Speciation generally involves a three-step process--range expansion, range fragmentation and the development of reproductive isolation between spatially separated populations. Speciation relies on cycling through these three steps and each may limit the rate at which new species form. We estimate phylogenetic relationships among all Himalayan songbirds to ask whether the development of reproductive isolation and ecological competition, both factors that limit range expansions, set an ultimate limit on speciation. Based on a phylogeny for all 358 species distributed along the eastern elevational gradient, here we show that body size and shape differences evolved early in the radiation, with the elevational band occupied by a species evolving later. These results are consistent with competition for niche space limiting species accumulation. Even the elevation dimension seems to be approaching ecological saturation, because the closest relatives both inside the assemblage and elsewhere in the Himalayas are on average separated by more than five million years, which is longer than it generally takes for reproductive isolation to be completed; also, elevational distributions are well explained by resource availability, notably the abundance of arthropods, and not by differences in diversification rates in different elevational zones. Our results imply that speciation rate is ultimately set by niche filling (that is, ecological competition for resources), rather than by the rate of acquisition of reproductive isolation.

The introduction of avian malaria (Plasmodium relictum) to Hawaii has provided a model system for studying the influence of exotic disease on naive host populations. Little is known, however, about the origin or the genetic variation of Hawaii's malaria and traditional classification methods have confounded attempts to place the parasite within a global ecological and evolutionary context. Using fragments of the parasite mitochondrial gene cytochrome b and the nuclear gene dihydrofolate reductase-thymidylate synthase obtained from a global survey of greater than 13000 avian samples, we show that Hawaii's avian malaria, which can cause high mortality and is a major limiting factor for many species of native passerines, represents just one of the numerous lineages composing the morphological parasite species. The single parasite lineage detected in Hawaii exhibits a broad host distribution worldwide and is dominant on several other remote oceanic islands, including Bermuda and Moorea, French Polynesia. The rarity of this lineage in the continental New World and the restriction of closely related lineages to the Old World suggest limitations to the transmission of reproductively isolated parasite groups within the morphological species.

The host specificity of blood parasites recovered from a survey of 527 birds in Cameroon and Gabon was examined at several levels within an evolutionary framework. Unique mitochondrial lineages of Haemoproteus were recovered from an average of 1.3 host species (maximum = 3) and 1.2 host families (maximum = 3) while lineages of Plasmodium were recovered from an average of 2.5 species (maximum = 27) and 1.6 families (maximum = 9). Averaged within genera, lineages of both Plasmodium and Haemoproteus were constrained in their host distribution relative to random expectations. However, while several individual lineages within both genera exhibited significant host constraint, host breadth varied widely among related lineages, particularly within the genus Plasmodium. Several lineages of Plasmodium exhibited extreme generalist host-parasitism strategies while other lineages appeared to have been constrained to certain host families over recent evolutionary history. Sequence data from two nuclear genes recovered from a limited sample of Plasmodium parasites indicated that, at the resolution of this study, inferences regarding host breadth were unlikely to be grossly affected by the use of parasite mitochondrial lineages as a proxy for biological species. The use of divergent host-parasitism strategies among closely related parasite lineages suggests that host range is a relatively labile character. Since host specificity may also influence parasite virulence, these results argue for considering the impact of haematozoa on avian hosts on a lineage-specific basis.

The degree to which haematozoan parasites can exploit a range of vectors and hosts has both ecological and evolutionary implications for their transmission and biogeography. Here we explore the extent to which closely related mosquito species share the same or closely related haematozoan parasites, and examine the overlap in parasite lineages with those isolated from avian hosts, Zosterops species, sampled across the same study sites. Mosquito samples were collected and analysed (14 species, n = 804) from four islands in Vanuatu and the main island of New Caledonia. Using polymerase chain reaction, 15.5% (14/90) of pooled mosquito (thoracic) samples showed positive amplifications. Subsequent phylogenetic analysis of the cytochrome b gene identified four genetically distinct Plasmodium and two Haemoproteus lineages from these samples, five of which were identical to parasite lineages (n = 21) retrieved from the avian hosts. We found that three Plasmodium lineages differing by a maximum of 0.9% sequence divergence were recovered from different species and genera of mosquitoes and two Haemoproteus lineages differing by 4.6% sequence divergence were carried by 10 distantly related (11-21% divergent) mosquito species. These data suggest a lack of both cospeciation and invertebrate host conservatism. Without experimental demonstration of the transmission cycle, it is not possible to establish whether these mosquitoes are the biological vectors of isolated parasite lineages, reflecting a limitation of a purely polymerase chain reaction-based approach. Nonetheless, our results raise the possibility of a new transmission pathway and highlight extensive invertebrate host shifts in an insular mosquito-parasite system.

Tissue samples from 699 birds from three regions of Asia (Myanmar, India, and South Korea) were screened for evidence of infection by avian parasites in the genera Plasmodium and Haemoproteus. Samples were collected from November 1994 to October 2004. We identified 241 infected birds (34.0%). Base-on-sequence data for the cytochrome b gene from 221 positive samples, 34 distinct lineages of Plasmodium, and 41 of Haemoproteus were detected. Parasite diversity was highest in Myanmar followed by India and South Korea. Parasite prevalence differed among regions but not among host families. There were four lineages of Plasmodium and one of Haemoproteus shared between Myanmar and India and only one lineage of Plasmodium shared between Myanmar and South Korea. No lineages were shared between India and South Korea, although an equal number of distinct lineages were recovered from each region. Migratory birds in South Korea and India originate from two different migratory flyways; therefore cross-transmission of parasite lineages may be less likely. India and Myanmar shared more host species and habitat types compared to South Korea. Comparison between low-elevation habitat in India and Myanmar showed a difference in prevalence of haematozoans.

We used screening techniques based on polymerase chain reaction (PCR) to explore the avian hematozoan parasites (Plasmodium spp. and Haemoproteus spp.) of two previously uninvestigated regions of continental South America. Comparisons of tropicalzone Guyana and temperate-zone Uruguay revealed that overall prevalence of Plasmodium and Haemoproteus species detected in a diverse sampling of potential hosts was significantly higher in Guyana. The difference in prevalence between the two geographic zones appears to be attributable to ecological differences rather than taxonomic sampling artifacts. Diversity of hematozoan haplotypes was also higher in Guyana. We found no relationship between hematozoan haplotype and host family sampled within or between regions. We found very few Plasmodium and no Haemoproteus haplotypes shared between the two regions, and evidence of geographic structuring of hematozoan haplotypes between the two regions. We suggest that a lack of hematozoan haplotype transmission between the two regions may be attributable to the migratory patterns of each region's avian hosts.

Aim (1) To describe the species-area relationships among communities of Plasmodium and Haemoproteus parasites in different island populations of the same host genus (Aves: Zosterops).(2) To compare distance-decay relationships (turnover) between parasite communities and those with potential avian and dipteran hosts, which differ with respect to their movement and potential to disperse parasite species over large distances.Location Two archipelagos in the south-west Pacific, Vanuatu and New Caledonia (c. 250 km west of Vanuatu) and its Loyalty Islands, with samples collected from a total of 16 islands of varying sizes (328-16,648 km 2 ).Methods We characterized parasite diversity and distribution via polymerase chain reaction (PCR) from avian (Zosterops) blood samples. Bayesian methods were used to reconstruct the parasite phylogeny. In accordance with recent molecular evidence, we treat distinct mitochondrial DNA lineages as equivalent to species in this study. Path analysis and parasite lineage accumulation curves were used to assess the confounding effect of inadequate sampling on the estimation of parasite richness. Species-area and species-distance relationships were assessed using linear regression: distance-decay relationships were assessed using Mantel tests.Results Birds and mosquito species and Plasmodium lineages exhibited significant species-area relationships. However, Plasmodium lineages showed the weakest 'species-area' relationship; no relationship was found for Haemoproteus lineages. Avian species richness influenced parasite lineage richness more than mosquito species richness did. Within individual avian host species, the species-area relationship of parasites showed differing patterns. Path analysis indicated that sampling effort was unlikely to have a confounding effect on parasite richness. Distance from mainland (isolation effect) showed no effect on parasite richness. Community similarity decayed significantly with distance for avifauna, mosquito fauna and Plasmodium lineages but not for Haemoproteus lineages.Main conclusions Plasmodium lineages and mosquito species fit the power-law model with steeper slopes than found for the avian hosts. The lack of speciesdistance relationship in parasites suggests that other factors, such as the competence of specific vectors and habitat features, may be more important than distance. The decay in similarity with distance suggests that the sampled Plasmodium lineages and their potential hosts were not randomly distributed, but rather exhibited spatially predictable patterns. We discuss these results in the context of the effects that parasite generality may have on distribution patterns.

The success of introduced species is frequently explained by their escape from natural enemies in the introduced region. We tested the enemy release hypothesis with respect to two well studied blood parasite genera (Plasmodium and Haemoproteus) in native and six introduced populations of the common myna Acridotheres tristis. Not all comparisons of introduced populations to the native population were consistent with expectations of the enemy release hypothesis. Native populations show greater overall parasite prevalence than introduced populations, but the lower prevalence in introduced populations is driven by low prevalence in two populations on oceanic islands (Fiji and Hawaii). When these are excluded, prevalence does not differ significantly. We found a similar number of parasite lineages in native populations compared to all introduced populations. Although there is some evidence that common mynas may have carried parasite lineages from native to introduced locations, and also that introduced populations may have become infected with novel parasite lineages, it may be difficult to differentiate between parasites that are native and introduced, because malarial parasite lineages often do not show regional or host specificity.