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We studied spatial variation in the effects of environment and network size on species positions and roles in multiple flea-mammal networks from four biogeographic realms. We asked whether species positions (measured as species strength [SS], the degree of interaction specialization [d'], and the eigenvector centrality [C]) or the roles of fleas and their hosts in the interaction networks: (a) are repeatable/conserved within a flea or a host species; (b) vary in dependence on environmental variables and/or network size; and (c) the effects of environment and network size on species positions or roles in the networks depend on species traits. The repeatability analysis of species position indices for 441 flea and 429 host species, occurring in at least two networks, demonstrated that the repeatability of SS, d', and C within a species was significant, although not especially high, suggesting that the indices' values were affected by local factors. The majority of flea and host species in the majority of networks demonstrated a peripheral role. A value of at least one index of species position was significantly affected by environmental variables or network size in 41 and 36, respectively, of the 52 flea and 52 host species that occurred in multiple networks. In both fleas and hosts, the occurrence of the significant effect of environment or network size on at least one index of species position, but not on a species' role in a network, was associated with some species traits., (© 2024 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

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.

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., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

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 CO 2 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., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

We investigated compositional and phylogenetic nestedness in the host assemblages of 26 host-generalist fleas across regions within the Palearctic. We asked the following questions: (i) are host assemblages exploited by a flea species compositionally or phylogenetically nested (=C-nested and P-nested, respectively) across regions?; (ii) if yes, what are the processes that generate nestedness, and does phylogenetic nestedness follow the same processes as compositional nestedness?; and (iii) are the biological traits of a flea species associated with its host assemblages' degree of nestedness? Nestedness was calculated for matrices with rows ordered either by decreasing region area (=a-matrices) or increasing distance from the centre of a flea's geographic range (d-matrices). Significant C-nestedness was found in either a- (three fleas) or d-matrices (three fleas) or both (10 fleas). Significant P-nestedness was detected in either a- (three fleas) or d-matrices (four fleas) or both (two fleas). In some but not other species, P-nestedness followed C-nestedness. The probability of C-nestedness to be significant, as well as its degree for d-matrices, was associated with a flea's morphoecological traits, whereas this was not the case for either a-matrices or the P-nestedness for either type of ordered matrices. We conclude that compositional, but not phylogenetic, nestedness is (i) generated by similar mechanisms in many flea species and (ii) may be simultaneously driven by different mechanisms in the same flea. In contrast, mechanisms promoting phylogenetic nestedness differ between flea species and seem to act separately., (Copyright © 2023 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Combining the biogeography and phylogenetic patterns of parasite-host associations allows a better understanding of the history of parasite–host interactions, which can be achieved via biogeographic regionalization incorporating phylogenetic information. Recently, the concepts of evoregions (regions where a majority of species evolved from one or several ancestors inhabiting these regions) and evolutionary transition zones (regions of high phylogenetic turnover) have been proposed, coupled with a classification approach for these concepts. We applied this approach to 206 flea species and 265 host species of the Palearctic and aimed to identify evoregions and evolutionary transition zones for both fleas and hosts and to understand whether these evoregions and transition zones match each other. We identified 5 evoregions with 3 transition zones for either fleas or hosts, but neither the positions and boundaries of the flea and host evoregions nor the transition zones coincided. Indications of multiple geographic centres of diversification of the same flea lineages suggested that (a) the common evolutionary history of fleas and hosts was characterized by multiple events other than codiversification and that (b) dispersal played an important role in flea and host assemblies. Barriers to dispersal could be represented by landscape features (deserts and mountain ranges) and/or climate differences.

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.)

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.

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.

The structure of ecological interaction networks is associated with evolutionary histories of the interacting species. This is reflected by the phylogenetic signals (PS) in these networks when closely related species interact with similar partners because some traits inherited from the ancestors may determine ecological interactions. We investigated PS for small mammalian hosts and fleas in 80 regional interaction networks from four biogeographic realms (the Palearctic, the Nearctic, the Afrotropics, and the Neotropics). We asked (i) whether the relative strength of PS in host-flea networks is similar between hosts and fleas and/or between realms; (ii) how environmental variation affects the PS of hosts and fleas in their interaction networks; and (iii) whether the PS for hosts or fleas is affected by the phylogenetic diversity of either hosts or fleas, respectively. We found that the PS for hosts was stronger than that for fleas in all realms. An environmental effect on the PS for hosts, but not for fleas, was found in three of the four realms (except the Neotropics). In the Palearctic and the Nearctic, a stronger PS was characteristic for cooler and/or drier regions, whereas the opposite was the case for the Afrotropics in regard to precipitation. The phylogenetic diversity of regional host and flea assemblages was not associated with the values of the respective PS in any realm. We conclude that the pattern of the relative strength of the PS for hosts and fleas in their interaction networks is similar in different biogeographic realms with vastly different host and flea faunas. However, the environmental effects on the PS are geographically variable and might be associated with the history of host-flea associations, as well as the spatial pattern of environmental variation, within a realm., (Copyright © 2022 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

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., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

An assemblage of species in a locality comprises two components, namely (i) species that are present (realised diversity) and (ii) species from the regional pool that may potentially inhabit this locality due to suitable ecological conditions, but that are absent (dark diversity). We investigated factors affecting the dark diversity of component communities of fleas parasitic on small mammals in the northern Palearctic at two scales. First, we considered the dark diversity of flea assemblages of the same host (for 13 host species) across regions and tested for the effects of environmental factors and the number of available host species on the dark diversity of within-region flea assemblages. Second, we considered the dark diversity of fleas across host species within a region (for 20 regions) and asked whether within-host dark diversity is associated with host phylogeny and/or traits. We found that the dark diversity of flea assemblages harboured by small mammals varied substantially (i) within the same host species across space (in 12 of 13 host species) and (ii) between host species within a region (in eight of 20 regions). The size of the dark diversity of flea assemblages of the same host across regions was generally affected by environmental factors (mainly by the amount of green vegetation), whereas the size of the dark diversity of flea assemblages of a host species within a region was affected by host traits (mainly by the degree of host sociality and the structure of its shelter and, to a lesser degree, by its geographic range size) but was not associated with host phylogenetic affinities. We conclude that application of the dark diversity concept to parasite communities across space or hosts allows a better understanding of the factors affecting the species richness and composition of these communities., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

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., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

We studied patterns of compositional, functional, and phylogenetic α- and β-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.

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.)

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.

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.

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.

To understand existence, patterns and mechanisms behind phylogenetic heritability in the geographic range size (GRS) of parasites, we measured phylogenetic signal (PS) in the sizes of both regional (within a region) and continental (within a continent) geographic ranges of fleas in five regions. We asked whether (a) GRS is phylogenetically heritable and (b) the manifestation of PS varies between regions. We also asked whether geographic variation in PS reflects the effects of the environment's spatiotemporal stability (e.g. glaciation disrupting geographic ranges) or is associated with time since divergence (accumulation differences among species over time). Support for the former hypothesis would be indicated by stronger PS in southern than in northern regions, whereas support for the latter hypothesis would be shown by stronger PS in regions with a large proportion of species belonging to the derived lineages than in regions with a large proportion of species belonging to the basal lineages. We detected significant PS in both regional and continental GRSs of fleas from Canada and in continental GRS of fleas from Mongolia. No PS was found in the GRS of fleas from Australia and Southern Africa. Venezuelan fleas demonstrated significant PS in regional GRS only. Local Indicators of Phylogenetic Association detected significant local positive autocorrelations of GRS in some clades even in regions in which PS has not been detected across the entire phylogeny. This was mainly characteristic of younger taxa.

We tested whether geographic range position of fleas parasitic on small mammals in the Palearctic is affected by environmental niche conservatism or geographic range conservatism by measuring phylogenetic signal in range centroids and boundaries. We predicted that stronger phylogenetic signal in latitudinal than longitudinal range positions would indicate the important role of niche conservatism as a driver of the evolution of fleas' geographic ranges. Phylogenetic signals in geographic range positions were measured across 120 species, as well as within five flea lineages (subfamily/family rank) of different evolutionary ages. To investigate the temporal pattern of the geographic range position's evolution, we fitted the phylogenetic patterns in the geographic coordinates of range centroids and border extremes to four models of trait evolution. We consistently detected significant phylogenetic signals in the latitudes of the range centroids and the northern range borders. The latitudes of the southern range borders and the longitudes of the eastern/western borders demonstrated phylogenetic signals less often, whereas no signal was found for the longitudes of the range centroids. The phylogenetic signal in range position was more pronounced in younger lineages. The phylogenetic signal indices mainly suggested the evolution of range positions according to the Brownian motion model, whereas the best fit was often provided by the Orstein-Uhlenbeck model. This contradiction forced us to invoke a parsimonious explanation that the phylogenetic signal in range positions results from the interplay between the footprint of the speciation pattern and limited dispersal from the ancestral ranges., (Copyright © 2018 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

We investigated the phylogenetic structure of flea assemblages collected from small mammals on opposite sides of and increasing distance from the former Bering Land Bridge (BLB) using crossed double principal coordinate analysis (crossed-DPCoA). Phylogenetic composition of flea assemblages differed substantially between continents, but phylogenetic similarity between the Nearctic and Palearctic assemblages was the highest in the regions closer to the BLB. Within continents, phylogenetic similarity of flea assemblages was lower between regions closer to the BLB and those farther from the BLB than among regions within each of these groups. The Palearctic assemblages were represented mainly by basal families, while the Nearctic assemblages were dominated by a derived family (Ceratophyllidae), suggesting predominantly eastward pre-glaciation movements. In contrast, within the youngest flea family (Ceratophyllidae), the basal clades were characteristic for the Nearctic, while some species of a few derived clades were characteristic for the Palearctic, suggesting that, at least, during glaciation, westward movements occurred as well. In addition, multiple within-family clades of fleas were represented on opposite sides of the BLB suggesting multiple colonization events. This study is the first attempt to apply modern analytical methods of community ecology to reveal patterns in historical biogeography.

We studied the effect of host gender and body mass on species richness of flea infracommunities in nine rodent host species from three biomes (temperate zone of central Europe, desert of the Middle East and the tropics of East Africa). Using season- and species-specific generalized linear mixed models and controlling for year-to-year variation, spatial clustering of rodent sampling and over-dispersion of the data, we found inconsistent associations between host characteristics and flea species richness. We found strong support for male-biased flea parasitism, especially during the reproductive period (higher species richness in male hosts than in females) in all considered European rodents (Apodemus agrarius, Myodes glareolus and Microtus arvalis) and in one rodent species from the Middle East (Dipodillus dasyurus). In contrast, two of three African rodent species (Lophuromys kilonzoi and Praomys delectorum) demonstrated a trend of female-biased flea species richness. Positive associations between body mass and the number of flea species were detected mainly in males (five of nine species: A. agrarius, M. glareolus, M. arvalis, D. dasyurus and Mastomys natalensis) and not in females (except for M. natalensis). The results of this study support earlier reports that gender-biased, in general, and male-biased, in particular, infestation by ectoparasites is not a universal rule. This suggests that mechanisms of parasite acquisition by an individual host are species-specific and have evolved independently in different rodent host-flea systems.

We investigated spatial variation in the phylogenetic structure of host spectra in fleas parasitic on small mammals. Measures of phylogenetic host specificity ((phylogenetic species clustering (PSC) and phylogenetic species variability (PSV)) varied significantly more between than within flea species, but the proportion of variation which accounted for among-species differences was low. In 13 of 18 common flea species, at least one of the indices of the phylogenetic structure of regional host spectra revealed a significantly positive association with the phylogenetic structure of regional host assemblage, while relationships between PSC or PSV of the regional host spectrum and the distance from either the region of a flea's maximal abundance or latitude were not supported. Overall, results of this study demonstrated that although the degree of phylogenetic host specificity in fleas can be considered as a true attribute of a flea species, it is highly spatially variable, with phylogenetic structure of the surrounding host pool being the main reason behind this variation.

The distribution of parasites among individual hosts is characterised by high variability that is believed to be a result of variations in host traits. To find general patterns of host traits affecting parasite abundance, we studied flea infestation of nine rodent species from three different biomes (temperate zone of central Europe, desert of Middle East and tropics of East Africa). We tested for independent and interactive effects of host sex and body mass on the number of fleas harboured by an individual host while accounting for spatial clustering of host and parasite sampling and temporal variation. We found no consistent patterns of the effect of host sex and body mass on flea abundance either among species within a biome or among biomes. We found evidence for sex-biased flea infestation in just five host species (Apodemus agrarius, Myodes glareolus, Microtus arvalis, Gerbillus andersoni, Mastomys natalensis). In six rodent species, we found an effect of body mass on flea abundance (all species mentioned above and Meriones crassus). This effect was positive in five species and negative in one species (Microtus arvalis). In M. glareolus, G. andersoni, M. natalensis, and M. arvalis, the relationship between body mass and flea abundance was mediated by host sex. This was manifested in steeper change in flea abundance with increasing body mass in male than female individuals (M. glareolus, G. andersoni, M. natalensis), whereas the opposite pattern was found in M. arvalis. Our findings suggest that sex and body mass are common determinants of parasite infestation in mammalian hosts, but neither of them follows universal rules. This implies that the effect of host individual characteristics on mechanisms responsible for flea acquisition may be manifested differently in different host species.

We investigated spatial variation in the phylogenetic structure (measured as a degree of phylogenetic clustering) of flea assemblages across the geographic ranges of 11 Palearctic species of small mammalian hosts and asked whether the phylogenetic structure of the flea assemblage of a host in a locality is affected by (i) distance of this locality from the centre of the host's geographic range, (ii) geographic position of the locality (distance to the equator) and (iii) phylogenetic structure of the entire flea assemblage of the locality. Our results demonstrated that the key factor underlying spatial variation of the phylogenetic structure of the flea assemblage of a host was the distance from the centre of the host's geographic range. However, the pattern of this spatial variation differed between host species and might be explained by their species-specific immunogenetic and/or distributional patterns. Local flea assemblages may also, to some extent, be shaped by environmental filtering coupled with historical events. In addition, the phylogenetic structure of a local within-host flea assemblage may mirror the phylogenetic structure of the entire across-host flea assemblage in that locality and, thus, be affected by the availability of certain phylogenetic lineages., (Copyright © 2013 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

Abstract Across different taxa, networks of mutualistic or antagonistic interactions show consistent architecture. Most networks are modular, with modules being distinct species subsets connected mainly with each other and having few connections to other modules. We investigate the phylogenetic relatedness of species within modules and whether a phylogenetic signal is detectable in the within- and among-module connectivity of species using 27 mammal-flea networks from the Palaearctic. In the 24 networks that were modular, closely related hosts co-occurred in the same module more often than expected by chance; in contrast, this was rarely the case for parasites. The within- and among-module connectivity of the same host or parasite species varied geographically. However, among-module but not within-module connectivity of host and parasites was somewhat phylogenetically constrained. These findings suggest that the establishment of host-parasite networks results from the interplay between phylogenetic influences acting mostly on hosts and local factors acting on parasites, to create an asymmetrically constrained pattern of geographic variation in modular structure. Modularity in host-parasite networks seems to result from the shared evolutionary history of hosts and by trait convergence among unrelated parasites. This suggests profound differences between hosts and parasites in the establishment and functioning of bipartite antagonistic networks.

We tested the hypothesis that compositional and/or phylogenetic dissimilarity of host assemblages affect compositional and/or phylogenetic dissimilarity of parasite assemblages, to different extents depending on scale, using regional surveys of fleas parasitic on small mammals from 4 biogeographical realms. Using phylogenetic community dissimilarity metric, we calculated the compositional and phylogenetic dissimilarity components between all pairs of host and parasite communities within realms and hemispheres. We then quantified the effect of compositional or phylogenetic dissimilarity in host regional assemblages, and geographical distance between assemblages, on the compositional or phylogenetic dissimilarity of flea regional assemblages within a realm, respectively. The compositional dissimilarity in host assemblages strongly affected compositional dissimilarity in flea assemblages within all realms and within both hemispheres. However, the effect of phylogenetic dissimilarity of host assemblages on that of flea assemblages was mostly confined to the Neotropics and Nearctic, but was detected in both the Old and New World at the higher scale, possibly because of phylogenetic heterogeneity in flea and host faunas between realms. The clearer effect of the compositional rather than the phylogenetic component of host community dissimilarity on flea community dissimilarity suggests important roles for host switching and ecological fitting during the assembly history of flea communities.

Fleas collected from rodents in the Negev Desert in southern Israel were molecularly screened for Bartonella species. A total of 1,148 fleas, collected from 122 rodents belonging to six species, were pooled in 245 pools based on flea species, sex, and rodent host species. Two Bartonella gene fragments, corresponding to RNA polymerase B (rpoB) and citrate synthase (gltA), were targeted, and 94 and 74 flea pools were found positive by PCR, respectively. The Bartonella 16S-23S internal transcribed spacer (ITS) region was also targeted, and 66 flea pools were found to be positive by PCR. Sixteen different Bartonella gltA genotypes were detected in 94 positive flea pools collected from 5 different rodent species, indicating that fleas collected from each rodent species can harbor several Bartonella genotypes. Based on gltA analysis, identified Bartonella genotypes were highly similar or identical to strains previously detected in rodent species from different parts of the world. A gltA fragment 100% similar to Bartonella henselae was detected in one flea pool. Another 2 flea pools contained gltA fragments that were closely related to B. henselae (98% similarity). The high sequence similarities to the zoonotic pathogen B. henselae warrant further investigation.

Different host species harbour parasite faunas that are anywhere from very similar to very different in species composition. A priori, the similarity in the parasite faunas of any two host species should decrease with increases in either the phylogenetic distance, the distinctness of the environments occupied or the geographical distance between these hosts. We tested these predictions using extensive data on the faunas of fleas (Insecta: Siphonaptera) and gamasid mites (Acari: Parasitiformes) parasitic on rodents across the Palaearctic. For each pair of host species, we computed the similarity in parasite faunas based on both species composition as well as the phylogenetic and/or taxonomic distinctness of parasite species. Phylogenetic distances between hosts were based on patristic distances through a rodent phylogeny, geographic distances were computed from geographic range data, and environmental dissimilarity was measured from the average climatic and vegetation scores of each host range. Using multiple regressions on distance matrices to assess the separate explanatory power of each of the three dependent variables, environmental dissimilarity between the ranges of host species emerged as the best predictor of dissimilarity between parasite faunas, especially for fleas; in the case of mites, phylogenetic distance between host species was also important. A closer look at the data indicates that the flea and mite faunas of two hosts inhabiting different environments are always different, whilst hosts living in similar environments can have either very similar or dissimilar parasite faunas. Additional tests showed that dissimilarity in flea or mite faunas between host geographic ranges was best explained by dissimilarity in vegetation, followed by dissimilarity in climatic conditions. Thus, external environmental factors may play greater roles than commonly thought in the evolution of host-parasite associations., ((c) 2009 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The strength of interspecific interactions varies over geographical scales, and can influence patterns of resource specialisation. Even with gene flow preventing local adaptation of a consumer to particular resources, we might expect that across its entire range, the consumer would show some specialisation for the resource types most likely to be encountered across the localities where it occurs. We tested the hypothesis that generalist fleas are more successful at exploiting small mammalian host species with which they co-occur frequently across their geographical range than host species that, though suitable, are encountered less frequently. This hypothesis was tested with data on 121 flea species compiled from field surveys across 35 regions of the Palaearctic. Using abundance (mean number of individual fleas per individual host) as a measure of flea success on a particular host species, positive correlations between flea abundance and the frequency of co-occurrence of a flea with each of its hosts amongst all regions surveyed were found in all but two of the flea species investigated, with one-fifth of these being significant. If overlap in geographical range between flea and host is used as a measure of frequency of encounters instead of the actual proportion of regions where they both occur, similar patterns are observed, though they are much weaker. In a comparative analysis across all flea species, there were significant relationships between the average abundance of fleas and average values of both measures of frequency of encounters (proportion of sites where they co-occur and range overlap), even when correcting for potential phylogenetic influences. The results suggest that for any given flea species, host species more commonly encountered throughout the spatial range of the flea are generally those on which the flea does best. Interaction frequency may be a key determinant of specialisation and abundance in host-parasite systems.

Among generalist parasites, some species exploit only hosts from one particular phylogenetic lineage, whereas others can use a broader phylogenetic range of hosts, often seemingly using a random subset of the locally available host species. The latter type of generalist parasites should have greater opportunities to expand their geographical range and should not be restricted to stable and predictable host species because they are less prone to extinction than generalist parasites limited to a phylogenetically narrower host spectrum. We analyzed the diversity skewness of the host spectrum of 21 flea species from South Africa and 39 flea species from northern North America. Diversity skewness measures the balance in the shape of the phylogenetic tree of a set of species: the greater the skew, the more one lineage is overrepresented in an assemblage. When compared to a null expectation, i.e., random selections from the locally available pool of host species, the host spectra of most fleas was not more or less skewed than that expected by chance, though there were a few exceptions. Across South African fleas, the diversity skewness of the host spectrum was strongly negatively correlated with the size of a flea's geographic range; this relationship was not seen among North American flea species. There was no evidence among either set of fleas that average host body mass (a surrogate measure of host life span) correlated with the diversity skewness of the host spectrum. These findings are discussed with respect to the evolution of host specificity, its measurement, and historical differences between the two geographic areas considered.

Population density is a fundamental property of a species and yet it varies among populations of the same species. The variation comes from the interplay between intrinsic features of a species that tend to produce repeatable density values across all populations of the same species and extrinsic environmental factors that differ among localities and thus tend to produce spatial variation in density. Is inter-population variation in density too large for density to be considered a true species character? We addressed this question using data on abundance (number of parasites per individual host, i.e. equivalent to density) of fleas ectoparasitic on small mammals. The data included samples of 548 flea populations, representing 145 flea species and obtained from 48 different geographical regions. Abundances of the same flea species on the same host species, but in different regions, were more similar to each other than expected by chance, and varied significantly among flea species, with 46% of the variation among samples accounted by differences between flea species. Thus, estimates of abundance are repeatable within the same flea species. The same repeatability was also observed, but to a lesser extent, across flea genera, tribes and subfamilies. Independently of the identity of the flea species, abundance values recorded on the same host species, or in the same geographical region, also showed significant statistical repeatability, though not nearly as strong as that associated with abundance values from the same flea species. There were also no strong indications that regional differences in abiotic variables were an important determinant of variation in abundance of a given flea species on a given host species. Abundance thus appears to be a true species trait in fleas, although it varies somewhat within bounds set by species-specific life history traits.

Host individuals not infested by parasites at a given time are either permanently free from parasites or could be infested at other times. We studied temporal variation in the presence or absence of fleas (Siphonaptera) on individual rodents of two species (Gerbillus dasyurus and Acomys cahirinus) and questioned if and how an individual rodent can change its infestation status temporally. Change in infestation status by fleas over time was found in 45.5% of G. dasyurus and 35.9% of A. cahirinus. In both host species, the proportion of individuals that either changed or did not change their infestation status did not differ from the null expectation of 1:1. No difference between the proportions of infested at two consecutive captures and uninfested at two consecutive captures animals was found in A. cahirinus, whereas the proportion of G. dasyurus infested by fleas at two consecutive captures was significantly greater than that of uninfested at two consecutive captures individuals. In both host species, similar proportions of individuals changed their infestation status either from being infested to being uninfested or vice versa. Among individuals of both species that were initially infested by fleas, similar proportions changed and did not change their subsequent infestation status. The same was true for the uninfested at the first capture A. cahirinus. However, among initially uninfested G. dasyurus, the proportion of individuals that remained uninfested was significantly greater than that of the individuals newly infested by fleas. In A. cahirinus, the probability to change the infestation status did not depend on the initial status of an individual. In G. dasyurus, the probability of the initially uninfested individuals to be subsequently found still uninfested was higher than to be subsequently found harboring fleas, whereas initially infested individuals could be subsequently found either still infested or flea-free with equal probability.

The ability of vector-borne diseases to persist and spread is closely linked to the ecological characteristics of the vector species they use. Yet there have been no investigations of how species used as vectors by pathogens such as the plague bacterium differ from closely related species that are not used as vectors. The plague bacterium uses mammals as reservoir hosts and fleas as vectors. The ability of different fleas to serve as vectors is assumed to depend on how likely they are to experience gut blockage following bacterial multiplication; the blockage causes fleas to regurgitate blood into a wound and thus inject bacteria into new hosts. Beyond these physiological differences, it is unclear whether there exist fundamental ecological differences between fleas that are effective vectors and those that are not. Here, using a comparative analysis, we identify clear associations between the ability of flea species to transmit plague and their ecological characteristics. First, there is a positive relationship between the abundance of flea species on their hosts and their potential as vectors. Second, although the number of host species exploited by a flea is not associated with its potential as a vector, there is a negative relationship between the ability of fleas to transmit plague and the taxonomic diversity of their host spectrum. This suggests a correlation between some ecological characteristics of fleas and their ability to develop the plague blockage. The plague pathogen thus uses mainly abundant fleas specialized on a narrow taxonomic range of mammals, features that should maximize the persistence of the disease in the face of high flea mortality, and its transmission to suitable hosts only. This previously unrecognized pattern of vector use is of importance for the persistence and transmission of the disease.

The number of species coexisting in a community may be regulated by local factors (e.g., competitive interactions), or by regional processes (e.g., dispersal from a regional species pool). The relative importance of local and regional processes can be inferred from the shape of the relationship between local and regional species richness. We investigated this relationship in communities of fleas parasitic on small mammals at two spatial scales: between the richness of fleas on individual hosts (infracommunities) and that of fleas on host populations (component communities), and between the richness of component communities and that of the entire regional species pool. We tested linearity (proportional sampling) versus curvilinearity with an asymptote (species saturation) by plotting "local" against "regional" species richness of fleas either among host species or within host species among populations. At the two spatial scales, we found consistent curvilinear relationships between species richness of the more "local" communities and richness of the more "regional" communities. This was true across all host species in the data set and for geographic subsets, even after controlling for the influence of sampling effort on estimates of species richness, and that of host phylogeny in interspecific analyses. We also tested for density compensation in species-poor communities. There was no strong evidence for density compensation at the infracommunity level, although its existence at the component community level appeared likely. Our results suggest that identical patterns in local-versus-regional species richness observed on two different spatial scales arise via different mechanisms: infracommunities appear saturated with flea species most likely because of local processes, such as host immune defenses, whereas component communities are saturated with species through interspecific competition, possibly among larval stages.

Evolutionary trends in the evolution of host specificity have been the focus of much discussion but little rigorous empirical testing. On the one hand, specialization is often presumed to lead irreversibly into evolutionary dead ends and little diversification; this would mean that generalists might evolve into specialists, but not vice versa. On the other hand, low host specificity may limit the risk of extinction and provide more immediate fitness benefits to parasites, such that selection may favour evolution toward a generalist strategy. Here, we test for directionality in the evolution of host specificity using a large data set and phylogenetic information on 297 species of fleas parasitic on small mammals. The analyses determined whether host specificity, measured both as the number of host species exploited and their taxonomic diversity, was related to clade rank of the flea species, or the number of branching events between an extant species and the root of the phylogenetic tree (i.e., the total path length from the root of the tree to the species). Based on regression analyses, we found positive relationships between the number of host species used and clade rank across all 297 species, as well as within one (Hystrichopsyllidae) of four large families and one of seven large genera investigated separately; in addition, we found a positive relationship between the taxonomic diversity of host species used and clade rank in another of the seven genera. These results suggest a slight evolutionary trend of decreasing host specificity. Using a much more conservative likelihood ratio test, however, a random walk, or null model, of evolution could not be discarded in favour of the directional trends in all cases mentioned above. Still, these results suggest that host specificity may have tended to decrease in many flea lineages, a process that could have been driven by the benefits of exploiting a wide range of host species.

Our aim was to determine which of four variables (number of host species exploited by the parasite, taxonomic distinctness of these hosts, geographic range of the principal host, and year of description of this host) was the best predictor of description date of fleas. The study used previously published data on 297 flea species parasitic on 197 species of small mammals from 34 different regions of the Holarctic and one region from the Neotropics. We used both simple linear and multiple regressions to evaluate the relationships between the four predictor variables and the year of flea description, on species values as well as on phylogenetically independent contrasts. Whether or not the analyses controlled for flea phylogeny, all predictor variables correlated significantly with year of flea description when tested separately. In multiple regressions, however, the number of exploited host species was the best predictor of the date of flea description, with the geographic range of the principal host species as well as the date of its description having a lesser, though significant, influence. Overall, our results indicate that a flea species is more likely to be discovered and described early if its biological characteristics (exploitation of many host species) and those of its hosts (long-known to science, broad geographic distributions) increase its chances of being included in a collection. Because the variables we investigated only explained about 10-11% of the variation in year of description among flea species, other factors must be important, such as temporal variability in the activity of flea taxonomists.

Animal species with larger local populations tend to be widespread across many localities, whereas species with smaller local populations occur in fewer localities. This pattern is well documented for free-living species and can be explained by the resource breadth hypothesis: the attributes that enable a species to exploit a diversity of resources allow it to attain a broad distribution and high local density. In contrast, for parasitic organisms, the trade-off hypothesis predicts that parasites exploiting many host species will achieve lower mean abundance on those hosts than more host-specific parasites because of the costs of adaptations against multiple defense systems. We test these alternative hypotheses with data on host specificity and abundance of fleas parasitic on small mammals from 20 different regions. Our analyses controlled for phylogenetic influences, differences in host body surface area, and sampling effort. In most regions, we found significant positive relationships between flea abundance and either the number of host species they exploited or the average taxonomic distance among those host species. This was true whether we used mean flea abundance or the maximum abundance they achieved on their optimal host. Although fleas tended to exploit more host species in regions with either larger number of available hosts or more taxonomically diverse host faunas, differences in host faunas between regions had no clear effect on the abundance-host specificity relationship. Overall, the results support the resource breadth hypothesis: fleas exploiting many host species or taxonomically unrelated hosts achieve higher abundance than specialist fleas. We conclude that generalist parasites achieve higher abundance because of a combination of resource availability and stability.

Parasites should make the same decisions that every animal makes regarding fitness reward. They can maximize reproductive success by selection of those habitats that guarantee the greatest fitness output. We consider the host population as a habitat of a parasite population. Consequently, hosts (=habitats) that differ quantitatively or qualitatively will support different numbers of parasites. The nature of habitat selection can be detected by isodars, lines along which habitat selection yields equivalent fitness reward. We applied this approach to study host selection of five fleas, each infesting two desert rodents. Xenopsylla conformis, Xenopsylla ramesis, Nosopsyllus iranus theodori and Stenoponia tripectinata medialis parasitize Gerbillus dasyurus and Meriones crassus. Synosternus cleopatrae pyramidis parasitizes Gerbillus andersoni allenbyi and Gerbillus pyramidum. Three fleas ( X. conformis, X. ramesis and S. c. pyramidis) were able to perceive quantitative (amount of the resource; e.g. organic matter in the nest for flea larvae) and/or qualitative (pattern of resource acquisition; e.g. host defensiveness) differences between hosts. Two other fleas did not perceive between-host differences. X. conformis was a density-dependent host selector that showed sharp selectivity at low density. X. ramesis and S. c. pyramidis were density-independent host selectors with a direct correspondence of density with habitat quality. N. i. theodori and S. t. medialis were non-selectors with no relationship at all between density and host quality. The results of the application of the isodar theory suggest that ectoparasites, like other animals, behave as if they are able to make choices and decisions that favour environments in which their reproductive benefit is maximized.

On any spatial scale, the species composition of a taxonomic group often departs from a phylogenetically random subset drawn from the pool of species available on a higher scale. Analysis of the uneven representation of related lineages in different assemblages can reveal the action of various forces shaping their diversification. For any assemblage, unequal diversification among lineages can be estimated using diversity skewness, an index of the balance of a phylogenetic tree whose values increase with increasing differences in diversification rates among tree branches. We tested for geographical patterns in the diversity skewness of flea assemblages parasitic on small mammals in 26 distinct geographic localities from the Palaearctic and 15 from the Nearctic. Overall, diversity skewness of the Nearctic flea assemblage was unexpectedly high compared to that of the global flea fauna, whereas that of the Palaearctic did not depart from the expectations of a null model. On a smaller scale, the diversity skewness of local flea assemblages was sometimes lower, sometimes higher, but, in most of the 41 localities, it did not differ significantly from that of random subsets taken from the species pool available on the larger spatial scale (either the world fauna or that of the biogeographical realm, i.e. Palaearctic or Nearctic). More importantly, among Palaearctic assemblages, diversity skewness increased with increasing latitude and/or decreasing mean air temperatures. The different patterns observed in the Palaearctic and Nearctic may be in part due the fact that flea diversification appears to have been more intense in the former than the latter, and to differences between them in relief and glacial history. Temperature-driven speciation rates may well explain the latitudinal gradient in diversity skewness in the Palaearctic. The results illustrate the action of various biogeographical processes in shaping the uneven differentiation of flea lineages on different spatial scales. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 807–814. [ABSTRACT FROM AUTHOR]