of the European roe deer (Capreolus capreolus) from the Alps, central Italy and Spain. Pooling these data with 70 published sequences from several European regions, we were able to identify patterns of divergence within the Italian peninsula, and in Europe in general. The results we obtained can be summarized as follows. First, the genetic structure of European roe deer populations is substantial (PhiST values around 0.6). Second, the divergence between some central Italian populations, the Alpine group (which is genetically close to the French, the Spanish and the Norwegian samples) and the Eastern European populations seems to reflect Upper Pleistocene subdivisions, possibly related to three southern European refugia. Third, a different group of central Italian individuals probably diverged more recently from the Alpine group, and their attribution to the subspecies C. c. italicus does not appear justified. Fourth, the analysis of mitochondrial DNA in the roe deer can be used to identify recently reintroduced animals in the western Alps which clearly cluster within the Eastern European group, thus providing an important tool for conservation and management strategies for this species.     

Substantial genetic substructuring in southeastern and alpine Australia revealed by molecular phylogeography of the Egernia whitii (Lacertilia: Scincidae) species group

Palaeoclimatic events and biogeographical processes since the mid-Tertiary are believed to have strongly influenced the evolution and distribution of the terrestrial vertebrate fauna of southeastern Australia. We examined the phylogeography of the temperate-adapted members of the Egernia whitii species group, a group of skinks that comprise both widespread low- to mid-elevation (E. whitii) and montane-restricted species (Egernia guthega, Egernia montana), in order to obtain important insights into the influence of past biogeographical processes on the herpetofauna of southeastern Australia. Sequence data were obtained from all six temperate-adapted species within the E. whitii species group, and specifically from across the distributional ranges of E. whitii, E. guthega and E. montana. We targeted a fragment of the ND4 mitochondrial gene (696 bp) and analysed the data using maximum likelihood and Bayesian methods. Our data reveal a deep phylogeographical break in the east Gippsland region of Victoria between 'northern' (Queensland, New South Wales, Australian Capital Territory) and 'southern' (Victoria, Tasmania, South Australia) populations of E. whitii. This divergence appears to have occurred during the late Miocene-Pliocene, with the Gippsland basin possibly forming a geographical barrier to dispersal. Substantial structuring within both the 'northern' and the 'southern' clades is consistent with the effects of Plio-Pleistocene glacial-interglacial cycles. Pleistocene glacial cycles also appear to have shaped the phylogeographical patterns observed in the alpine species, E. guthega and E. montana. We used our results to examine the biogeographical process that led to the origin and subsequent diversification of the lowland and alpine herpetofauna of southeastern Australia.     

Is the last glaciation the only relevant event for the present genetic population structure of the meadow brown butterfly Maniola jurtina (Lepidoptera: Nymphalidae)?

Phylogeographical studies are available for a considerable number of European species, but few analyses exist for temperate species with very large and fairly continuous populations that are also absent from Northern Europe. Therefore, we studied the butterfly Maniola jurtina as a model for this group. The species has two major genetic lineages (mean genetic distance between lineages: 0.033; F _CT: 0.052), most probably evolving in glacial differentiation centres in the western and eastern Mediterranean. The onset of this differentiation might have been the beginning of the last glacial stage maximum some 40 kyr bp . A hybrid zone between these two lineages exists in western Central Europe. No genetic substructures have been found within the two lineages ( F _SC: 0.017) and average genetic distances are very small. Therefore, it is highly probable that postglacial expansion was of the phalanx type. There is, at most, very limited differentiation at regional and local scales. However, the genetic diversity within populations is high (means: A : 2.68; H _E: 17.2%; P : 78%), as would be predicted for such a common species. Comparison of these results with a published allozyme analysis revealed a similar phylogeographical pattern, but lower genetic diversity in the latter. Morphological patterns of wings and genitalia show similar geographical patterns as allozyme data.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 419–431.     

Here be dragons: a phylogenetic and biogeographical study of the Smaug warreni species complex (Squamata: Cordylidae) in southern Africa

Taxonomy of the Smaug warreni species complex remains contentious despite known morphological differences and geographical separation of the various taxa. This study uses an 11‐gene dataset to recover phylogenetic relationships between the seven nominal members of the S. warreni complex. Eight well‐supported clades were returned, with S. warreni barbertonensis found to be paraphyletic. A time‐calibrated analysis of molecular data indicates that all eight clades in the S. warreni complex separated in the late Miocene, much earlier than the date suggested by the existing hypothesis of vicariance through the ingression of Kalahari sands. Ecological niche modelling indicates that although all clades are allopatric, a slight decrease in temperature could potentially render them sympatric, supporting an hypothesis of range expansion through climatic change. © 2014 The Linnean Society of London     

中国片尺蛾属分类订正并记三新纪录种(鳞翅目：尺蛾科：灰尺蛾亚科)

记录中国分布的片尺蛾属6种,并记述中国3新纪录种：黑片尺蛾Fascellina porphyreofusa Hampson, 1895; F. inornata Warren, 1893; F. rectimarginata Warren, 1894。文中提供了属种描述和鉴别特征,给出了种检索表及成虫和外生殖器图。     

Limited hybridization along a large contact zone between two genetic lineages of the butterfly Erebia medusa (Satyrinae, Lepidoptera) in Central Europe

Genetic lineages evolving during glacial isolation frequently come into contact as the result of postglacial range expansions. Hybridization often occurs along these contact zones. In Europe, the high mountain systems of the Alps and Pyrenees are well known for their hybrid belts. This article studies the contact zone of the Woodland Ringlet Erebia medusa in a Hercynian mountain area in the Czech-German border region not exceeding 1500 m a.s.l. Hybrid populations between an eastern and a western genetic lineage were detected by amova -based tests, principal component analysis and neighbour joining analysis. Over most of the range of the Czech-German Border Mts, the ridges separate the western and the eastern genetic lineage from each other. However, two important hybrid areas were detected: (1) the watershed of the Ohře river in the north-west of that area, a major valley system passing through these mountains and (2) the high plateaux of the Šumava Mts in the south-east, an extended area of high elevation. The location of hybrid populations in geographical vicinity to non-hybrid populations and the generally low F _IS (2.1%) make reduced fitness of hybrid individuals little likely. The hybrid populations have intermediate genetic diversity between the genetically poor western and the genetically rich eastern lineage populations.     

TErritorial defence and mating success in males of the small heath butterfly, coenonympha pamphilus L. (Lepidoptera: Satyridae)

It has long been recognized that territorial defence in male butterflies must be some sort of mating strategy and that the territories are used as mating stations. However, so far, no systematic study has established the adaptive significance of territorial behaviour. This study is an attempt to fill this gap. By comparing the distribution of territories with the distribution of locations where wild and released virgin females mated this study shows that, in the small heath butterfly (Coenonympha pamphilus), males in territories have higher mating success than males outside territories. This supported the hypotheses that the function of territorial behaviour is to secure more matings and that the territories are mating stations. Wing length measurements suggest the same. Large males residing in territories tended to mate more often than small males, which were usually found outside territories. Since resident males were larger than non-resident males, this size difference was used to see how territorial occupancy influenced longevity. Mark-recapture of measured males revealed no significant correlation between wing length and the further life expectancy of males, strongly suggesting that the mating success of males in territories is also higher when measured over their whole lifetime.     

Taxonomic revision of the marbled skink (Cyclodina oliveri,Reptilia: Scincidae) species complex,with a description of a new species

Abstract

We have completed a taxonomic revision of the New Zealand marbled skink (Cyclodina oliveri) species complex. Morphological analyses and mitochondrial sequence data (ND2, ND4, Cytochrome b; Total 1933 bp) are used to describe a new taxon (commonly known as the “Mokohinau” skink) and redefine C. oliveri. The morphological and molecular data indicate that C. oliveri is distributed on the Poor Knights Islands, Mercury Islands and Aldermen Islands. The new species is restricted to the Mokohinau Islands, Hen and Chickens group, Little Barrier Island and Great Barrier Island. Our data demonstrate that there is no support for the separation of the Poor Knights Islands population of C. oliveri from those on the Mercury Islands and Aldermen Islands. The genetic data indicate that C. whitakeri is part of the C. oliveri species group. Divergence time estimates indicate that the C. oliveri species complex diverged during the late‐Miocene, with further divergences among island groups in C. oliveri including the origin of the new taxon during the late‐Pliocene and mid‐Pleistocene. We present a diagnostic key for Cyclodina.     

The influence of history and contemporary stream hydrology on the evolution of genetic diversity within species: an examination of microsatellite DNA variation in bull trout,Salvelinus confluentus (Pisces: Salmonidae)

An understanding of the relative roles of historical and contemporary factors in structuring genetic variation is a fundamental, but understudied aspect of geographic variation. We examined geographic variation in microsatellite DNA allele frequencies in bull trout (Salvelinus confluentus, Salmonidae) to test hypotheses concerning the relative roles of postglacial dispersal (historical) and current landscape features (contemporary) in structuring genetic variability and population differentiation. Bull trout exhibit relatively low intrapopulation microsatellite variation (average of 1.9 alleles per locus, average He = 0.24), but high levels of interpopulation divergence (F(ST) = 0.39). We found evidence of historical influences on microsatellite variation in the form of a decrease in the number of alleles and heterozygosities in populations on the periphery of the range relative to populations closer to putative glacial refugia. In addition, one region of British Columbia that was colonized later during deglaciation and by more indirect watershed connections showed less developed and more variable patterns of isolation by distance than a similar region colonized earlier and more directly from refugia. Current spatial and drainage interconnectedness among sites and the presence of migration barriers (falls and cascades) within individual streams were found to be important contemporary factors influencing historical patterns of genetic variability and interpopulation divergence. Our work illustrates the limited utility of equilibrium models to delineate population structure and patterns of genetic diversity in recently founded populations or those inhabiting highly heterogeneous environments, and it highlights the need for approaches incorporating a landscape context for population divergence. Substantial microsatellite DNA divergence among bull trout populations may also signal divergence in traits important to population persistence in specific environments.     

The complete mitochondrial genome of the endangered Apollo butterfly,Parnassius apollo (Lepidoptera: Papilionidae) and its comparison to other Papilionidae species

The Apollo butterfly, Parnassius apollo is a representative species of the butterfly subfamily Parnassiinae. This charming species is one of the most endangered butterfly species in the world. In this study, we sequenced its complete mitochondrial genome (mitogenome), with the aim of accumulating genetic information for further studies of population genetics and mitogenome evolution in the Papilionidae. The 15,404-bp long mitogenome harbors a typical set of 37 genes and is the largest butterfly mitogenome determined, except for Papilio maraho (16,094 bp). Like many other sequenced lepidopteran species, one tRNA^Trp-like and one tRNA^Leu(UUR)-like sequences were detected in the AT-rich region. A total of 164 bp of non-coding sequences are dispersed in 14 regions throughout the genome. The longest intergenic spacer (68 bp) is located between tRNA^Ser(AGN) and tRNA^Glu, and is the largest spacer at this location among Papilionidae species. This spacer may have resulted from an 8-fold repetition of a TTTCTTCT motif or a 4-fold repetition of a CTTTATTT motif.     

Tracing the radiation of Maniola (Nymphalidae) butterflies: new insights from phylogeography hint at one single incompletely differentiated species complex

The use of DNA sequence data often leads to the recognition of cryptic species within putatively well‐known taxa. The opposite case, detecting less diversity than originally described, has, however, far more rarely been documented. Maniola jurtina, the Meadow Brown butterfly, occurs all over Europe, whereas all other six species in the genus Maniola are restricted to the Mediterranean area. Among them, three are island endemics on Sardinia, Cyprus, and Chios, respectively. Maniola species are almost indistinguishable morphologically, and hybridization seems to occur occasionally. To clarify species boundaries and diversification history of the genus, we reconstructed the phylogeography and phylogeny of all seven species within Maniola analyzing 138 individuals from across its range using mitochondrial and nuclear genetic markers. Examination of variation in mitochondrial and nuclear DNA surprisingly revealed a case of taxonomic “oversplitting”. The topology of the recovered phylogenetic tree is not consistent with accepted taxonomy, but rather reveals haplotype clades that are incongruent with nominal species boundaries: instead of seven species, we recognized only two major, yet incompletely segregated, lineages. Our results are consistent with the hypothesis that Maniola originated in Africa. We suggest that one lineage dispersed over the Strait of Gibraltar and the Iberian Peninsula to the west of Europe, while the other lineage spreads eastward through Asia Minor and over the Bosporus to Eastern Europe.     

The presence of a large cercopithecine (cf. Theropithecus sp.) in the ‘Ubeidiya formation (Early Pleistocene, Israel)

This study presents the discovery of a right cercopithecine calcaneus from the site of ‘Ubeidiya, Israel, dated to ca. 1.6 Ma. The fossil is described and statistically compared to bones of modern and fossil cercopithecids. The specimen can be attributed to a large-bodied cercopithecine and represents a new primate taxon previously unidentified in the Early Pleistocene of the Southern Levant. Among extant genera, it is most clearly similar to calcanei of Theropithecus. However, it could also represent Paradolichopithecus, but this alternative is unlikely due to the morphological uniqueness of the latter taxon.The finding of an African taxon in the Levant suggests a circum-Mediterranean dispersal route for the taxon out of Africa, and emphasizes the importance of the Levantine corridor as a biogeographic dispersal route between Africa and Eurasia during the Early Pleistocene. Evidence for the biogeography of large-bodied primates is essential for the understanding of the dispersal routes of “Out of Africa I” taxa and can help elucidate Homo dispersal patterns in the Early Pleistocene.     

The species of the genus Micrambe Thomson 1863, from South Africa (Coleoptera: Cryptophagidae)

The 22 species of the genus Micrambe Thomson known from South Africa are analyzed and revised. Micrambe danielssoni sp. nov is described. Following a detailed study of the type material in different entomological collections, the status of two taxa is revised and two new synonymies are proposed: Micrambe kolbei Grouvelle 1908, syn. nov. for Micrambe minuta Grouvelle 1908, and Micrambe similis Bruce 1957, syn. nov. for Micrambe peringueyi Grouvelle 1908. A lectotypus is designated for one taxon: Micrambe capensis (Redtenbacher 1867). A key to all the species occurring in South Africa is presented.     

Comparative phylogeography of two sibling species of forest‐dwelling rodent (Praomys rostratus and P. tullbergi) in West Africa: different reactions to past forest fragmentation

Two sibling species of the rodent genus Praomys occur in West African forests: P. tullbergi and P. rostratus. By sampling across their geographical ranges (459 individuals from 77 localities), we test the hypothesis that climatic oscillations during the Quaternary made an impact on the observed pattern of cytochrome b sequence variation. We show that, although these two species have parapatric geographical distributions, their phylogeographical histories are dissimilar, which could be related to their distinct ecological requirements. Since the arid phases of the Pleistocene were characterized by isolated forest patches, and intervening wetter periods by forest expansion, these changes in forest cover may be the common mechanism responsible for the observed phylogeographical patterns in both of these species. For example, in both species, most clades had either allopatric or parapatric geographical distributions; however, genetic diversity was much lower in P. tullbergi than in P. rostratus. The genetic pattern of P. tullbergi fits the refuge hypothesis, indicating that a very small number of populations survived in distinct forest blocks during the arid phases, then expanded again with forest recovery. In contrast, a number of populations of P. rostratus appear to have survived during the dry periods in more fragmented forest habitats, with varying levels of gene flow between these patches depending on climatic conditions and forest extent. In addition, historical variations of the West African hydrographic network could also have contributed to the pattern of genetic differentiation observed in both species.     

The effects of island area, isolation and source population size on the presence of the grayling butterfly Hipparchia semele (L.) (Lepidoptera: Satyrinae) on British and Irish offshore islands

Records of Hipparchia semele on British and Irish islands have been modelled against island area, isolation (sea and land distance) and the size of the nearest potential source populations. All three variables have been found to contribute significantly to the presence or absence of H. semele on the islands. Isolation is a more significant predictor than island area. This result differs from the multiple species case where area was found to be a more important influence than isolation. Records on islands are also shown to depend on the size of populations at the nearest sources; this underpins the relationships identified for the multiple species case, first, between the number of species on islands and at nearest sources and, second, between the incidence of species on islands and at nearest sources. There are clear indications that smaller islands may become increasingly marginalized for H.semele; with ongoing habitat loss, because isolation increases and source populations become sparser, the probability of H. semele recolonizing islands also decreases.