Immature hard ticks infected with Rickettsia amblyommatis on breeding birds from Pantanal.

Immature hard ticks from the genus Amblyomma feed on blood from a wide range of Neotropical avian hosts. They serve as vectors for pathogens of medical and veterinary importance, such as Rickettsia agents of the spotted fever group (SFG). Hence, determining ecological factors that increase encounter rates between immature ticks and their avian hosts may contribute to the understanding of tick-borne diseases transmission. Here, we used 720 individual birds from 96 species surveyed in the Brazilian Pantanal to test whether host breeding season influenced tick infestation probabilities. Additionally, collected ticks were screened for Rickettsia agents to describe new avian-tick-bacteria associations. Our models revealed that the probability of an individual bird being infested with immature ticks was similar during the breeding and pre-breeding season, but higher loads of immature tick stages were found during the breeding season. Host sex did not predict infestation probability, but Rickettsia agents recovered from ticks were more prevalent during the pre-breeding season. The new records of host usage by larvae and nymphs of Amblyomma in Pantanal and the growing body of tick surveys in Neotropical avian communities, suggest that immature ticks may benefit from avian blood sources during their annual cycle. The low number of infected ticks with Rickettsia agents on Pantanal birds suggest that this vertebrate group are likely not acting as reservoirs for these microorganisms. However, long-term surveys at the same site are imperative to determine which tick species are acting as reservoirs for Rickettsia agents in Pantanal and determine whether birds are playing a role in dispersing ticks and tick-borne pathogens.

Beta diversity, prevalence, and specificity of avian haemosporidian parasites throughout the annual cycle of Chilean Elaenia (Elaenia chilensis), a Neotropical austral migrant.

Migratory birds are implicated in dispersing haemosporidian parasites over great geographic distances. However, their role in sharing these vector-transmitted blood parasites with resident avian host species along their migration flyway is not well understood. We studied avian haemosporidian parasites in 10 localities where Chilean Elaenia, a long-distance Neotropical austral migrant species, spends part of its annual cycle to determine local parasite transmission among resident sympatric host species in the elaenia's distributional range across South America. We sampled 371 Chilean Elaenias and 1,818 birds representing 243 additional sympatric species from Brazilian wintering grounds to Argentinian breeding grounds. The 23 haemosporidian lineages found in Chilean Elaenias exhibited considerable variation in distribution, specialization, and turnover across the 10 avian communities in South America. Parasite lineage dissimilarity increased with geographic distance, and infection probability by Parahaemoproteus decreased in localities harbouring a more diverse haemosporidian fauna. Furthermore, blood smears from migrating Chilean Elaenias and local resident avian host species did not contain infective stages of Leucocytozoon, suggesting that transmission did not take place in the Brazilian stopover site. Our analyses confirm that this Neotropical austral migrant connects avian host communities and transports haemosporidian parasites along its distributional range in South America. However, the lack of transmissive stages at stopover site and the infrequent parasite lineage sharing between migratory host populations and residents at breeding and wintering grounds suggest that Chilean Elaenias do not play a significant role in dispersing haemosporidian parasites, nor do they influence local transmission across South America.

Host foraging behavior and nest type influence prevalence of avian haemosporidian parasites in the Pantanal.

Avian haemosporidians from the genera Plasmodium and Haemoproteus are vector transmitted parasites. A growing body of evidence suggests that variation in their prevalence within avian communities is correlated with a variety of avian ecological traits. Here, we examine the relationship between infection probability and diversity of haemosporidian lineages and avian host ecological traits (average body mass, foraging stratum, migratory behavior, and nest type). We used molecular methods to detect haemosporidian parasites in blood samples from 642 individual birds of 149 species surveyed at four localities in the Brazilian Pantanal. Based on cytochrome b sequences, we recovered 28 lineages of Plasmodium and 17 of Haemoproteus from 31 infected avian species. Variation in lineage diversity among bird species was not explained by avian ecological traits. Prevalence was heterogenous across avian hosts. Bird species that forage near the ground were less likely to be infected by Haemoproteus, whereas birds that build open cup nests were more likely infected by Haemoproteus. Furthermore, birds foraging in multiple strata were more likely to be infected by Plasmodium. Two other ecological traits, often related to host resistance (body mass and migratory behavior), did not predict infection probability among birds sampled in the Pantanal. Our results suggest that avian host traits are less important determinants of haemosporidian diversity in Pantanal than in other regions, but reinforces that host attributes, related to vector exposure, are to some extent important in modulating infection probability within an avian host assemblage.

Tropical deforestation induces thresholds of reproductive viability and habitat suitability in Earth's largest eagles.

Apex predators are threatened globally, and their local extinctions are often driven by failures in sustaining prey acquisition under contexts of severe prey scarcity. The harpy eagle Harpia harpyja is Earth's largest eagle and the apex aerial predator of Amazonian forests, but no previous study has examined the impact of forest loss on their feeding ecology. We monitored 16 active harpy eagle nests embedded within landscapes that had experienced 0 to 85% of forest loss, and identified 306 captured prey items. Harpy eagles could not switch to open-habitat prey in deforested habitats, and retained a diet based on canopy vertebrates even in deforested landscapes. Feeding rates decreased with forest loss, with three fledged individuals dying of starvation in landscapes that succumbed to 50-70% deforestation. Because landscapes deforested by > 70% supported no nests, and eaglets could not be provisioned to independence within landscapes > 50% forest loss, we established a 50% forest cover threshold for the reproductive viability of harpy eagle pairs. Our scaling-up estimate indicates that 35% of the entire 428,800-km2 Amazonian 'Arc of Deforestation' study region cannot support breeding harpy eagle populations. Our results suggest that restoring harpy eagle population viability within highly fragmented forest landscapes critically depends on decisive forest conservation action.

Host movement and time of year influence tick parasitism in Pantanal birds.

Ticks are among the best studied parasitic groups as they spread important pathogens of medical and veterinary importance worldwide. Migratory birds can play an important role in transporting ticks infected with pathogens across wide geographic regions. It is therefore important to understand which factors promote tick parasitism rates across their avian hosts and the associated potential for disease spread. Here, we identified the host attributes of infestation probability of ticks from the genus Amblyomma in 955 birds from Pantanal, Brazil. Infestation rates exhibited considerable variation across the 129 avian species surveyed and were explained by both host ecological traits and evolutionary history. The probability of an individual bird being infested with immature ticks (larvae and/or nymphs) was higher across resident bird species that forage at ground level and during the wet season. Bird species that feed on vertebrates were less likely to be infested by ticks. Other ecological traits known to promote tick exposure (age, body mass, social behavior, and sex) did not predict infestation probability. Our findings demonstrate that tick occurrence in Pantanal birds is determined by avian host attributes, but tick occurrence throughout the year constrains exposure to host-seeking ticks. Moreover, the ecology of the avian host might prevent the potential spread of tick-borne diseases outside Pantanal as migratory hosts are likely less infested by ticks.

Low host specificity and lack of parasite avoidance by immature ticks in Brazilian birds.

Ticks are ectoparasites that feed on blood of a broad taxonomic range of terrestrial and flying vertebrates and are distributed across a wide range of environmental conditions. Here, we explore the biotic and abiotic factors on infestation probability of ticks of the genus Amblyomma and assess the degree of host specificity based on analysis of 1028 birds surveyed across Brazil. We show that tick infestation rates exhibited considerable variation across the 235 avian species analyzed and that the probability of an individual bird being parasitized by immature ticks (larvae and nymphs) increased with annual precipitation. Host phylogeny and two host ecological traits known to promote tick exposure (body mass and foraging behavior) did not predict infestation probability. Moreover, immature ticks displayed a low degree of host specificity at the family level. Lastly, tick occurrence in birds carrying infection with avian malaria and related parasites did not differ from those free of these haemosporidian parasites, indicating a lack of parasite avoidance by immature ticks. Our findings demonstrate that tick occurrence in birds across Brazilian biomes responds to environmental factors rather than ecological and evolutionary host attributes.

An inverse latitudinal gradient in infection probability and phylogenetic diversity for Leucocytozoon blood parasites in New World birds.

Geographic variation in environmental conditions as well as host traits that promote parasite transmission may impact infection rates and community assembly of vector-transmitted parasites. Identifying the ecological, environmental and historical determinants of parasite distributions and diversity is therefore necessary to understand disease outbreaks under changing environments. Here, we identified the predictors and contributions of infection probability and phylogenetic diversity of Leucocytozoon (an avian blood parasite) at site and species levels across the New World. To explore spatial patterns in infection probability and lineage diversity for Leucocytozoon parasites, we surveyed 69 bird communities from Alaska to Patagonia. Using phylogenetic Bayesian hierarchical models and high-resolution satellite remote-sensing data, we determined the relative influence of climate, landscape, geography and host phylogeny on regional parasite community assembly. Infection rates and parasite diversity exhibited considerable variation across regions in the Americas. In opposition to the latitudinal gradient hypothesis, both the diversity and prevalence of Leucocytozoon parasites decreased towards the equator. Host relatedness and traits known to promote vector exposure neither predicted infection probability nor parasite diversity. Instead, the probability of a bird being infected with Leucocytozoon increased with increasing vegetation cover (NDVI) and moisture levels (NDWI), whereas the diversity of parasite lineages decreased with increasing NDVI. Infection rates and parasite diversity also tended to be higher in cooler regions and higher latitudes. Whereas temperature partially constrains Leucocytozoon diversity and infection rates, landscape features, such as vegetation cover and water body availability, play a significant role in modulating the probability of a bird being infected. This suggests that, for Leucocytozoon, the barriers to host shifting and parasite host range expansion are jointly determined by environmental filtering and landscape, but not by host phylogeny. Our results show that integrating host traits, host ancestry, bioclimatic data and microhabitat characteristics that are important for vector reproduction are imperative to understand and predict infection prevalence and diversity of vector-transmitted parasites. Unlike other vector-transmitted diseases, our results show that Leucocytozoon diversity and prevalence will likely decrease with warming temperatures.

Environmental structure affects taxonomic diversity but not functional structure of understory birds in the southwestern Brazilian Amazon

Changes in environmental characteristics can affect habitat use by birds, impacting the number of individuals, number of species, and changing species composition and functional structure of assemblages. Metrics that evaluate the functional structure of biological assemblages constitute a complementary tool to the traditional taxonomic approach, because they quantify the differences between species by means of functional traits. We assessed the effect of environmental characteristics on the taxonomic diversity (species richness, species composition and number of individuals) and functional structure (functional richness, functional evenness, Rao's functional diversity, and community-weighted mean traits) of bird assemblages in northern Mato Grosso state, in the southwestern Brazilian Amazon. We sampled birds in 32 plots. At each plot birds were captured using mist nets, and eight environmental variables were measured: canopy openness, leaf litter, elevation, number of trees in three classes of DBH, soil clay content, and distance to nearest stream. To evaluate functional structure, we measured seven morphological traits from individuals of each bird species. Habitat variables had a significant effect on taxonomic diversity. However, the general functional structure was not affected. Elevation and distance to nearest stream were the main variables driving changes in taxonomic diversity and had a minor effect on functional richness. The other metrics of functional structure were not significantly affected by the set of environmental variables. Our results suggest that the sampled bird assemblages exhibit some functional tolerance (redundancy) to small-scale environmental variation, implying certain resilience to ecosystem modification. (AU)

Avian host composition, local speciation and dispersal drive the regional assembly of avian malaria parasites in South American birds.

Identifying the ecological factors that shape parasite distributions remains a central goal in disease ecology. These factors include dispersal capability, environmental filters and geographic distance. Using 520 haemosporidian parasite genetic lineages recovered from 7,534 birds sampled across tropical and temperate South America, we tested (a) the latitudinal diversity gradient hypothesis and (b) the distance-decay relationship (decreasing proportion of shared species between communities with increasing geographic distance) for this host-parasite system. We then inferred the biogeographic processes influencing the diversity and distributions of this cosmopolitan group of parasites across South America. We found support for a latitudinal gradient in diversity for avian haemosporidian parasites, potentially mediated through higher avian host diversity towards the equator. Parasite similarity was correlated with climate similarity, geographic distance and host composition. Local diversification in Amazonian lineages followed by dispersal was the most frequent biogeographic events reconstructed for haemosporidian parasites. Combining macroecological patterns and biogeographic processes, our study reveals that haemosporidian parasites are capable of circumventing geographic barriers and dispersing across biomes, although constrained by environmental filtering. The contemporary diversity and distributions of haemosporidian parasites are mainly driven by historical (speciation) and ecological (dispersal) processes, whereas the parasite community assembly is largely governed by host composition and to a lesser extent by environmental conditions.

Host associations and turnover of haemosporidian parasites in manakins (Aves: Pipridae).

Parasites of the genera Plasmodium and Haemoproteus (Apicomplexa: Haemosporida) are a diverse group of pathogens that infect birds nearly worldwide. Despite their ubiquity, the ecological and evolutionary factors that shape the diversity and distribution of these protozoan parasites among avian communities and geographic regions are poorly understood. Based on a survey throughout the Neotropics of the haemosporidian parasites infecting manakins (Pipridae), a family of Passerine birds endemic to this region, we asked whether host relatedness, ecological similarity and geographic proximity structure parasite turnover between manakin species and local manakin assemblages. We used molecular methods to screen 1343 individuals of 30 manakin species for the presence of parasites. We found no significant correlations between manakin parasite lineage turnover and both manakin species turnover and geographic distance. Climate differences, species turnover in the larger bird community and parasite lineage turnover in non-manakin hosts did not correlate with manakin parasite lineage turnover. We also found no evidence that manakin parasite lineage turnover among host species correlates with range overlap and genetic divergence among hosts. Our analyses indicate that host switching (turnover among host species) and dispersal (turnover among locations) of haemosporidian parasites in manakins are not constrained at this scale.

Rickettsial infection in ticks from wild birds from Cerrado and the Pantanal region of Mato Grosso, midwestern Brazil.

The involvement of different species of ticks and wild animals, such as birds, play an important role in the epidemiology of tick-borne diseases. Birds may serve as reservoirs for some tick-borne diseases, and may carry and spread hematophagous ectoparasites mechanically. This study aimed to show the diversity of ticks on birds and molecular detection of rickettsial infection in ticks from Pantanal and Cerrado, two similar Brazilian biomes characterized by hydrological seasons. During two years, August 2012 to May 2014, ticks were collected from birds and from the environment in total of 14 visits for collecting samples, distributed in all hydrological seasons. A total of 674 birds were captured representing 113 species from 26 families. In total, 71 birds were parasitized (10.5%), and 155 ticks of the following 7 tick species (in decreasing order of prevalence) were identified: Amblyomma longirostre Koch, Amblyomma nodosum Neumann, Amblyomma cajennense Fabricius sensu lato (s.l.), Amblyomma calcaratum Neumann, Ornithodoros mimon Kohls, Clifford & Jones, Amblyomma ovale Koch, and Amblyomma triste Koch. Among free-living ticks collected in the environment, A. cajennense s.l. was the most common. This is the first occurrence of O. mimon on birds, and of A. triste on Passeriformes in Brazil. Molecular analyses revealed that 6 A. longirostre ticks were infected by 'Candidatus Rickettsia amblyommii', whereas 1 A. nodosum was infected by a Rickettsia parkeri-like agent, previously reported as Rickettsia sp. strain NOD. Spotted fever group (SFG) agents were, for the first time, reported in ticks from birds captured in Pantanal biome, and the potential involvement of these agents as human pathogens should be considered in further studies.