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

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

Searching for common patterns in parasite ecology: species and host contributions to beta-diversity in helminths of South African ungulates and fleas of South American rodents.

We searched for common patterns in parasite ecology by investigating species and host contributions to the beta-diversity of infracommunities (=assemblages of parasites harboured by a host individual) in helminths of three species of South African ungulates and fleas of 11 species of South American rodents, assuming that a comparison of patterns in distinctly different parasites and hosts would allow us to judge the generality or, at least, commonness of these patterns. We used data on species' composition and numbers of parasites and asked whether (i) parasite species' attributes (life cycle, transmission mode, and host specificity in helminths; possession of sclerotized combs, microhabitat preference, and host specificity in fleas) or their population structure (mean abundance and/or prevalence) and (ii) host characteristics (sex and age) affect parasite and host species' contributions to parasite beta-diversity (SCBD and HCBD, respectively). We found that parasite species' morphological and ecological attributes were mostly not associated with their SCBD. In contrast, parasite SCBD, in both ungulates and rodents, significantly increased with either parasite mean abundance or prevalence or both. The effect of host characteristics on HCBD was detected in a few hosts only. In general, parasite infracommunities' beta-diversity appeared to be driven by variation in parasite species rather than the uniqueness of the assemblages harboured by individual hosts. We conclude that some ecological patterns (such as the relationships between SCBD and parasite abundance/prevalence) appear to be common and do not differ between different host-parasite associations in different geographic regions, whereas other patterns (the relationships between SCBD and parasite species' attributes) are contingent and depend on parasite and host identities.

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

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

Climatic gradients and forest composition shape bat communities in Eastern Mediterranean pine plantations.

Biotic and abiotic factors can act as filters for determining the species composition of biological communities. We aimed to identify abiotic factors driving the assembly of bat communities in Eastern Mediterranean pine plantations along a north-south climatic gradient, as they are crucial forest habitats for the assessment and conservation of these communities. We expected that bat communities are predominantly shaped by environmental filtering. We conducted acoustic sampling in 35 pine plantations in Israel and analyzed recordings for species identification. We used the ESLTP analysis, an extension of the three-table ordination (RLQ analysis), to explore relationships between environmental characteristics, species occurrences, and functional traits of species while accounting for phylogenetic relationships between species and spatial distribution of the communities. Communities showed phylogenetic and trait clustering. Climatic conditions and forest vegetation composition shaped communities of bats, affecting the distribution of traits related to foraging behaviors, vegetation clutter, and the ability of bats to maneuver in it. Maneuverable species were associated with the northern Mediterranean climatic zone, with a scarce cover of drought-tolerant small shrubs and grassland. Fast flyers were associated with the center-south semi-arid area, with abundant drought-tolerant small shrubs and grassland. These forces might have a predominant role in the assembly of these communities, presumably due to the stressful climatic conditions of the study area. The ESLTP approach can be extended to other taxa and environments to predict species responses to disturbance and environmental changes and give insights into environmental management.

Environment and traits affect parasite and host species positions but not roles in flea-mammal networks.

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.

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

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

Evoregions of fleas and their small mammalian hosts: Do they coincide?

Combining the biogeography and phylogenetic patterns of parasite-host associations allows a better understanding of the history of parasite­host 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.

Documenting the microbiome diversity and distribution in selected fleas from South Africa with an emphasis on the cat flea, Ctenocephalides f. felis.

The factors that influence parasite associated bacterial microbial diversity and the geographic distributions of bacteria are not fully understood. In an effort to gain a deeper understanding of the relationship between the bacterial diversity of Ctenocephalides fleas and host species and the external environment, we conducted a metagenetic analysis of 107 flea samples collected from 8 distinct sampling sites in South Africa. Pooled DNA samples mostly comprising of 2 or 3 individuals sampled from the same host, and belonging to the same genetic cluster, were sequenced using the Ion PGM™ Hi-Q™ Kit and the Ion 316™ Chip v2. Differences were detected in the microbiome compositions between Ctenocephalides felis, Ctenocephalides canis and Ctenocephalides connatus. Although based on a small sample, C. connatus occurring on wildlife harboured a higher bacterial richness when compared to C. felis on domestic animals. Intraspecific differences in the microbial OTU diversity were detected within C. f. felis that occurred on domestic cats and dogs. Different genetic lineages of C. f. felis were similar in microbial compositions but some differences exist in the presence or absence of rare bacteria. Rickettsia and Bartonella OTU's identified in South African cat fleas differ from those identified in the USA and Australia. Intraspecific microbial compositions also differ across geographic sampling sites. Generalized dissimilarity modelling showed that temperature and humidity are potentially important environmental factors explaining the pattern obtained.

Nestedness and beta diversity of gastrointestinal helminth communities in common warthogs, Phacochoerus africanus (Suidae), at 2 localities in South Africa.

Few studies have investigated the ecological interactions between wild species of Suidae and their parasites, leaving our knowledge concerning this host­parasite system fragmented. In the present study, we applied network studies to analyse community nestedness in helminth assemblages of common warthogs, Phacochoerus africanus (Gmelin) (Suidae). Helminth data were compiled from 95 warthogs, including young and adult males and females, from 2 different conservation areas in Mpumalanga and Limpopo Provinces, South Africa, collected monthly over a period of 1 year each. The aim was to study the effect of host sex, age and season of sampling on the structure of helminth infracommunities harboured by the warthogs and to search for non-random structural patterns in the warthog­helminth interaction networks. Furthermore, we investigated the influence of a warthog's age, sex and season of sampling on beta diversity and dark diversity of their helminth infracommunities. Lastly, we asked whether the effects of host sex, age and sampling season on helminth communities differed between the 2 localities. We found that helminth communities of warthogs were nested and host­parasite interactions were influenced by all 3 factors as well as combinations thereof. However, the resulting patterns differed at the 2 localities, indicating that local environmental processes are important drivers of community structure.

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

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

Compositional and phylogenetic nestedness of host assemblages exploited by generalist ectoparasites across their geographic ranges: drivers and associations with ectoparasite traits.

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.

Ectoparasites associated with the Bushveld gerbil (Gerbilliscus leucogaster) and the role of the host and habitat in shaping ectoparasite diversity and infestations.

Rodents are known hosts for various ectoparasite taxa such as fleas, lice, ticks and mites. South Africa is recognized for its animal diversity, yet little is published about the parasite diversity associated with wild rodent species. By focusing on a wildlife-human/domestic animal interface, the study aims to record ectoparasite diversity and levels of infestations of the Bushveld gerbil, Gerbilliscus leucogaster, and to establish the relationship between ectoparasite infestation parameters and host- and habitat factors. Rodents (n = 127) were trapped in 2 habitat types (natural and agricultural) during 2014­2020. More than 6500 individuals of 32 epifaunistic species represented by 21 genera and belonging to 5 taxonomic groups (fleas, sucking lice, ticks, mesostigmatan mites and trombiculid mites) were collected. Mesostigmatan mites and lice were the most abundant and fleas and mesostigmatan mites the most prevalent groups. Flea and mesostigmatan mite numbers and mesostigmatan mite species richness was significantly higher on reproductively active male than female rodents. Only ticks were significantly associated with habitat type, with significantly higher tick numbers and more tick species on rodents in the natural compared to the agricultural habitat. We conclude that the level of infestation by ectoparasites closely associated with the host (fleas and mites) was affected by host-associated factors, while infestation by ectoparasite that spend most of their life in the external environment (ticks) was affected by habitat type.

Free-living gerbils with higher testosterone take fewer risks.

Among the physiological differences between the sexes are circulating androgen levels. Testosterone (T) is an androgen that has been linked to aggression and risk-taking in male vertebrates, so that males with higher T are generally more aggressive and take more risks. In females, T is not often measured, and its relationship with behaviour has been less studied. The costs of elevated T are assumed to be higher for reproductive females, while the benefits higher for males. Here, we tested the association between endogenous T and risk-taking behaviours in both males and females under well-studied experimental settings in free-living Baluchistan gerbils (Gerbillus nanus; Gn). In addition, we experimentally elevated Gn T levels using implants and measured risk-taking behaviour. Surprisingly, we found that there were no differences in the association between T and risk-taking behaviours between males and females, and that in both sexes, Gn with higher T levels took fewer risks. We also found that Gn spent equal time foraging between risky (open habitat) and safe (under a bush) experimental food patches. We expected Gn, which are nocturnal, to take fewer risks during full moon nights, but found that Gn were more active during moon lit nights than during dark (new moon) nights. This study demonstrates that T has many functions, and that its effects are complex and often unpredictable. It also shows that hypotheses regarding the propensity to take risks under specific coverage and light regimes are not universal, and likely include variables such as species, environment, context, and predator-specific behavioural strategies.

Rtapas: An R Package to Assess Cophylogenetic Signal between Two Evolutionary Histories.

Cophylogeny represents a framework to understand how ecological and evolutionary process influence lineage diversification. The recently developed algorithm Random Tanglegram Partitions provides a directly interpretable statistic to quantify the strength of cophylogenetic signal and incorporates phylogenetic uncertainty into its estimation, and maps onto a tanglegram the contribution to cophylogenetic signal of individual host-symbiont associations. We introduce Rtapas, an R package to perform Random Tanglegram Partitions. Rtapas applies a given global-fit method to random partial tanglegrams of a fixed size to identify the associations, terminals, and internal nodes that maximize phylogenetic congruence. This new package extends the original implementation with a new algorithm that examines the contribution to phylogenetic incongruence of each host-symbiont association and adds ParaFit, a method designed to test for topological congruence between two phylogenies, to the list of global-fit methods than can be applied. Rtapas facilitates and speeds up cophylogenetic analysis, as it can handle large phylogenies (100+ terminals) in affordable computational time as illustrated with two real-world examples. Rtapas can particularly cater for the need for causal inference in cophylogeny in two domains: (i) Analysis of complex and intricate host-symbiont evolutionary histories and (ii) assessment of topological (in)congruence between phylogenies produced with different DNA markers and specifically identify subsets of loci for phylogenetic analysis that are most likely to reflect gene-tree evolutionary histories. [Cophylogeny; cophylogenetic signal; gene tree incongruence; phylogenetic congruence; phylogenomics.].

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

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

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

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

Latitudinal gradients in body size and sexual size dimorphism in fleas: males drive Bergmann's pattern.

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.

Impact of host sex and age on the diversity of endoparasites and structure of individual-based host-parasite networks in nyalas (Tragelaphus angasii Angas) from three game reserves in KwaZulu-Natal province, South Africa.

In recent years, numerous studies have examined the effect of host sex and age on the structure of parasite communities in several host taxa under various environmental conditions and in different geographic regions. However, the influence of such factors on the structure of host-parasite networks has received less attention, and remarkably few studies have been carried out on large terrestrial mammals. In this study, we investigated the effects of host age and sex on the parasite infra- and component communities of nyalas (Tragelaphus angasii) and on the structure of individual-based nyala-endoparasite networks. We also aimed to evaluate to what extent these effects vary spatially and if they are mediated by conservation management. Based on a data set of internal macroparasites of 74 nyalas from three game reserves in KwaZulu-Natal province, we found that host age strongly influenced parasite community structure as well as the structure of parasite-nyala networks, whereas host sex played a minor role. However, the effects of both host sex and age were mediated by environmental conditions and thus led to different patterns at the three localities. Our findings highlight that host-parasite communities from different localities should not be pooled when conducting host-parasite network and community studies as this may bias results and mask patterns that are typical for a given locality.

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

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

Phylogenetic signals in flea-host interaction networks from four biogeographic realms: differences between interactors and the effects of environmental factors.

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.