Testing for biogeographic mechanisms promoting divergence in Caribbean crickets (genus Amphiacusta)

Aim This work examines whether the history of diversification of Amphiacusta (Orthoptera, Gryllidae) in the Caribbean corresponds to a vicariant or a dispersalist model. Location The Greater Antillean islands of the Caribbean region. Methods The phylogenetic relationships among species were estimated using a procedure that directly estimates the underlying species tree from independent loci (in this case, one mitochondrial and one nuclear locus). This tree was then used to test for topological congruence with a vicariant model, and to estimate divergence times. Results The analyses based on the expected pattern of species divergence (i.e. species‐tree topology) support a vicariant model. With the notable exception of a dispersal event marking the colonization of Jamaica, the timing of the events are generally consistent with a vicariant scenario, given the current taxon sampling and potential errors with dating the divergence events. Main conclusions The tendency of species to co‐segregate by island suggests that intra‐island diversification is common. Despite their flightlessness, species of Amphiacusta are apparently capable of long‐distance dispersal, such as colonization from the Puerto Rican/Virgin Island bank to Jamaica. The topology of the species tree is consistent with a vicariant model of divergence, and the dates of divergence between island groups are generally consistent with an island–island vicariance model. A strict island–island vicariance scenario can, however, be rejected because of inferred dispersal events such as the colonization of Jamaica. Nevertheless, the biogeographic tests suggest that most of the diversity was generated under a combination of intra‐island diversification and island–island vicariance. Additional sampling of taxa will be needed to verify this hypothesized scenario. Our findings indicate that Amphiacusta presents an ideal opportunity for examining the role of sexual selection in promoting diversification, which would complement the large number of studies focused on adaptive divergence of Caribbean taxa.     

Towards a panbiogeography of the seas

A contrast is drawn between the concept of speciation favoured in the Darwin–Wallace biogeographic paradigm (founder dispersal from a centre of origin) and in panbiogeography (vicariance or allopatry). Ordinary ecological dispersal is distinguished from founder dispersal. A survey of recent literature indicates that ideas on many aspects of marine biology are converging on a panbiogeographic view. Panbiogeographic conclusions supported in recent work include the following observations: fossils give minimum ages for groups and most taxa are considerably older than their earliest known fossil; Pacific/Atlantic divergence calibrations based on the rise of the Isthmus of Panama at 3 Ma are flawed; for these two reasons most molecular clock calibrations for marine groups are also flawed; the means of dispersal of taxa do not correlate with their actual distributions; populations of marine species may be closed systems because of self‐recruitment; most marine taxa show at least some degree of vicariant differentiation and vicariance is surprisingly common among what were previously assumed to be uniform, widespread taxa; mangrove and seagrass biogeography and migration patterns in marine taxa are best explained by vicariance; the Indian Ocean and the Pacific Ocean represent major biogeographic regions and diversity in the Indo‐Australian Archipelago is related to Indian Ocean/Pacific Ocean vicariance; distribution in the Pacific is not the result of founder dispersal; distribution in the south‐west Pacific is accounted for by accretion tectonics which bring about distribution by accumulation and juxtaposition of communities; tectonic uplift and subsidence can directly affect vertical distribution of marine communities; substantial parallels exist between the biogeography of terrestrial and marine taxa; biogeographically and geologically composite areas are tractable using panbiogeographic analysis; metapopulation models are more realistic than the mainland/island dispersal models used in the equilibrium theory of island biogeography; and regional biogeography is a major determinant of local community composition. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 675–723.     

Panmixia supports divergence with gene flow in Darwin's small ground finch, Geospiza fuliginosa, on Santa Cruz, Galápagos Islands

The divergence‐with‐gene‐flow model of speciation has a strong theoretical basis with a growing number of plausible examples in nature, but remains hotly debated. Darwin’s finches of the Galápagos Archipelago have played an important role in our understanding of speciation processes. Recent studies suggest that this group may also provide insights into speciation via divergence with gene flow. On the island of Santa Cruz, recent studies found evidence for adaptive divergence in Darwin’s small ground finch, Geospiza fuliginosa, between ecologically contrasting arid and humid zones. Despite the short geographical distance between these zones, strong disruptive selection during low rainfall periods is expected to generate and maintain adaptive divergence. Conversely, during high rainfall periods, when disruptive selection is predicted to be weakened, population divergence in adaptive traits is expected to break down. Because periods of low and high rainfall irregularly alternate, the geographical pattern of adaptive divergence can be assumed to break down and, importantly, regenerate in situ. Here, we use microsatellite allele frequency data to assess the genetic population structure of G. fuliginosa on Santa Cruz. We sample 21 sites and four ecological zones across the island. We reject hypotheses of population substructure linked to ecological and geographical differences among sites in favour of a single panmictic population. Panmixia implies high levels of gene flow within Santa Cruz, which favours selection over genetic drift as a valid process generating phenotypic divergence in G. fuliginosa on Santa Cruz. We discuss how our findings may support classic adaptation, phenotypic plasticity, matching habitat choice or any combination of these three processes.     

Parallel habitat specialization within the wolf spider genus Hogna from the Galápagos

Within most island archipelagos, such as the Galápagos, similar ecological gradients are found on geographically isolated islands. Species radiations in response to these ecological gradients may follow different scenarios being (i) a single habitat specialization event followed by secondary colonization of each ecotype on the different islands or (ii) repeated and parallel habitat specialization on each island separately. This latter scenario has been considered less likely as gene flow might hamper such ecotypic differentiation. At least for the Galápagos, the extent to which this process is involved in species radiations remains yet poorly understood. Within the wolf spider genus Hogna, seven species are described that can be divided into three different ecotypes based on general morphology and habitat preference i.e. species that inhabit the pampa vegetation in the highlands, species that occur in coastal dry habitats and one generalist species. Comparison of the species phylogeny based on one mitochondrial (COI) and one nuclear (28S) gene fragment convincingly demonstrates that ‘pampa’ and ‘coastal dry’ species evolved in parallel on the islands Santa Cruz and San Cristóbal. Despite the observation that allozymes analysis indicated that each species forms a distinct genetic cluster, phylogenetic divergence within these species complexes was very low and paraphyletic and most likely due to hybridization rather than incomplete lineage sorting, as demonstrated for the Santa Cruz species complex. This suggests that within‐island speciation occurred under low levels of gene flow. Species phylogeny in general did not follow the progression of island emergence as a molecular clock analysis suggested that island endemic species may have diverged after as well as before the emergence of the islands. This represents the first clear example of parallel and within‐island speciation because of habitat specialization on the Galápagos and that such divergence most likely occurred under historic gene flow.     

Alula size signals male condition and predicts reproductive performance in an Arctic‐breeding passerine

While studies of achromatic plumage signaling are scarce relative to chromatic ornaments, achromatic ornaments have the potential to act as an efficient form of visual communication due to the highly conspicuous contrast between black and white body regions. Recently, achromatic plumage reflectance has been shown to indicate condition, yet the condition‐dependence of achromatic patch size remains unstudied. Here we show the first evidence that alula size, an achromatic plumage patch, has the potential to signal a male’s condition and predict reproductive performance. In Arctic‐breeding snow buntings Plectrophenax nivalis, the size of the alula simultaneously predicted pre‐breeding physiological health and the number of offspring produced, through an intermediate variable (lay date). Snow buntings appear to pair assortatively; males and females arriving earlier pair together, and changes in body condition over the breeding season are positively related within pairs. We suggest that simple achromatic plumage patches, like alula size, have the potential to act as condition‐dependent signals. Consequently, females may benefit from assessing these signals to reliably evaluate a male’s condition and reproductive potential as a means of maximizing their reproductive success.     

Phylogenomics unravels Quaternary vicariance and allopatric speciation patterns in temperate‐montane plant species: A case study on the Ranunculus auricomus species complex

The time frame and geographical patterns of diversification processes in European temperate‐montane herbs are still not well understood. We used the sexual species of the Ranunculus auricomus complex as a model system to understand how vicariance versus dispersal processes in the context of Pleistocene climatic fluctuations have triggered speciation in temperate‐montane plant species. We used target enrichment sequence data from about 600 nuclear genes and coalescent‐based species tree inference methods to resolve phylogenetic relationships among the sexual taxa of the complex. We estimated absolute divergence times and, using ancestral range reconstruction, we tested if speciation was enhanced by vicariance or by dispersal processes. Phylogenetic relationships among taxa were fully resolved with some incongruence in the position of the tetraploid R. marsicus. Speciation events took place in a very short time at the end of the Mid‐Pleistocene Transition (830–580 thousand years ago [ka]). A second wave of intraspecific geographical differentiation occurred at the end of the Riss glaciation or during the Eemian interglacial between 200 and 100 ka. Ancestral range reconstruction suggests a widespread European ancestor of the R. auricomus complex. Vicariance has triggered allopatric speciation in temperate‐montane plant species during the climatic deterioration that occurred during the last phase of the Mid‐Pleistocene Transition. Vegetation restructuring from forest into tundra could have confined these forest species into isolated glacial macro‐ and microrefugia. During subsequent warming periods, range expansions of these species could have been hampered by apomictic derivatives and by other congeneric competitors in the same habitat.     

Population structure on breeding grounds of Lake Malawi’s ‘twilight zone’ cichlid fishes

Aim Free‐ranging benthopelagic fishes often have large population sizes and high rates of dispersal. These traits can act to homogenize population structure across the distributional range of a species and to reduce the likelihood of allopatric speciation. The apparent absence of any barriers to gene flow among populations, together with prior molecular evidence for panmixia across the ranges of three species, has resulted in Diplotaxodon, a genus of benthopelagic cichlid fishes of Lake Malawi, being proposed as a candidate case of sympatric speciation. Our aim was to further investigate this possibility by testing for intraspecific genetic subdivision among breeding populations, and intraspecific differences in breeding habitat. Location Lake Malawi, central‐east Africa. Methods We analysed eight microsatellite DNA loci to test for spatial genetic differences among populations on breeding grounds of eight Diplotaxodon species. We also tested for temporal population genetic differences within breeding grounds of three species. Records of ripe Diplotaxodon encountered during sampling were analysed to test if spatial variation in assemblage structure was linked to nearshore water depth and geographic proximity of sampling sites. Results Consistent with previous molecular evidence, within four of the eight species tested we found no evidence of spatial genetic structuring among breeding populations. However, within the other four species we found slight yet significant spatial genetic differences, indicating restricted gene flow among breeding grounds. There was no evidence of temporal genetic differences within sites. Analyses of the distributions of ripe Diplotaxodon revealed differences in assemblage structure linked to nearshore water depth. Main conclusions Together, these results demonstrate both the evolution of fidelity to deep‐water breeding locations in some Diplotaxodon species, and differences in breeding habitat among species. These findings are consistent with a role for divergence of breeding habitat in speciation of these cichlids, possibly promoted by dispersal limitation among geographically segregated spawning aggregations.     

Ecological divergence combined with ancient allopatry in lizard populations from a small volcanic island

Population divergence and speciation are often explained by geographical isolation, but may also be possible under high gene flow due to strong ecology‐related differences in selection pressures. This study combines coalescent analyses of genetic data (11 microsatellite loci and 1 Kbp of mtDNA) and ecological modelling to examine the relative contributions of isolation and ecology to incipient speciation in the scincid lizard Chalcides sexlineatus within the volcanic island of Gran Canaria. Bayesian multispecies coalescent dating of within‐island genetic divergence of northern and southern populations showed correspondence with the timing of volcanic activity in the north of the island 1.5–3.0 Ma ago. Coalescent estimates of demographic changes reveal historical size increases in northern populations, consistent with expansions from a volcanic refuge. Nevertheless, ecological divergence is also supported. First, the two morphs showed non‐equivalence of ecological niches and species distribution modelling associated the northern morph with mesic habitat types and the southern morph with xeric habitat types. It seems likely that the colour morphs are associated with different antipredator strategies in the different habitats. Second, coalescent estimation of gene copy migration (based on microsatellites and mtDNA) suggest high rates from northern to southern morphs demonstrating the strength of ecology‐mediated selection pressures that maintain the divergent southern morph. Together, these findings underline the complexity of the speciation process by providing evidence for the combined effects of ecological divergence and ancient divergence in allopatry.     

Interspecific comparison of the fecal microbiota structure in three Arctic migratory bird species

The gut microbiota of birds is known to be characterized for different species, although it may change with feeding items. In this study, we compared the gut microbiota of birds with different feeding behaviors in the same habitat. We collected fecal samples from three Arctic species, snow buntings Plectrophenax nivalis, sanderlings Calidris alba, and pink‐footed geese Anser brachyrhynchus that are phylogenetically quite distant in different families to evaluate effects of diet on gut microbiota. Also, we characterized the prevalence of fecal bacteria using the Illumina MiSeq platform to sequence bacterial 16S rRNA genes. Our NMDS results showed that fecal bacteria of snow buntings and sanderlings were significantly distant from those of pink‐footed geese. Although all three birds were occupied by three bacterial phyla, Proteobacteria, Firmicutes, and Bacteroidetes, dominant taxa still varied among the species. Our bacterial sequences showed that snow buntings and sanderlings were dominated by Firmicutes and Bacteroidetes, while pink‐footed geese were dominated by Proteobacteria. In addition, the bacterial diversity in snow buntings and sanderlings was significantly higher than that in pink‐footed geese. Our results suggest that insectivorous feeding diet of snow buntings and sanderlings could be responsible for the similar bacterial communities between the two species despite the distant phylogenetic relationship. The distinctive bacterial community in pink‐footed geese was discussed to be related with their herbivorous diet.     

Pliocene climatic change in insular Southeast Asia as an engine of diversification in Ficedula flycatchers

Aims Insular Southeast Asia and adjacent regions are geographically complex, and were dramatically affected by both Pliocene and Pleistocene changes in climate, sea level and geology. These circumstances allow the testing of several biogeographical hypotheses regarding species distribution patterns and phylogeny. Avian species in this area present a challenge to biogeographers, as many are less hindered by barriers that may block the movements of other species. Widely distributed Southeast Asian avian lineages, of which there are many, have been generally neglected. Ficedula flycatchers are distributed across Eurasia, but are most diverse within southern Asia and Southeast Asian and Indo‐Australian islands. We tested the roles of vicariance, dispersal and the evolution of migratory behaviours as mechanisms of speciation within the Ficedula flycatchers, with a focus on species distributed in insular Southeast Asia. Methods Using a published molecular phylogeny of Ficedula flycatchers, we reconstructed ancestral geographical areas using dispersal vicariance analysis, weighted ancestral area analysis, and a maximum likelihood method. We evaluated the evolution of migratory behaviours using maximum likelihood ancestral character state reconstruction. Speciation timing estimates were calculated via local molecular clock methods. Results Ficedula originated in southern mainland Asia, c. 6.5 Ma. Our analyses indicate that two lineages within Ficedula independently and contemporaneously colonized insular Southeast Asia and Indo‐Australia, c. 5 Ma. The potential impact of vicariance due to rising sea levels is difficult to assess in these early colonization events because the ancestral areas to these clades are reconstructed as oceanic islands. Within each of these clades, inter‐island dispersal was critical to species’ diversification across oceanic and continental islands. Furthermore, Pliocene and Pleistocene climatic change may have caused the disjunct island distributions between several pairs of sister taxa. Both vicariance and dispersal shaped the distributions of continental species. Main conclusions This study presents the first evaluation, for Ficedula, of the importance of vicariance and dispersal in shaping distributions, particularly across insular Southeast Asia and Indo‐Australia. Although vicariant speciation may have initially separated the island clades from mainland ancestors, speciation within these clades was driven primarily by dispersal. Our results contribute to the emerging body of literature concluding that dynamic geological processes and climatic change throughout the Pliocene and Pleistocene have been important factors in faunal diversification across continental and oceanic islands.     

The last bastion? X chromosome genotyping of Anopheles gambiae species pair males from a hybrid zone reveals complex recombination within the major candidate ‘genomic island of speciation’

Speciation with gene flow may be aided by reduced recombination helping to build linkage between genes involved in the early stages of reproductive isolation. Reduced recombination on chromosome X has been implicated in speciation within the Anopheles gambiae complex, species of which represent the major Afrotropical malaria vectors. The most recently diverged, morphologically indistinguishable, species pair, A. gambiae and Anopheles coluzzii, ubiquitously displays a ‘genomic island of divergence’ spanning over 4 Mb from chromosome X centromere, which represents a particularly promising candidate region for reproductive isolation genes, in addition to containing the diagnostic markers used to distinguish the species. Very low recombination makes the island intractable for experimental recombination studies, but an extreme hybrid zone in Guinea Bissau offers the opportunity for natural investigation of X‐island recombination. SNP analysis of chromosome X hemizygous males revealed: (i) strong divergence in the X‐island despite a lack of autosomal divergence; (ii) individuals with multiple‐recombinant genotypes, including likely double crossovers and localized gene conversion; (iii) recombination‐driven discontinuity both within and between the molecular species markers, suggesting that the utility of the diagnostics is undermined under high hybridization. The largely, but incompletely protected nature of the X centromeric genomic island is consistent with a primary candidate area for accumulation of adaptive variants driving speciation with gene flow, while permitting some selective shuffling and removal of genetic variation.     

Historical biogeography of the endemic Campanulaceae of Crete

Aim The clade Campanulaceae in the Cretan area is rich in endemics, with c. 50% of its species having restricted distributions. These species are analysed in the context of a larger phylogeny of the Campanulaceae. Divergence times are calculated and hypotheses of vicariance and dispersal are tested with the aim of understanding whether Cretan lineages represent remnants of an older continental flora. Location The Cretan area: Crete and the Karpathos Islands (Greece). Methods We obtained chloroplast DNA sequence data from rbcL, atpB and matK genes for 102 ingroup taxa, of which 18 are from the Cretan area, 11 are endemics, and two have disjunct, bi‐regional distributions. We analysed the data using beast , a Bayesian approach that simultaneously infers the phylogeny and divergence times. We calibrated the tree by placing a seed fossil in the phylogeny, and used published age estimates as a prior for the root. Results The phylogenetic reconstruction shows that all Campanula species fall within a well‐supported campanuloid clade; however, Campanula is highly polyphyletic. The Cretan endemics do not form a monophyletic group, and species are scattered throughout the campanuloid clade. Therefore, the Cretan taxa did not evolve following a single vicariance or dispersal event. Most Cretan lineages represent remnants of an older continental flora, with the exception of one clade that radiated in situ after island isolation, and one lineage that appears to have arrived by dispersal. Main conclusions Most Cretan species were present in the islands at the time of their isolation, and very little long‐distance dispersal to Crete and diversification within Crete has occurred since then. Endemism is probably driven by loss of species on the mainland after island isolation. Species on the islands may have been more widespread in the past, but they are now restricted to often inaccessible areas, probably as a result of human pressure.     

Interactions with Great Skuas Stercorarius skua as a factor in the long‐term decline of an Arctic Skua Stercorarius parasiticus population

Many of the UK’s seabird species have displayed high variation in breeding success since the 1980s, largely due to changes in the availability of Lesser Sandeels Ammodytes marinus, their main prey. During this time, Arctic Skuas Stercorarius parasiticus experienced a rapid decline in the UK and the species has subsequently been placed on the Red List of birds of conservation concern. Although shortage of Lesser Sandeels is likely to be an influential factor, the Arctic Skua’s breeding range overlaps with that of the Great Skua Stercorarius skua, a larger bird with a more varied diet, and interspecific interactions for nesting habitat may exert an additional pressure on Arctic Skua breeding populations. Results from four censuses, spanning 21 years, were used to model habitat use and analyse distributional change in nesting Arctic Skuas at a major colony located on Fetlar, Shetland, Scotland. The decline in Arctic Skuas was not uniform across the island and competition with Great Skuas for nest‐sites appears to have influenced localized breeding distribution. By 2006, Arctic Skuas had been almost entirely excluded from shrub heath, blanket bog and coastal heath habitats, which were identified as preferred habitat in 1986. In 2006, Arctic Skua breeding territories were mainly restricted to one core area of preferred habitat where over 90% nested in high density as this habitat became increasingly occupied by Great Skuas. The more generalist foraging habit of the Great Skua allowed the population to grow rapidly as numbers of the more specialist Arctic Skua decreased during times of low sandeel availability. Our model suggests that both interspecific competition for territories with Great Skuas and food limitation have played important roles in the decline of Arctic Skuas on Fetlar.     

The comparative biogeography of Philippine geckos challenges predictions from a paradigm of climate‐driven vicariant diversification across an island archipelago

A primary goal of biogeography is to understand how large‐scale environmental processes, like climate change, affect diversification. One often‐invoked but seldom tested process is the “species‐pump” model, in which repeated bouts of cospeciation are driven by oscillating climate‐induced habitat connectivity cycles. For example, over the past three million years, the landscape of the Philippine Islands has repeatedly coalesced and fragmented due to sea‐level changes associated with glacial cycles. This repeated climate‐driven vicariance has been proposed as a model of speciation across evolutionary lineages codistributed throughout the islands. This model predicts speciation times that are temporally clustered around the times when interglacial rises in sea level fragmented the islands. To test this prediction, we collected comparative genomic data from 16 pairs of insular gecko populations. We analyze these data in a full‐likelihood, Bayesian model‐choice framework to test for shared divergence times among the pairs. Our results provide support against the species‐pump model prediction in favor of an alternative interpretation, namely that each pair of gecko populations diverged independently. These results suggest the repeated bouts of climate‐driven landscape fragmentation have not been an important mechanism of speciation for gekkonid lizards across the Philippine Archipelago.     

Plio-Pleistocene climatic oscilations, Holarctic biogeography and speciation in an avian subfamily

Aim In this paper, I discuss the temporal and spatial aspects of historical biogeography and speciation in a widely distributed Holarctic subfamily of birds (Tetraoninae). Location Northern Holarctic. Results Using dated fossils, I calibrated the molecular clock for the mitochondrial control region at 7.23 ± 1.58% nucleotide divergence (maximum likelihood corrected) per million years. The data suggest that grouse (Tetraoninae) originated in the Middle Pliocene, 6.3 Ma. Grouse apparently originated in the northern part of western Nearctic, and Palearctic was colonized independently three times, first by the ancestor of all grouse in the Middle Pliocene, then by the ancestor of forest (Falcipennis, Tetrao and Lyrurus) and prairie (Centrocercus, Dendragapus and Tympanuchus) grouse in the Late Pliocene, and finally by the ancestral Lagopus in the Early Pleistocene. Only once Nearctic was colonized from Palearctic by a common ancestor of forest grouse. Sympatry and range symmetry were positively correlated with molecular divergence. These correlations suggest that peripatric isolation was the predominant mode of speciation throughout grouse history. Main conclusions Speciation events in grouse were driven by climatic oscillations of the Pliocene and Pleistocene. Isolation of small peripheral populations from widely distributed ancestors was the dominant mode of speciation in grouse. Isolations during interglacials both across Beringia, and in southern mountain areas when boreal habitats were restricted to high elevations, suggest an important role for vicariance in grouse speciation.     

Local endemism and within‐island diversification of shrews illustrate the importance of speciation in building Sundaland mammal diversity

Island systems are important models for evolutionary biology because they provide convenient, discrete biogeographic units of study. Continental islands with a history of intermittent dry land connections confound the discrete definitions of islands and have led zoologists to predict (i) little differentiation of terrestrial organisms among continental shelf islands and (ii) extinction, rather than speciation, to be the main cause of differences in community composition among islands. However, few continental island systems have been subjected to well‐sampled phylogeographic studies, leaving these biogeographic assumptions of connectivity largely untested. We analysed nine unlinked loci from shrews of the genus Crocidura from seven mountains and two lowland localities on the Sundaic continental shelf islands of Sumatra and Java. Coalescent species delimitation strongly supported all currently recognized Crocidura species from Sumatra (six species) and Java (five species), as well as one undescribed species endemic to each island. We find that nearly all species of Crocidura in the region are endemic to a single island and several of these have their closest relative(s) on the same island. Intra‐island genetic divergence among allopatric, conspecific populations is often substantial, perhaps indicating species‐level diversity remains underestimated. One recent (Pleistocene) speciation event generated two morphologically distinct, syntopic species on Java, further highlighting the prevalence of within‐island diversification. Our results suggest that both between‐ and within‐island speciation processes generated local endemism in Sundaland, supplementing the traditional view that the region's fauna is relictual and primarily governed by extinction.     

Evolutionary history of the scorpionfly Dicerapanorpa magna (Mecoptera,Panorpidae)

Climate changes can have fundamental impacts on the distributional patterns of montane species, and range shifts frequently lead to allopatric divergence followed by the establishment of secondary contact zones. Many European and North American organisms have retreated to southern refugia during glacial periods and colonized northward during postglacial periods, but little is known about the evolutionary response of cold‐adapted insects to Pleistocene climate changes in eastern Asia. The scorpionfly Dicerapanorpa magna (Chou), with cold temperate habitat preference and weak dispersal ability, provides a good model system to explore how climate changes have influenced the distribution and divergence of cold‐adapted insects in eastern Asia. This study reconstructed the demographic dynamics and evolutionary history of D. magna with phylogeographic approaches, and predicted the species’ suitable areas under the Last Glacial Maximum (LGM) and current scenarios with the ecological niche modelling analysis. The mitochondrial cytochrome c oxidase subunit I resolved three phylogenetic lineages in D. magna dating back to Pleistocene, corresponding well with the geographically isolated Qinling, Bashan and Minshan Mountains. The ecological niche modelling recovered the suitable habitats for D. magna were the Qinling and Bashan Mountains under LGM and current conditions. The three lineages of D. magna might be in a process of incipient speciation, and likely derived their current distribution from separate glacial origins, followed by vicariance and divergence.     

Biogeographical basis of recent phenotypic divergence among birds: a global study of subspecies richness

Theory predicts that biogeographic factors should play a central role in promoting population divergence and speciation. Previous empirical studies into biogeography and diversification have been relatively restricted in terms of the geographical area, phylogenetic scope, and the range of biogeographic factors considered. Here we present a global analysis of allopatric phenotypic divergence (measured as subspecies richness) across more than 9600 bird species. The main aim of this study was to examine the extent to which biogeographical factors can explain patterns of phenotypic divergence. Analysis of the taxonomic distribution of subspecies among species suggests that subspecies formation and extinction have occurred at a considerably faster rate than has species formation. However, the observed distribution departs from the expectation under a random birth-death model of diversification. Across 19 phylogenetic trees, we find no significant linear relationship between species age and subspecies richness, implying that species age is a poor predictor of subspecies richness. Both subspecies richness and subspecies diversification rate are found to exhibit low phylogenetic signal, meaning that closely related species do not tend to possess similar numbers of subspecies. As predicted by theory, high subspecies richness was associated with large breeding range size, island dwelling, inhabitation of montane regions, habitat heterogeneity, and low latitude. Of these factors, breeding range size was the variable that explained the most variation. Unravelling whether species that have invaded previously glacial areas have more or fewer subspecies than expected proves to be complicated due to a covariation between the postglacial colonization, latitude, geographic range size, and subspecies richness. However, the effect of postglacial colonization on subspecies richness appears to be small. Mapping the distribution of species' subspecies richness globally reveals geographical patterns that correspond to many of the predictions of the statistical models, but may also reflect geographical variation in taxonomic practice. Overall, we demonstrate that biogeographic models can explain about 30% of the global variation in subspecies richness in birds.     

ADAPTIVE RADIATION AND ECOLOGICAL DIVERSIFICATION OF SULAWESI'S ANCIENT LAKE SHRIMPS

Ancient lakes are natural laboratories for the study of adaptive radiation. Recently, two lake systems on the Indonesian island Sulawesi have emerged as promising new model systems. A species flock of atyid freshwater shrimps in the Malili lake system comprises 15 colorful endemic taxa. Mitochondrial DNA data suggest two independent colonizations by riverine ancestors. Only one colonization event led to subsequent radiation into 14 species, while the second clade comprises just one species. The vast majority of species (n= 12) are habitat specialists, which are confined to the larger Malili clade and include all taxa with species‐specific color patterns and a restricted distribution within the five connected Malili lakes. Morphological, genetic, and ecological data are consistent with the existence of an adaptive radiation in the Malili lakes, involving the habitat‐specific diversification of trophic morphology. In addition to testing criteria for the recognition of an adaptive radiation, an ancestral state reconstruction reveals an equal probability for either a riverine generalist or a lacustrine specialist as ancestor of the large Malili clade, which is interpreted as indicative of an early stage of habitat specialization within this radiation. Finally, our results suggest that species diversification may have been primarily driven by ecological specialization and allopatric speciation.     

Evolution of the tribe Tropheini from Lake Tanganyika: synchronized explosive speciation producing multiple evolutionary parallelism

One of the most surprising outcomes of recent molecular studies on cichlid fishes of the three Great East African Lakes Victoria, Malawi and Tanganyika, was the stunning rapidity of speciation and cladogenesis at early stages of adaptive radiation. Despite their rapid pace, speciation events were so far intuitively assumed to proceed in a bifurcating and tree-like fashion, even if they could not be resolved by gene phylogenies due to a lack of resolution. On the basis of phylogenetic analyses of the Tropheini, a lineage of endemic rock-dwelling cichlid fishes from Lake Tanganyika, we suggest a pathway of explosive speciation that accounts for a non-bifurcating manner of cladogenesis. This pattern is likely to be the result of the contemporaneous origin of a multitude of founder populations in geographically isolated rock habitats among which gene flow was interrupted simultaneously by a major change of the lake habitat in the form of a rapid rise of the lake level. As a consequence, all new species arising from that vicariance event must exhibit almost equal genetic distances to each other, within the scope of genetic diversity of the founder population(s), even if the actual processes of subsequent speciation and eco-morphological diversification followed independent routes. Our phylogeny also suggests a high frequency of parallel evolution of equivalent trophic specialization in the Tropheini. This phenomenon seems to be an inherent feature of this pathway of speciation, due to the action of similar selective forces on the same set of species colonizing isolated habitats of the same type. Explosive speciation via synchronization of genetic divergence triggered by rapid environmental changes seems to be particularly likely to occur at advanced stages of adaptive radiation, when species are already adapted to particular habitats and have a reduced ability for dispersal.