Host phylogeography and beta diversity in avian haemosporidian (Plasmodiidae) assemblages of the Lesser Antilles

1. We estimated the correlation between host phylogeographical structure and beta diversity of avian haemosporidian assemblages of passerine birds to determine the degree to which parasite communities change with host evolution, expressed as genetic divergence between island populations, and we investigated whether differences among islands in the haemosporidia of a particular host species reflect beta diversity in the entire parasite assemblage, beta diversity in vectors, turnover of bird species and/or geographical distance. 2. We used Mantel tests to assess the significance of partial correlations between host nucleotide difference (based on cytochrome b) and haemosporidian (Haemoproteus spp. and Plasmodium spp.) mitochondrial lineage beta diversity within a given host species and between Plasmodium mitochondrial lineage beta diversity and mosquito and bird species beta diversity (or turnover). Three abundant and widespread host species (Tiaris bicolor, Coereba flaveola and Loxigilla noctis/barbadensis) were included in the study. Haemosporidian lineage beta diversity among nine islands was assessed using the Chao-Jaccard, Chao-S?rensen and Morisita-Horn indices of community similarity. Beta diversity indices of mosquito species and turnover of bird species were calculated from data in published records and field guides. 3. In Loxigilla spp., we found a positive correlation with geographical distance and an unexpected negative correlation between haemosporidian beta diversity and host genetic distance. Tiaris bicolor exhibited a significant positive correlation between haemosporidian beta diversity and beta diversity within the entire parasite assemblage. We did not find significant correlations between parasite beta diversity and mosquito beta diversity or bird species turnover. 4. Host phylogeographical structure does not appear to drive within-host beta diversity of haemosporidian lineages. Instead, the array of parasites on one host can reflect the haemosporidian assemblage on other hosts.     

The parasites of Anolis lizards the northern Lesser Antilles

Summary The communities of parasitic helminths from ten species of lizards on seven islands in the Caribbean were examined to ascertain the relative importance of predictable deterministic factors and unpredicatable colonization or extinction events in determining the structure of the parasite community. A simple graphical model of community structure is used as a null model to describe the features of a community that are dependent only upon the size of the host population and features of the life histories of the constituent parasite species. This model predicts that parasite species will exhibit a nested pattern of local and global relative abundance. The observed data correspond fairly well to this pattern. The absences of individual parasite species from communities where they might be expected to be present emphasizes the role of stochastic colonization and extinction events in delineating the constituent members of the community on any island.Statistical analysis of the distribution of parasite species per host illustrates that this pattern is random in habitats where parasite species diversity is low, but decreasingly variable in habitats where more diverse parasite communities occur. Increased parasite diversity also leads to an increase in the proportion of hosts that contain mixed species infections. Comparisons of worm burdens from single and mixed species infections within individual hosts suggest that interactions between parasite species only rarely leads to reduced worm burdens.     

The parasites of Anolis lizards in the northern Lesser Antilles

Summary The helminth communities from ten species of lizard on seven islands in the Caribbean were sampled by collecting one hundred specimens of each species. Nine genera of parasites were identified; these included six nematodes, two digeneans and an acanthocephalan. No relationship was discernible between parasite density or abundance and island area or altitude, although dry islands tend to have fewer species of parasites. Anolis lizards of the bimaculatus and wattsi series share similar parasites with four out of nine species common to both series. The parasite community of lizards on these islands is depauperate with respect to similar surveys on the larger islands of the Greater Antilles.On three of the islands lizards were sub-sampled by collecting from moist woodland and more xeric habitats. These data suggest that differences between habitats are as significant as differences between islands in determining parasite burdens. Worm burdens of the commonest parasite species, T. cubensis, increased monotonically with host body size and no evidence was found to suggest that these parasites affect either host survival or fecundity. The sex-ratio of this species correlated with mean abundance of the parasite, with females the dominant sex on islands or in habitats where the parasite was common. This pattern may reflect haplodiploid sexual determination in this species.     

Host specialization and geographic localization of avian malaria parasites: a regional analysis in the Lesser Antilles

We recovered 26 genetically distinct avian malaria parasite lineages, based on cytochrome b sequences, from a broad survey of terrestrial avifauna of the Lesser Antilles. Here we describe their distributions across host species within a regional biogeographic context. Most parasite lineages were recovered from a few closely related host species. Specialization on one host species and distribution across many hosts were both rare. Geographic patterns of parasite lineages indicated limited dispersal and frequent local extinction. The central islands of the archipelago share similar parasite lineages and patterns of infection. However, the peripheral islands harbor well-differentiated parasite communities, indicating long periods of isolation. Nonetheless, 20 of 26 parasite lineages were recovered from at least one of three other geographic regions, the Greater Antilles, North America, and South America, suggesting rapid dispersal relative to rate of differentiation. Six parasite lineages were restricted to the Lesser Antilles, primarily to endemic host species. Host differences between populations of the same parasite lineage suggest that host preference may evolve more rapidly than mitochondrial gene sequences. Taken together, distributions of avian malarial parasites reveal evidence of coevolution, host switching, extinction, and periodic recolonization events resulting in ecologically dynamic as well as evolutionarily stable patterns of infection.     

Migrant birds disperse haemosporidian parasites and affect their transmission in avian communities

Migration has an important impact on the transmission of pathogens. Migratory birds disperse parasites through their routes and may consequently introduce them to new areas and hosts. Hence, haemosporidian parasites, which are among the most prevalent, diverse and important bird pathogens, are potentially dispersed when infecting migrant hosts. Further, migrant hosts could enhance local parasite prevalence and richness by transporting new parasite strains to new areas. Here, we hypothesize and aim to evaluate if 1) migratory birds spread parasite lineages along their routes, and 2) localities crossed by more migratory birds have greater prevalence and richness of haemosporidians. For the first hypothesis, we tested whether parasite lineages found 1) in both migrants and residents, and 2) only in residents, differ in their frequencies of occurrence among localities. For the second hypothesis, we tested for a relationship among localities between the overall local haemosporidian parasite richness and prevalence, and the proportion of migratory bird individuals present in a locality. We combined a dataset on 13 200 bird samples with additional data from the MalAvi database (total: ~2800 sequenced parasites comprising 675 distinct lineages, from 506 host species and 156 localities) from South America, and used Bayesian multi-level models to test our hypotheses. We demonstrate that parasites shared between resident and migratory species are the most spatially widespread, highlighting the potential of migrants to carry and transmit haemosporidians. Further, the presence of migrants in a locality was negatively related to local parasite richness, but not associated with local prevalence. Here, we confirm that migrants can contribute to parasite dispersal and visiting migrants are present in regions with lower Plasmodium prevalence. Also, we observed their presence might raise Haemoproteus community prevalence. Therefore, we demonstrate migrants enhance pathogens spread and their presence may influence parasite community transmission.     

Island and taxon effects in parasitism revisited: avian malaria in the Lesser Antilles

We identify and describe the distribution of 12 genetically distinct malaria parasite lineages over islands and hosts in four common passerine birds in the Lesser Antilles. Combined parasite prevalence demonstrates strong host effects, little or no island effect, and a significant host-times-island interaction, indicating independent outcomes of host-parasite infections among island populations of the same host species. Host- and/or island-specific parasite lineages do not explain these host-parasite associations; rather, individual lineages themselves demonstrate the same type of independent interactions. Unlike overall prevalence, individual parasite lineages show considerable geographic structure (i.e., island effects) as well as species effects indicating that parasite lineages are constrained in their ability to move between hosts and locations. Together, our results suggest an upper limit to the number of host individuals that malaria parasites, as a community, can infect. Within this limit, however, the relative frequency of the different lineages varies reflecting fine scale interactions between host and parasite populations. Patterns of host-parasite associations within this system suggest both historical co-evolution and ecologically dynamic and independent host-parasite interactions.     

Prevalence of avian haemosporidian parasites and their host fidelity in the central Philippine islands

We examined the prevalence and host fidelity of avian haemosporidian parasites belonging to the genera Haemoproteus, Leucocytozoon and Plasmodium in the central Philippine islands by sampling 23 bird families (42 species). Using species-specific PCR assays of the mitochondrial cytochrome b gene (471base pairs, bp), we detected infections in 91 of the 215 screened individuals (42%). We also discriminated between single and multiple infections. Thirty-one infected individuals harbored a single Haemoproteus lineage (14%), 18 a single Leucocytozoon lineage (8%) and 12 a single Plasmodium lineage (6%). Of the 215 screened birds, 30 (14%) presented different types of multiple infections. Intrageneric mixed infections were generally more common (18 Haemoproteus/Haemoproteus, 3 Leucocytozoon/Leucocytozoon, and 1 Plasmodium/Plasmodium) than intergeneric mixed infections (7 Haemoproteus/Leucocytozoon and 1 Haemoproteus/Leucocytozoon/Plasmodium). We recovered 81 unique haemosporidian mitochondrial haplotypes. These clustered in three strongly supported monophyletic clades that correspond to the three haemosporidian genera. Related lineages of Haemoproteus and Leucocytozoon were more likely to derive from the same host family than predicted by chance; however, this was not the case for Plasmodium. These results indicate that switches between host families are more likely to occur in Plasmodium. We conclude that Haemoproteus has undergone a recent diversification across well-supported host-family specific clades, while Leucocytozoon shows a longer association with its host(s). This study supports previous evidence of a higher prevalence and stronger host-family specificity of Haemoproteus and Leucocytozoon compared to Plasmodium.     

Migratory birds have higher prevalence and richness of avian haemosporidian parasites than residents

Individuals of migratory species may be more likely to become infected by parasites because they cross different regions along their route, thereby being exposed to a wider range of parasites during their annual cycle. Conversely, migration may have a protective effect since migratory behaviour allows hosts to escape environments presenting a high risk of infection. Haemosporidians are one of the best studied, most prevalent and diverse groups of avian parasites, however the impact of avian host migration on infection by these parasites remains controversial. We tested whether migratory behaviour influenced the prevalence and richness of avian haemosporidian parasites among South American birds. We used a dataset comprising ~ 11,000 bird blood samples representing 260 bird species from 63 localities and Bayesian multi-level models to test the impact of migratory behaviour on prevalence and lineage richness of two avian haemosporidian genera (Plasmodium and Haemoproteus). We found that fully migratory species present higher parasite prevalence and higher richness of haemosporidian lineages. However, we found no difference between migratory and non-migratory species when evaluating prevalence separately for Plasmodium and Haemoproteus, or for the richness of Plasmodium lineages. Nevertheless, our results indicate that migratory behaviour is associated with an infection cost, namely a higher prevalence and greater variety of haemosporidian parasites.     

Phylofloristics: an example from the Lesser Antilles

Aims The last decade has seen many plant ecologists integrating phylogenetic analysis into ecology to explain patterns of species co-occurrence and compositional similarity across assemblages. Despite the advances in this area, there are still some challenges that need to be addressed. One challenge is that most of the phylogenetic studies of plant assemblages have focused only on a small proportion of all of the vascular plants that co-occur (e.g. woody plants), while much of the remaining co-occurring flora has been ignored.Methods Here we introduce an analytical approach that we term phylofloristics that analyzes the compositional similarity of floras in relation to spatial and environmental gradients to understand their assembly. As an illustration, we assembled a large phylogenetic tree for the flora of the Lesser Antilles and evaluated the patterns of floristic and phylofloristic similarity among the island-specific floras. We analyzed the relationship of these similarities with spatial and environmental distance and compared the results for non-endemic and endemic lineages.Important findings The results show a major influence of environmental heterogeneity on the assembly of island floras and far less evidence for the importance of dispersal limitation of lineages and species. This study highlights the importance of incorporating broader taxonomic sampling to improve our understanding of assembly processes in ecology. We expect future phylofloristic studies will improve the approach we have taken by generating more refined phylogenetic trees and by incorporating phylogeographic information.     

Food production and population size in the Lesser Antilles

This article poses the question of whether aboriginal population size in the Lesser Antilles was limited by the total calories obtainable from manioc, the amount of protein obtainable from animal sources, or both of these factors. Estimates of cultivable land and productivity show that calories obtainable from manoic are not likely to have limited population size. Estimates of the amount of animal food that would have been needed to provide sufficient protein for populations of varying size, based on studies of faunal remains, indicate that protein is more likely than calories to have been a limiting factor in population size because the amount of animal protein required to support a population of several thousand is unlikely to have been consistently obtainable. A method is suggested for estimating the maximum human population supportable by animal protein resources found on Antillean islands.     

The prevalence of avian haemosporidian parasites in an invasive bird is lower in urban than in non-urban environments

Urbanization affects the ecology of wildlife diseases and although it has been suggested that there are lower risks of infection in urban areas, there have been no experiments to support this conclusion. We assessed haemosporidian prevalence and intensity in House Sparrows Passer domesticus using field and experimental data under contrasting conditions (i.e. urban vs. non-urban). For experimental data, we kept 32 male House Sparrows in captivity as a proxy of stress, and for field data we sampled 49 House Sparrows (17 females). We made use of microscopy to determine the relative intensity and used the polymerase chain reaction to estimate infection prevalence. We obtained total leucocyte counts, leucocyte differentials, heterophil/lymphocyte ratio (H/L) as a measure of stress, and the Polychromatic Index as a measure of physiological condition (erythropoiesis). We identified a total of 10 haemosporidian lineages. For field samples (both males and females), we found a significantly higher prevalence of infection in non-urban House Sparrows than in urban ones. Under experimental conditions, non-urban House Sparrows showed a higher prevalence than urban ones both before and after captivity, with a significant increase in parasite intensity. The number of infected birds increased after captivity for both urban (~ 32%) and non-urban House Sparrows (~ 19%), indicating either a recrudescence of chronic and relapses of latent infections or the appearance of infections that had been acquired earlier. The H/L ratio was significantly higher for non-urban than for urban male House Sparrows before captivity. No difference in H/L was found for urban House Sparrows before and after captivity, indicating tolerance to city stressors. Our results showed a significant decrease in H/L for non-urban birds after captivity, suggesting higher stress in the non-urban agricultural environment. Haemosporidian infections were not associated with the H/L ratio. Our study provides evidence that highly urbanized areas within cities represent lower haemosporidian infection risks than do non-urban areas for House Sparrows.     

The distribution of seagrasses in Dominica, Lesser Antilles

Seagrass beds are the largest organism-built marine habitat in Dominica, yet have only been surveyed since 2007. Standardized examinations along a depth gradient between 0 and 24 m, focusing on magnoliophyte species composition and benthic cover of shoots at 17 seagrass bed sites, were carried out between September 10 and December 7, 2008. The Cymodoceaceae Syringodium filiforme (Kuetzing 1860) and Halodule wrightii (Ascherson 1868), as well as the Hydrocharitaceae Halophila decipiens (Ostenfeld 1902), H. stipulacea (Fosskal & Ascherson 1867) and Thalassia testudinum (Banks ex K?nig 1805) displayed distinct regional and horizontal distribution patterns. Syringodium filiforme is the island's dominant seagrass along the western and northern coasts, occurring at depths between 2 and 18 m and with a mean benthic cover ranging from 0.9-10% along the West coast. Along the North coast it grew between 0.2 and 1 m depth with a mean maximum benthic cover of 48.9%. Halodule wrightii grew along the North and West coasts, in depths between 1 and 14m in areas of recent and chronic disturbances. Its delicate morphology and sparse benthic cover (< 0.1%) did not constitute seagrass beds. Halophila decipiens grew along the deep, shallow and lateral margins of west coast S. filiforme beds and monospecifically in depths between 3 and 24m. Halophila stipulacea, an invasive species, was widespread along 45km of the West coast and was found in depths between 5 and 24m. Both Halophila species formed extensive beds at depths beyond the survey limit of 24m thus playing a potentially important role in the resettlement of shallow areas after storms. H. decipiens and H. stipulacea are currently the second and third most common seagrasses on the island respectively, despite their absence along the North coast. T. testudinum was confined to North coast's sheltered reef flats at depths Im or less with mean a benthic cover ranging from 2 to 76%. It grew monospecifically in the most turbulent and in the calmest locations, yet intermixed with S. filiforme in areas of moderate turbulence. Strong surge along the West coast (October 15-16, 2008), associated with Hurricane Omar, caused uprooting and burial of seagrass beds in varying degrees, in particular along the shallow margins between 2 and 10m depth. This event also demonstrated the dynamic nature of Dominica's shallow seagrass bed margins and the resistance level of individual beds to storm disturbances.     

The Onychophora of Trinidad, Tobago and the Lesser Antilles

This paper brings together the scattered literature on the Onychophora from Trinidad, Tobago and the Lesser Antilles. The 11 known species from this area are described, incorporating new material collected by the author. Scanning electron micrographs of the dorsal integument are presented for all but one species. Parthenogenesis is reported in a population of Epiperipatus imthurni from Trinidad.     

Host specificity of avian haemosporidian parasites is unrelated among sister lineages but shows phylogenetic signal across larger clades

Parasites can vary in the number of host species they infect, a trait known as “host specificity”. Here we quantify phylogenetic signal—the tendency for closely related species to resemble each other more than distantly related species—in host specificity of avian haemosporidian parasites (genera Plasmodium, Haemoproteus and Leucocytozoon) using data from MalAvi, the global avian haemosporidian database. We used the genetic data (479 base pairs of cytochrome b) that define parasite lineages to produce genus level phylogenies. Combining host specificity data with those phylogenies revealed significant levels of phylogenetic signal while controlling for sampling effects; phylogenetic signal was higher when the phylogenetic diversity of hosts was taken into account. We then tested for correlations in the host specificity of pairs of sister lineages. Correlations were generally close to zero for all three parasite genera. These results suggest that while the host specificity of parasite sister lineages differ, larger clades may be relatively specialised or generalised.     

Fibropapillomatosis Prevalence and Distribution in Immature Green Turtles (Chelonia mydas) in Martinique Island (Lesser Antilles)

Fibropapillomatosis (FP) threatens the survival of green turtle (Chelonia mydas) populations at a global scale, and human activities are regularly pointed as causes of high FP prevalence. However, the association of ecological factors with the disease’s severity in complex coastal systems has not been well established and requires further studies. Based on a set of 405 individuals caught over ten years, this preliminary study provides the first insight of FP in Martinique Island, which is a critical development area for immature green turtles. Our main results are: (i) 12.8% of the individuals were affected by FP, (ii) FP has different prevalence and temporal evolution between very close sites, (iii) green turtles are more frequently affected on the upper body part such as eyes (41.4%), fore flippers (21.9%), and the neck (9.4%), and (iv) high densities of individuals are observed on restricted areas. We hypothesise that turtle’s aggregation enhances horizontal transmission of the disease. FP could represent a risk for immature green turtles’ survival in the French West Indies, a critical development area, which replenishes the entire Atlantic population. Continuing scientific monitoring is required to identify which factors are implicated in this panzootic disease and ensure the conservation of the green turtle at an international scale.

     

Shell-Filled Burrows in the Upper Oligocene Antigua Formation,Antigua, Lesser Antilles

Limestones of the Upper Oligocene Antigua Formation of Antigua contain unusual burrows filled with the densely packed debris of shelly benthos. Unlined burrows (Planolites?) in deep-water biofacies at Half Moon Bay, parish of Saint Philip, are packed with a monospecific assemblage of large benthic foraminiferans (Lepidocyclina canelli Lemoine &; Douvillé) and a single brachiopod valve, Tichosina sp. A similar burrow in shallower-water biofacies at Hughes Point, parish of Saint Philip, is packed with echinoderm debris and, particularly, test fragments of the spatangoid echinoid Lovenia n. sp. Fragments of the same echinoid fill a conical burrow, Bergaueria isp. The latter is probably a physical accumulation, the common fragments of echinoid being washed into an empty burrow. In contrast, the infill of Planolites? isp. is more likely to be an accumulation mediated by the burrower.