Molecular architecture of Pipistrellus pipistrellus/Pipistrellus pygmaeus complex (Chiroptera: Vespertilionidae): further cryptic species and Mediterranean origin of the divergence

Previous genetic analyses have demonstrated that two phonic types of one of the most common European bats, the Common pipistrelle, belong to distinct species, although they are almost identical morphologically (45 kHz Pipistrellus pipistrellus and 55 kHz Pipistrellus pygmaeus). To reconstruct the history of the species complex and explain the codistribution of both forms in Europe and the Mediterranean, we performed phylogenetic analysis based on a 402-bp portion of the cytochrome b gene. Particular attention was paid to the eastern and southern parts of the range where no data were available. We found further distinctive allopatric haplotypes from Libya and Morocco. The difference of about 6-7% described in the Libyan population suggests the occurrence of a new species in the southern Mediterranean. The species status of Moroccan population is also discussed. The phylogeographic patterns obtained and analysis of fossil records support the hypothesis of expansion of both species into Europe from the Mediterranean region during the Holocene. The allopatric speciation model fits our data best. The paleobiographic scenario envisaged is corroborated also by molecular clock estimations and correlations with Late Neogene environmental changes in the Mediterranean region which ended with the Messinian salinity crisis.     

Song learning accelerates allopatric speciation

The songs of many birds are unusual in that they serve a role in identifying conspecific mates, yet they are also culturally transmitted. Noting the apparently high rate of diversity in one avian taxon, the songbirds, in which song learning appears ubiquitous, it has often been speculated that cultural transmission may increase the rate of speciation. Here we examine the possibility that song learning affects the rate of allopatric speciation. We construct a population-genetic model of allopatric divergence that explores the evolution of genes that underlie learning preferences (predispositions to learn some songs over others). We compare this with a model in which mating signals are inherited only genetically. Models are constructed for the cases where songs and preferences are affected by the same or different loci, and we analyze them using analytical local stability analysis combined with simulations of drift and directional sexual selection. Under nearly all conditions examined, song divergence occurs more readily in the learning model than in the nonlearning model. This is a result of reduced frequency-dependent selection in the learning models. Cultural evolution causes males with unusual genotypes to tend to learn from the majority of males around them, and thus develop songs compatible with the majority of the females in the population. Unusual genotypes can therefore be masked by learning. Over a wide range of conditions, learning therefore reduces the waiting time for speciation to occur and can be predicted to accelerate the rate of speciation.     

Proximate sources of marine biodiversity

When temperature and other kinds of barrier divide formerly continuous populations and confine them to more restricted geographical areas, there is an evolutionary reaction that will, over time, result in the formation of endemic species. In such cases, an allopatric speciation process is considered to have taken place because reproductive isolation was caused by physical means instead of by natural selection. In contrast, when populations exist in a very high-diversity area and remain undivided by physical events, they exhibit a tendency to speciate by means of sympatry (or parapatry). This process, sometimes called competitive or ecological speciation, does involve reproductive isolation by means of natural selection. Populations that exist in geographical provinces bounded by physical barriers add to the overall diversity through the production of endemic species. This increase by species packing is relatively slow due to the very gradual tempo of the allopatric speciation process. Populations existing in centres of origin add to the general diversity through the production of species that are dominant in terms of their ability to spread over large parts of the world. It is proposed that such species are usually formed by sympatric speciation, a process that can be c. 20 times faster than species formation by allopatry. It is not suggested that sympatry is exclusive to centres of origin, nor that allopatry is confined to peripheral provinces. Both processes are widespread, but there do appear to be distinctive geographical concentrations. Considering that numbers of widespread species produced by centres of origin may eventually become subdivided by barriers, and thus give rise to descendants by allopatry, it is difficult to say how much of our present species diversity has come from one source or the other. Both speciation by sympatry from centres of origin and speciation by allopatry in peripheral provinces appear to be important sources of marine biodiversity.     

Reproductive character displacement and environmental filtering shape floral variation between sympatric sister taxa

Divergence in reproductive traits between closely related species that co‐occur contributes to speciation by reducing interspecific gene flow. In flowering plants, greater floral divergence in sympatry than allopatry may reflect reproductive character displacement (RCD) by means of divergent pollinator‐mediated selection or mating system evolution. However, environmental filtering (EF) would prevail for floral traits under stronger selection by abiotic factors than pollination, and lead to sympatric taxa being more phenotypically similar. We determine whether floral UV pigmentation and size show signatures of RCD or EF using a biogeographically informed sister taxa comparison. We determine whether 35 sister pairs in the Potentilleae tribe (Rosaceae) are allopatric or sympatric and confirm that sympatric sisters experience more similar bioclimatic conditions, an assumption of the EF hypothesis. We test whether interspecific differences are greater in allopatry or sympatry while accounting for divergence time. For UV pigmentation, sympatric sisters are more phenotypically similar than allopatric ones. For flower size, sympatric sisters show increased divergence with time since speciation but allopatric ones do not. We conclude that floral UV pigmentation shows a signature of EF, whereas flower size shows a signature of RCD. Discordant results between the traits suggest that the dominant selective agent differs between them.     

Canid hybridization: contemporary evolution in human‐modified landscapes

Contemporary evolution through human‐induced hybridization occurs throughout the taxonomic range. Formerly allopatric species appear especially susceptible to hybridization. Consequently, hybridization is expected to be more common in regions with recent sympatry owing to human activity than in areas of historical range overlap. Coyotes ( Canis latrans) and gray wolves ( C. lupus) are historically sympatric in western North America. Following European settlement gray wolf range contracted, whereas coyote range expanded to include eastern North America. Furthermore, wolves with New World (NW) mitochondrial DNA (mtDNA) haplotypes now extend from Manitoba to Québec in Canada and hybridize with gray wolves and coyotes. Using mtDNA and 12 microsatellite markers, we evaluated levels of wolf‐coyote hybridization in regions where coyotes were present (the Canadian Prairies, n = 109 samples) and absent historically (Québec, n = 154). Wolves with NW mtDNA extended from central Saskatchewan (51°N, 69°W) to northeastern Québec (54°N, 108°W). On the Prairies, 6.3% of coyotes and 9.2% of wolves had genetic profiles suggesting wolf‐coyote hybridization. In contrast, 12.6% of coyotes and 37.4% of wolves in Québec had profiles indicating hybrid origin. Wolves with NW and Old World ( C. lupus) mtDNA appear to form integrated populations in both regions. Our results suggest that hybridization is more frequent in historically allopatric populations. Range shifts, now expected across taxa following climate change and other human influence on the environment, might therefore promote contemporary evolution by hybridization.     

The marine East Indies: diversity and speciation

Aim To discuss the impact of new diversity information and to utilize recent findings on modes of speciation in order to clarify the evolutionary significance of the East Indies Triangle. Location The Indo‐Pacific Ocean. Methods Analysis of information on species diversity, distribution patterns and speciation for comparative purposes. Results Information from a broad‐scale survey of Indo‐Pacific fishes has provided strong support for the theory that the East Indies Triangle has been operating as a centre of origin. It has become apparent that more than two‐thirds of the reef fishes inhabiting the Indo‐Pacific are represented in the Triangle. An astounding total of 1111 species, more than are known from the entire tropical Atlantic, were reported from one locality on the small Indonesian island of Flores. New information on speciation modes indicates that the several unique characteristics of the East Indian fauna are probably due to the predominance of competitive (sympatric) speciation. Main conclusions It is proposed that, within the East Indies, the high species diversity, the production of dominant species, and the presence of newly formed species, are due to natural selection being involved in reproductive isolation, the first step in the sympatric speciation process. In contrast, speciation in the peripheral areas is predominately allopatric. Species formed by allopatry are the direct result of barriers to gene flow. In this case, reproductive isolation may be seen as a physical process that does not involve natural selection. Allopatric species formation often takes millions of years, while the sympatric process is generally much faster. Following species formation, dispersal from the East Indies appears to take place according to the centrifugal hypothesis.     

ON THE COYNE AND ORR‐IGIN OF SPECIES: EFFECTS OF INTRINSIC POSTZYGOTIC ISOLATION,ECOLOGICAL DIFFERENTIATION,X CHROMOSOME SIZE,AND SYMPATRY ON DROSOPHILA SPECIATION

Coyne and Orr found that mating discrimination (premating isolation) evolves much faster between sympatric than allopatric Drosophila species pairs. Their meta‐analyses established that this pattern, expected under reinforcement, is common and that Haldane's rule is ubiquitous in Drosophila species divergence. We examine three possible contributors to the reinforcement pattern: intrinsic postzygotic isolation, dichotomized as to whether hybrid males show complete inviability/sterility; host‐plant divergence, as a surrogate for extrinsic postzygotic isolation; and X chromosome size, whether roughly 20% or 40% of the genome is X‐linked. We focus on “young” species pairs with overlapping ranges, contrasted with allopatric pairs. Using alternative criteria for “sympatry” and tests that compare either level of prezygotic isolation in sympatry or frequency of sympatry, we find no statistically significant effects associated with X chromosome size or our coarse quantifications of intrinsic postzygotic isolation or ecological differentiation. Although sympatric speciation seems very rare in animals, the pervasiveness of the reinforcement pattern and the commonness of range overlap for close relatives indicate that speciation in Drosophila is often not purely allopatric. It remains to determine whether increased premating isolation with sympatry results from secondary contact versus parapatric speciation and what drives this pattern.     

Breeding ecology and nest site selection in allopatric mainland Citril Finches Carduelis[citrinella] citrinella and insular Corsican Finches Carduelis[citrinella] corsicanus

The breeding ecology and nest site selection of mainland Citril Finches and insular Corsican Finches have been studied throughout their limited range. For many years both endemic forms were considered to be two sub-species; however, based on evidence from more recent molecular studies they have been split into two species. This study provides data on the variations in breeding ecology and nest site selection in the different sub-populations of these little studied species. A secondary aim was to search out evidence of ecological differentiation between mainland Citril Finches and insular Corsican Finches. We found that the studied sub-populations of both species largely overlapped in breeding ecology. Our data confirms the close similarity of Citril Finches and Corsican Finches, both which are, similar to mountain birds, well adapted to the local habitat conditions of their different mountain systems. Several differences were identified within the studied sub-populations of the two (sub-) species with respect to nest site selection, probably caused by environmental conditions and local predators. One of the main differences between the two species is that Citril Finches breed mainly in half-open conifer forests (especially pine forests), while Corsican Finches breed in the more open scrubby mountains of the Mediterranean islands dominated mainly by the Tree Heath as an adaptation to the different landscapes on the islands. In contrast to Citril Finches, this preference of the Corsican Finches for Tree Heath as nesting plants – even if suitable pines are available – is typical of the species. These behavioural changes result in a niche expansion into open habitats at lower altitudes. We suggest that the observed niche expansion and behavioural variations are not suitable criteria for taxonomic status, a proposal in contrast to that of Sangster [Ibis 142:487–490 (2000)]. We further suggest that the few ecological differences found in this study between the two (sub-)species are the result of the so-called insular syndrome, which includes changes in life history traits such as morphology, demography and behaviour.