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.

Intraspecific variation of body size in fleas: effects of host sex and flea phenology.

We investigated the effects of host sex and flea phenology (estimated as periods of high versus low abundance) on individual body size in four fleas of small mammals. Amalaraeus penicilliger and Ctenophthalmus uncinatus are ectoparasites of the bank vole Myodes glareolus, whereas Doratopsylla dasycnema and Palaeopsylla soricis are ectoparasites of the common shrew Sorex araneus. We found significant effects of host sex and phenology on the body size of all flea species, although there was no general trend in the directions of these effects. Larger A. penicilliger were found on female hosts, whereas larger P. soricis were found on male hosts. In the remaining species, larger fleas were collected from male hosts during periods of high abundance (male C. uncinatus and female D. dasycnema) and from female hosts during periods of low abundance (male C. uncinatus). Regarding phenology, larger fleas were recorded during periods of either high (A. penicilliger, C. uncinatus, D. dasycnema) or low (C. uncinatus, P. soricis) abundance, but this depended on flea and/or host sex. We conclude that the directions of the host sex and phenology effects varied between flea species. Furthermore, the direction of the host sex effect was mediated by the effect of phenology and vice versa.

Species and site contributions to ß-diversity in fleas parasitic on the Palearctic small mammals: ecology, geography and host species composition matter the most.

The ß-diversity of fleas parasitic on small mammals in 45 regions of the Palearctic was partitioned into species [species contributions to ß-diversity (SCBD)] and site ( = assemblage) contributions [local contributions to ß-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.