Climate change in the Arctic: Testing the poleward expansion of ticks and tick-borne diseases.

Climate change is most strongly felt in the polar regions of the world, with significant impacts on the species that live there. The arrival of parasites and pathogens from more temperate areas may become a significant problem for these populations, but current observations of parasite presence often lack a historical reference of prior absence. Observations in the high Arctic of the seabird tick Ixodes uriae suggested that this species expanded poleward in the last two decades in relation to climate change. As this tick can have a direct impact on the breeding success of its seabird hosts and vectors several pathogens, including Lyme disease spirochaetes, understanding its invasion dynamics is essential for predicting its impact on polar seabird populations. Here, we use population genetic data and host serology to test the hypothesis that I. uriae recently expanded into Svalbard. Both black-legged kittiwakes (Rissa tridactyla) and thick-billed murres (Uria lomvia) were sampled for ticks and blood in Kongsfjorden, Spitsbergen. Ticks were genotyped using microsatellite markers and population genetic analyses were performed using data from 14 reference populations from across the tick's northern distribution. In contrast to predictions, the Spitsbergen population showed high genetic diversity and significant differentiation from reference populations, suggesting long-term isolation. Host serology also demonstrated a high exposure rate to Lyme disease spirochaetes (Bbsl). Targeted PCR and sequencing confirmed the presence of Borrelia garinii in a Spitsbergen tick, demonstrating the presence of Lyme disease bacteria in the high Arctic for the first time. Taken together, results contradict the notion that I. uriae has recently expanded into the high Arctic. Rather, this tick has likely been present for some time, maintaining relatively high population sizes and an endemic transmission cycle of Bbsl. Close future observations of population infestation/infection rates will now be necessary to relate epidemiological changes to ongoing climate modifications.

Evolutionary Genetics of Borrelia.

The genus Borrelia consists of evolutionarily and genetically diverse bacterial species that cause a variety of diseases in humans and domestic animals. These vector-borne spirochetes can be classified into two major evolutionary groups, the Lyme borreliosis clade and the relapsing fever clade, both of which have complex transmission cycles during which they interact with multiple host species and arthropod vectors. Molecular, ecological, and evolutionary studies have each provided significant contributions towards our understanding of the natural history, biology and evolutionary genetics of Borrelia species; however, integration of these studies is required to identify the evolutionary causes and consequences of the genetic variation within and among Borrelia species. For example, molecular and genetic studies have identified the adaptations that maximize fitness components throughout the Borrelia lifecycle and enhance transmission efficacy but provide limited insights into the evolutionary pressures that have produced them. Ecological studies can identify interactions between Borrelia species and the vertebrate hosts and arthropod vectors they encounter and the resulting impact on the geographic distribution and abundance of spirochetes but not the genetic or molecular basis underlying these interactions. In this review we discuss recent findings on the evolutionary genetics from both of the evolutionarily distinct clades of Borrelia species. We focus on connecting molecular interactions to the ecological processes that have driven the evolution and diversification of Borrelia species in order to understand the current distribution of genetic and molecular variation within and between Borrelia species.

Changes in Bacterial Diversity, Composition and Interactions During the Development of the Seabird Tick Ornithodoros maritimus (Argasidae).

Characterising within-host microbial interactions is essential to understand the drivers that shape these interactions and their consequences for host ecology and evolution. Here, we examined the bacterial microbiota hosted by the seabird soft tick Ornithodoros maritimus (Argasidae) in order to uncover bacterial interactions within ticks and how these interactions change over tick development. Bacterial communities were characterised through next-generation sequencing of the V3-V4 hypervariable region of the bacterial 16S ribosomal RNA gene. Bacterial co-occurrence and co-exclusion were determined by analysing networks generated from the metagenomic data obtained at each life stage. Overall, the microbiota of O. maritimus was dominated by four bacterial genera, namely Coxiella, Rickettsia, Brevibacterium and Arsenophonus, representing almost 60% of the reads. Bacterial diversity increased over tick development, and adult male ticks showed higher diversity than did adult female ticks. Bacterial networks showed that co-occurrence was more frequent than co-exclusion and highlighted substantial shifts across tick life stages; interaction networks changed from one stage to the next with a steady increase in the number of interactions through development. Although many bacterial interactions appeared unstable across life stages, some were maintained throughout development and were found in both sexes, such as Coxiella and Arsenophonus. Our data support the existence of a few stable interactions in O. maritimus ticks, on top of which bacterial taxa accumulate from hosts and/or the environment during development. We propose that stable associations delineate core microbial interactions, which are likely to be responsible for key biological functions.

A high gene flow in populations of Amblyomma ovale ticks found in distinct fragments of Brazilian Atlantic rainforest.

The genetic structure of populations of the tick Amblyomma ovale from five distinct areas of the Brazilian Atlantic rainforest was evaluated via DNA sequencing and associated with the presence of domestic dogs acting as hosts at the edge of forest fragments. Ticks were collected from domestic dogs and from the environment between 2015 and 2017. Four collection areas were located in the surroundings and within the Serra do Mar State Park, São Paulo State (23°37'21"S, 45°24'43"W), where dogs were bimonthly monitored along 2 years using camera traps and GSM trackers. To determine the spatial limits of genetic structure, ticks collected upon dogs living near the Serra do Baturié, Ceará State (4°15'40"S, 38°55'54"W) were included as well. A total of 39 haplotypes of 16S rRNA and Cox 1 mitochondrial genes sequences were observed, with 27 of them coming from areas within the Serra do Mar State Park. No haplotype was shared between the Serra do Mar and the Serra do Baturié indicating isolation of tick populations at the scale of 2000 km. Although three different haplotype lineages of A. ovale occurred within the Serra do Mar State Park, no genetic structure was found across the study sites within this park, suggesting high tick gene flow across a range of 45 km. Monitoring data from domestic dogs and wild carnivores showed that these species share the same habitats at the forest edge, with dogs playing a likely limited role in tick dispersal. Our findings have important implications for understanding the genetic structure of wide spread A. ovale along Brazilian rainforest remnants, which can further be associated to tick-borne infectious agents, such as Rickettsia parkeri, and used for predicting future patterns of tick diversity in the Brazilian Atlantic rainforest.

Gene flow and adaptive potential in a generalist ectoparasite.

BACKGROUND: In host-parasite systems, relative dispersal rates condition genetic novelty within populations and thus their adaptive potential. Knowledge of host and parasite dispersal rates can therefore help us to understand current interaction patterns in wild populations and why these patterns shift over time and space. For generalist parasites however, estimates of dispersal rates depend on both host range and the considered spatial scale. Here, we assess the relative contribution of these factors by studying the population genetic structure of a common avian ectoparasite, the hen flea Ceratophyllus gallinae, exploiting two hosts that are sympatric in our study population, the great tit Parus major and the collared flycatcher Ficedula albicollis. Previous experimental studies have indicated that the hen flea is both locally maladapted to great tit populations and composed of subpopulations specialized on the two host species, suggesting limited parasite dispersal in space and among hosts, and a potential interaction between these two structuring factors. RESULTS: C. gallinae fleas were sampled from old nests of the two passerine species in three replicate wood patches and were genotyped at microsatellite markers to assess population genetic structure at different scales (among individuals within a nest, among nests and between host species within a patch and among patches). As expected, significant structure was found at all spatial scales and between host species, supporting the hypothesis of limited dispersal in this parasite. Clustering analyses and estimates of relatedness further suggested that inbreeding regularly occurs within nests. Patterns of isolation by distance within wood patches indicated that flea dispersal likely occurs in a stepwise manner among neighboring nests. From these data, we estimated that gene flow in the hen flea is approximately half that previously described for its great tit hosts. CONCLUSION: Our results fall in line with predictions based on observed patterns of adaptation in this host-parasite system, suggesting that parasite dispersal is limited and impacts its adaptive potential with respect to its hosts. More generally, this study sheds light on the complex interaction between parasite gene flow, local adaptation and host specialization within a single host-parasite system.

100 million years of multigene family evolution: origin and evolution of the avian MHC class IIB.

BACKGROUND: Gene duplication has led to a most remarkable adaptation involved in vertebrates' host-pathogen arms-race, the major histocompatibility complex (MHC). However, MHC duplication history is as yet poorly understood in non-mammalian vertebrates, including birds. RESULTS: Here, we provide evidence for the evolution of two ancient avian MHC class IIB (MHCIIB) lineages by a duplication event prior to the radiation of all extant birds >100 million years ago, and document the role of concerted evolution in eroding the footprints of the avian MHCIIB duplication history. CONCLUSIONS: Our results suggest that eroded footprints of gene duplication histories may mimic birth-death evolution and that in the avian MHC the presence of the two lineages may have been masked by elevated rates of concerted evolution in several taxa. Through the presence of a range of intermediate evolutionary stages along the homogenizing process of concerted evolution, the avian MHCIIB provides a remarkable illustration of the erosion of multigene family duplication history.

The Recent Evolution of a Maternally-Inherited Endosymbiont of Ticks Led to the Emergence of the Q Fever Pathogen, Coxiella burnetii.

Q fever is a highly infectious disease with a worldwide distribution. Its causative agent, the intracellular bacterium Coxiella burnetii, infects a variety of vertebrate species, including humans. Its evolutionary origin remains almost entirely unknown and uncertainty persists regarding the identity and lifestyle of its ancestors. A few tick species were recently found to harbor maternally-inherited Coxiella-like organisms engaged in symbiotic interactions, but their relationships to the Q fever pathogen remain unclear. Here, we extensively sampled ticks, identifying new and atypical Coxiella strains from 40 of 58 examined species, and used this data to infer the evolutionary processes leading to the emergence of C. burnetii. Phylogenetic analyses of multi-locus typing and whole-genome sequencing data revealed that Coxiella-like organisms represent an ancient and monophyletic group allied to ticks. Remarkably, all known C. burnetii strains originate within this group and are the descendants of a Coxiella-like progenitor hosted by ticks. Using both colony-reared and field-collected gravid females, we further establish the presence of highly efficient maternal transmission of these Coxiella-like organisms in four examined tick species, a pattern coherent with an endosymbiotic lifestyle. Our laboratory culture assays also showed that these Coxiella-like organisms were not amenable to culture in the vertebrate cell environment, suggesting different metabolic requirements compared to C. burnetii. Altogether, this corpus of data demonstrates that C. burnetii recently evolved from an inherited symbiont of ticks which succeeded in infecting vertebrate cells, likely by the acquisition of novel virulence factors.

Evolutionary changes in symbiont community structure in ticks.

Ecological specialization to restricted diet niches is driven by obligate, and often maternally inherited, symbionts in many arthropod lineages. These heritable symbionts typically form evolutionarily stable associations with arthropods that can last for millions of years. Ticks were recently found to harbour such an obligate symbiont, Coxiella-LE, that synthesizes B vitamins and cofactors not obtained in sufficient quantities from blood diet. In this study, the examination of 81 tick species shows that some Coxiella-LE symbioses are evolutionarily stable with an ancient acquisition followed by codiversification as observed in ticks belonging to the Rhipicephalus genus. However, many other Coxiella-LE symbioses are characterized by low evolutionary stability with frequent host shifts and extinction events. Further examination revealed the presence of nine other genera of maternally inherited bacteria in ticks. Although these nine symbionts were primarily thought to be facultative, their distribution among tick species rather suggests that at least four may have independently replaced Coxiella-LE and likely represent alternative obligate symbionts. Phylogenetic evidence otherwise indicates that cocladogenesis is globally rare in these symbioses as most originate via horizontal transfer of an existing symbiont between unrelated tick species. As a result, the structure of these symbiont communities is not fixed and stable across the tick phylogeny. Most importantly, the symbiont communities commonly reach high levels of diversity with up to six unrelated maternally inherited bacteria coexisting within host species. We further conjecture that interactions among coexisting symbionts are pivotal drivers of community structure both among and within tick species.

Host dispersal as the driver of parasite genetic structure: a paradigm lost?

Understanding traits influencing the distribution of genetic diversity has major ecological and evolutionary implications for host-parasite interactions. The genetic structure of parasites is expected to conform to that of their hosts, because host dispersal is generally assumed to drive parasite dispersal. Here, we used a meta-analysis to test this paradigm and determine whether traits related to host dispersal correctly predict the spatial co-distribution of host and parasite genetic variation. We compiled data from empirical work on local adaptation and host-parasite population genetic structure from a wide range of taxonomic groups. We found that genetic differentiation was significantly lower in parasites than in hosts, suggesting that dispersal may often be higher for parasites. A significant correlation in the pairwise genetic differentiation of hosts and parasites was evident, but surprisingly weak. These results were largely explained by parasite reproductive mode, the proportion of free-living stages in the parasite life cycle and the geographical extent of the study; variables related to host dispersal were poor predictors of genetic patterns. Our results do not dispel the paradigm that parasite population genetic structure depends on host dispersal. Rather, we highlight that alternative factors are also important in driving the co-distribution of host and parasite genetic variation.

The High Diversity and Global Distribution of the Intracellular Bacterium Rickettsiella in the Polar Seabird Tick Ixodes uriae.

Obligate intracellular bacteria of the Rickettsiella genus are emerging as both widespread and biologically diverse in arthropods. Some Rickettsiella strains are highly virulent entomopathogenic agents, whereas others are maternally inherited endosymbionts exerting very subtle manipulations on host phenotype to promote their own spread. Recently, a variety of Rickettsiella strains have been reported from ticks, but their biology is entirely unknown. In the present study, we examined the incidence and diversity of Rickettsiella in 11 geographically distinct populations of the polar seabird tick Ixodes uriae. We found Rickettsiella in most tick populations with a prevalence ranging from 3 to 24 %. 16S ribosomal RNA (rRNA) and GroEL gene sequences revealed an unexpected diversity of Rickettsiella, with 12 genetically distinct Rickettsiella strains present in populations of I. uriae. Phylogenetic investigations further revealed that these Rickettsiella strains do not cluster within a tick-specific clade but rather exhibit distinct evolutionary origins demonstrating frequent horizontal transfers between distantly related arthropod species. Tick rearing further showed that Rickettsiella are present in eggs laid by infected females with no evidence of abortive development. Using this data set, we discuss the potential biological significance of Rickettsiella in seabird ticks. Most notably, we suggest that these organisms may not be pathogenic forms but rather use more subtle adaptive strategies to persist within tick populations.

The role of seabirds of the Iles Eparses as reservoirs and disseminators of parasites and pathogens.

The role of birds as reservoirs and disseminators of parasites and pathogens has received much attention over the past several years due to their high vagility. Seabirds are particularly interesting hosts in this respect. In addition to incredible long-distance movements during migration, foraging and prospecting, these birds are long-lived, site faithful and breed in dense aggregations in specific colony locations. These different characteristics can favor both the local maintenance and large-scale dissemination of parasites and pathogens. The Iles Eparses provide breeding and feeding grounds for more than 3 million breeding pairs of seabirds including at least 13 species. Breeding colonies on these islands are relatively undisturbed by human activities and represent natural metapopulations in which seabird population dynamics, movement and dispersal can be studied in relation to that of circulating parasites and pathogens. In this review, we summarize previous knowledge and recently-acquired data on the parasites and pathogens found in association with seabirds of the Iles Eparses. These studies have revealed the presence of a rich diversity of infectious agents (viruses, bacteria and parasites) carried by the birds and/or their local ectoparasites (ticks and louse flies). Many of these agents are widespread and found in other ecosystems confirming a role for seabirds in their large scale dissemination and maintenance. The heterogeneous distribution of parasites and infectious agents among islands and seabird species suggests that relatively independent metacommunities of interacting species may exist within the western Indian Ocean. In this context, we discuss how the patterns and determinants of seabird movements may alter parasite and pathogen circulation. We conclude by outlining key aspects for future research given the baseline data now available and current concerns in eco-epidemiology and biodiversity conservation.

Rickettsia spp. in seabird ticks from western Indian Ocean islands, 2011-2012.

We found a diversity of Rickettsia spp. in seabird ticks from 6 tropical islands. The bacteria showed strong host specificity and sequence similarity with strains in other regions. Seabird ticks may be key reservoirs for pathogenic Rickettsia spp., and bird hosts may have a role in dispersing ticks and tick-associated infectious agents over large distances.

Massive infection of seabird ticks with bacterial species related to Coxiella burnetii.

Seabird ticks are known reservoirs of bacterial pathogens of medical importance; however, ticks parasitizing tropical seabirds have received less attention than their counterparts from temperate and subpolar regions. Recently, Rickettsia africae was described to infect seabird ticks of the western Indian Ocean and New Caledonia, constituting the only available data on bacterial pathogens associated with tropical seabird tick species. Here, we combined a pyrosequencing-based approach with a classical molecular analysis targeting bacteria of potential medical importance in order to describe the bacterial community in two tropical seabird ticks, Amblyomma loculosum and Carios (Ornithodoros) capensis. We also investigated the patterns of prevalence and host specificity within the biogeographical context of the western Indian Ocean islands. The bacterial community of the two tick species was characterized by a strong dominance of Coxiella and Rickettsia. Our data support a strict Coxiella-host tick specificity, a pattern resembling the one found for Rickettsia spp. in the same two seabird tick species. Both the high prevalence and stringent host tick specificity suggest that these bacteria may be tick symbionts with probable vertical transmission. Detailed studies of the pathogenicity of these bacteria will now be required to determine whether horizontal transmission can occur and to clarify their status as potential human pathogens. More generally, our results show that the combination of next generation sequencing with targeted detection/genotyping approaches proves to be efficient in poorly investigated fields where research can be considered to be starting from scratch.

Tracing the colonization and diversification of the worldwide seabird ectoparasite Ixodes uriae.

Historical patterns of dispersal and population isolation are key components shaping contemporary genetic diversity across landscapes and require explicit consideration when examining the relative role of different factors in driving the evolution of host specificity in parasitic organisms. In this study, we investigate the worldwide colonization history of a common ectoparasite of seabirds, the tick Ixodes uriae. This tick has a circumpolar distribution across both hemispheres but has repeatedly formed host-specific races within different regions. By combining mitochondrial and nuclear data, we infer how this species spread to its present-day distribution and how the colonization process may have affected the geographic and host-associated structure of this tick within regions. We demonstrate that I. uriae is highly structured at a global scale and isolates into four genetic groups that correspond to well-defined geographical regions. Molecular dating suggests that the diversification of I. uriae began in the early Miocene (22 Myr) and that this tick colonized most of the southern hemisphere before moving into northern latitudes via two independent routes. However, no relationship between the degree of host race divergence and colonization history was evident, supporting previous hypotheses that host specialization evolves relatively rapidly in this parasite, but does not typically lead to speciation. We discuss the possible historical and contemporary mechanisms of large-scale dispersal for this ectoparasite and how its biological characteristics may condition current patterns of genetic diversity. More generally, our results illustrate how combining broad-scale sampling and modern molecular tools can help disentangle complex patterns of diversification in widespread parasites.

An experimental test of host specialization in a ubiquitous polar ectoparasite: a role for adaptation?

The evolution of host specificity is considered to be an essential mechanism driving parasite diversity. It may be governed by adaptive constraints that lead to host-dependent fitness trade-offs. Alternatively, specificity may arise via transmission constraints that isolate parasite populations, without necessarily involving adaptation per se. Here, we ask whether the repeated observation of host-associated genetic races across the worldwide distribution of the seabird ectoparasite Ixodes uriae is associated with host adaptation. We conducted a field-based experiment to test for adaptive specialisation in host races of I. uriae. We transferred unengorged ticks of two life stages (nymphs and adults) originating from three host species (black-legged kittiwake, common guillemot and Atlantic puffin) onto young kittiwake nestlings and followed attraction and attachment rates, engorgement times and feeding success of the transplanted ticks. All ticks were also typed genetically to match exploitation patterns with genetic differences among races. Ticks from atypical hosts were significantly less attracted to nestlings than ticks from the typical host, and showed lower feeding success and higher mortality. The degree of host specificity matched patterns of neutral genetic variation among races, with puffin ticks being more specific than guillemot ticks. Differences in specificity were also apparent among tick life stages, suggesting that nymphal ticks may be less discriminating of host type than adult ticks. Our results indicate that the genetic divergence previously observed among sympatric I. uriae host races is at least partially linked to adaptive specialisation to the host species and not simply to host-mediated transmission. They also suggest that the adaptation process may evolve differently in different life stages based on trade-offs with physiological constraints. The identification of the selective forces acting in host specialization will now be necessary to better characterize these patterns and to understand how transmission interacts with the adaptation process to generate parasite biodiversity.

A new species of the feather mite genus Rhinozachvatkinia (Acari: Avenzoariidae) from Calonectris shearwaters (Procellariiformes: Procellariidae): integrating morphological descriptions with DNA barcode data.

Rhinozachvatkinia calonectris sp. n., a new species of the feather mite genus Rhinozachvatkinia Mironov, 1989 (Avenzoariidae: Bonnetellinae), is described from two species of shearwaters in the North-East of the Atlantic Ocean, Calonectris edwardsii (Oustalet) (type host) and Calonectris borealis (Cory) (Procellariiformes: Procellariidae). We completed the morphological description of this new feather mite species with sequence data on the mitochondrial cytochrome c oxidase subunit I gene fragment (COI). The full generic status of Rhinozachvatkinia, originally established as a subgenus of Zachvatkinia Dubinin, 1949, is formally fixed and its systematic relationships are briefly discussed.

Among-colony variation in plastic ingestion by Yellow-legged gulls (Larus michahellis) across the western Mediterranean basin.

The Mediterranean region is both a hotspot for biodiversity and for the accumulation of plastic pollution. Many species are exposed to this pollution while feeding, including a wide diversity of seabirds. Our objective was to investigate spatial variation in the quantity and types of plastic ingested by Yellow-legged gulls using information obtained from regurgitated pellets collected in 11 colonies. Anthropogenic debris, and particularly plastic, was found in pellets from all colonies, but the amount varied considerably. This among-colony difference was stable over the two years of study. The presence of marine prey and the proportion of agricultural area around the colonies significantly influenced the number of ingested plastics. As landfills close and garbage management improves, the availability of anthropogenic waste should decline. Following the response of gulls to these changes will be particularly useful for monitoring plastic pollution and for understanding the response of opportunistic wildlife to environmental modifications.

Spatial variation of mercury contamination in yellow-legged gulls (Larus michahellis) in the Western Mediterranean.

Mercury (Hg) is a global pollutant of major concern in marine and coastal environments. In the Mediterranean Sea, Hg concentrations in biota are higher than in other seas, even when seawater concentrations are similar. Seabirds, as marine top predators, can reflect Hg contamination on a large spatial scale. By sampling seabirds at 17 different breeding colonies, we evaluated Hg concentrations of yellow-legged gulls (Larus michahellis) in the occidental Mediterranean basin in 2021 and 2022. More specifically, we investigated spatial variation of Hg contamination in both chicks and adults as well as associated toxicological risks through the use of blood and feathers, which reflect contamination over different periods of the year. The highest concentrations in chicks were found in Djerba (Tunisia) with blood Hg values of (mean ± SD) 1.69 ± 0.51 µg g-1 dry weight (dw). Adults were most contaminated in Djerba and Dragonera (Balearic Islands, Spain) with blood Hg concentrations of respectively 3.78 ± 2.54 and 5.25 ± 3.73 µg g-1 dw. Trophic ecology was investigated using stable isotope analyses (δ13C, δ15N and δ34S as proxies of feeding habitat and diet), and showed that spatial variation in Hg was mainly driven by foraging habitat in both chicks and adults. Low Hg concentrations were related to the use of anthropogenic food sources. An effect of colony location was also found, suggesting spatial differences in local environmental pollution transfer up to seabirds. Our results also supported the use of δ34S to discriminate between marine and continental foraging habitats in generalist seabirds. This study provides new insights onto the spatial distribution of Hg contamination in a widespread seabird, reporting some of the highest Hg values recorded for this species. Populations with highest concentrations are of potential concern regarding toxicological risks.

Mercury contamination and potential health risk to French seabirds: A multi-species and multi-site study.

Mercury (Hg) is a naturally occurring highly toxic element which circulation in ecosystems has been intensified by human activities. Hg is widely distributed, and marine environments act as its main final sink. Seabirds are relevant bioindicators of marine pollution and chicks are particularly suitable for biomonitoring pollutants as they reflect contamination at short spatiotemporal scales. This study aims to quantify blood Hg contamination and identify its drivers (trophic ecology inferred from stable isotopes of carbon (δ13C) and nitrogen (δ15N), geographical location, chick age and species) in chicks of eight seabird species from 32 French sites representing four marine subregions: the English Channel and the North Sea, the Celtic Sea, the Bay of Biscay and the Western Mediterranean. Hg concentrations in blood ranged from 0.04 µg g-1 dry weight (dw) in herring gulls to 6.15 µg g-1 dw in great black-backed gulls. Trophic position (δ15N values) was the main driver of interspecific differences, with species at higher trophic positions showing higher Hg concentrations. Feeding habitat (δ13C values) also contributed to variation in Hg contamination, with higher concentrations in generalist species relying on pelagic habitats. Conversely, colony location was a weak contributor, suggesting a relatively uniform Hg contamination along the French coastline. Most seabirds exhibited low Hg concentrations, with 74% of individuals categorized as no risk, and < 0.5% at moderate risk, according to toxicity thresholds. However, recent work has shown physiological and fitness impairments in seabirds bearing Hg burdens considered to be safe, calling for precautional use of toxicity thresholds, and for studies that evaluate the impact of Hg on chick development.

Changing distributions of ticks: causes and consequences.

Today, we are witnessing changes in the spatial distribution and abundance of many species, including ticks and their associated pathogens. Evidence that these changes are primarily due to climate change, habitat modifications, and the globalisation of human activities are accumulating. Changes in the distribution of ticks and their invasion into new regions can have numerous consequences including modifications in their ecological characteristics and those of endemic species, impacts on the dynamics of local host populations and the emergence of human and livestock disease. Here, we review the principal causes for distributional shifts in tick populations and their consequences in terms of the ecological attributes of the species in question (i.e. phenotypic and genetic responses), pathogen transmission and disease epidemiology. We also describe different methodological approaches currently used to assess and predict such changes and their consequences. We finish with a discussion of new research avenues to develop in order to improve our understanding of these host-vector-pathogen interactions in the context of a changing world.