Dissimilarity in flea and host assemblages and their interaction networks along a spatial distance gradient: different patterns revealed by different network dissimilarity metrics.

We investigated the distance-decay pattern (an increase in dissimilarity with increasing geographic distance) in regional assemblages of fleas and their small mammalian hosts, as well as their interaction networks, in four biogeographic realms. Dissimilarity of assemblages (ßtotal) was partitioned into species richness differences (ßrich) and species replacement (ßrepl) components. Dissimilarity of networks was assessed using two metrics: (a) whole network dissimilarity (ßWN) partitioned into species replacement (ßST) and interaction rewiring (ßOS) components and (b) D statistics, measuring dissimilarity in the pure structure of the networks, without using information on species identities and calculated for hosts-shared-by-fleas networks (Dh) and fleas-shared-by-hosts networks (Df). We asked whether the distance-decay pattern (a) occurs among interactor assemblages or their interaction networks; (b) depends on the network dissimilarity metric used; and (c) differs between realms. The ßtotal and ßrepl of flea and host assemblages increased with distance in all realms except for host assemblages in the Afrotropics. ßrich for flea and host assemblages increased with distance in the Nearctic only. In networks, ßWN and ßST demonstrated a distance-decay pattern, whereas ßOS was mainly spatially invariant except in the Neotropics. Correlations of Dh or Df and geographic distance were mostly non-significant. We conclude that investigations of dissimilarity in interaction networks should include both types of dissimilarity metrics (those that consider partner identities and those that consider the pure structure of networks). This will allow elucidating the predictability of some facets of network dissimilarity and the unpredictability of other facets.

Relationships between functional alpha and beta diversities of flea parasites and their small mammalian hosts.

We studied the relationships between functional alpha and beta diversities of fleas and their small mammalian hosts in 4 biogeographic realms (the Afrotropics, the Nearctic, the Neotropics and the Palearctic), considering 3 components of alpha diversity (functional richness, divergence and regularity). We asked whether (a) flea alpha and beta diversities are driven by host alpha and beta diversities; (b) the variation in the off-host environment affects variation in flea alpha and beta diversities; and (c) the pattern of the relationship between flea and host alpha or beta diversities differs between geographic realms. We analysed alpha diversity using modified phylogenetic generalized least squares and beta diversity using modified phylogenetic generalized dissimilarity modelling. In all realms, flea functional richness and regularity increased with an increase in host functional richness and regularity, respectively, whereas flea functional divergence correlated positively with host functional divergence in the Nearctic only. Environmental effects on the components of flea alpha diversity were found only in the Holarctic realms. Host functional beta diversity was invariantly the best predictor of flea functional beta diversity in all realms, whereas the effects of environmental variables on flea functional beta diversity were much weaker and differed between realms. We conclude that flea functional diversity is mostly driven by host functional diversity, whereas the environmental effects on flea functional diversity vary (a) geographically and (b) between components of functional alpha diversity.

Nestedness of flea assemblages harboured by small mammalian hosts revisited: phylogenetic and functional nestedness do not follow compositional nestedness.

We studied compositional, phylogenetic, and functional nestedness in the flea assemblages of 14 host species across regions. Our main questions were (a) are a host's flea assemblages compositionally, phylogenetically, or functionally nested? (b) Do similar processes drive these nestedness facets? (d) Are a host's biological traits associated with nestedness of its flea assemblages? Rows of host matrices were ordered by decreasing species richness/the sum of the branch lengths of a phylogenetic tree/functional dendrogram or by decreasing region area or by increasing distance from the centre of a host's geographic range. None of the matrices sorted by species richness/sum of branch lengths were nested from a compositional perspective, but they were significantly nested from phylogenetic and functional perspectives. Compositional, phylogenetic, and functional nestedness of matrices sorted by region area or by distance from the host's geographic range centre varied between hosts. In some hosts, flea assemblages were nested from all three perspectives independently of how matrix rows were sorted, whereas in other hosts, the occurrence of significant nestedness depended on the order of the matrix rows. The degree of phylogenetic and functional nestedness for matrices sorted by the sum of branch lengths was associated with a host species' morphoecological traits and the latitude of its geographic range. We conclude that consideration of nestedness based solely on species composition does not allow a comprehensive understanding of the patterns of parasite community structure. Nestedness should also be considered from phylogenetic and functional perspectives.

Functional similarity affects similarity in partner composition in flea-mammal networks.

Functional signal in an interaction network is a phenomenon in which species resembling each other in their traits interact with similar partners. We tested the functional signal concept in realm-specific and regional flea-host networks from four biogeographic realms and asked whether the species composition of (a) host spectra and (b) flea assemblages is similar between functionally similar flea and host species, respectively. Analogously to testing for phylogenetic signal, we applied Mantel tests to investigate the correlation between flea or host functional distances calculated from functional dendrograms and dissimilarities in sets of interacting partners. In all realm-specific networks, functionally similar fleas tended to exploit similar hosts often belonging to the same genus, whereas functionally similar hosts tended to harbour similar fleas, again often belonging to the same genus. The strength of realm-specific functional signals and the frequency of detecting a significant functional signal in the regional networks differed between realms. The frequency of detecting a significant functional signal in the regional networks correlated positively with the network size for fleas and with the number of hosts in a network for hosts. A functional signal in the regional networks was more frequently found for hosts than for fleas. We discuss the mechanisms behind the functional signal in both fleas and their hosts, relate geographic functional signal patterns to the historic biogeography of fleas and conclude that functional signals in the species composition of host spectra for fleas and of flea assemblages for hosts result from the interplay of evolutionary and ecological processes.

Phylogenetic patterns in regional flea assemblages from 6 biogeographic realms: strong links between flea and host phylogenetic turnovers and weak effects of phylogenetic originality on host specificity.

We investigated phylogenetic patterns in flea assemblages from 80 regions in 6 biogeographic realms and asked whether (a) flea phylogenetic turnover is driven by host phylogenetic turnover, environmental dissimilarity or geographic distance; (b) the relative importance of these drivers differs between realms; and (c) the environmental drivers of flea phylogenetic turnover are similar to those of host phylogenetic turnover. We also asked whether the phylogenetic originality of a flea species correlates with the degree of its host specificity and whether the phylogenetic originality of a host species correlates with the diversity of its flea assemblages. We found that host phylogenetic turnover was the best predictor of flea phylogenetic turnover in all realms, whereas the effect of the environment was weaker. Environmental predictors of flea phylogenetic turnover differed between realms. The importance of spatial distances as a predictor of the phylogenetic dissimilarity between regional assemblages varied between realms. The responses of host turnover differed from those of fleas. In 4 of the 6 realms, geographic distances were substantially better predictors of host phylogenetic turnover than environmental gradients. We also found no general relationship between flea phylogenetic originality and its host specificity in terms of either host species richness or host phylogenetic diversity. We conclude that flea phylogenetic turnover is determined mainly by the phylogenetic turnover of their hosts rather than by environmental gradients. Phylogenetic patterns in fleas are manifested at the level of regional assemblages rather than at the level of individual species.

Metabolic rate and ecological traits of ectoparasites: a case study with seven flea species from the Negev Desert.

We studied the relationship between fleas' metabolic rate and their ecological traits, using data on standard metabolic rate (SMR), mean abundance, host specificity, and geographic range size in males and females of seven desert flea species. SMR was measured via mass-specific CO2 emission, whereas host specificity was measured as (a) the mean number of host species used by a flea per region in regions where this flea was recorded; (b) the total number of host species a flea exploited across its geographic range; and (c) the phylogenetic diversity of the flea's hosts. To control for confounding effects of phylogeny when analysing data on multiple species, we applied the Phylogenetic Generalised Least Squares (PGLS) model. We found that the only ecological trait significantly correlating with flea SMR was the phylogenetic diversity of hosts utilized by a flea across its geographic range. The strength of the association between SMR and host phylogenetic diversity was higher in male than in female fleas. We explain the relationship between flea SMR and their host specificity by the necessity of host-opportunistic species to compensate for the high energetic cost of neutralizing multiple defences from multiple hosts by increased SMR.

The species composition of local flea assemblages at a small scale in two South American regions is predominantly driven by niche-based mechanisms.

We applied a step-down factor analysis (SDFA) and multi-site generalised dissimilarity modelling (MS-GDM) to local flea communities harboured by small mammals (i.e., collected at small sampling sites over a short time period) in two South American regions (Patagonia and the Northwestern Argentina) with the aim of understanding whether these communities were assembled via niche-based or dispersal-based processes. The SDFA allows us to determine whether clusters of flea assemblages across different types of climates, vegetation and soils can be distinguished (suggesting niche-based assembly). MS-GDM allows us to determine whether a substantial proportion of the variation in flea species turnover is explained by specific climate-associated, vegetation-associated and soil-associated variables (indicating niche-based assembly) or host turnover (indicating dispersal-based assembly). Mapping of assemblages on climate, vegetation and soil maps, according to their loadings on axis 1 or axis 2 of the SDFA, did not provide clear-cut results. Clusters of similar loadings could be recognized within some, but not other, climate, vegetation and soil types. However, MS-GDM demonstrated that the effect of environmental variables (especially air temperature) on flea compositional turnover was much stronger than that of host turnover, indicating the predominance of niche-based processes in local community assembly. A comparison of our results with those on the mechanisms that drive species assembly in regional communities allows us to conclude that local and regional communities result from the joint action of niche-based and dispersal-based processes, with the former more important at a smaller spatial scale and the latter at a larger spatial scale.

Regional flea and host assemblages form biogeographic, but not ecological, clusters: evidence for a dispersal-based mechanism as a driver of species composition.

We used data on the species composition of regional assemblages of fleas and their small mammalian hosts from 6 biogeographic realms and applied a novel method of step-down factor analyses (SDFA) and cluster analyses to identify biogeographic (across the entire globe) and ecological (within a realm across the main terrestrial biomes) clusters of these assemblages. We found that, at the global scale, the clusters of regional assemblage loadings on SDFA axes reflected well the assemblage distribution, according to the biogeographic realms to which they belong. At the global scale, the cluster topology, corresponding to the biogeographic realms, was similar between flea and host assemblages, but the topology of subtrees within realm-specific clusters substantially differed between fleas and hosts. At the scale of biogeographic realms, the distribution of regional flea and host assemblages did not correspond to the predominant biome types. Assemblages with similar loadings on SDFA axes were often situated in different biomes and vice versa. The across-biome, within-realm distributions of flea vs host assemblages suggested weak congruence between these distributions. Our results indicate that dispersal is a predominant mechanism of flea and host community assembly across large regions.

Host phylogeny and ecology, but not host physiology, are the main drivers of (dis)similarity between the host spectra of fleas: application of a novel ordination approach to regional assemblages from four continents.

We investigated the patterns of phylogenetic and functional (dis)similarity in the species composition of host spectra between co-habitating generalist flea species in regional assemblages from four continents (Europe, Asia, North America and Africa) using a recently developed ordination approach (Double Similarity Principal Component Analysis). From the functional perspective, we considered physiological [body mass and basal metabolic rate (BMR)] and ecological (shelter depth and complexity) host traits. We asked (a) whether host phylogeny, physiology or ecology is the main driver of (dis)similarities between flea host spectra and (b) whether the patterns of phylogenetic and functional (dis)similarity in host spectra vary between flea assemblages from different continents. Phylogenetic similarity between the host spectra was highest in Africa, lowest in North America and moderate in Europe and Asia. In each assemblage, phylogenetic clusters of hosts dominating in the host spectra could be distinguished. The functional similarity between the host spectra of co-occurring fleas was low for shelter structure in all assemblages and much higher for body mass and BMR in three of the four assemblages (except North America). We conclude that host phylogeny and shelter structure are the main drivers of (dis)similarity between the host spectra of co-habitating fleas. However, the effects of these factors on the patterns of (dis)similarity varied across continents.

Fitness consequences of host colonization in two generalist fleas: Context-dependency and the effect of spatial co-occurrence.

We studied the fitness consequences of colonizing a novel host by experimental lines of fleas (Synosternus cleopatrae and Xenopsylla ramesis) maintained for 18-22 generations on the principal or novel (sympatric or allopatric) hosts via number, developmental success and size of the offspring of the fleas exploiting these hosts. We asked whether (a) fitness on non-principal hosts increases after prolonged maintenance; (b) the colonization success depends on the spatial co-occurrence of a flea and a host and (c) colonization of a novel host is accompanied by a decreased ability to exploit an original host. The ability of fleas to colonize novel hosts differed between species, with S. cleopatrae, but not X. ramesis, increasing its offspring production on novel hosts. Spatial co-occurrence did not affect colonization success. Maintenance on an alternative host was not accompanied by decreased adaptation to the original host. When fleas returned to the original host, their reproductive output was higher than that of their ancestors. We conclude that the success of colonizing a novel host is (a) context-dependent and varies between flea and host species and (b) not accompanied by the loss of ability to exploit an ancestral host but may lead to an increase in this ability.

Temporal variation of metacommunity structure in arthropod ectoparasites harboured by small mammals: the effects of scale and climatic fluctuations.

We applied the elements of metacommunity structure (EMS) approach and studied the temporal dynamics of metacommunity structure in arthropod ectoparasites (fleas, gamasid mites and ixodid ticks) harboured by six small mammalian hosts sampled for three decades in the same locality in Western Siberia at three hierarchical scales (inframetacommunities, component metacommunities and a compound metacommunity). All metacommunities were positively coherent. Inframetacommunity structures varied across sampling periods in all host species. The main structural pattern in an inframetacommunity of the same host varied across sampling times but was mostly characterized by clumped species distributions (Clementsian, Gleasonian and their quasi-versions). Component metacommunities in five of the six host species were characterized by either a Clementsian or a quasi-Clementsian distribution. In four of the six host species, this pattern was driven by mite distribution. The temporal structure of compound metacommunity was characterized by a Clementsian pattern. In contrast to the majority of component metacommunities, this pattern was driven by fleas, whereas the temporal structure of gamasid mite compound metacommunities demonstrated a Gleasonian distribution. The temporal gradient in infracommunity composition was not associated with temporal changes in either air temperature or precipitation, whereas the precipitation gradient was positively correlated with the structure of component (in five host species) and compound metacommunities. In conclusion, the best-fit metacommunity structure of ectoparasites varies temporally due to temporal changes in distribution patterns that can be associated with year-to-year climatic variation, affecting both hosts and parasites.

Dark host specificity in two ectoparasite taxa: repeatability, parasite traits, and environmental effects.

We applied the concept of dark diversity (species that may potentially inhabit a locality but are absent) to the host spectrum of a parasite and defined it as dark host specificity (DHS). We studied the trait-associated and geographic patterns of dark host specificity in fleas and gamasid mites parasitic on small mammals, asking the following questions: (a) Is dark host specificity repeatable across populations of the same species? (b) Is it associated with morphological and/or ecological species traits? (c) What are the factors associated with geographical variation in the DHS among populations of the same species? The DHS was repeatable within species with a large proportion of variance among samples, accounted for by differences between species. The average DHS of fleas, but not mites, was affected by parasite traits, with the DHS being higher in fleas with larger geographic ranges, higher characteristic abundance levels, and summer reproduction peaks. In the majority of ectoparasites, the regional DHS decreased with an increase in either structural or phylogenetic host specificity. The associations between the DHS and the environmental or host-associated characteristics of a region were revealed in a few species (eight of 22 fleas and three of 12 mites). The DHS decreased with (a) an increase in air temperature in two fleas, (b) a decrease in precipitation in two fleas, and (c) an increase in regional host species richness (in three fleas and three mites). Overall, our results suggest that dark host specificity in arthropod ectoparasites is a species-specific character associated, to a large extent, with the breadth of their host-related niches, while the influences of parasite traits and local environmental conditions are minor.

Dispersal-based versus niche-based processes as drivers of flea species composition on small mammalian hosts: inferences from species occurrences at large and small scales.

Biological communities may be assembled by both niche-based and dispersal-based (= historic) processes with the relative importance of these processes in community assembly being scale- and context-dependent. To infer whether (a) niche-based or dispersal-based processes play the main role in the assembly of flea communities parasitic on small mammals and whether (b) the main processes of flea community assembly are scale-dependent, we applied a novel permutation-based algorithm (PER-SIMPER) and the dispersal-niche continuum index (DNCI), to data on the species incidence of fleas and their hosts at two spatial scales. At the larger (continental) scale, we analysed flea communities in four biogeographic realms across adjacent continental sections. At the smaller (local) scale, we considered flea communities across two main regions (lowlands and mountains) and seven habitat types within Slovakia. Our analyses demonstrated that species composition of fleas and their small mammalian hosts depended predominantly on historical processes (dispersal) at both scale. This was true for the majority of biogeographic realms at continental scale (except the Nearctic) and both regions at local scale. Nevertheless, strong niche-based assembly mechanism was found in the Nearctic assemblages. At local scale, the intensity of dispersal processes was weaker and niche-driven processes were stronger between habitats within a region than between mountain and lowland regions. We provide historical and ecological explanations for these patterns. We conclude that the assembly of compound flea communities is governed, to a great extent, by the dispersal processes acting on their hosts and, to a lesser extent, by the niche-based processes.

Spatial and temporal variation of compositional, functional, and phylogenetic diversity in ectoparasite infracommunities harboured by small mammals.

We studied patterns of compositional, functional, and phylogenetic α- and ß-diversity in flea and gamasid mite infracommunities of small Siberian mammals, taking into account host-associated (species) and environmental (biome or sampling period) factors. We asked: (a) How do these factors and their interactions affect infracommunity diversity? (b) Does infracommunity composition, in terms of species, traits, and phylogenetic lineages, deviate from random? (c) Are species, traits, and phylogenetic lineages in infracommunities clustered or overdispersed?, and (d) Do patterns of diversity differ between the three diversity facets and/or the two ectoparasite taxa? We found that the α-diversity of infracommunities was strongly affected by host species, biome, and sampling period. The highest proportion of infracommunity diversity in both taxa was associated with the interaction between either host species and biome or host species and sampling period. Infracommunities of both taxa within, as well as between, host species, biomes, and sampling periods were characterized by the clustering of species, traits and lineages. The patterns of the effects of host species, biome, and sampling period on infracommunity diversity were congruent among the three diversity facets in both fleas and mites. We conclude that the assembly patterns in ectoparasite infracommunities mirror those characteristics of component and compound communities.

Colonization of a novel host by fleas: changes in egg production and egg size.

We studied the success of fleas, Synosternus cleopatrae and Xenopsylla ramesis, in switching to a novel host by establishing experimental lines maintained on different hosts for 18 generations. Fleas fed on principal (P-line) or novel hosts, either sympatric with (S-line) or allopatric to (A-line) a flea and its principal host, then we assessed their reproductive performance via the number and size of eggs. We compared reproductive performance between hosts within a line and between lines within a host asking: (a) whether fleas adapt to a novel host species after multiple generations; (b) if yes, whether the pattern of adaptation differs between novel host species sympatric with or allopatric to a flea and its principal host; and (c) adaptation to a novel host is accompanied with a loss of success in exploitation of an original host. Fleas from the S- and A-lines increased their egg production on a novel host (except X. ramesis from the S-line). S. cleopatrae from the S-line but not the A-line increased egg size on a novel host, whereas X. ramesis from the A-line but not the S-line produced larger eggs from a novel host. We found no indication of a loss of reproductive performance on the original host while adapting to a novel host. We conclude that fleas are able to switch rapidly to a new host with the pattern of a switch to either sympatric or an allopatric host depending on the identities of both flea and host species.

Parasite counts or parasite incidences? Testing differences with four analyses of infracommunity modelling for seven parasite-host associations.

One of the challenges in studies of parasite community ecology is whether the input data for analyses should be parasite abundances/counts, i.e. count data (CD), or parasite incidences (presences/absences), i.e. incidence data (ID). We analysed species responses to environmental factors and species associations in the infracommunities of helminths and ectoparasites in four hosts from Europe (Sorex araneus and Myodes glareolus) and South Africa (Rhabdomys pumilio and Rhabdomys dilectus) and compared the results of four analyses [redundancy analysis (RD), RLQ analysis, joint species distribution modelling (JSDM) and Markov random fields (MRF)] that used either CD or ID as an input. In addition, we compared the differences between the CD and ID results of two analyses (JSDM and MRF) across parasite species between (a) host species within helminths and ectoparasites; (b) helminths and ectoparasites within a host species; and (c) parasite species with contrasting levels of intensity. The results of most analyses for the majority of parasite-host associations were qualitatively similar. However, models based on the ID input performed better than models based on the CD input in three out of four types of analyses (RDA, JSDM and MRF). The differences between the CD and ID models varied between host species (being the lowest in R. pumilio for JSDM and in S. araneus for MRF). However, they were not affected by the level of parasite intensity.

Harrison's rule scales up to entire parasite assemblages but is determined by environmental factors.

Harrison's rule states that parasite body size and the body size of their hosts tend to be positively correlated. After it was proposed a century ago, a number of studies have investigated this trend, but the support level has varied greatly between parasite/host associations. Moreover, while the rule has been tested at the individual species level, we still lack knowledge on whether Harrison's rule holds at the scale of parasite and host communities. Here, we mapped flea (parasites) and rodent (hosts) body sizes across Mongolia and asked whether Harrison's rule holds for parasite/host assemblages (i.e. whether a parasite's average body size in a locality is positively correlated with its host's average body size). In addition, we attempted to disentangle complex relationships between flea size, host size and environmental factors by testing alternative hypotheses for the determinants of fleas' body size variation. We gathered occurrence data for fleas and rodents from 2,370 sites across Mongolia, constructed incidence matrices for both taxa and calculated the average body sizes of fleas and their hosts over half-degree cells. Then, we applied a path analysis, accounting for spatial autocorrelation, trying to disentangle the drivers of the correlation between parasite and host body sizes. We found a strong positive correlation between average flea and host size across assemblages. Surprisingly though, we found that environmental factors simultaneously affected the body sizes of both fleas and hosts in the same direction, leading to a most likely deceptive correlation between parasite and host size across assemblages. We suggest that environmental factors may, to a great extent, reflect the environmental conditions inside the hosts' burrows where fleas develop and attain their adult body size, thus influencing their larval growth. Similarly, rodent body size is strongly influenced by air temperature, in the direction predicted by Bergmann's rule. If our findings are valid in other host-parasite associations, this may explain the dissenting results of both support and lack thereof for Harrison's rule.

Patterns of zeta diversity in ectoparasite communities harboured by small mammals at three hierarchical scales: taxon-invariance and scale-dependence.

We studied compositional turnover in communities of fleas and mites harboured by small mammals using zeta diversity metric (similarity between multiple communities) and asked whether the patterns of zeta diversity decline with an increase in the number of communities differ between taxa and hierarchical scales [infracommunities (parasite assemblages on individual hosts), component communities (parasite assemblages harboured by host populations), and compound communities (all parasite species in a locality)]. The average number of shared species declined with an increasing number of communities (zeta order). It attained zero at higher orders in infracommunities of both taxa with the shape of the zeta decline being best fitted by the negative exponential function, and the retention rate curves being modal. In contrast, zeta diversity values for compound communities of mites and fleas did not attain zero at higher zeta orders, and the form of the zeta decline was best fitted by the power-law function, whereas the retention rate curves were asymptotic. In component communities, the form of zeta decline was best fitted by either exponential or power-law function in dependence of whether communities were considered within a host across localities or across hosts within a locality and whether ubiquitous species were taken into account. Our main conclusions are that (a) the rules governing compositional turnover in parasite communities for the lowest and the highest hierarchical scales are taxon-invariant but scale-dependent and (b) species composition of infracommunities is mainly driven by stochastic assembly processed, whereas that of compound communities is mainly driven by niche-based processes.

Feeding performance on a novel host: no adaptation over generations and differential patterns in two flea species.

To model the colonization of a novel host by fleas, Synosternus cleopatrae and Xenopsylla ramesis, we established experimental lines maintained for 15 generations on a principal or a novel host (either co-occurring with a flea or not). We compared the blood meal size and the energy expended for digestion by fleas from the 15th generation of each line on these hosts between hosts within a line and between lines within a host asking (a) whether fleas adapt to a novel host (increased blood consumption/decreased energy expended for digestion); (b) if yes, whether this adaptation leads to the loss of ability to exploit an original host, and (c) whether the success of adaptation to a novel host depends on its ecological co-occurrence with a flea. The blood consumption and digestion energetics of fleas fed on the principal host differed from those on other hosts. The effect of the principal host on feeding performance differed between fleas, with S. cleopatrae consuming less blood and expending more energy for digestion on the principal than on any other host, whereas the opposite was true for X. ramesis. No changes in feeding performance on a novel host over generations were found. We propose several explanations for the lack of adaptation to a novel host over time. We explain the poor performance of S. cleopatrae on its principal host via its immune response mounting pattern. We argue that the principal host of a parasite is not necessarily the host on which the parasite demonstrates the best performance.

Flea infestation, social contact, and stress in a gregarious rodent species: minimizing the potential parasitic costs of group-living.

Both parasitism and social contact are common sources of stress that many gregarious species encounter in nature. Upon encountering such stressors, individuals secrete glucocorticoids and although short-term elevation of glucocorticoids is adaptive, long-term increases are correlated with higher mortality and deleterious reproductive effects. Here, we used an experimental host-parasite system, social rodents Acomys cahirinus and their characteristic fleas Parapulex chephrenis, in a fully-crossed design to test the effects of social contact and parasitism on stress during pregnancy. By analysing faecal glucocorticoid metabolites, we found that social hierarchy did not have a significant effect on glucocorticoid concentration. Rather, solitary females had significantly higher glucocorticoid levels than females housed in pairs. We found a significant interaction between the stressors of parasitism and social contact with solitary, uninfested females having the highest faecal glucocorticoid metabolite levels suggesting that both social contact and infestation mitigate allostatic load in pregnant rodents. Therefore, the increased risk of infestation that accompanies group-living could be outweighed by positive aspects of social contact within A. cahirinus colonies in nature.