Rejecting strictly allopatric speciation on a continental island: prolonged postdivergence gene flow between Taiwan (Leucodioptron taewanus,Passeriformes Timaliidae) and Chinese (L. canorum canorum) hwameis

Allopatry is conventionally considered the geographical mode of speciation for continental island organisms. However, strictly allopatric speciation models that assume the lack of postdivergence gene flow seem oversimplified given the recurrence of land bridges during glacial periods since the late Pliocene. Here, to evaluate whether a continental island endemic, the Taiwan hwamei (Leucodioptron taewanus, Passeriformes Timaliidae) speciated in strict allopatry, we used weighted‐regression‐based approximate Bayesian computation (ABC) to analyse the genetic polymorphism of 18 neutral nuclear loci (total length: 8500 bp) in Taiwan hwamei and its continental sister species, the Chinese hwamei (L. canorum canorum). The nonallopatry model was found to fit better with observed genetic polymorphism of the two hwamei species (posterior possibility = 0.82). We also recovered unambiguous signals of nontrivial bidirectional postdivergence gene flow (N_em » 1) between Chinese hwamei and Taiwan hwamei until 0.5 Ma. Divergence time was estimated to be 3.5 to 2 million years earlier than that estimated from mitochondrial cytochrome b sequences. Finally, using the inferred nonallopatry model to simulate genetic variation at 24 nuclear genes examined showed that the adiponectin receptor 1 gene may be under divergent adaptation. Our findings imply that the role of geographical barrier may be less prominent for the speciation of continental island endemics, and suggest a shift in speciation studies from simply correlating geographical barrier and genetic divergence to examining factors that facilitate and maintain divergence, e.g. differential selection and sexual selection, especially in the face of interpopulation gene flow.     

Adaptation and speciation of Dolichopoda cave crickets (Orthoptera, Rhaphidophoridae): geographic variation of morphometric indices and allozyme frequencies

Morphometric analyses were carried out for 14 cave dwelling populations of Dolichopoda (Orthoptera, Rhaphidophoridae) from peninsular Italy belonging to the laetitiae-geniculata complex. The selected set of variables includes linear measures of body parts and appendages whose variation is expected to be adaptive and to reflect the degree of specialization to the cave environment.
Inter-population morphometric variation was compared with the patterns of allozyme variation, investigated electrophoretically in a previous paper, by means of ordination and classification multivariate analyses. Results from this comparison reject the hypothesis of parallel geographic variation of morphometric indices and allele frequencies, revealing independent trends for the two sets of characters.     

Microdistribution within a population ofAcrocnida brachiata (Montagu), (Echinodermata: Ophiuroidea) in Little Killary, west coast of Ireland

Acrocnida brachiata (Montagu), like many other infaunal ophiuroids, has a penchant for gregariousness. In Little Killary, a small embayment on the west coast of Ireland, it colonises an extensive tract of sandy inshore ground (ca 7 m depth), at densities of 150–200 individuals/m². In the course of a two-year study, variation in regular suction-sampling returns prompted a detailed study of the species distribution pattern. Towards this end, 256 contiguous and stratified core samples were taken along a line of transect, in May 1995. These showed randomness with respect to the population at large, but exposed an aggregated distribution for juveniles and adults separately, at different levels within the sediment. Juveniles formed wide, ‘loose’ patches in the surficial deposit, while adults formed smaller, denseer patches deeper within the sediment. It remains to be seen what this approach may expose, when applied in different seasons, or during different phases of the species life cycle.     

Evolution of oviposition strategies and speciation in the globeflower flies Chiastocheta spp. (Anthomyiidae)

Trollius europaeus (Ranunculaceae) is involved in an intimate interaction with several species of Chiastocheta flies (Anthomyiidae) that are both seed predators and pollinators. In this paper, we analyse the oviposition strategy of the six Chiastocheta species found to coexist on T. europaeus in 19 populations from the French Alps. We show that the species are not equivalent in their oviposition behaviour: C. rotundiventris usually deposits no more than one egg per flower in first-day flowers whereas C. dentifera aggregates its eggs on fruits and thus does not contribute to pollination at all; the four remaining species deposit eggs sequentially during the flowering period from the 2nd to the 7th day. Hence, the outcomes of the interaction in terms of net seed production for the plant greatly depend on the Chiastocheta species visiting it, ranging from a mutualistic to a purely parasitic interaction. We assessed mitochondrial divergence between Chiastocheta spp. by sequencing a 1320-bp mitochondrial DNA fragment. The low divergence observed between species (0–4.15%) suggests that genus diversification took place recently. Unlike in other plant–insect systems where diversification is usually thought to be driven by cospeciation or host shifts, we propose that Chiastocheta speciation took place within the host plant. Basal separation of a particularly mutualistic species provided favourable conditions for plant specialization on this seed-parasite as a pollinator early in the evolution of the association. The parasitic species ovipositing on fruits derived from a species ovipositing on flowers. Diversification of the intermediate strategies probably occurred in relation with the Pleistocene climatic events, reproductive isolation between species being reinforced by niche partitioning for oviposition and/or sexual selection.     

Evolution and speciation of cave-dwelling Fulgoroidea in the Canary Islands (Homoptera: Gixiidae and Meenoplidae)

Ten new cavernicolous Fulgoroidea species of the families Cixiidae and Meenoplidae are described from the Canary Islands: Cixius palmeros, C. pinarcoladus, C. ratonicus, C. tacandus, Memoplus claustrophilus from La Palma, C. ariadne, C. nycticolus, M. charon from El Hierro, and Tachycixius crypticus, T. retrusus from Tenerife. Notes on their ecology and distribution are given, and where possible, phylogenetic affinities to existing epigean species of the corresponding families are discussed. Despite the present-day relict status of the majority of cavernicolous species, it cannot be concluded with certainty whether their evolution has followed allopatric or parapatric modes of speciation.     

Random interbreeding between cryptic lineages of the Common Raven: evidence for speciation in reverse

DNA sequence studies frequently reveal evidence of cryptic lineages in morphologically uniform species, many of which turn out to be evolutionarily distinct species. The Common Raven (Corvus corax) includes two deeply divergent mtDNA lineages: one lineage seems restricted to western North America and the other is Holarctic in distribution. These deep clades hint of the possibility of cryptic species in the western United States. We tested this hypothesis in a population consisting of an equal proportion of both mtDNA clades, by quantifying mating patterns and associated fitness consequences with respect to mtDNA. We also tested for morphological, behavioural and ecological correlates of sex and mtDNA clade membership. Mate pairings were random with respect to mtDNA clades, and there were no differences in reproductive success between assortatively and nonassortatively mated pairs. We found no differences in survival or resource use between clades. There were no differences in morphological or behavioural characters between mtDNA clades, except one clade trended towards greater mobility. These results suggest there are no barriers to gene flow between mtDNA clades and argue that the mtDNA clades have remerged in this population, likely due to a lack of ecological or signal differentiation between individuals in each lineage. Hence, in Common Ravens, phylogeographic structure in mtDNA is a reflection of likely past isolation rather than currently differentiated species.     

Assemblages of sea stars (Echinodermata: Asteroidea) and brittle stars (Echinodermata: Ophiuroidea) in the Weddell Sea (Antarctica) and off Northeast Greenland (Arctic): a comparison of diversity and abundance

Composition and distribution of asteroid and ophiuroid assemblages were investigated by means of Agassiz trawl catches at 34 stations in 220- to 1,200-m depth in the Weddell Sea and at 17 stations in 90- and 830-m depth off Northeast Greenland. A total of 86 species (48 sea stars, 38 brittle stars) were identified in the Weddell Sea whereas off Northeast Greenland a total of 26 species (16 sea stars, 10 brittle stars) were recorded. In both study areas, brittle stars were numerically more important than sea stars, and abundances generally decreased with water depths. Multivariate analyses revealed a conspicuous depth zonation of sea and brittle stars off Greenland. Very high abundances of Ophiocten sericeum and Ophiura robusta characterized the assemblages on shallow shelf banks whereas in greater depths Ophiopleura borealis, Ophioscolex glacialis and Ophiacantha bidentata became dominant, albeit at significantly lower densities. Mass occurrences of brittle stars, such as those recorded on Greenlandic shelf banks, have not been discovered in the Weddell Sea, where distinct assemblages were discriminated in deep shelf trenches as well as on the eastern and southern shelf. Ophioplocus incipiens, Ophiurolepis martensi and Ophiurolepis brevirima were the most prominent species on the eastern shelf. Ophiacantha antarctica, Ophiurolepis gelida and Ophionotus victoriae on the southern shelf, and Ophiosparte gigas as well as the asteriod Hymenaster sp., in the shelf trenches. Overall, the Weddell Sea housed conspicuously more asterozoan species than the waters off Greenland. Higher species diversity was also evident at both a regional and local scale, especially for the eastern Weddell Sea shelf. However, because many species from the Weddell Sea are closely related, the Weddell Sea assemblages were not significantly different from the Greenland ones in terms of taxonomic diversity and distinctness. Received: 29 April 1996/Accepted: 10 June 1996     

A young clade repeating an old pattern: diversity in Nothonotus darters (Teleostei: Percidae) endemic to the Cumberland River

Hypotheses of diversification in eastern North American freshwater fishes have focused primarily on allopatric distributions of species between disjunct highland areas and major river systems. However, these hypotheses do not fully explain the rich diversity of species within highland regions and river systems. Relatively old diversification events at small geographic scales have been observed in the Barcheek Darter subclade that occurs in the Cumberland River drainage (CRD) in Kentucky and Tennessee, United States of America, but it is unknown if this pattern is consistent in other darter subclades. We explored phylogeographic diversity in two species of Nothonotus darters, N. microlepidus and N. sanguifluus, endemic to the CRD to compare phylogenetic patterns between Barcheek Darters and species of Nothonotus. We collected sequence data for a mitochondrial gene (cytb) and three nuclear genes (MLL, S7 and RAG1) from 19 N. microlepidus and 35 N. sanguifluus specimens. Gene trees were estimated using maximum likelihood and Bayesian methods, and a 'species tree' was inferred using a Bayesian method. These trees indicate that species diversity in Nothonotus is underestimated. Five distinct lineages were evident, despite retained ancestral polymorphism and unsampled extirpated populations. Comparison of chronograms for Barcheek Darters and Nothonotus revealed that microendemism resulting from species diversification at small geographic scales in the CRD is a consistent pattern in both old and young darter subclades. Our analyses reveal that geographic isolating mechanisms that result in similar phylogeographic patterns in the CRD are persistent through long expanses of evolutionary time.     

Catalogue of neritic-benthonic echinoderms (Echinodermata: Asteroidea and Ophiuroidea) from the Cuban Archipelago

This is an updated taxonomic catalogue of neritic-benthonic Echinodermata (Asteroidea and Ophiuroidea) from Cuba, that includes locality, range, habitat and depth where each species is found. The species list is based on the collections of the Oceanology Institute; Havana; the "Felipe Poey" Museum (Havana University) and the National Museum of Natural History, Smithsonian Institution, Washington D.C. Specimens for those collections were captured by a variety of field techniques, narcotized with Cloral hydrate, fixed in formalin and stored in 70-80% alcohol. A total of 75 species have been identified from Cuban waters: Asteroidea (20 species) and Ophiuroidea (55 species).     

Comparative phylogeography of two coastal polychaete tubeworms in the Northeast Atlantic supports shared history and vicariant events

The historic processes which have led to the present-day patterns of genetic structure in the marine coastal fauna of the Northeast Atlantic are still poorly understood. While tectonic uplifts and changes in sea level may have caused large-scale vicariance, warmer conditions during glacial maxima may have allowed pockets of diversity to persist to a much wider extent than in the Northwestern Atlantic. The large-scale geographic distribution of deeply divergent lineages of the coastal polychaete tubeworms Pectinaria koreni (two clades) and Owenia fusiformis (three clades) were compared using a fragment of the mitochondrial cytochrome oxidase I gene (mtCOI). All lineages were present along the biogeographic transition zone on the north coast of Brittany (France) and we found evidence pointing towards congruence in the timing of cladogenic events between Pectinaria sp. (P. auricoma/P. belgica and P. koreni) and Owenia sp., suggesting a shared history of vicariant events. More conserved 16SrRNA sequences obtained from four species of Pectinariidae together with mtCOI sequences of P. koreni seem consistent with an initial establishment of pectinariids in the north, and a southward colonization of the Northeast Atlantic. Phylogeographic patterns in O. fusiformis were also consistent with a north/south pattern of lineage splitting and congruent levels of divergence were detected between lineages of both species. We observed signatures of both persistence in small northern glacial refugia, and of northwards range expansion from regions situated closer to the Mediterranean. However, whether the recolonization of the Northeast Atlantic by both species actually reflects separate interglacial periods is unclear with regards to the lack of molecular clock calibration in coastal polychaete species.     

Hybridization, ecological races and the nature of species: empirical evidence for the ease of speciation

Species are generally viewed by evolutionists as 'real' distinct entities in nature, making speciation appear difficult. Charles Darwin had originally promoted a very different uniformitarian view that biological species were continuous with 'varieties' below the level of species and became distinguishable from them only when divergent natural selection led to gaps in the distribution of morphology. This Darwinian view on species came under immediate attack, and the consensus among evolutionary biologists today appears to side more with the ideas of Ernst Mayr and Theodosius Dobzhansky, who argued 70 years ago that Darwin was wrong about species. Here, I show how recent genetic studies of supposedly well-behaved animals, such as insects and vertebrates, including our own species, have supported the existence of the Darwinian continuum between varieties and species. Below the level of species, there are well-defined ecological races, while above the level of species, hybridization still occurs, and may often lead to introgression and, sometimes, hybrid speciation. This continuum is evident, not only across vast geographical regions, but also locally in sympatry. The existence of this continuum provides good evidence for gradual evolution of species from ecological races and biotypes, to hybridizing species and, ultimately, to species that no longer cross. Continuity between varieties and species not only provides an excellent argument against creationism, but also gives insight into the process of speciation. The lack of a hiatus between species and ecological races suggests that speciation may occur, perhaps frequently, in sympatry, and the abundant intermediate stages suggest that it is happening all around us. Speciation is easy!     

Micro-evolution in the weevil genus Larinus: the formation of host biotypes and speciation

Data are presented on allozyme variation between 15 populations of the stenophagous capitulum weevil, Larinus cynarae , and three populations of its congener, L. latus , that had been collected throughout the northern mediterranean range of these species. A phenetic analysis of these data revealed no direct relationship between genetic variation and host-plant association within L. cynarae , but there was a strong geographical structuring of allozyme patterns. Most of the genetic variation was due to differences between geographical regions and variation within these was small. Wright's F _ST values showed that Italian and Greek populations of L. cynarae were most distinct from L. latus , with southern Iberian, northern Spanish and French populations increasingly less so. This pattern was associated with a cline in the frequencies of certain alleles along this geographical arc from France to Greece. A phenogram of Nei's genetic distances indicated the close genetic relationship between the two species of Larinus and separated the populations of L. cynarae into three allopatric groups. These groups have different host-plant spectra - dominated by Cynara cardunculus in Italy and Greece, Cynara humilis/Onopordum in southern Iberia and Onopordum spp. in France/Northern Spain - and can be considered to be host biotypes of L. cynarae. L. latus , which occurs in Greece and further east is also an Onopordum specialist. An analysis of the phylogeny of this group of Larinus indicates a primary separation into eastern ( L. latus ) and western ( L. cynarae ) taxa, with further branching of the L. cynarae lineage into the putative host-biotypes. An hypothesis for the evolution of these taxa is given, based on the evolutionary history of host-plant taxa and geographical constraints.     

Hybridization among cryptic species of the cellar fungus Coniophora puteana (Basidiomycota)

In this study we have analysed the genetic variation and phylogeography in a global sample of the cellar fungus Coniophora puteana, which is an important destroyer of wooden materials indoor. Multilocus genealogies of three DNA regions (beta tubulin, nrDNA ITS and translation elongation factor 1alpha) revealed the occurrence of three cryptic species (PS1-3) in the morphotaxon C. puteana. One of the lineages (PS3) is apparently restricted to North America while the other two (PS1-2) have wider distributions on multiple continents. Interspecific hybridization has happened between two of the lineages (PS1 and PS3) in North America. In three dikaryotic isolates, two highly divergent beta tubulin alleles coexisted, one derived from PS1 and one from PS3. Furthermore, one isolate included a recombinant ITS sequence, where ITS1 resembled the ITS1 version of PS3 while ITS2 was identical to a frequent PS1 ITS2 version. This pattern must be due to hybridization succeeded by intralocus recombination in ITS. The results further indicated that introgression has happened between subgroups appearing in PS1. We hypothesize that the observed reticulate evolution is due to previous allopatric separation followed by more recent reoccurrence in sympatry, where barriers to gene flow have not yet evolved. A complex phylogeographical structure is observed in the morphotaxon C. puteana caused by (i) cryptic speciation; (ii) the interplay between natural migration and distribution patterns and probably more recent human mediated dispersal events; and (iii) hybridization and introgression.     

Molecular and morphological differentiation between steganacarid mites (Acari: Oribatida) from the Canary islands

Two steganacarid species, Steganacarus ( Steganacarus ) tenerifensis and S . ( S .) carlosi , the first endemic to Tenerife and the second to La Gomera and Tenerife, were found to have such wide morphological variability as to cast doubt on their specific limits. Cluster analysis and MDS ordering were used to quantify morphological variation, and biochemical procedures were used to assess genetic variation in the two entities. The results revealed disagreement between the morphological and genetic data as reported in previous studies on steganacarid mites. Morphological traits were highly heterogeneous in the Canary island populations and genetic structure was homogeneous with very low heterozygosity. The separation of the two Canary island species, not to mention the phenotypes close to carlosi , is difficult to justify due to their very high genetic identity. The presence of morphological intermediates suggests the same conclusion. However, the sympatry of some phenotypes and previous data on this animal group are a source of doubt and suggest that further investigations are needed before conclusions can be reached in this regard. Correlations between morphological differentiation and known palaeogeographic events are suggested in the light of the possible evolutionary history of the steganacarid mites on the Canary islands.     

Defining reproductively isolated units in a cryptic and syntopic species complex using mitochondrial and nuclear markers: the brooding brittle star, Amphipholis squamata (Ophiuroidea)

At a time when biodiversity is threatened, we are still discovering new species, and particularly in the marine realm. Delimiting species boundaries is the first step to get a precise idea of diversity. For sympatric species which are morphologically undistinguishable, using a combination of independent molecular markers is a necessary step to define separate species. Amphipholis squamata , a cosmopolitan brittle star, includes several very divergent mitochondrial lineages. These lineages appear totally intermixed in the field and studies on morphology and colour polymorphism failed to find any diagnostic character. Therefore, these mitochondrial lineages may be totally interbreeding presently. To test this hypothesis, we characterized the genetic structure of the complex in the French Mediterranean coast using sequences of mitochondrial DNA (16S) and for the first time, several nuclear DNA markers (introns and microsatellites). The data revealed six phylogenetic lineages corresponding to at least four biological species. These sibling species seem to live in syntopy. However, they seem to display contrasted levels of genetic diversity, suggesting they have distinct demographic histories and/or life-history traits. Genetic differentiation and isolation-by-distance within the French Mediterranean coasts are revealed in three lineages, as expected for a species without a free larval phase. Finally, although recombinant nuclear genotypes are common within mitochondrial lineages, the data set displays a total lack of heterozygotes, suggesting a very high selfing rate, a feature likely to have favoured the formation of the species complex.     

Speciation and anagenesis in the genus Cyprinella of Mexico (Teleostei: Cyprinidae): a case study of Model III allopatric speciation

Allozyme variation at 42 presumptive gene lociis presented for members of the C. formosa species group. This group iscorroborated as a monophyletic assemblage whosecommon ancestor occupied pluvial Lake Palomasof the Guzman Basin. With increasingaridity during the Pleistocene this basin andassociated populations of this commonancestor were fragmented into several lineagesthat diverged independently of oneanother. The pattern of relationships andlevels of anagenetic change observed inindependent lineages for this clade are notconsistent with expectations of the mostcommon modes of speciation, Model I large-scalevicariance or Model II peripheralisolation. Rather, divergence in these fishlineages is consistent with the rarely observedModel III allopatric speciation. Consistentwith predictions of this model, thephylogenetic pattern recovered revealspolychotomous relationships (a hard polytomy)and varied rates of anagenetic change acrossexamined lineages.     

Geographic distribution of Ophiothela brittle stars (Echinodermata: Ophiuroidea): substrate use plasticity and implications for the silent invasion of O. mirabilis in the Atlantic

Hydrobiologia - This study aimed to address the Ophiothela species distribution worldwide, report invasive populations, and investigate their association with benthic organisms through an...     

Molecular genetic and quantitative trait divergence associated with recent homoploid hybrid speciation: a study of Senecio squalidus (Asteraceae)

Hybridization is increasingly seen as a trigger for rapid evolution and speciation. To quantify and qualify divergence associated with recent homoploid hybrid speciation, we compared quantitative trait (QT) and molecular genetic variation between the homoploid hybrid species Senecio squalidus and its parental species, S. aethnensis and S. chrysanthemifolius, and also their naturally occurring Sicilian hybrids. S. squalidus originated and became invasive in the United Kingdom following the introduction of hybrid plants from Mount Etna, Sicily, about 300 years ago. We recorded considerable molecular genetic differentiation between S. squalidus and its parents and their Sicilian hybrids in terms of both reduced genetic diversity and altered allele frequencies, potentially due to the genetic bottleneck associated with introduction to the United Kingdom. S. squalidus is also distinct from its parents and Sicilian hybrids for QTs, but less so than for molecular genetic markers. We suggest that this is due to resilience of polygenic QTs to changes in allele frequency or lack of selection for hybrid niche divergence in geographic isolation. While S. squalidus is intermediate or parental-like for most QTs, some trangressively distinct traits were observed, which might indicate emerging local adaptation in its invasive range. This study emphasizes the important contribution of founder events and geographic isolation to successful homoploid hybrid speciation.     

The geography and ecology of plant speciation: range overlap and niche divergence in sister species

A goal of evolutionary biology is to understand the roles of geography and ecology in speciation. The recent shared ancestry of sister species can leave a major imprint on their geographical and ecological attributes, possibly revealing processes involved in speciation. We examined how ecological similarity, range overlap and range asymmetry are related to time since divergence of 71 sister species pairs in the California Floristic Province (CFP). We found that plants exhibit strikingly different age-range correlation patterns from those found for animals; the latter broadly support allopatric speciation as the primary mode of speciation. By contrast, plant sisters in the CFP were sympatric in 80% of cases and range sizes of sisters differed by a mean of 10-fold. Range overlap and range asymmetry were greatest in younger sisters. These results suggest that speciation mechanisms broadly grouped under ‘budding’ speciation, in which a larger ranged progenitor gives rise to a smaller ranged derivative species, are probably common. The ecological and reproductive similarity of sisters was significantly greater than that of sister–non-sister congeners for every trait assessed. However, shifts in at least one trait were present in 93% of the sister pairs; habitat and soil shifts were especially common. Ecological divergence did not increase with range overlap contrary to expectations under character displacement in sympatry. Our results suggest that vicariant speciation is more ubiquitous in animals than plants, perhaps owing to the sensitivity of plants to fine-scale environmental heterogeneity. Despite high levels of range overlap, ecological shifts in the process of budding speciation may result in low rates of fine-scale spatial co-occurrence. These results have implications for ecological studies of trait evolution and community assembly; despite high levels of sympatry, sister taxa and potentially other close relatives, may be missing from local communities.