The effect of deer management on the abundance of Ixodes ricinus in Scotland

The management of wildlife hosts for controlling parasites and disease has a history of mixed success. Deer can be important hosts for ticks, such as Ixodes ricinus, which is the primary vector of disease-causing zoonotic pathogens in Europe. Deer are generally managed by culling and fencing for forestry protection, habitat conservation, and commercial hunting, and in this study we test whether these deer management methods can be useful for controlling ticks, with implications for tick-borne pathogens. At different spatial scales and habitats we tested the hypotheses that tick abundance is reduced by (1) culling deer and (2) deer exclusion using fencing. We compared abundance indices of hosts and questing I. ricinus nymphs using a combination of small-scale fencing experiments on moorland, a large-scale natural experiment of fenced and unfenced pairs of forests, and cross-sectional surveys of forest and moorland areas with varying deer densities. As predicted, areas with fewer deer had fewer ticks, and fenced exclosures had dramatically fewer ticks in both large-scale forest and small-scale moorland plots. Fencing and reducing deer density were also associated with higher ground vegetation. The implications of these results on other hosts, pathogen prevalence, and disease risk are discussed. This study provides evidence of how traditional management methods of a keystone species can reduce a generalist parasite, with implications for disease risk mitigation.     

Influence of host migration between woodland and pasture on the population dynamics of the tick Ixodes ricinus: A modelling approach

Ticks act as vectors of pathogens that can be harmful to animals and/or humans. Epidemiological models can be useful tools to investigate the potential effects of control strategies on diseases such as tick-borne diseases. The modelling of tick population dynamics is a prerequisite to simulating tick-borne diseases and the corresponding spread of the pathogen. We have developed a dynamic model to simulate changes in tick density at different stages (egg, larva, nymph and adult) under the influence of temperature. We have focused on the tick Ixodes ricinus, which is widespread in Europe. The main processes governing the biological cycles of ticks were taken into account: egg laying, hatching, development, host (small, mainly rodents, or large, like deer and cattle, mammals) questing, feeding and mortality. This model was first applied to a homogeneous habitat, where simulations showed the ability of the model to reproduce the general patterns of tick population dynamics. We considered thereafter a multi-habitat model, where three different habitats (woodland, ecotone and meadow) were connected through host migration. Based on this second application, it appears that migration from woodland, via the ecotone, is necessary to sustain the presence of ticks in the meadow. Woodland can therefore be considered as a source of ticks for the meadow, which in turn can be regarded as a sink. The influence of woodland on surrounding tick densities increases in line with the area of this habitat before reaching a plateau. A sensitivity analysis to parameter values was carried out and demonstrated that demographic parameters (sex ratio, development, mortality during feeding and questing, host finding) played a crucial role in the determination of questing nymph densities. This type of modelling approach provides insight into the influence of spatial heterogeneity on tick population dynamics.     

Multiple causes of variable tick burdens on small-mammal hosts

Blood meals by blacklegged ticks (Ixodes scapularis) on vertebrate hosts serve to transmit the agents of several zoonotic diseases, including Lyme disease, human babesiosis, and human granulocytic anaplasmosis, between host and tick. If ticks are aggregated on hosts, a small proportion of hosts may be responsible for most transmission events. Therefore, a key element in understanding and controlling the transmission of these pathogens is identifying the group(s) or individuals feeding a disproportionate number of ticks. Previous studies of tick burdens, however, have focused on differences in mean annual burdens between one or a few groups of hosts, ignoring both the strong seasonal dynamics of I. scapularis and their aggregation on hosts. We present a statistical modeling framework that predicts burdens on individual hosts throughout the year as a function of temporal-, site-, and individual-specific attributes, as well as the degree of aggregation in a negative binomial distribution. We then fit alternate versions of this model to an 11-year data set of I. scapularis burdens on white-footed mice (Peromyscus leucopus) and eastern chipmunks (Tamias striatus) to explore which factors are important to predicting tick burdens. We found that tick burdens are a complex function of many extrinsic and intrinsic factors, including seasonality. Specifically: (1) burdens on mice and chipmunks increased with densities of host-seeking ticks in a manner that suggests hosts become saturated. (2) Chipmunks draw larval ticks away from mice, which are efficient reservoirs of the Lyme disease bacterium, and mice draw nymphs away from chipmunks, which are key nymphal hosts. (3) While individual correlates were statistically important, the relationships were complex, and no group or correlate (sex, age, mass) could explain which hosts fed a disproportionate number of ticks. (4) Ticks were strongly aggregated on hosts within and across groups suggesting that some undiscovered quality of individual hosts was responsible for the aggregation. (5) Those individuals that fed more nymphs than expected, and are thus more likely to be infected with the Lyme disease agent, also tend to feed and infect more larvae than expected. Predicting which individuals those are is not yet possible.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Managing the Abundance and Diversity of Breeding Bird Populations through Manipulation of Deer Populations

Abstract: Deer densities in forests of eastern North America are thought to have significant effects on the abundance and diversity of forest birds through the role deer play in structuring forest understories. We tested the ability of deer to affect forest bird populations by monitoring the density and diversity of vegetation and birds for 9 years at eight 4-ha sites in northern Virginia, four of which were fenced to exclude deer. Both the density and diversity of understory woody plants increased following deer exclosure. The numerical response of the shrubs to deer exclosure was significantly predicted by the soil quality (ratio of organic carbon to nitrogen) at the sites. Bird populations as a whole increased following exclosure of deer, particularly for ground and intermediate canopy species. The diversity of birds did not increase significantly following exclosure of deer, however, primarily because of replacement of species as understory vegetation proceeded through successional processes. Changes in understory vegetation accounted for most of the variability seen in the abundance and diversity of bird populations. Populations of deer in protected areas are capable of causing significant shifts in the composition and abundance of bird communities. These shifts can be reversed by increasing the density and diversity of understory vegetation, which can be brought about by reducing deer density.     

Lyme Disease and Conservation

Lyme disease is a tick-borne illness that is widespread in North America, especially in the northeastern and northcentral United States. This disease could negatively influence efforts to conserve natural populations in two ways: (1) the disease could directly affect wild animal health; and (2) tick control efforts could adversely affect natural populations and communities. Lyme disease affects several domestic animals, but symptoms have been reported in only a few wild species. Direct effects of Lyme disease on wild animal populations have not been reported, but the disease should be considered as a possible cause in cases of unexplained population declines in endemic areas. Methods available to manage ticks and Lyme disease include human self-protection techniques, manipulation of habitats and host species populations, biological control, and pesticide applications. The diversity of available techniques allows selection of approaches to minimize environmental effects by (1) emphasizing personal protection techniques, (2) carefully targeting management efforts to maximize efficiency, and (3) integrating environmentally benign techniques to improve management while avoiding broad-scale environmentally destructive approaches. The environmental effects of Lyme disease depend, to a large extent, on the methods chosen to minimize human exposure to infected ticks. Conservation biologists can help design tick management programs that effectively lower the incidence of human Lyme disease while simultaneously minimizing negative effects on natural populations.     

Modelling landscape effects on density-contact rate relationships of deer in eastern Alberta: Implications for chronic wasting disease

Managing wildlife diseases requires an understanding of disease transmission, which may be strongly affected by host population density and landscape features. Transmission models are typically fit from time-series disease prevalence data and modelled based on how the contact rate among hosts is affected by density, which is often assumed to be a linear (density-dependent transmission) or constant (frequency-dependent transmission) relationship. However, long-term time-series data is unavailable for emerging diseases, and this approach cannot account for independent effects of landscape. We developed a mechanistic model based on ecological data to empirically derive the contact rate-density relationship in white-tailed and mule deer in an enzootic region of chronic wasting disease (CWD) in Alberta, Canada and to determine whether it was affected by landscape. Using data collected from aerial surveys and GPS-telemetry, we developed empirical relationships predicting deer group size, home range size, and habitat selection to iteratively simulate deer distributions across a range of densities and landscapes. We calculated a relative measure of total per-capita contact rate, which is proportional to the number of other deer contacted per individual per unit time, for each distribution as the sum of pairwise contact rates between a target deer and all other individuals. Each pairwise contact rate was estimated from an empirical relationship developed from GPS-telemetry data predicting pairwise contact rates as a function of home range overlap and landscape structure. Total per-capita contact rates increased as a saturating function of density, supporting a transmission model intermediate between density- and frequency-dependent transmission. This pattern resulted from group sizes that reached an asymptote with increasing deer density, although this relationship was mediated by tree and shrub coverage in the landscape, such that in heavily wooded areas, the contact rate saturated at much lower densities. These results suggest that CWD management based on herd reductions, which require a density-dependent contact rate to be effective, may have variable effects on disease across a single management region. The novel mechanistic approach we employed for estimating effects of density and landscape on transmission is a powerful complement to typical data-fitting approaches for modelling disease transmission.     

Species diversity reduces parasite infection through cross-generational effects on host abundance

With growing interest in the effects of biodiversity on disease, there is a critical need for studies that empirically identify the mechanisms underlying the diversity-disease relationship. Here, we combined wetland surveys of host community structure with mechanistic experiments involving a multi-host parasite to evaluate competing explanations for the dilution effect. Sampling of 320 wetlands in California indicated that snail host communities were strongly nested, with competent hosts for the trematode Ribeiroia ondatrae predominating in low-richness assemblages and unsuitable hosts increasingly present in more diverse communities. Moreover, competent host density was negatively associated with increases in snail species richness. These patterns in host community assembly support a key prerequisite underlying the dilution effect. Results of multigenerational mesocosm experiments designed to mimic field-observed community assemblages allowed us to evaluate the relative importance of host density and diversity in influencing parasite infection success. Increases in snail species richness (from one to four species) had sharply negative effects on the density of infected hosts (-90% reduction). However, this effect was indirect; competition associated with non-host species led to a 95% reduction in host density (susceptible host regulation), owing primarily to a reduction in host reproduction. Among susceptible hosts, there were no differences in infection prevalence as a function of community structure, indicating a lack of support for a direct effect of diversity on infection (encounter reduction). In monospecific conditions, higher initial host densities increased infection among adult hosts; however, compensatory reproduction in the low-density treatments equalized the final number of infected hosts by the next generation, underscoring the relevance of multigenerational studies in understanding the dilution effect. These findings highlight the role of interspecific competition in mediating the relationship between species richness and parasite infection and emphasize the importance of field-informed experimental research in understanding mechanisms underlying the diversity-disease relationship.     

Refuge sharing network predicts ectoparasite load in a lizard

Living in social groups facilitates cross-infection by parasites. However, empirical studies on indirect transmission within wildlife populations are scarce. We investigated whether asynchronous overnight refuge sharing among neighboring sleepy lizards, Tiliqua rugosa, facilitates indirect transmission of its ectoparasitic tick, Amblyomma limbatum. We fitted 18 neighboring lizards with GPS recorders, observed their overnight refuge use each night over 3 months, and counted their ticks every fortnight. We constructed a transmission network to estimate the cross-infection risk based on asynchronous refuge sharing frequencies among all lizards and the life history traits of the tick. Although self-infection was possible, the network provided a powerful predictor of measured tick loads. Highly connected lizards that frequently used their neighbors’ refuges were characterized by higher tick loads. Thus, indirect contact had a major influence on transmission pathways and parasite loads. Furthermore, lizards that used many different refuges had lower cross- and self-infection risks and lower tick loads than individuals that used relatively fewer refuges. Increasing the number of refuges used by a lizard may be an important defense mechanism against ectoparasite transmission in this species. Our study provides important empirical data to further understand how indirectly transmitted parasites move through host populations and influence individual parasite loads.     

Acceleration of Exotic Plant Invasion in a Forested Ecosystem by a Generalist Herbivore

Abstract: The successful invasion of exotic plants is often attributed to the absence of coevolved enemies in the introduced range (i.e., the enemy release hypothesis). Nevertheless, several components of this hypothesis, including the role of generalist herbivores, remain relatively unexplored. We used repeated censuses of exclosures and paired controls to investigate the role of a generalist herbivore, white‐tailed deer (Odocoileus virginianus), in the invasion of 3 exotic plant species (Microstegium vimineum, Alliaria petiolata, and Berberis thunbergii) in eastern hemlock (Tsuga canadensis) forests in New Jersey and Pennsylvania (U.S.A.). This work was conducted in 10 eastern hemlock (T. canadensis) forests that spanned gradients in deer density and in the severity of canopy disturbance caused by an introduced insect pest, the hemlock woolly adelgid (Adelges tsugae). We used maximum likelihood estimation and information theoretics to quantify the strength of evidence for alternative models of the influence of deer density and its interaction with the severity of canopy disturbance on exotic plant abundance. Our results were consistent with the enemy release hypothesis in that exotic plants gained a competitive advantage in the presence of generalist herbivores in the introduced range. The abundance of all 3 exotic plants increased significantly more in the control plots than in the paired exclosures. For all species, the inclusion of canopy disturbance parameters resulted in models with substantially greater support than the deer density only models. Our results suggest that white‐tailed deer herbivory can accelerate the invasion of exotic plants and that canopy disturbance can interact with herbivory to magnify the impact. In addition, our results provide compelling evidence of nonlinear relationships between deer density and the impact of herbivory on exotic species abundance. These findings highlight the important role of herbivore density in determining impacts on plant abundance and provide evidence of the operation of multiple mechanisms in exotic plant invasion.     

Attractive and repellent host odours guide ticks to their respective feeding sites

Summary. We have studied on-host behaviour of adults of the brown ear tick (Rhipicephalus appendiculatus Neumann, 1901) and the red-legged tick (R. evertsi Neumann, 1897), which prefer to feed mainly inside the ears and the anal regions of bovids respectively. Both species were found to be relatively successful in orienting toward and locating their respective feeding sites from different parts of the host body. Our observations suggested the operation of both avoidance (closer to the feeding site of the other) and attraction (closer to its own feeding site) responses of the ticks. In the laboratory, odour trapped from cattle ears attracted R. appendiculatus but repelled R. evertsi, whereas that from the anal region had an opposite effect. This odour-based push-pull pair of stimuli may largely account for efficient orientation behaviour of the two tick species to their respective feeding sites. We propose that such concurrent deployment of repulsive and attractive cues may be quite widespread among arthropods and related organisms that specialise on specific hosts or microenvironments in the performance of their biological functions.     

Deer herbivory alters forest response to canopy decline caused by an exotic insect pest.

Hemlock woolly adelgid (HWA; Adelges tsugae) infestations have resulted in the continuing decline of eastern hemlock (Tsuga canadensis) throughout much of the eastern United States. While the initial impacts of HWA infestations have been documented, our understanding of forest response to this disturbance remains incomplete. HWA infestation is not occurring in isolation but within a complex ecological context. The role of potentially important interacting factors, such as elevated levels of white-tailed deer herbivory, is poorly understood. Despite the potential for herbivory to alter forest successional trajectories following a canopy disturbance, little is known about herbivory-disturbance interactions, and herbivory is rarely considered in assessing forest response to a co-occurring disturbance. We used repeated censuses of deer exclosures and paired controls (400 paired plots) to quantify the impact of deer herbivory on tree seedling species abundance in 10 eastern hemlock ravines that span a gradient in HWA-induced canopy decline severity. Use of a maximum likelihood estimation framework and information theoretics allowed us to quantify the strength of evidence for alternative models developed to estimate the impacts of herbivory on tree seedling abundance as a function of varying herbivore density and canopy decline severity. The exclusion of deer herbivory had marked impacts on the abundance of the studied seedling species: Acer rubrum, Acer saccharum, Betula lenta, Nyssa sylvatica, Quercus montana, and Tsuga canadensis. For all six species, the relationship between seedling abundance and deer density was either exponential or saturating. Although the functional form of the response varied among seedling species, the inclusion of both deer density and canopy decline severity measures consistently resulted in models with substantially greater support. Canopy decline resulted in higher proportional herbivory impacts and altered the ranking of herbivory impacts by seedling species. Our results suggest that, by changing species' competitive abilities, white-tailed deer herbivory alters the trajectory of forest response to this exotic insect pest and has the potential to shift future overstory composition.     

Parasite transmission in complex communities: predators and alternative hosts alter pathogenic infections in amphibians

While often studied in isolation, host-parasite interactions are typically embedded within complex communities. Other community members, including predators and alternative hosts, can therefore alter parasite transmission (e.g., the dilution effect), yet few studies have experimentally evaluated more than one such mechanism. Here, we used data from natural wetlands to design experiments investigating how alternative hosts and predators of parasites mediate trematode (Ribeiroia ondatrae) infection in a focal amphibian host (Pseudacris regilla). In short-term predation bioassays involving mollusks, zooplankton, fish, larval insects, or newts, four of seven tested species removed 62-93% of infectious stages. In transmission experiments, damselfly nymphs (predators) and newt larvae (alternative hosts) reduced infection in P. regilla tadpoles by -50%, whereas mosquitofish (potential predators and alternative hosts) did not significantly influence transmission. Additional bioassays indicated that predators consumed parasites even in the presence of alternative prey. In natural wetlands, newts had similar infection intensities as P. regilla, suggesting that they commonly function as alternative hosts despite their unpalatability to downstream hosts, whereas mosquitofish had substantially lower infection intensities and are unlikely to function as hosts. These results underscore the importance of studying host-parasite interactions in complex communities and of broadly linking research on predation, biodiversity loss, and infectious diseases.     

Interannual dynamics of antibody levels in naturally infected long-lived colonial birds

Little is known about the long-term persistence of specific antibodies (Ab) in natural host-parasite systems despite its potential epidemiological and ecological importance. In long-lived species, knowledge of the dynamics of individual immunological profiles can be important not only for interpreting serology results, but also for assessing transmission dynamics and the potential selective pressures acting on parasites. The aim of this paper was to investigate temporal variation in levels of specific Ab against the bacterium Borrelia burgdorferi sensu lato in adults of a long-lived colonial seabird, the Black-legged Kittiwake Rissa tridactyla. In wild populations, adults are naturally exposed each breeding season to a Borrelia vector, the tick Ixodes uriae. Breeding birds were captured during four consecutive breeding seasons, and parasite infestation quantified. Using enzyme-linked immunosorbent assay (ELISA) and immunoblots, we found that the immunological profiles of anti-Borrelia Ab were highly repeatable among years, reflecting the interannual persistence of Ab levels. We nevertheless also observed that year-to-year changes of Ab levels were related to exposure to ticks in the previous year. The long-term persistence of Ab levels may be an important mechanism of individual protection against future exposure to the microparasite. It will also affect the availability of susceptible hosts, and thus the transmission dynamics of the bacterium. These results illustrate the need to consider the dynamics of the immune response in order to better understand the evolutionary ecology of host-parasite interactions in natural populations.     

Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

An epidemiological model of East Coast Fever in African livestock

An epidemiological model of the dynamics of East Coast Fever (ECF) in East Africa caused by the protozoan parasite Theileria parva and transmitted by the brown-ear tick Rhipicephalus appendiculatus was developed. In the model, ticks are assigned to either on-host or off-host categories both of which differ in their capacity to receive and transmit the disease. Cattle are assigned to categories of susceptible, infected and infectious as well as recovered animals having immunity to the disease. The parameters of the model were estimated from data reported in the literature.A mathematical analysis of the ECF/tick/cattle model with and without disease was conducted. Depending on the ratio between fecundity and mortality rates in cattle and in the absence of disease, different scenarios emerge including extinction of ticks, coexistence of ticks and cattle and total extinction of ticks and cattle. Furthermore, the analysis of the model with the disease yielded threshold conditions for the existence and the persistence of stable coexistence equilibria for the epidemiological system that may lead to enzootic stability.The model was used to identify critical aspects of the dynamics required to develop management strategies: (i) tick control in areas where the disease is absent, (ii) threshold-based tick and disease control, and (iii) conditions permitting the establishment of enzootic stability of the ECF/tick/cattle system. The analysis also identifies critical areas requiring further field investigation, sets the basis for developing a realistic model for field implementation, and provides a tool for project development and evaluation in the context of international research.     

Network structure and parasite transmission in a group living lizard,the gidgee skink, <Emphasis Type="Italic">Egernia stokesii</Emphasis>

Gidgee skinks (Egernia stokesii) form large social aggregations in rocky outcrops across the Flinders Ranges in South Australia. Group members share refuges (rock crevices), which may promote parasite transmission. We measured connectivity of individuals in networks constructed from patterns of common crevice use and observed patterns of parasitism by three blood parasites (Hemolivia, Schellackia and Plasmodium) and an ectoparasitic tick (Amblyomma vikirri). Data came from a 1-year mark-recapture study of four populations. Transmission networks were constructed to represent possible transmission pathways among lizards. Two lizards that used the same refuge within an estimated transmission period were considered connected in the transmission network. An edge was placed between them, directed towards the individual that occupied the crevice last. Social networks, a sub-set of same-day only associations, were small and highly fragmented compared with transmission networks, suggesting that non-synchronous crevice use leads to more transmission opportunities than direct social association. In transmission networks, lizards infested by ticks were connected to more other tick-infested lizards than uninfected lizards. Lizards infected by ticks and carrying multiple blood parasite infections were in more connected positions in the network than lizards without ticks or with one or no blood parasites. Our findings suggest higher levels of network connectivity may increase the risk of becoming infected or that parasites influence lizard behaviour and consequently their position in the network. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Wildlife Loss through Domestication: the Case of Endangered Key Deer

Abstract:  Wildlife extinction represents the ultimate failure of wildlife conservation. It has many causes, some of them natural, but is increasingly tied to anthropogenic factors. Wildlife loss via domestication, however, is rarely considered. We evaluated the potential for inadvertent domestication of wildlife by determining the effect of feeding and watering on Key deer ( Odocoileus virginianus clavium ) density, group size, and distribution. Key deer sightings were significantly higher in areas (42 ha) surrounding the households that provided food and water (0.18 deer/m;  n = 8   ) than in randomly selected areas (0.03 deer/m;  t = 3.82, 14  df, p = 0.002   ). Average distance to a household providing food and water decreased logistically as group size increased, and large groups (>2 individuals each) were observed more frequently in areas where food and water were provided (27.5%) than in the randomly selected areas (7.5%). The incidence of large groups outside feeding areas (7.5%), however, was similar to the incidence of large groups during early urbanization (5.1%; 1968–1973). Our results suggest illegal feeding caused changes in density, group size, and distribution indicative of domestication. Because fresh water and food were primary selective pressures for Key deer before illegal feeding and watering, genetic changes may occur in the future. For those who value "wildness" in wildlife, domestication of wildlife species is a serious problem that must be addressed.     

Modification of hosts' behavior by a parasite: field evidence for adaptive manipulation

Parasites relying on trophic transmission to complete their life cycles often induce modifications of their host's behavior in ways that may increase their susceptibility to predation by final hosts. These modifications have often been interpreted as parasite adaptations, but very few studies have demonstrated that host manipulation has fitness benefits for the parasite. The aim of the present study was to address the adaptive significance of parasite manipulation by coupling observations of behavioral manipulation to estimates of trophic transmission to the definitive host in the natural environment. We show that the acanthocephalan parasite Pomphorhynchus laevis manipulates the drifting behavior of one of its intermediate hosts, the amphipod Gammarus pulex, but not of a sympatric host, the introduced amphipod Gammarus roeseli. We found a 26.3-28.3 times higher proportion of infected G. pulex in the stomach content of one of the definitive hosts of P. laevis, the bullhead Cottus gobio, than in the benthos. No such trend was observed for G. roeseli. The bell-shaped curve of mean parasite abundance (MPA) relative to host size observed in G. pulex also supported an increased predation mortality of P. laevis-infected individuals compared to uninfected amphipods. Again, no such pattern was observed in G. roeseli. Furthermore, our results indicate that the modifications induced by P. laevis are specific to the definitive host and do not increase the risk of predation by inappropriate hosts, here the adult edible frog Rana esculenta. Overall, our study is original in that it establishes, under field conditions, a direct link between parasitic manipulation and increased transmission to the definitive host, and more importantly, identifies the specificity of the manipulation both in the intermediate host species and toward the definitive host.