Flea Communities Research Articles

Deconstructing spatial patterns in species composition of ectoparasite communities: the relative contribution of host composition, environmental variables and geography

ABSTRACTAim We determined whether dissimilarity in species composition between parasite communities depends on geographic distance, environmental dissimilarity or host faunal dissimilarity, for different subsets of parasite species with different levels of host specificity.Location Communities of fleas parasitic on small mammals from 28 different regions of the Palaearctic.Method Dissimilarities in both parasite and host species composition were computed between each pair of regions using the Bray–Curtis index. Geographic distances between regions were also calculated, as were measures of environmental dissimilarity consisting of the pairwise Euclidean distances between regions derived from elevation, vegetation and climatic variables. The 136 flea species included in the dataset were divided into highly host‐specific species (using 1–2 host species per region, on average), moderately host‐specific species (2.2–4 hosts per region) and generalist species (>4 hosts per region). The relative influence of geographic distance, host faunal dissimilarity and environmental dissimilarity on dissimilarity of flea species composition among all regions was analysed for the entire set of flea species as well as for the three above subsets using multiple regressions on distance matrices.Results When including all flea species, dissimilarity in flea species composition was affected by all three independent variables, although the pure effect of dissimilarity in host species composition was the strongest. Results were different when the subsets of fleas differing in host specificity were treated separately. In particular, dissimilarity in species composition of highly host‐specific fleas increased solely with environmental dissimilarity, whereas dissimilarity for both moderately specific and non‐specific fleas increased with both geographic distance and dissimilarity in host species composition.Main conclusions Host specificity seems to dictate which of the three factors considered is most likely to affect the dissimilarity between flea communities. Counter‐intuitively, environmental dissimilarity played a key role in determining dissimilarity in species composition of highly host‐specific fleas, possibly because, although their presence in a region relies on the occurrence of particular host species, their abundance is itself mostly determined by climatic conditions. Our results show that deconstructing communities into subsets of species with different traits can make it easier to uncover the mechanisms shaping geographic patterns of diversity.

FLEA ABUNDANCE, DIVERSITY, AND PLAGUE IN GUNNISON'S PRAIRIE DOGS (CYNOMYS GUNNISONI) AND THEIR BURROWS IN MONTANE GRASSLANDS IN NORTHERN NEW MEXICO

Plague, a flea-transmitted infectious disease caused by the bacterium Yersinia pestis, is a primary threat to the persistence of prairie dog populations (Cynomys spp.). We conducted a 3-yr survey (2004-2006) of fleas from Gunnison's prairie dogs (Cynomys gunnisoni) and their burrows in montane grasslands in Valles Caldera National Preserve in New Mexico. Our objectives were to describe flea communities and identify flea and rodent species important to the maintenance of plague. We live-trapped prairie dogs and conducted burrow sweeps at three colonies in spring and summer of each year. One hundred thirty prairie dogs and 51 golden-mantled ground squirrels (Spermophilus lateralis) were captured over 3,640 trap nights and 320 burrows were swabbed for fleas. Five flea species were identified from prairie dogs and ground squirrels and four were identified from burrow samples. Oropsylla hirsuta was the most abundant species found on prairie dogs and in burrows. Oropsylla idahoensis was most common on ground squirrels. Two colonies experienced plague epizootics in fall 2004. Plague-positive fleas were recovered from burrows (O. hirsuta and Oropsylla tuberculata tuberculata) and a prairie dog (O. hirsuta) in spring 2005 and summer 2006. Three prairie dogs collected in summer 2005 and 2006 had plague antibody. We found a significant surge in flea abundance and prevalence, particularly within burrows, following plague exposure. We noted an increased tendency for flea exchange opportunities in the spring before O. hirsuta reached its peak population. We hypothesize that the role of burrows as a site of flea exchange, particularly between prairie dogs and ground squirrels, may be as important as summer conditions that lead to buildup in O. hirsuta populations for determining plague outbreaks.

Modeling Susceptible Infective Recovered Dynamics and Plague Persistence in California Rodent–Flea Communities

Plague persists as an enzootic in several very different rodent-flea communities around the world. In California, a diversity of rodent-flea communities maintains the disease, and a single-host reservoir seems unlikely. Logistic regression of plague presence on climate and topographic variables predicts plague in many localities where it is absent. Thus, a dynamic community-based analysis was needed. Deterministic Susceptible Infective Recovered (SIR) models were adapted for plague and analyzed with an eye for insights concerning disease persistence. An R simulation program, Plaguesirs, was developed incorporating multihost and multivector SIR dynamics, demographic and environmental stochasticity, density dependence, and seasonal variation in birth and death. Flea-rodent utilization matrices allowed us to get transmission rates as well as flea carrying capacities. Rodent densities allowed us to estimate host carrying capacities, while maximum birth rates were mainly approximated through an examination of litter phenology and demography. We ran a set of simulations to assess the role of community structure in maintaining plague in a simulated version of Chuchupate campground in Ventura County. Although the actual campground comprises 10 rodent and 19 flea species, we focused on a subset suspected to act as a reservoir community. This included the vole Microtus californicus, the deer mouse Peromyscus maniculatus, the Ceratophyllid fleas Aetheca wagneri and Malareus telchinum, and the Leptopsyllid flea Peromyscopsylla hesperomys. The dynamics of 21 subsets of this community were simulated for 20 years. Single-rodent communities showed much lower disease persistence than two-rodent communities. However, so long as Malareus was present, endemicity was enhanced; removal of the other two fleas slightly increased disease persistence. Two critical features improved disease persistence: (1) host breeding season heterogeneity and (2) host population augmentation (due to two similar host species instead of one). Voles are winter-spring breeders compared to the spring-summer deer mice. While host synchronicity may enhance epidemics, host asynchronicity favors endemics.

Ectoparasite communities on Tupaia belangeri (Scandentia: Tupaiidae) in surrounding areas of Erhai Lake in Yunnan, China

Objective To investigate the ectoparasite communities on Tupaia belangeri (T. belangeri) in the surrounding areas of Erhai Lake and their potential importance in medical and veterinary science. Methods Different locations in surrounding areas of Erhai Lake in Yunnan were selected as the investigation spots. T. belangeri were randomly captured with baited mouse cages and then brought to the laboratory for identification according to their body shape, size and color, and measurements such as the lengths of their body, ear and hind feet. Ectoparasites on them were collected and identified to determine the species under microscope after specimens were mounted on slides, with the clearing and drying process completed. The constituent ratio (C), prevalence (P) and average ectoparasite abundance (A) were used to test the prevalence and density of ectoparasites; Non-parametric Mann-Whitney {/-tests were used to test the differences in the abundance and richness of ectoparasites between female and male hosts; Spearman correlation analysis was used to analyze the relationship between ectoparasites and the body parameters of hosts. Results There was very high species diversity of ectoparasites found on T. belangeri and some species of ectoparasites were reported associated with human diseases. A high proportion (88%) of 107 T. belangeri was found to be infested with ectoparasites. A total of 75 species of ectoparasites on them were collected, including 58 species of chigger mite, 12 species of gamasid mite, 4 species of flea and 1 species of sucking louse. Within this ectoparasite complex, 13 species were previously reported to be vectors of human disease agents. Non-parametric Mann-Whitney U-tests showed significant differences between male and female T. belangeri in terms of the abundance and richness of total ectoparasites. However, the species richness and abundance of chigger mites, sucking lice, fleas or gamasid mites presented no significant differences between male and female hosts. Spearman correlation analysis showed that none of correlations were observed between host body parameters with abundance and richness of total ectoparasites, chigger mites, sucking lice, fleas or gamatid mites. Conclusion The results suggest that there is very high species diversity of ectoparasites on T. belangeri, consisting mainly of communities of sucking lice, fleas, chiggers and gamasid mites.

Diversité du cortège en Siphonaptères chez le tamia de Sibérie (Tamias sibiricusLaxmann) introduit en Forêt de Sénart (Ile-de-France)

We examined the fleas community in an introduced population of Siberian chipmunks, Tamias sibiricus, between 2005 and 2007, in the Forest of Sénart (Essonne, France). We collected and identified 383 fleas on 463 chipmunks (total: 1,891 captures on 471 chipmunks). In 2005, 120 fleas were also collected on 65 bank voles, Clethrionomys glareolus, and on 25 wood mice, Apodemus sylvaticus, trapped within the same area. Ceratophyllus sciurorum sciurorum formed 73.6% of the chipmunks' flea community, with an annual prevalence (P) ranging between 8 and 13% and a mean intensity (I) ranging between 1.1 and 1.6 fleas per individual. Among the six other species infecting this Sciurid, Ctenophthalmus agyrtes impavidus constituted 17.2% (P: 1.6-2.2%; I: 1.1-2.6), and Megabothris turbidus 8.1% (P: 0.8-1.9%; I: 1.0-1.4) of the flea community, respectively. These last two species represented respectively 60.8% and 36.6% of the flea community on the bank vole and the wood mouse. Originated from Asia, chipmunks did not import any flea species to Sénart, probably because they were used as pets before their release in the wild. Abundance in C. s. sciurorum increased with adult chipmunk density and with juvenile density in summer. On adult chipmunks C. s. sciurorum tented to decrease with increasing abundance of red squirrels (Sciurus vulgaris]. Moreover, the two other flea species mainly infected young chipmunks during the fall, and their number was not related to chipmunk density. However, the distribution of species within the flea community became more balanced with increase juvenile chipmunk density. Overall, these results indicate that the close phyletic relationship between chipmunks and red squirrels contributed in the acquisition and the spread of fleas by chipmunks. Primary and secondary hosts densities, their habitat use, and more specifically burrowing activities and tree canopy use, also played a role in the spread of fleas on chipmunks.

Bacterial Communities of Bartonella -Positive Fleas: Diversity and Community Assembly Patterns

We investigated the bacterial communities of nine Bartonella-positive fleas (n = 6 Oropsylla hirsuta fleas and n = 3 Oropsylla montana fleas), using universal primers, clone libraries, and DNA sequencing. DNA sequences were used to classify bacteria detected in a phylogenetic context, to explore community assembly patterns within individual fleas, and to survey diversity patterns in dominant lineages.

Fleas parasitizing domestic dogs in Spain

In addition to their importance to veterinary clinical practice as ectoparasites, fleas of domestic dogs are of special concern because they can be vectors of disease, including zoonoses. Flea assemblages parasitizing domestic dogs usually comprise several flea species whose distribution is determined by factors acting at several scales. Knowledge of these factors will aid in assessment of the distribution patterns of flea parasitism, and is an important tool in developing control strategies and in evaluation of flea-borne disease risk in dogs and humans. In this survey we used data from 744 domestic dogs from 79 localities in Spain to explore the associations between the abundance of flea species, host-dependent factors (sex and age), and host habitat factors including abode (farm, house with garden, apartment), location (urban or rural), the presence of other pets, and dog activity (measured as the frequency with which dogs left their abode). We also considered environmental factors including the time of year and mean annual temperature and rainfall. Variations in flea community structure at infracommunity and component community levels were also explored. Four flea species were found parasitizing dogs. Ctenocephalides felis was the most abundant (88.02% of fleas identified), followed by Ctenocephalides canis (10.38%), Pulex irritans (1.47%) and Echidnophaga gallinacea (0.13%). Overall flea abundance was higher on dogs living on farms than in apartments, as was the abundance of Ct. felis, Ct. canis and P. irritans. Ct. felis was more abundant on dogs living in houses than in apartments, but the reverse was found for P. irritans. Overall flea abundance and Ct. canis abundance were highest in rural areas, whereas the presence of other pets sharing the abode was associated with higher overall flea abundance and Ct. felis abundance. Only P. irritans abundance was positively related to the activity of dogs. Ct. canis and P. irritans abundances were higher during the warm period of the year. Mean annual temperature was negatively correlated with overall, Ct. canis and P. irritans abundances, but positively related to Ct. felis abundance. Annual rainfall was negatively correlated with Ct. canis and P. irritans abundances. Variations in the number of flea species found on a dog reflected the abundance distribution patterns for each species and their associations with host habitat and environmental factors. At the component community level, flea species richness was inversely related to annual mean temperature. The structure of flea assemblages on dogs was mainly associated with host habitat and environmental variables, and not with host-dependent variables. However, a large amount of variation in flea abundance remained unexplained, suggesting the effect of other non-controlled factors.

Are ectoparasite communities structured? Species co‐occurrence, temporal variation and null models

1. We studied temporal variation in the structure of flea communities on small mammalian hosts from eastern Slovakia using null models. We asked (a) whether flea co-occurrences in infracommunities (in the individual hosts) in different hosts as well as in the component communities (in the host species) demonstrate a non-random pattern; (b) whether this pattern is indicative of either positive or negative flea species interactions; (c) whether this pattern varies temporally; and (d) whether the expression of this pattern is related to population size of either fleas or hosts or both. 2. We constructed a presence/absence matrix of flea species for each temporal sample of a host species and calculated four metrics of co-occurrence, namely the C-score, the number of checkerboard species pairs, the number of species combinations and the variance ratio (V-ratio). Then we compared these metrics with the respective indices calculated for 5000 null matrices that were assembled randomly using two algorithms, namely fixed-fixed (FF) and fixed-equiprobable (FE). 3. Most co-occurrence metrics calculated for real data did not differ significantly from the metrics calculated for simulated matrices using the FF algorithm. However, the indices observed for 42 of 75 presence/absence matrices differed significantly from the null expectations for the FE models. Non-randomness was detected mainly by the C-score and V-ratio metrics. In all cases, the direction of non-randomness was the same, namely the aggregation, not competition, of flea species in host individuals and host species. 4. The inclusion or exclusion of the uninfested hosts in the FE models did not affect the results for individual host species. However, exclusion of the uninfested host species led to the acceptance of the null hypothesis for only six of 13 temporal samples of the component flea communities for which non-randomness was detected when the uninfested hosts were included in the analysis. 5. In most host species, the absolute values of the standardized size effect of both the C-score and V-ratio increased with an increase in host density and a concomitant decrease in flea abundance and prevalence. 6. Results of this study demonstrated that (a) flea assemblages on small mammalian hosts were structured at some times, whereas they appeared to be randomly assembled at other times; (b) whenever non-randomness of flea co-occurrences was detected, it suggested aggregation but never segregation of flea species in host individuals or populations; and (c) the expression of structure in flea assemblages depended on the level of density of both fleas and hosts.

Host Specificity, Parasite Community Size and the Relation between Abundance and its Variance

We investigated the empirical relationship between mean abundance and its variance, known as Taylor’s power law, in fleas parasitic on small mammals. It has been suggested that the exponent of this function, b, represents a true biological character of a species and, dependent on the level of host specificity, varies among species. Other empirical and theoretical studies suggest that exponent b depends on interspecific competition and varies intraspecifically. We tested these hypotheses using data from central and eastern Slovakia. We demonstrate that the slope of Taylor’s relationship (a) is repeatable within a flea species, i.e. the slope represents a true species character; (b) increases with an increase of the degree of flea host specificity; and (c) decreases with an increase in flea community size. We discuss our results with the idea that the host can mediate interactions among and within flea species.

Relationships between local and regional species richness in flea communities of small mammalian hosts: saturation and spatial scale

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.

Aggregation and species coexistence in fleas parasitic on small mammals

The aggregation model of coexistence states that species coexistence is facilitated if interspecific aggregation is reduced relative to intraspecific aggregation. We investigated the relationship between intraspecific and interspecific aggregation in 17 component communities (the flea assemblage of a host population) of fleas parasitic on small mammals and hypothesized that interspecific interactions should be reduced relative to intraspecific interactions, facilitating species coexistence. We predicted that the reduction of the level of interspecific aggregation in relation to the level of intraspecific aggregation would be positively correlated with total flea abundance and species richness of flea assemblages. We also expected that the higher degree of facilitation of flea coexistence would be affected by host parameters such as body mass, basal metabolic rate (BMR) and depth and complexity of burrows. Results of this study supported the aggregation model of coexistence and demonstrated that, in general, a) conspecific fleas were aggregated across their hosts; b) flea assemblages were not dominated by negative interspecific interactions; and c) the level of interspecific aggregation in flea assemblages was reduced in relation to the level of intraspecific aggregation. Intraspecific aggregation tended to be correlated positively to body mass, burrow complexity and mass‐independent BMR of a host. Positive interspecific associations of fleas tended to occur more frequently in species‐rich flea assemblages and/or in larger hosts possessing deep complex burrows. Intraspecific aggregation increased relative to interspecific aggregation when species richness of flea infracommunities (the flea assemblage of a host individual) and component communities increased. We conclude that the pattern of flea coexistence is related both to the structure of flea communities and affinities of host species.

Abundance and distribution of fleas on desert rodents: linking Taylor's power law to ecological specialization and epidemiology

We investigated variation in the abundance-prevalence relationships of fleas among 17 different flea-host associations as well as among different species of hosts and fleas in the Negev desert. We explored variation in the value of exponent of Taylor's power relationship with changes in flea community size and flea specialization (host specificity and seasonal pattern of activity). We tested if a simple epidemiological model can reproduce the pattern of the abundance-prevalence relationship. We confirmed aggregated distribution of fleas within a population of host species as well as across a whole host community and the existence of a positive relationship between local flea abundance and their prevalence. Prevalence, mean abundance and variance of abundance were significantly higher in host specific than host opportunistic fleas. When ecological specialization was considered, based on a seasonal pattern of activity, these parameters were higher in year-round-active than seasonal fleas. The degree of flea specialization and flea community richness affected the pattern of the relationship between mean abundance and its variance. Power law slopes decreased with increasing richness of flea community. A simple epidemiological model based on mean flea abundance and degree of aggregation, corrected for host sample size, can predict the observed pattern of prevalence. In some cases, observed flea prevalence was higher than that predicted from the epidemiological model. The discrepancy of the observed prevalence from that predicted by the model can be explained by either a relatively low negative effect of flea parasitism on a host (at least, in terms of pathology) or strong resistance of a host to flea parasitism or both.

Abundance patterns and coexistence processes in communities of fleas parasitic on small mammals

The abundance of a given species in a community is likely to depend on both the total abundance and diversity of other species making up that community. A large number of co‐occurring individuals or co‐occurring species may decrease the abundance of any given species via diffuse competition; however, indirect interactions among many co‐occurring species can have positive effects on a focal species. The existence of diffuse competition and facilitation remain difficult to demonstrate in natural communities. Here, we use data on communities of fleas ectoparasitic on small mammals from 27 distinct geographical regions to test whether the abundance of any given flea species in a community is affected by either the total abundance of all other co‐occurring flea species, or the species richness and/or taxonomic diversity of the flea community. At all scales of analysis, i.e. whether we compared the same flea species on different host species, or different flea species, two consistent results emerged. First, the abundance of a given flea species correlates positively with the total abundance of all other co‐occurring flea species in the community. Second, the abundance of any given flea species correlates negatively with either the species richness or taxonomic diversity of the flea community. The results do not support the existence of diffuse competition in these assemblages, because the more individuals of other flea species are present on a host population, the more individuals of the focal species are there as well. Instead, we propose explanations involving either apparent facilitation among flea species via suppression of host immune defenses, or niche filtering processes acting to restrict the taxonomic composition and abundance of flea assemblages.

THE COMPARISON BETWEEN FLEA COMMUNITIES ON TEN SPECIES OF SMALL MAMMALS IN THE FOCI OF HUMAN PLAGUE IN YUNNAN CHINA

Abstract Fleas on the body surface of a certain species of small mammal form a special flea community. On the basis of a series of calculations, a comparison of flea communities on ten species of small mammals in the foci of human plague in Yunnan Province, China, was carried out through a fuzzy clustering analysis. The three flea communities on Rattus flavipectus, Rattus nitidus and Rattus rattus sladeni, which belong to the same genus (genus Rattus) showed a higher similarity than others and they clustered into a group in the phenogram of the clustering analysis. Other flea communities on Anourosorez squamipes Suncus murinus Crocidura attenuata Apodemus draca Eothenomys eleusis, Niviventer confucianus and Mus pahari showed lower similarities between each other. The result suggests that the small mammal hosts may be the most important factor which strongly influence the similarity of ectoparasitic flea communities. When the hosts have a close affinity in the taxonomy, the flea communities on their body surface tend to be similar. The hosts in different genera in this case study tend to have different types of ectoparasitic flea communities.

Unsaturated Fish Communities in African Rivers

Unsaturated Fish Communities in African Rivers