Dramatic intraspecific differences in migratory routes,stopover sites and wintering areas,revealed using light-level geolocators

Migratory divides are contact zones between breeding populations that use divergent migratory routes and have been described in a variety of species. These divides are of major importance to evolution, ecology and conservation but have been identified using limited band recovery data and/or indirect methods. Data from band recoveries and mitochondrial haplotypes suggested that inland and coastal Swainson''s thrushes (Catharus ustulatus) form a migratory divide in western North America. We attached light-level geolocators to birds at the edges of this contact zone to provide, to our knowledge, the first direct test of a putative divide using data from individual birds over the entire annual cycle. Coastal thrushes migrated along the west coast to Mexico, Guatemala and Honduras. Some of these birds used multiple wintering sites. Inland thrushes migrated across the Rocky Mountains, through central North America to Columbia and Venezuela. These birds migrated longer distances than coastal birds and performed a loop migration, navigating over the Gulf of Mexico in autumn and around this barrier in spring. These findings support the suggestion that divergent migratory behaviour could contribute to reproductive isolation between migrants, advance our understanding of their non-breeding ecology, and are integral to development of detailed conservation strategies for this group.     

The evolutionary forces maintaining a wild polymorphism of Littorina saxatilis: model selection by computer simulations

Two rocky shore ecotypes of Littorina saxatilis from north-west Spain live at different shore levels and habitats and have developed an incomplete reproductive isolation through size assortative mating. The system is regarded as an example of sympatric ecological speciation. Several experiments have indicated that different evolutionary forces (migration, assortative mating and habitat-dependent selection) play a role in maintaining the polymorphism. However, an assessment of the combined contributions of these forces supporting the observed pattern in the wild is absent. A model selection procedure using computer simulations was used to investigate the contribution of the different evolutionary forces towards the maintenance of the polymorphism. The agreement between alternative models and experimental estimates for a number of parameters was quantified by a least square method. The results of the analysis show that the fittest evolutionary model for the observed polymorphism is characterized by a high gene flow, intermediate-high reproductive isolation between ecotypes, and a moderate to strong selection against the nonresident ecotypes on each shore level. In addition, a substantial number of additive loci contributing to the selected trait and a narrow hybrid definition with respect to the phenotype are scenarios that better explain the polymorphism, whereas the ecotype fitnesses at the mid-shore, the level of phenotypic plasticity, and environmental effects are not key parameters.     

Does bill size polymorphism affect courtship song characteristics in the African finch Pyrenestes ostrinus?

Vocal characteristics of bill morphs of the African finch Pyrenestes ostrinus were investigated to see whether there were differences between the morphs that could potentially lead to assortative mating. Morphological differences between bill morphs are of a scale that could change resonance characteristics, with as a physical consequence, a change in acoustic characteristics. Song and variation in acoustic characteristics are very likely to be an important factor in mate choice. We analysed recordings of large and small-billed birds, and measured 11 acoustic characteristics focusing on frequency use and possibly correlated temporal features. In addition, we investigated in more detail the energy distribution within the frequency limits. We found no differences between bill morphs in acoustic characteristics of courtship song. Our findings contrast with other empirical studies which show an impact of the suprasyringeal vocal tract on song output. One possible explanation could be that the morphological changes affect resonance characteristics in multiple ways which do not operate in concert. Beside proximate aspects, we discuss the role of song and the bill polymorphism in the context of sympatric speciation.     

Lacustrine spawning: is this a new reproductive strategy among ‘large’ African cyprinid fishes?

Changes in the gonado‐somatic index and abundance of the different Labeobarbus species in the mouths of four major afferent rivers of Lake Tana, Ethiopia, were monitored monthly during 1999 and 2000. Riverine spawning was characteristic for seven of Lake Tana's 15 contemporary Labeobarbus species. These seven did not show spatial segregation among afferent rivers but significant temporal segregation occurred in aggregating in the river mouths and migrating towards the upstream spawning areas during the breeding season (June–October). Among the eight other species, peak gonad development occurred generally in the same period as in the riverine spawners. These species, however, did not aggregate in the river mouths during the breeding period and were absent from the upstream spawning areas. A derived, novel strategy, lacustrine spawning was hypothesized for these eight Labeobarbus species. This hypothesis was further supported by observations of running female fishes in the littoral zones distant from any of the afferent rivers. This derived strategy is only common among the littoral‐dwelling Labeobarbus species with restricted distribution patterns. At present it is thought that sequential waves of speciation and habitat divergence followed by trophic specialization, shaped the diversity of Lake Tana labeobarbs.     

Genetic and ecological divergence of a monophyletic cichlid species pair under fully sympatric conditions in Lake Ejagham, Cameroon

Although there is mounting evidence that speciation can occur under sympatric conditions, unambiguous examples from nature are rare and it is almost always possible to propose alternative allopatric or parapatric scenarios. To identify an unequivocal case of sympatric speciation it is, therefore, necessary to analyse natural settings where recent monophyletic species flocks have evolved within a small and confined spatial range. We have studied such a case with a cichlid species flock that comprises five Tilapia forms endemic to a tiny lake (Lake Ejagham with a surface area of approximately 0.49 km2) in Western Cameroon. Analysis of mitochondrial D-Loop sequences shows that the flock is very young (approximately 10(4) years) and has originated from an adjacent riverine founder population. We have focused our study on a particular pair of forms within the lake that currently appears to be in the process of speciation. This pair is characterized by an unique breeding colouration and specific morphological aspects, which can serve as synapomorphic characters to prove monophyly. It has differentiated into a large inshore and a small pelagic form, apparently as a response to differential utilization of food resources. Still, breeding and brood care occurs in overlapping areas, both in time and space. Analysis of nuclear gene flow on the basis of microsatellite polymorphisms shows a highly restricted gene flow between the forms, suggesting reproductive isolation between them. This reproductive isolation is apparently achieved by size assortative mating, although occasional mixed pairs can be observed. Our findings are congruent with recent theoretical models for sympatric speciation, which show that differential ecological adaptations in combination with assortative mating could easily lead to speciation in sympatry.     

Clinal variation in morphology and assortative mating in the soldier beetle, Chauliognathus pennsylvanicus (Goleoptera: Cantharidae)

In north Georgia populations of the soldier beetle, Chauliognathus pennsylvanicus , the length of the elytral spot varies clinally. At the southern end of a 200 km cline the distribution of spot length is unimodal with longer spot lengths predominating while at the northern end of the cline the distribution is bimodal but with shorter spot lengths being more frequent. North of the cline only short elytral spot lengths are observed, while the converse is true south of the cline. The strength of assortative mating on the basis of elytral spot length increases from south to north along the cline, resulting in complete pre-mating isolation between short and long spot length morphs at the north end of the cline. Laboratory mate choice tests indicate that assortative mating in the field is not the result of differential timing of activity or microhabitat choice but rather that it represents a real behavioural preference. Individuals from monomorphic populations on either side of the cline do not mate assortatively in the laboratory, indicating that reproductive isolation has evolved on the cline.     

Testing alternative models for sexual isolation in natural populations of Littorina saxatilis: indirect support for by‐product ecological speciation?

Two ecotypes of the rough periwinkle Littorina saxatilis occur at different shore levels, showing assortative mating for size and partial reproductive isolation when they meet at the mid-shore. This system represents a putative case of incomplete speciation in sympatry. Two processes contribute to the assortative mating: morph-specific microhabitat aggregation and mate choice. The estimation of mate choice coefficients in nature and a simulation of the aggregation effects on sexual isolation were used to disentangle these processes as well as to test alternative mechanisms of mate choice. Mate choice significantly increased the frequency of within-morph pairs and significantly decreased the frequency of between-morph pairs, whereas those pairs including at least one hybrid morph mated randomly. These results allow us to reject a discriminant mate choice and support a model of evolution of sexual isolation as a side-effect of size-assortative mating in a context of divergent natural selection for size in the population. This mechanism is more compatible with a model of incomplete by-product ecological speciation, as suggested by previous evidence.     

Site-specific genetic divergence in parallel hybrid zones suggests nonallopatric evolution of reproductive barriers

The evolution of reproductive isolation in the presence of gene flow is supported by theoretical models but rarely by data. Empirical support might be gained from studies of parallel hybrid zones between interbreeding taxa. We analysed gene flow over two hybrid zones separating ecotypes of Littorina saxatilis to test the expectation that neutral genetic markers will show site-specific differences if barriers have evolved in situ. Distinct ecotypes found in contrasting shore habitats are separated by divergent selection and poor dispersal, but hybrid zones appear between them. Swedish islands formed by postglacial uplift 5000 years ago provide opportunities to assess genetic structure in a recently evolved system. Each island houses a discrete population containing subpopulations of different ecotypes. Hybrid zones between ecotypes may be a product of ecological divergence occurring on each island or a consequence of secondary overlap of ecotypes of allopatric origin that have spread among the islands. We used six microsatellite loci to assess gene flow and genetic profiles of hybrid zones on two islands. We found reduced gene flow over both hybrid zones, indicating the presence of local reproductive barriers between ecotypes. Nevertheless, subpopulations of different ecotypes from the same island were genetically more similar to each other than were subpopulations of the same ecotype from different islands. Moreover, neutral genetic traits separating the two ecotypes across hybrid zones were site-specific. This supports a scenario of in situ origin of ecotypes by ecological divergence and nonallopatric evolution of reproductive barriers.     

An analysis of genetic differentiation among assortatively mating Drosophila melanogaster in Zimbabwe

African Drosophila melanogaster populations, and those from Zimbabwe in particular, have attracted much interest recently. African flies differ genetically from 'cosmopolitan' populations and were found to exhibit discriminative mating behaviour against individuals from 'cosmopolitan' populations. It has therefore been proposed that Zimbabwean and some other African populations are in an 'incipient stage of speciation'. However, whether the mating behaviour is an effective barrier against gene flow from other populations, and whether intra-population genetic differentiation has already evolved in sympatry is not known. Here, we took a population-based approach to test whether the well-characterized mating type differences have resulted in a genome-wide differentiation at the population level. Using 122 polymorphic microsatellite loci mapping to the third chromosome, we demonstrate a significant genetic differentiation between Zimbabwean flies differing in their mating behaviour. We also provide evidence to suggest that this difference is unlikely to be attributable to population structure within Zimbabwe. However, the analysis of individual microsatellite loci did not indicate more loci differentiating these two groups than expected by chance. Our data suggest that the 'Z'-'M' mating behaviour is strong enough to result in a small but significant genetic differentiation. Thus, future studies based on a larger population sample of flies characterized for their mating behaviour and using more markers are expected to provide more information on the genetic basis of the mating traits in the Zimbabwe flies.     

Recent speciation between sympatric Tanganyikan cichlid colour morphs

Lake Tanganyika, Africa's oldest lake, harbours an impressive diversity of cichlid fishes. Although diversification in its radiating groups is thought to have been initially rapid, cichlids from Lake Tanganyika show little evidence for ongoing speciation. In contrast, examples of recent divergence among sympatric colour morphs are well known in haplochromine cichlids from Lakes Malawi and Victoria. Here, we report genetic evidence for recent divergence between two sympatric Tanganyikan cichlid colour morphs. These Petrochromis morphs share mitochondrial haplotypes, yet microsatellite loci reveal that their sympatric populations form distinct genetic groups. Nuclear divergence between the two morphs is equivalent to that which arises geographically within one of the morphs over short distances and is substantially smaller than that among other sympatric species in this genus. These patterns suggest that these morphs diverged only recently, yet that barriers to gene flow exist which prevent extensive admixture despite their sympatric distribution. The morphs studied here provide an unusual example of active diversification in Lake Tanganyika's generally ancient cichlid fauna and enable comparisons of speciation processes between Lake Tanganyika and other African lakes.     

Assortative mating among rock-dwelling cichlid fishes supports high estimates of species richness from Lake Malawi

It has been estimated that Lake Malawi, Africa, contains 500–650 endemic species of cichlid fishes, the largest number of vertebrate species endemic to any comparable sized area on the planet. As many of these putative species cannot be distinguished anatomically, these estimates of species richness depend to a great extent on the assumption that sympatrically occurring male colour morphs represent biological species. We have tested this assumption using a combination of behavioural observations of courtship and microsatellite DNA analysis for six putative species of the Pseudotropheus ( Tropheops ) complex and three of the Pseudotropheus ( Maylandia ) complex occurring sympatrically at Nkhata Bay. We were unable to demonstrate assortative courtship for the species pairs Pseudotropheus ( Maylandia ) zebra / P . 'gold zebra' or P. ( Tropheops ) 'band'/ P. ( T .) 'rust' because we were unable to distinguish between the females of these taxa. All other taxa showed clear assortative courtship, except for P. ( T .) 'deep', a deep-water species which was rarely observed. Fixation indices (θ_ST for the infinite allele model, and R _ST for the stepwise mutation model) calculated from six microsatellite DNA loci demonstrated significant deviations from panmixia in all pairwise comparisons of putative species, indicating little or no gene flow between populations. All taxa showed high levels of allelic diversity providing evidence that genetic bottlenecking may have been of limited importance in the speciation process. Assortative mating among taxa differing only in male colouration is supportive of theories that speciation in these fishes has been driven by sexual selection by female choice.