Effect of the spatial context along the invasion process: “Hierarchical spatial” or “Host-switching spatial” hypotheses?

Very little is known about how spatial effects influence invasive species throughout the invasion sequence. We propose here two mechanisms to explain the changes in spatial effects throughout the stages of invasion, using the soybean aphid (Aphis glycines) as a model. First, the “hierarchical spatial effect” hypothesis, based on a change in the relative importance of the spatial scales throughout the invasion process, with main effect at broad scale during the first years of invasion, and main effect at local scale during the subsequent years. Second, the “host-switching spatial effect” hypothesis, stating that the spatial effect is driven by a switch in the effect of the host/habitat throughout the invasion process, from effect of main summer host/habitat during the first years of invasion to effect of overwintering host/habitat during the subsequent years. Data from governmental archives and field samplings enabled to investigate the spatial effects on aphid density at three scales (regional, landscape, local) during a 7 year period (2006–2012). Our results demonstrate that the hierarchical spatial effect hypothesis is not an adequate model for the soybean aphid, aphid density being more affected by landscape-scale factors irrespective of years. In contrast, our results are in accordance with the host-switching spatial hypothesis, with positive effect of the main summer host/habitat (soybean) during the first steps of invasion (2006–2008), followed by a positive effect of overwintering habitats (buckthorn, woodland) during the subsequent years (2010–2012). Overall, investigating these hypotheses in other systems would determine whether the same tendency is observed for other invasive species.     

Helminth communities of four commercially important fish species from Chetumal Bay, Mexico

The relative importance of ecology and evolution as factors determining species richness and composition of the helminth communities of fish is a matter of current debate. Theoretical studies use host-parasite lists, but these do not include studies on a temporal or spatial scale. Local environmental conditions and host biological characteristics are shown to influence helminth species richness and composition in four fish species (Eugerres plumieri, Hexanematichthys assimilis, Oligoplites saurus, and Scomberomorus maculatus) in Chetumal Bay, Mexico. With the exception of H. assimilis, the helminth communities had not been previously studied and possible associations between environmental and host biological characteristics as factors determining helminth species richness and composition using redundancy analysis (RDA) are described. Thirty-four helminth species are identified, with the highest number of species (19 total (mean = 6.3 +/- 2.1)) and the lowest (9 (4.0 +/- 1.0)) occurring in H. assimilis and S. maculatus, respectively. The larval nematodes Contracaecum sp. and Pseudoterranova sp. were not only the helminth species shared by all four host species but also were the most prevalent and abundant. Statistical associations between helminth community parameters and local ecological variables such as host habitat use, feeding habits, mobility, and time of residence in coastal lagoons are identified. Phylogeny is important because it clearly separates all four host species by their specialist parasites, although specific habitat and feeding habits also significantly influence the differentiation between the four fish species.     

Species and sex biases in ectoparasitism of dragonflies by mites

An important problem in understanding the evolution of parasite host range is determining the extent to which parasite fitness varies among host species and the factors affecting that fitness variation. We present a detailed investigation on the patterns of host use and successful parasitism of two dragonfly species by the ectoparasitic water mite, Limnochares americana Lundblad. In our field surveys, we found both species biases and sex biases in parasitism by mites, which appear explained by differences in exposure. Differential habitat use by dragonflies helped explain male biases in parasitism in both host species, but was not useful in explaining species biases in parasitism. Species biases in parasitism may be explained by more subtle variation in habitat use not explored in this study, or perhaps by differences in timing of emergence, as we found for the two dragonfly species. Despite species differences in parasitism in nature, we found that mites attached equally successfully to both dragonfly species during experimental infestations. However, mites failed to engorge more often on the dragonfly species less often used as a host in nature. This host species also was more likely to have dead mites in natural infestations as compared to the other host species, which was more often and more heavily parasitized. Our results are consistent with previous research suggesting parasites are less successful on less often used hosts. Such research has implications for understanding determinants of host range for animal parasites.     

Host-specialization and species diversity in fish parasites: phylogenetic conservatism?

The pattern of parasite species diversification and specialization, appreciated by host range, is investigated in fish parasites. We test whether host range is linked with phylogeny at a high taxonomic level, and if there is a relationship between host range and host species diversification. For this purpose we used two sets of data, one on macro-parasites of marine fishes of the Mediterranean Sea and the other on macro-parasites of marine and freshwater fishes of Canada. Similar patterns of host range among parasitic groups were found. Our findings suggest that habitat (marine vs freshwater) and geographic localization (Canada vs Mediterranean region) play little role in determining the observed patterns of host range. We highlight the potential influence of phylogeny (high-taxonomic level) on the level host range in parasites. We find that parasites with free-swimming larval stages and with direct life cycles have a narrower range of host species than do parasites with indirect life cycle, even if we cannot control for phylogenetic effects because of the lack of variation of life cycles within each parasitic group. Finally, a positive relationship was found between the number of known hosts and parasite species diversity in the case of Mediterranean parasite species. The relationship between host range and species diversification should be related to the mechanism of cospeciation.     

Intra‐specific variability in life‐cycle synchronization of an ectoparasitic fly to its avian host

The role of environmental and host‐associated factors in synchronization of host–parasite life‐cycles is an important question of evolutionary ecology. Yet, only handsome of studies examined this question at the intraspecific level. Here we explore how host‐associated traits, such as breeding phenology and host breeding habitat, can influence parasite phenology and co‐occurrence at different spatial scales. We studied the system comprised of a generalist ectoparasitic fly Carnus hemapterus and one of its avian hosts, the European roller Coracias garrulus. Inter‐annual variation in phenology was larger for parasites than hosts. Host predictability in terms of occurrence and phenological regularity was moderate, suggesting that this resource can be difficult to be tracked by the parasite. A large proportion of flies consistently emerged before the appearance of suitable host resources at both the nest and population level. Consequently, we revealed low and highly variable inter‐annual host–parasite synchronization rates. Nevertheless, we found that parasites from nests of early and progressively earlier breeding European rollers were more synchronized with their hosts than parasites from nests of late and progressively later breeding hosts, respectively. Temporal trends in host suitability and parasite emergence at the population scale suggest that other mechanisms, such as dispersal or exploitation of other host species, ensure parasites access to resources and counteract asynchrony with the host at the nest scale.     

Linking community assembly and structure across scales in a wild mouse parasite community

Understanding what processes drive community structure is fundamental to ecology. Many wild animals are simultaneously infected by multiple parasite species, so host–parasite communities can be valuable tools for investigating connections between community structures at multiple scales, as each host can be considered a replicate parasite community. Like free‐living communities, within‐host–parasite communities are hierarchical; ecological interactions between hosts and parasites can occur at multiple scales (e.g., host community, host population, parasite community within the host), therefore, both extrinsic and intrinsic processes can determine parasite community structure. We combine analyses of community structure and assembly at both the host population and individual scales using extensive datasets on wild wood mice (Apodemus sylvaticus) and their parasite community. An analysis of parasite community nestedness at the host population scale provided predictions about the order of infection at the individual scale, which were then tested using parasite community assembly data from individual hosts from the same populations. Nestedness analyses revealed parasite communities were significantly more structured than random. However, observed nestedness did not differ from null models in which parasite species abundance was kept constant. We did not find consistency between observed community structure at the host population scale and within‐host order of infection. Multi‐state Markov models of parasite community assembly showed that a host's likelihood of infection with one parasite did not consistently follow previous infection by a different parasite species, suggesting there is not a deterministic order of infection among the species we investigated in wild wood mice. Our results demonstrate that patterns at one scale (i.e., host population) do not reliably predict processes at another scale (i.e., individual host), and that neutral or stochastic processes may be driving the patterns of nestedness observed in these communities. We suggest that experimental approaches that manipulate parasite communities are needed to better link processes at multiple ecological scales.     

Water temperature,not fish morph,determines parasite infections of sympatric Icelandic threespine sticklebacks (Gasterosteus aculeatus)

Parasite communities of fishes are known to respond directly to the abiotic environment of the host, for example, to water quality and water temperature. Biotic factors are also important as they affect the exposure profile through heterogeneities in parasite distribution in the environment. Parasites in a particular environment may pose a strong selection on fish. For example, ecological differences in selection by parasites have been hypothesized to facilitate evolutionary differentiation of freshwater fish morphs specializing on different food types. However, as parasites may also respond directly to abiotic environment the parasite risk does not depend only on biotic features of the host environment. It is possible that different morphs experience specific selection gradients by parasites but it is not clear how consistent the selection is when abiotic factors change. We examined parasite pressure in sympatric morphs of threespine stickleback (Gasterosteus aculeatus) across a temperature gradient in two large Icelandic lakes, Myvatn and Thingvallavatn. Habitat‐specific temperature gradients in these lakes are opposite. Myvatn lava rock morph lives in a warm environment, while the mud morph lives in the cold. In Thingvallavatn, the lava rock morph lives in a cold environment and the mud morph in a warm habitat. We found more parasites in fish living in higher temperature in both lakes, independent of the fish morph, and this pattern was similar for the two dominating parasite taxa, trematodes and cestodes. However, at the same time, we also found higher parasite abundance in a third morph living in deep cold–water habitat in Thingvallavatn compared to the cold‐water lava morph, indicating strong effect of habitat‐specific biotic factors. Our results suggest complex interactions between water temperature and biotic factors in determining the parasite community structure, a pattern that may have implications for differentiation of stickleback morphs.     

Linking ecology with parasite diversity in Neotropical fishes

A comparative analysis was performed to seek large-scale patterns in the relationships between a set of fish species traits (body size, type of environment, trophic level, schooling behaviour, depth range, mean habitat temperature, geographical range, ability to enter brackish waters and capability of migration) and the diversity of their metazoan parasite assemblages among 651 Neotropical fish species. Two measurements of parasite diversity are used: the species richness and the taxonomic distinctness of a fish's parasite assemblage, including all metazoan parasites, ectoparasites only, or endoparasites only. The results showed that, on this scale, the average taxonomic distinctness of parasite assemblages was clearly more sensitive to the influence of host traits than parasite species richness. Differences in the taxonomic diversification of the parasite assemblages of different fish species were mainly related to the fish's environment (higher values in benthic–demersal species), trophic level (positive correlation with increasing level), temperature (positive correlation with temperature in marine ectoparasites, negative in endoparasites; positive for all groups of parasites in freshwater fishes) and oceanic distribution (higher values in fish species from the Pacific Ocean than those of the Atlantic). The results suggest that, among Neotropical fish species, only certain key host traits have influenced the processes causing the taxonomic diversification of parasite assemblages.     

Linking fish population dynamics to habitat conditions: insights from the application of a process-oriented approach to several Great Lakes species

One of the major challenges facing fishery scientists and managers today is determining how fish populations are influenced by habitat conditions. Many approaches have been explored to address this challenge, all of which involve modeling at one level or another. In this paper, we explore a process-oriented model approach whereby the critical population processes of birth and death rates are explicitly linked to habitat conditions. Application of this approach to five species of Great Lakes fishes including: walleye (Sander vitreus), lake trout (Salvelinus namaycush), smallmouth bass (Micropterus dolomieu), yellow perch (Perca flavescens), and rainbow trout (Onchorynchus mykiss), yielded a number of insights into the modeling process. One of the foremost insights is that processes determining movement and transport of fish are critical components of such models since these processes largely determine the habitats fish occupy. Because of the importance of fish location, an individual-based model appears to be a nearly inescapable modeling requirement. There is, however, a paucity of field-based data directly relating birth, death, and movement rates to habitat conditions experienced by individual fish. There is also a paucity of habitat information at a fine temporal and spatial scale for many important habitat variables. Finally, the general occurrence of strong ontogenetic changes in the response of different life stages to habitat conditions emphasizes the need for a modeling approach that considers all life stages in an integrated fashion.     

Speciation by host switch and adaptive radiation in a fish parasite genus Gyrodactylus (Monogenea,Gyrodactylidae)

Four hundred Gyrodactylus species have been formally described, but the estimated number of species in this fish ectoparasite genus of Monogenean Platyhelminthes is more than 20,000. The unusually high species richness has lead to the hypotheses of speciation and adaptive radiation via host switching. These hypotheses were tested by reconstructing a molecular phylogeny for the subgenus G. (Limnonephrotus) which is a group of freshwater parasites, including five species infecting wild and farmed salmonids. The highly variable ITS1 and ITS2 segments and the conservative 5.8S ribosomal gene were sequenced in 22 species plus two species representing the subgenus G. (Paranephrotus) as an outgroup. The phylogeny was compared with host systematics: the species were collected from six fish families (Cyprinidae, Salmonidae, Percidae, Esocidae, Gasterosteidae, and Gobitidae). The phylogenetic analysis demonstrated that G. (Limnonephrotus) is a monophyletic group that was originally hosted by cyprinids. The speciation has occurred in two episodes, the older one manifested in genetic distances 25-33% (4-6 Myr BP). The latter speciation burst occurred in one clade only, perhaps one million years ago. This clade has been morphologically identified as a wageneri species group. It is a monophyletic group of 18 species [studied here] and contains all five salmonid parasites, but also parasites, on cyprinids, percids, esocids, and gasterosteids. In G. (Limnonephrotus), eight host switches crossing the host family barrier were observed, and at least three of them were followed by repetitive speciation. Seven host-switch events were statistically confirmed by bootstrapping. The suggested model of speciation by host switch was accepted, and interestingly the adaptive radiation seems to be a consequence of host switch to a new family (key innovation model). The molecular and ecological evolution rate of Gyrodactylus parasites is manyfold in comparison to host species, and the phylogenies are largely independent and disconnected.     

Speciation and diversification of parasite lineages: an analysis of congeneric parasite species in vertebrates

The evolutionary diversification of living organisms is a central research theme in evolutionary ecology, and yet it remains difficult to infer the action of evolutionary processes from patterns in the distribution of rates of diversification among related taxa. Using data from helminth parasite communities in 76 species of birds and 114 species of mammals, the influence of four factors that may either be associated with or modulate rates of parasite speciation were examined in a comparative analysis. Two measures of the relative number of congeneric parasite species per host species were used as indices of parasite diversification, and related to host body mass, host density, latitude, and whether the host is aquatic or terrestrial. The occurrence of congeneric parasites was not distributed randomly with respect to these factors. Aquatic bird species tended to harbour more congeneric parasites than terrestrial birds. Large-bodied mammal species, or those living at low latitudes, harboured more congeneric parasites than small-bodied mammals, or than those from higher latitudes. Host density had no apparent association with either measures of parasite diversification. These patterns, however, reflect only the present-day distribution of parasite diversification among host taxa, and not the evolutionary processes responsible for diversification, because the apparent effects of the factors investigated disappeared once corrections were made for host phylogeny. This indicates that features other than host body size, host density, latitude, and whether the habitat is terrestrial or aquatic, have been the key driving forces in the diversification of parasitic helminth lineages. This revised version was published online in July 2006 with corrections to the Cover Date.     

Can parasites synchronise the population fluctuations of sympatric tetraonids? –examining some minimum conditions

Sympatric populations of tetraonid birds tend to fluctuate in synchrony, at least on local scales. If shared parasites among sympatric populations of different tetraonid species are to operate as a local, synchronizing factor for population fluctuations at least two conditions should be met: i) the host species should share the same (or similar) parasite species, and ii) geographical location should contribute significantly more to the variation in the parasite species composition and abundance than differences among host species. We examined these conditions among subpopulations of sympatric willow ptarmigan and rock ptarmigan and found that host species shared a common pool of parasite species, and geographic location was more important than host species in determining parasite abundance across locations. There was no time lag between density oscillations in the two hosts, suggesting a symmetrical pattern of transmission and maintenance of parasites within habitats governed by the density of hosts and the environment. These findings are consistent with the idea that parasites may play a role in generating synchronous density fluctuations, but large scale experiments are needed to verify this hypothesis.     

Phylogenetic signals in host–parasite associations for Neotropical bats and Nearctic desert rodents

Hosts and their parasites have strong ecological and evolutionary relationships, with hosts representing habitats and resources for parasites. In the present study, we use approaches developed to evaluate the statistical dependence of species trait values on phylogenetic relationships to determine whether host–parasite relationships (i.e. parasite infections) are contingent on host phylogeny. If host–parasite relationships are contingent on the ability of hosts to provide habitat or resources to parasites, and if host phylogeny is an effective surrogate for among‐host variation in habitat and resource quality, host–parasite relationships should evince phylogenetic signals (i.e. be contingent on host phylogeny). Because the strength of ecological relationships between parasites and their hosts may affect the likelihood of phylogenetic signals occurring in host–parasite relationships, we hypothesized that (1) host specificity would be positively correlated with the strength of phylogenetic signals and (2) the strength of phylogenetic signals will be greater for parasites that rely more on their host throughout their life cycle. Analyses were conducted for ectoparasites from tropical bats and for ectoparasites, helminths, and coccidians from desert rodents. Phylogenetic signals were evaluated for parasite presence and for parasite prevalence. The frequency of phylogenetic signal occurrence was similar for parasite presence and prevalence, with a signal detected in 24–27% of cases at the species level and in 67% and 15% of cases at the genus level for parasites of bats and rodents, respectively. No differences in signal strength or the likelihood of detecting a signal existed between groups of parasites. Phylogenetic signal strength was correlated with host specificity, suggesting that mechanisms increasing host specificity also increase the likelihood of a phylogenetic signal in host use by parasites. Differences in the transmission mode did not affect signal strength or the likelihood of detecting a signal, indicating that variation in host switching opportunities associated with the transmission mode does not affect signal strength.     

Spatial heterogeneity in recruitment of larval trematodes to snail intermediate hosts

Spatial variation in parasitism is commonly observed in intermediate host populations. However, the factors that determine the causes of this variation remain unclear. Increasing evidence has suggested that spatial heterogeneity in parasitism among intermediate hosts may result from variation in recruitment processes initiated by definitive hosts. I studied the perching and habitat use patterns of wading birds, the definitive hosts in this system, and its consequences for the recruitment of parasites in snail intermediate hosts. Populations of the mangrove snail, Cerithidea scalariformis, collected from mangrove swamps on the east coast of central Florida are parasitized by a diverse community of trematode parasites. These parasites are transmitted from wading birds, which frequently perch on dead mangrove trees. I tested the hypothesis that mangrove perches act as transmission foci for trematode infections of C. scalariformis and that the spatial variation of parasitism frequently observed in this system is likely to emanate from the distribution of wading birds. On this fine spatial scale, definitive host behaviors, responding to a habitat variable, influenced the distribution, abundance and species composition of parasite recruitment to snails. This causal chain of events is supported by regressions between perch density, bird abundance, bird dropping density and ultimately parasite prevalence in snails. Variation between prevalence of parasites in free-ranging snails versus caged snails shows that while avian definitive hosts initiate spatial patterns of parasitism in snails through their perching behaviors, these patterns may be modified by the movement of snail hosts. Snail movement could disperse their associated parasite populations within the marsh, which may potentially homogenize or further increase parasite patchiness initiated by definitive hosts.     

Conservation implications of competition between generalist and specialist rodents in Mediterranean afforested landscape

Density dependent habitat selection at the community level is regarded as a major determinant of biodiversity at the local scale, and data on these processes and how they are affected by human activities is highly applicable to conservation. By studying the competitive relationships between a specialist and a generalist we can acquire valuable insights about how different environmental elements determine species abundance and distribution and consequently biodiversity. Here we describe a study of density dependent processes that determine the community structure of two rodents: a specialist—the broad toothed mouse (Apodemus mystacinus), and a generalist—the common spiny mouse (Acomys cahirinus) in a Mediterranean maqui habitat, and how this structure is impacted by anthropogenic planting of pine stands. We carried out two field experiments: The first, based on open field trapping, looking at how rodent communities change with habitat structure. The second experiment was an enclosure study aimed at validating the habitat preferences and competitive relationship between the specialist and the generalist. We identified asymmetric competition relationships in which the specialist was dominant over the generalist. Competition intensity was lower in maqui with >10% oak cover, although both species abundances were high. Competition was found only during the limiting season (summer). Based on these findings we produced management recommendations to keep indigenous small mammals’ biodiversity high. Density dependent habitat selection processes play a central role in determining biodiversity, and understanding the mechanisms motivating these processes is needed if alterations in biodiversity in response to human disturbance are to be understood.     

The underrated importance of predation in transmission ecology of direct lifecycle parasites

Most parasites with complex life cycles exploit trophic webs to pass from host to host in order to develop and, eventually, reproduce. Thus predation constitutes the necessary route for transmission. Conversely, the transmission of parasites that use a single host to develop and reproduce should be, in principle, not particularly affected by host trophic ecology. Here I challenge this view, showing that predation may be relevant also for direct lifecycle parasites. I used a large dataset of fish trophic interactions to investigate if the degree of monogenean species overlap in predators and prey deviated from randomness. I demonstrated that predators and prey often share more monogenean parasite genera than explained by host habitat ecology, geographical distribution and phylogeny. This suggests that predation may play an important role in promoting monogenean host range expansion. In addition, a non‐negligible proportion of considered prey–predator pairs showed a significantly high overlap in their monogenean parasites at the species level. This may indicate a tendency of some monogenean parasites to evolve transmission strategies targeted towards host interactions. If this hypothesis is true, these monogenean parasites would be much more vulnerable to co‐extinction than previously thought. Synthesis Predation is not expected to play an important role in the ecology and evolution of simple life cycle parasites. Yet, several predator fish tend to share with their prey more monogenean parasites than one would expect predicted from their geographical distribution, habitat preference, and or phylogenetic relationships. This suggests that some monogenean parasites have evolved transmission strategies more targeted towards host interactions than towards species‐specific traits. If this hypothesis is supported, it would have strong implications on host–parasite evolutionary ecology, primarily, suggesting the existence of peculiar situations where some parasites have evolved high specialized host finding behaviors to expand their host range.     

Ecology,not distance,explains community composition in parasites of sky-island Audubon’s Warblers

Haemosporidian parasites of birds are ubiquitous in terrestrial ecosystems, but their coevolutionary dynamics remain poorly understood. If species turnover in parasites occurs at a finer scale than turnover in hosts, widespread hosts would encounter diverse parasites, potentially diversifying as a result. Previous studies have shown that some wide-ranging hosts encounter varied haemosporidian communities throughout their range, and vice-versa. More surveys are needed to elucidate mechanisms that underpin spatial patterns of diversity in this complex multi-host multi-parasite system. We sought to understand how and why a community of avian haemosporidian parasites varies in abundance and composition across elevational transects in eight sky islands in southwestern North America. We tested whether bird community composition, environment, or geographic distance explain haemosporidian parasite species turnover in a widespread host that harbors a diverse haemosporidian community, the Audubon’s Warbler (Setophaga auduboni). We tested predictors of infection using generalized linear models, and predictors of bird and parasite community dissimilarity using generalized dissimilarity modeling. Predictors of infection differed by parasite genus: Parahaemoproteus was predicted by elevation and climate, Leucocytozoon varied idiosyncratically among mountains, and Plasmodium was unpredictable, but rare. Parasite turnover was nearly three-fold higher than bird turnover and was predicted by elevation, climate, and bird community composition, but not geographic distance. Haemosporidian communities vary strikingly at fine spatial scales (hundreds of kilometers), across which the bird community varies only subtly. The finer scale of turnover among parasites implies that their ranges may be smaller than those of their hosts. Avian host species should encounter different parasite species in different parts of their ranges, resulting in spatially varying selection on host immune systems. The fact that parasite turnover was predicted by bird turnover, even when considering environmental characteristics, implies that host species or their phylogenetic history plays a role in determining which parasite species will be present in a community.     

Local maladaptation in the soft scale insect Saissetia coffeae (Hemiptera: Coccidae)

Local adaptation has often been documented in herbivorous insects. The potential for local maladaptation in phytophagous insects, however, has not been widely considered. I performed a two-generation reciprocal cross-transplant experiment with the generalist soft scale insect Saissetia coffeae (Hemiptera: Coccidae) on two common species of host plants in rain forest habitat in Costa Rica. In this system, S. coffeae showed significant local maladaptation at the level of the host species. Lineages originally collected from Witheringia enjoyed a strong advantage over those collected from Lomariopsis when both sets of lineages were placed on Lomariopsis; however, when both sets of lineages were raised on Witheringia, their fitnesses were statistically indistinguishable. While some aspects of the biology of S. coffeae may impair its ability to adapt to local selection pressures, scale insects are often locally adapted on fine spatial scales, and local maladaptation is therefore especially surprising. Other documented cases of local maladaptation in parasites appear to be due to evolution on the part of the host. The possibility that hosts or natural enemies may place local genotypes at a disadvantage, producing a pattern of local maladaptation, is one that deserves more consideration in the context of plant-insect interactions.     

Some unsolved aspects of the parasitism problem

Various interpretations of the parasitism phenomenon exist. In this work data supporting the ecological understanding of its nature are presented. For parasitic species formed in the process of evolution the host organism has become the habitat (nutrition, multiplication), i.e., the environment, where interactions between the host and parasites are governed by ecological regularities. The consequences of this interactions are different (disease, asymptomatic infection), being secondary they reflect the result of concrete interactions. For this reason parasitism should not be identified in terms of only one consequence of such interaction--the development of pathologic processes.     

Host behaviour drives parasite genetics at multiple geographic scales: population genetics of the chewing louse,Thomomydoecus minor

Pocket gophers and their symbiotic chewing lice form a host–parasite assemblage known for a high degree of cophylogeny, thought to be driven by life history parameters of both host and parasite that make host switching difficult. However, little work to date has focused on determining whether these life histories actually impact louse populations at the very fine scale of louse infrapopulations (individuals on a single host) at the same or at nearby host localities. We used microsatellite and mtDNA sequence data to make comparisons of chewing‐louse (Thomomydoecus minor) population subdivision over time and over geographic space where there are different potential amounts of host interaction surrounding a zone of contact between two hybridizing pocket‐gopher subspecies. We found that chewing lice had high levels of population isolation consistent with a paucity of horizontal transmission even at the very fine geographic scale of a single alfalfa field. We also found marked genetic discontinuity in louse populations corresponding with host subspecies and little, if any, admixture in the louse genetic groups even though the lice are closely related. The correlation of louse infrapopulation differentiation with host interaction at multiple scales, including across a discontinuity in pocket‐gopher habitat, suggests that host behaviour is the primary driver of parasite genetics. This observation makes sense in light of the life histories of both chewing lice and pocket gophers and provides a powerful explanation for the well‐documented pattern of parallel cladogenesis in pocket gophers and chewing lice.