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Transitional zones, such as edge habitat, are key landscapes for investigating biodiversity. "Soft edges" are permeable corridors that hosts can cross, while "hard edges" are impermeable borders that hosts cannot pass. Although pathogen transmission in the context of edges is vital to species conservation, drivers of host-parasite relationships in ecological edges remain poorly understood. Thus, we defined a framework for testing hypotheses of host-parasite interactions in hard and soft edges by (1) characterizing hard and soft edges from both the host and parasite perspectives, (2) predicting the types of parasites that would be successful in each type of edge, and (3) applying our framework to species invasion fronts as an example of host-parasite relationships in a soft edge. Generally, we posited that parasites in soft edges are more likely to be negatively affected by habitat fragmentation than their hosts because they occupy higher trophic levels but parasite transmission would benefit from increased host connectivity. Parasites along hard edges, however, are at higher risk of local extinction due to host population perturbations with limited opportunity for parasite recolonization. We then used these characteristics to predict functional traits that would lead to parasite success along soft and hard edges. Finally, we applied our framework to invasive species fronts to highlight predictions regarding host connectivity and parasite traits in soft edges. We anticipate that our work will promote a more complete discussion of habitat connectivity using a common framework and stimulate empirical research into host-parasite relationships within ecological edges and transitional zones.

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.

We tested whether and how the maternal environment (i.e. host species exploited by a mother), rearing conditions (i.e. host species exploited by her offspring) or both (i.e. matches and mismatches in host species exploited by a mother and her offspring) affect reproductive performance in the offspring. We experimentally manipulated maternal and rearing environments in two generalist fleas (Xenopsylla conformis and Xenopsylla ramesis) implementing a factorial cross-rearing design. Mothers exploited either the principal host (PH) or auxiliary hosts that were either closely (CAH) or distantly related (DAH) to the PH. After six generations of infesting a given host species, we cross-reared fleas within and between host species. These fleas reproduced and we measured their reproductive performance both quantitatively (i.e. egg number) and qualitatively (i.e. egg size, development time, body size of the next generation). We found that identity of the host a flea was reared on (=actual host) had the strongest effect on its performance. Individuals reared on the PH performed considerably better than those reared on either auxiliary host. Moreover, fleas reared on a CAH performed better than those reared on a DAH. Actual host identity also had a stronger effect on reproductive performance in X. ramesis than in X. conformis. Nevertheless, there was no difference in performance between match and mismatch maternal and actual host identities. We conclude that rearing environment has the strongest effect on fitness in generalist parasites. Moreover, phylogenetic distance between an auxiliary host and the PH determines the level of suitability of the former.

We tested whether biogeographic patterns characteristic of species diversity and composition may also apply to community assembly by investigating geographic variation in the pattern (PSA) (aggregation versus segregation) and strength of species associations (SSA) in flea and mite communities harbored by small mammalian hosts in Western Siberia. We asked whether (a) there is a relationship between latitude and PSA or SSA and (b) similarities in PSA or SSA follow a distance decay pattern or if they are better explained by variation in environmental factors (altitude, amount of vegetation, precipitation, and air temperature). We used a sign of a co-occurrence metric (the C-score) as an indicator of PSA and its absolute standardized value as a measure of SSA. We analyzed data using logistic and linear models, generalized dissimilarity modeling (GDM), and a logistic version of the multiple regression on distance matrices (MRM). The majority of the C-scores of the observed presence/absence matrices indicated a tendency to species aggregation rather than segregation. No effect of latitude on PSA or SSA was found. The dissimilarity in PSA was affected by environmental dissimilarity in mite compound communities only. A relatively large proportion of the deviance of spatial variation in SSA was explained by the GDMs in infracommunities, but not component communities, and in only three (of seven) and two (of eight) host species of fleas and mites, respectively. The best predictors of dissimilarity in SSA in fleas differed between host species, whereas the same factor (precipitation) was the best predictor of dissimilarity in SSA in mites. We conclude that PSA and SSA in parasite communities rarely conform to biogeographic rules. However, when a biogeographic pattern is detected, its manifestation differs among hosts and between ectoparasite taxa.

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.

We studied the effects of variation in environmental, host-associated and spatial factors on variation in compositional, phylogenetic/taxonomic and functional facets of beta-diversity in fleas and gamasid mites parasitic on small mammals and asked whether (a) the importance of these factors as drivers of beta-diversity differs among its multiple facets and (b) the effects of variation in environment, hosts and space on beta-diversity variation differ between the two ectoparasite taxa. To understand the relative effects of each group of predictors, we used a distance-based redundancy analysis and variation partitioning. The greatest portions of variation in the compositional beta-diversity of fleas were equally explained by host-associated and spatial predictors, whereas variation in host species composition contributed the most to variation in the compositional beta-diversity of mites. Variation in the phylogenetic (i.e. based on phylogenetic tree) beta-diversity of fleas was mainly due to variation in the phylogenetic composition of host communities, while the taxonomic (i.e. based on Linnean taxonomy) beta-diversity of mites was influenced by environmental variation. Unique contributions of spatial and environmental variation explained most of the variation in functional beta-diversity and its species replacement (= turnover) component (i.e. beta-diversity explained by replacement of species alone) in fleas and mites, respectively. Variation in the richness difference component (i.e. beta-diversity explained by species loss/gain alone) of functional beta-diversity was mainly affected by either variation in the functional composition of host assemblages (fleas) or its joint action with environmental variables (mites). We conclude that the pattern of the relative effects of environmental, host-associated and spatial factors on beta-diversity is context-dependent and may differ among different facets of beta-diversity, among different beta-diversity components and also among taxa dependent on biological affinities.

Sex-biased parasitism highlights potentially divergent approaches to parasite resistance resulting in differing energetic trade-offs for males and females; however, trade-offs between immunity and self-maintenance could also depend on host body condition. We investigated these relationships in the big brown bat, Eptesicus fuscus, to determine if host sex or body condition better predicted parasite resistance, if testosterone levels predicted male parasite burdens, and if immune parameters could predict male testosterone levels. We found that male and female hosts had similar parasite burdens and female bats scored higher than males in only one immunological measure. Top models of helminth burden revealed interactions between body condition index and agglutination score as well as between agglutination score and host sex. Additionally, the strength of the relationships between sex, agglutination, and helminth burden is affected by body condition. Models of male parasite burden provided no support for testosterone predicting helminthiasis. Models that best predicted testosterone levels did not include parasite burden but instead consistently included month of capture and agglutination score. Thus, in our system, body condition was a more important predictor of immunity and worm burden than host sex.

Nested subsets occur in ecological communities when species-poor communities are subsets of larger, species-rich communities. Understanding this pattern can help elucidate species colonization abilities, extinction risks, and general structuring of biological communities. Here, we evaluate nestedness in a poorly studied host–parasite system, bats and their helminths, across the Japanese archipelago and within its different bioclimatic regions. We hypothesized that (1) if helminth communities are nested across geographic sites at the level of the archipelago, then broad-scale processes, like colonization-extinction dynamics, mainly structure parasite assemblages; (2) if helminth communities are nested across geographic sites at the level of the bioclimatic region, then fine-scale environmental variation plays a significant role in species nestedness; (3) if helminth community nestedness mirrors host species nestedness, then communities are nested because the habitats they occupy are nested; and (4) if nestedness does not occur or if it is not correlated with any geographical or host data, then passive sampling could be responsible for the patterns of parasite assemblage in our sample. We found that helminth communities were nested across host species throughout the archipelago but, when considering each bioclimatic region, helminths in only one region were significantly more nested than the null model. Helminth communities were also nested across sites within all four bioclimatic regions. These results suggest that helminths form nested subsets across the archipelago due to broad-scale processes that reflect the overall lineages of their mammalian hosts; however, at the regional scale, environmental processes related to nestedness of their habitats drive parasite community nestedness.

Parents may alter offspring phenotype depending on the type of environment they encounter. Parasitism is a common stressor; therefore, maternal reproductive investment could change in response to parasitic infection. However, few experiments have investigated the relationship between parasitism and maternal investment, whereas earlier field studies provided contradictory evidence. We investigated number, sex ratio, and growth of offspring in two rodent species, solitary altricial Meriones crassus and social precocial Acomys cahirinus, exposed to parasitism by fleas Xenopsylla ramesis and Parapulex chephrenis. No effect of treatment on litter size or sex ratio of a litter was found in either rodent species. Flea parasitism was found to affect pre-weaning body mass gain in M. crassus, but not in A. cahirinus pups. Furthermore, it appeared that female M. crassus invested resources into their offspring differently in dependence of litter size. In small litters (1–3 offspring), pups from infested females gained more body mass before weaning than pups from uninfested mothers. However, this trend was reversed in females with large litters indicating that parasitized females have a finite amount of resources with which to provision their young. Thus, M. crassus mothers parasitized by fleas seemed to receive some sort of external cues (e.g., stress caused by infestation) that prompted them to alter offspring provisioning, depending on species-specific possibilities and constraints. Therefore, parasites could be a mediator of environmentally induced maternal effects and offspring provisioning may have adaptive value against parasitism.

Evaluating host resistance via parasite fitness helps place host-parasite relationships within evolutionary and ecological contexts; however, few studies consider both these processes simultaneously. We investigated how different levels of parasite pressure affect parasite mortality and reproductive success in relationship to host defense efforts, using the rodent Gerbillus nanus and the flea Xenopsylla conformis as a host-parasite system. Fifteen immune-naive male rodents were infested with 20, 50, or 100 fleas for four weeks. During this time number of new imagoes produced per adult flea (our flea reproductive output metric), flea mortality, and change in circulating anti-flea immunoglobulin G (our measure of adaptive immune defense) were monitored. Three hypotheses guided this work: (1) increasing parasite pressure would heighten host defenses; (2) parasite mortality would increase and parasite reproductive output would decrease with increasing investment in host defense; and (3) hosts under high parasite pressure could invest in behavioral and/or immune responses. We predicted that at high infestation levels (a) parasite mortality would increase; (b) flea reproductive output per individual would decrease; and (c) host circulating anti-flea antibody levels would increase. The hypotheses were partially supported. Flea mortality significantly increased and flea reproductive output significantly decreased as flea pressure increased. Host adaptive immune defense did not significantly change with increasing flea pressure. Therefore, we inferred that investment in host behavioral defense, either alone or in combination with density-dependent effects, may be more efficient at increasing flea mortality and decreasing flea reproductive output than antibody production during initial infestation in this system.