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We tested for the effects of latitude and geographic range size (GRS) on body size, leg length, and sexual size dimorphism (SSD) across 103 species of fleas, taking into account phylogenetic between-species relationships. When the data on body size were combined for males and females, the positive correlation between body size and latitude, but not GRS, was revealed. When the analysis was restricted to one sex only, the effect of latitude appeared to be non-significant for females, whereas male body size increased with an increase in latitude. Intraspecific body size variation was not associated with either the latitude or the latitudinal span of the geographic range, independently of which data subset was analyzed. No evidence of association between size-independent tibia length and latitude was found for either females, males, or both sexes combined. The degree of SSD decreased with a decrease in latitude but was not affected by GRS. We conclude that macroecological patterns might be manifested differently in males and females. This should be kept in mind while searching for these patterns., (© 2022 The Authors. Integrative Zoology published by International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Biotic and abiotic stressors impose various fitness costs on individuals across a variety of taxa. In vertebrates, these stressors typically trigger complex neuroendocrine responses that stimulate glucocorticoid (GC) secretion from the adrenal cortex. Short-term elevation of GCs can be adaptive as it shifts energy toward physiological processes that cope with acute stressors; however, chronic increases in GC levels could have detrimental effects on fitness. Parasitism can be considered an important biotic stressor in nature and a possible cause of reproductive failure that could substantially affect an individual's fitness. Thus, we aimed to test the effects of parasitism and maternal stress, as measured by GCs, during pregnancy and the relationship between these variables and measures of reproductive output using a rodent-flea system. Female Egyptian spiny mice (Acomys cahirinus) were randomly assigned to flea (Parapulex chephrenis) infested or uninfested treatments before and during pregnancy. The offspring of these females were flea-free. Feces were collected at five time points during the experiment to determine maternal fecal glucocorticoid metabolite (FGCM) concentrations. Overall, infested females had lower FGCM levels during gestation but higher FGCM levels post-parturition and larger mass changes than uninfested females. Additionally, models related to pup quality and quantity often included some measure of maternal investment or body condition moderating relationships between infestation and stress. This suggests that flea parasitism or high GC levels alone might not significantly impact host reproduction but rather females can experience different effects depending on their level of investment, which could be limited by body condition and/or the number of pups present in a litter., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

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., (© 2020 British Ecological Society.)

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.

The β-diversity of fleas parasitic on small mammals in 45 regions of the Palearctic was partitioned into species [species contributions to β-diversity (SCBD)] and site ( = assemblage) contributions [local contributions to β-diversity (LCBD)]. We asked what are the factors affecting SCBD and LCBD and tested whether (a) variation in ecological, morphological, life history and geographic traits of fleas can predict SCBD and (b) variation in flea and host community metrics, off-host environmental factors, host species composition of flea assemblages can predict LCBD. We used spatial variables to describe geographic distribution of flea assemblages with various LCBD values. SCBD significantly increased with an increase in abundance and a decrease in phylogenetic host specificity of a flea as well as with size and latitude of its geographic range, but was not associated with any morphological/life history trait. LCBD of flea assemblages did not depend on either flea or host species richness or environmental predictors, but was significantly affected by compositional uniqueness ( = LCBD) of regional host assemblages and variables describing their species composition. In addition, variation in LCBD was also explained by broad-to-moderate-scale spatial variables. We conclude that SCBD of fleas could be predicted via their ecological and geographic traits, whereas LCBD of their assemblages could be predicted via host composition.

We studied the relationships between body size and (a) abundance and (b) host specificity in fleas parasitic on small mammals (rodents and shrews) in the Palearctic taking into account the confounding effect of phylogeny. We tested these relationships both across 127 flea species and within separate phylogenetic clades, predicting higher abundance and lower host specificity (in terms of the number or diversity of hosts used by a flea) in smaller species. We also tested for the relationships between body size and abundance separately for species that spend most of their lives on a host's body (the "body" fleas) and species that spend most of their lives in a host's burrow or nest (the "nest" fleas). A significant phylogenetic signal in body size was detected across all fleas, as well as in five of six separate clades. Across all fleas and in majority of phylogenetic clades, mean abundance significantly increased with an increase in body size. The same pattern was found for both the "body" and the "nest" fleas, although the slope of the relationship appeared to be steeper in the former than in the latter. Neither measure of host specificity demonstrated a significant correlation with body size regardless of the subset of flea species analysed. We explain higher abundance attained by larger flea species by higher fecundity and/or competitive advantage upon smaller species at larval stage. We conclude that the macroecological patterns reported to date in parasites are far from being universal.

The aims of this study were to determine whether sexual size dimorphism in fleas and gamasid mites (i) conforms to Rensch's rule (allometry of sexual size dimorphism) and (ii) covaries with sex ratio in infrapopulations (conspecific parasites harboured by an individual host), xenopopulations (conspecific parasites harboured by a population of a given host species in a locality) and suprapopulations (conspecific parasites harboured by an entire host community in a locality). Rensch's rule in sexual size dimorphism was tested across 150 flea and 55 mite species, whereas covariation between sexual size dimorphism and sex ratio was studied using data on ectoparasites collected from small mammalian hosts in Slovakia and western Siberia. For fleas, we controlled for the confounding effect of phylogeny. The slope of the linear regression of female size on male size was significantly smaller than 1 in fleas, but did not differ from 1 in mites. The proportion of males in flea infrapopulations significantly increased with an increase in the female-to-male body size ratio. The same was true for obligatory haematophagous mites. No relationship between sex ratio and sexual size dimorphism was found for xenopopulations of either taxon or for mite suprapopulations. However, when controlling for the confounding effect of phylogeny, a significant negative correlation between sex ratio and sexual size dimorphism was revealed for flea suprapopulations. We conclude that (i) some macroecological patterns differ between ectoparasite taxa exploiting the same hosts (allometry in sexual size dimorphism), whereas other patterns are similar (sexual size dimorphism-sex ratio relationship in infrapopulations), and (ii) some patterns are scale-dependent and may demonstrate the opposite trends in parasite populations at different hierarchical levels., (Copyright © 2018 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)