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.

Survey of flea infestation in cats and dogs in Western Andalusia, Spain: Seasonality and other risk factors for flea infestation.

This epidemiological survey aims to provide an update on the main flea species that parasitize domestic animals in the Western Andalusia assessing several ecological features that could be considered as possible risk factors for flea infestation. Over a 19-month period (June 2021 to January 2023), we obtained a total of 802 flea samples from 182 dogs (Canis lupus familiaris, Carnivora: Canidae, Linnaeus, 1758) and 78 cats (Felis silvestris catus, Carnivora: Felidae, Schreber, 1775). For each parasitized host, an epidemiological survey was completed, including the following information: geographical origin, age, sex, rural or urban habitat, type of animal's lifestyle (domestic or non-domestic), health status, cohabiting or not with other animals and the total number of collected fleas. The most common species was Ctenocephalides felis (Siphonaptera: Pulicidae) (Bouché, 1835) with a total of 713 specimens, which accounted for 89% of the total fleas. The second most abundant species was Pulex irritans (Siphonaptera: Pulicidae) (Linnaeus, 1758) with a total of 46 collected fleas (6% of the total). The remaining species identified were Archaeopsylla erinacei (Siphonaptera: Pulicidae) (Bouché, 1835) (25 specimens), Spilopsyllus cuniculi (Siphonaptera: Pulicidae) (Dale, 1878) (12 specimens) and Ctenocephalides canis (Siphonaptera: Pulicidae) (Curtis, 1826) (6 specimens), which accounted for 3%, 1% and 1%, respectively, of the total fleas collected. The months with the highest number of collected fleas were, in ascending order, May 2022, September 2021 and July 2021. Dogs had a greater diversity of flea species, and flea sex ratios were female biased in all identified species and among all studied hosts. Finally, we identified some potential host risk factors that promoted higher flea intensities, such as living in rural areas, or presenting other pathologies.

Evidence of castrations in fleas of small mammals (Chiroptera and Rodentia) from Argentina.

Morphological abnormalities in fleas seem to be common in nature and are under reported in Argentina. In this note, we describe anomalies in two males and one female of Alectopsylla unisetosa Mahnert (Ischnopsyllidae) and one of Polygenis sp. (Rhopalopsyllidae) male collected from small mammals in the provinces of Neuquen and Salta, Argentina. In all specimens, the anomalies were observed at the level of the genitalia recognized as partial castration. The structures mainly affected were the modified abdominal segments, the aedeagus (in male), and the spermatheca (in female). The present communication is the first one devoted exclusively to teratogenous fleas in Argentina.

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.

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.

Patterns and drivers of vector-borne microparasites in a classic metapopulation.

Many organisms live in fragmented populations, which has profound consequences on the dynamics of associated parasites. Metapopulation theory offers a canonical framework for predicting the effects of fragmentation on spatiotemporal host­parasite dynamics. However, empirical studies of parasites in classical metapopulations remain rare, particularly for vector-borne parasites. Here, we quantify spatiotemporal patterns and possible drivers of infection probability for several ectoparasites (fleas, Ixodes trianguliceps and Ixodes ricinus) and vector-borne microparasites (Babesia microti, Bartonella spp., Hepatozoon spp.) in a classically functioning metapopulation of water vole hosts. Results suggest that the relative importance of vector or host dynamics on microparasite infection probabilities is related to parasite life-histories. Bartonella, a microparasite with a fast life-history, was positively associated with both host and vector abundances at several spatial and temporal scales. In contrast, B. microti, a tick-borne parasite with a slow life-history, was only associated with vector dynamics. Further, we provide evidence that life-history shaped parasite dynamics, including occupancy and colonization rates, in the metapopulation. Lastly, our findings were consistent with the hypothesis that landscape connectivity was determined by distance-based dispersal of the focal hosts. We provide essential empirical evidence that contributes to the development of a comprehensive theory of metapopulation processes of vector-borne parasites.

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.

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.

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.

Predictive factors for flea occurrence in red foxes (Vulpes vulpes) from semi-arid Mediterranean environments.

The role of red fox as host for a wide range of parasites, particularly fleas and other arthropods causing vector-borne diseases, in combination with its capability to adapt to anthropized environments, makes this wild canid an epidemiologically remarkable species at the wildlife-domestic-human interface, especially in the present time of rise of emerging and re-emerging diseases. This study evaluated the prevalence and parasite intensity of fleas in 88 foxes from Murcia Region (Southeastern Spain) and determined the geographic distribution of areas with the highest potential risk of flea presence. Pulex irritans, Ctenocephalides felis, Spilopsyllus cuniculi and Nosopsyllus fasciatus were identified. The overall prevalence was 76.13%. This is the first time that N. fasciatus has been reported in foxes from Murcia Region. The predictive model established a certain pattern to determine the areas with the highest risk of acquiring fleas. Positive correlation of daily potential evapotranspiration (ET0 ) in winter and the opposite effect occurring for ET0 in summer were obtained, as well as positive correlations for mean daily temperature (Tmean ) in summer and mean precipitation (Pmean ) in winter and summer. The model was also found positively correlated in the forest habitat ecotone areas and the anthropized areas.

Host specificity of flea parasites of mammals from the Andean Biogeographic Region.

Host specificity of fleas affects their biodiversity that plays a major role in determining the potential transmission routes by pathogens through vertebrate hosts, including humans. In the Biogeographic Andean region, numerous systematic and ecological studies have been conducted, revealing a high diversity of flea taxa of mammals and the presence of pathogenic organisms transmitted by fleas; however, the degree of preference with which each flea species associates with a mammal host remains poorly understood in this region. Herein, host specificity in mammal fleas from the Andean region was analysed. We employed the number of host species for each flea species and the index of host specificity STD *. Following the literature, 144 species and 13 subspecies of fleas (31 genera and 10 families) have been described in the Andean biogeographic region; 76 taxa are endemic to this region. To carry out the analyses of host specificity, we considered 1759 records of fleas collected from 124 species and 59 genera of wild and domestic mammals, mostly rodent species (85.9%). Our results indicate that typical Andean fleas are genus or family host specific (mostly STD * less than 3.0). More diverse mammal hosts are parasitized by more diverse flea genera and families and these hosts are phylogenetically related. Otherwise, these hosts are associated with different flea lineages, suggesting the interaction of ecological and evolutionary mechanisms (host-switching, ecological adaptations and co-evolutionary alternation). The fields of disease ecology and One Health are considering the host specificity of arthropod vectors as an important point to understand the mechanisms of emergence and re-emergence of diseases. Our results allow us to estimate the risk of diseases involving fleas in the Andean region.

Seasonality and other risk factors for fleas infestations in domestic dogs and cats.

Fleas in the genus Ctenocephalides are the most clinically important parasitic arthropods of dogs and cats worldwide yet risk factors that might increase the risk of infestation in small animals remains unclear. Here we developed a supervised text mining approach analysing key aspects of flea epidemiology using electronic health records from domestic cats and dogs seen at a sentinel network of 191 voluntary veterinary practices across Great Britain between March 2014 and July 2020. Our methods identified fleas as likely to have been present during 22,276 of 1,902,016 cat consultations (1.17%) and 12,168 of 4,844,850 dog consultations (0.25%). Multivariable logistic regression modelling found that animals originating from areas of least deprivation were associated with 50% reductions in odds of veterinary-recorded flea infestation compared to the most deprived regions in England. Age of the animal was significantly associated with flea presentation in both cats and dogs, with cases peaking before animals reached 12 months. Cases were recorded through each study years, peaking between July and October, with fluctuations between each year. Our findings can be used towards healthcare messaging for veterinary practitioners and owners.

Vector-borne protozoan and bacterial pathogen occurrence and diversity in ectoparasites of the Egyptian Rousette bat.

Bats are known reservoir hosts for a wide variety of parasites and pathogens, including bacteria and protozoans. Some of these pathogens are vector-borne, and although their role is poorly studied, ectoparasites may contribute significantly to their transmission. The aim of this study was to molecularly detect the presence of vector-borne microorganisms in bat-associated ectoparasites to explore their diversity and distribution in these insects. We tested the presence of Bartonella spp., Polychromophilus spp., and Trypanosoma spp. in bat flies and bat fleas collected from 56 Egyptian Rousette bats (Rousettus aegyptiacus), using conventional PCR. We found a high prevalence of 43.9% (47/107) of Bartonella spp. in bat flies, but a low prevalence of 6.6% (4/61) in bat fleas. Polychromophilus and Trypanosoma DNA were absent in both bat flies and bat fleas. Furthermore, we found novel gltA Bartonella sequences, as well as genotypes that are highly similar to recently described and potentially zoonotic ones. Our results show high diversity of Bartonella in bat flies, however, their role in pathogen transmission is still unknown and should be further explored.

Host body mass, not sex, affects ectoparasite loads in yellow-necked mouse Apodemus flavicollis.

We investigated the presence and potential causes of sex bias in ectoparasite infestations in the yellow-necked mouse Apodemus flavicollis. We compared the natural tick and flea burdens of male and female mice in a temperate beech forest and assessed whether the observed differences were driven by host sex or body mass. We found that males were more heavily infested by ticks compared to female mice. However, this difference was driven by host body mass, and not sex itself. Host body mass positively correlated with flea loads, but there was no evidence of sex bias in flea abundance. In addition, the abundance of both ticks and fleas infesting yellow-necked mice changed over time, both seasonally (month to month) and annually (year to year). Our results underscore the importance of the sexual size dimorphism and the parasite taxon as the primary factors that influence the occurrence of sex-biased parasitism in small mammals.

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.

Ecological factors shaping the ectoparasite community assembly of the Azara's Grass Mouse, Akodon azarae (Rodentia: Cricetidae).

Parasites are integral members of the global biodiversity. They are useful indicators of environmental stress, food web structure and diversity. Ectoparasites have the potential to transmit vector-borne diseases of public health and veterinary importance and to play an important role in the regulation and evolution of host populations. The interlinkages between hosts, parasites and the environment are complex and challenging to study, leading to controversial results. Most previous studies have been focused on one or two parasite groups, while hosts are often co-infected by different taxa. The present study aims to assess the influence of environmental and host traits on the entire ectoparasite community composition of the rodent Akodon azarae. A total of 278 rodents were examined and mites (Mesostigmata), lice (Phthiraptera), ticks (Ixodida) and fleas (Siphonaptera) were determined. A multi-correspondence analysis was performed in order to analyze interactions within the ectoparasite community and the influence of environmental and host variables on this assembly. We found that environmental variables have a stronger influence on the composition of the ectoparasite community of A. azarae than the host variables analyzed. Minimum temperature was the most influential variable among the studied. In addition, we found evidence of agonistic and antagonistic interactions between ticks and mites, lice and fleas. The present study supports the hypothesis that minimum temperature plays a major role in the dynamics that shape the ectoparasite community of A. azarae, probably through both direct and indirect processes. This finding becomes particularly relevant in a climate change scenario.

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.

Host phylogeny and ecology, but not host physiology, are the main drivers of (dis)similarity between the host spectra of fleas: application of a novel ordination approach to regional assemblages from four continents.

We investigated the patterns of phylogenetic and functional (dis)similarity in the species composition of host spectra between co-habitating generalist flea species in regional assemblages from four continents (Europe, Asia, North America and Africa) using a recently developed ordination approach (Double Similarity Principal Component Analysis). From the functional perspective, we considered physiological [body mass and basal metabolic rate (BMR)] and ecological (shelter depth and complexity) host traits. We asked (a) whether host phylogeny, physiology or ecology is the main driver of (dis)similarities between flea host spectra and (b) whether the patterns of phylogenetic and functional (dis)similarity in host spectra vary between flea assemblages from different continents. Phylogenetic similarity between the host spectra was highest in Africa, lowest in North America and moderate in Europe and Asia. In each assemblage, phylogenetic clusters of hosts dominating in the host spectra could be distinguished. The functional similarity between the host spectra of co-occurring fleas was low for shelter structure in all assemblages and much higher for body mass and BMR in three of the four assemblages (except North America). We conclude that host phylogeny and shelter structure are the main drivers of (dis)similarity between the host spectra of co-habitating fleas. However, the effects of these factors on the patterns of (dis)similarity varied across continents.

Diversity and distribution of ectoparasite taxa associated with Micaelamys namaquensis (Rodentia: Muridae), an opportunistic commensal rodent species in South Africa.

South Africa boasts a rich diversity of small mammals of which several are commensal and harbour parasites of zoonotic importance. However, limited information is available on the parasite diversity and distribution associated with rodents in South Africa. This is particularly relevant for Micaelamys namaquensis (Namaqua rock mouse), a regionally widespread and locally abundant species that is often commensal. To address the paucity of data, the aims of the study were to record the ectoparasite diversity associated with M. namaquensis and develop distribution maps of lice and mites associated with M. namaquensis and other rodents in South Africa. Micaelamys namaquensis individuals (n = 216) were obtained from 12 localities representing multiple biomes during 2017­2018. A total of 5591 ectoparasites representing 5 taxonomic groups ­ fleas, lice, mesostigmatid mites, chiggers and ticks was recorded. These consisted of at least 57 taxa of which ticks were the most speciose (20 taxa). Novel contributions include new host and locality data for several ectoparasite taxa and undescribed chigger species. Known vector species were recorded which included fleas (Ctenocephalides felis, Dinopsyllus ellobius and Xenopsylla brasiliensis) and ticks (Haemaphysalis elliptica, Rhipicephalus appendiculatus and Rhipicephalus simus). Locality records indicate within-taxon geographic differences between the 2 louse species and the 2 most abundant mite species. It is clear that M. namaquensis hosts a rich diversity of ectoparasite taxa and, as such, is an important rodent species to monitor in habitats where it occurs in close proximity to humans and domestic animals.

Fitness consequences of host colonization in two generalist fleas: Context-dependency and the effect of spatial co-occurrence.

We studied the fitness consequences of colonizing a novel host by experimental lines of fleas (Synosternus cleopatrae and Xenopsylla ramesis) maintained for 18-22 generations on the principal or novel (sympatric or allopatric) hosts via number, developmental success and size of the offspring of the fleas exploiting these hosts. We asked whether (a) fitness on non-principal hosts increases after prolonged maintenance; (b) the colonization success depends on the spatial co-occurrence of a flea and a host and (c) colonization of a novel host is accompanied by a decreased ability to exploit an original host. The ability of fleas to colonize novel hosts differed between species, with S. cleopatrae, but not X. ramesis, increasing its offspring production on novel hosts. Spatial co-occurrence did not affect colonization success. Maintenance on an alternative host was not accompanied by decreased adaptation to the original host. When fleas returned to the original host, their reproductive output was higher than that of their ancestors. We conclude that the success of colonizing a novel host is (a) context-dependent and varies between flea and host species and (b) not accompanied by the loss of ability to exploit an ancestral host but may lead to an increase in this ability.