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.

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.

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 2014–2020. 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.

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.

Chigger mites (Trombiculidae) are temporary habitat-specific ectoparasites that often occur on rodents. Little ecological data are available on chiggers associated with rodents in South Africa. The study aims were to (1) record the chigger species associated with rodents in the savanna, (2) assess if chigger species display parasitope preference on the rodent body and (3) compare the distribution of chigger species in natural, agricultural and urban habitats. Rodents (n = 314) belonging to eight genera were trapped in the savanna biome during 2014 and 2015. Twelve chigger species, of which five are recently described species, were recorded from 161 rodent hosts. The data include three new country locality records. Microtrombicula mastomyia was the most prevalent species across sampling seasons and habitat types. Significant parasitope preference was recorded for two species, with the ear, face and tail base some of the preferred attachment sites. Sampling season and habitat type had a significant effect on chigger communities with summer and agricultural habitats recording the highest species richness, while the highest species diversity was recorded in natural habitats. The study contributes to our current knowledge regarding rodent-associated chigger diversity and distribution in South Africa and further highlights the importance of environmental characteristics in shaping chigger communities.

The African penguin (Spheniscus demersus) is a critically endangered species endemic to southern Africa. Limited information is available on the parasite diversity associated with the species in natural settings. This study explores the diversity and incidence of parasites associated with African penguins and their nests, and records the effect of host and environmental factors on parasite infestation. Ecto-, haemo- and helminth parasites were recorded from 210 adult birds, 583 chicks and 628 nests across five colonies (two mainland and three islands) along the south-western coast of South Africa, in 2016 and 2017. Mean nest density (total and active nests) and climate variables (temperature and precipitation) were obtained for each colony. Parapsyllus humboldti was the most abundant and prevalent ectoparasite on penguins and in nests (69.10 and 57.80%, respectively), while Piroplasmorida/Haemospororida (33.51%) and Cardiocephaloides spp. (56.17%) were the most prevalent haemo- and helminth parasites of penguins, respectively. In general parasite abundance and prevalence was significantly affected by penguin age (chicks vs adults), location (mainland vs islands), nest density (total and active nests) and season (spring vs autumn/winter). It is concluded that parasite infestations are structured and that penguin chicks at mainland colonies are more susceptible to parasite infestations during spring.

To reveal factors responsible for spatial variation in parasite community composition we studied patterns of similarity in helminth species composition in two closely-related rodents (Rhabdomys pumilio and Rhabdomys dilectus) that differ in their social and spatial behaviour and live under different environmental conditions across 20 localities in South Africa. We asked whether the two hosts harbour similar assemblages, whether these are more dissimilar between than within hosts and if host social structure, behaviour or environment affects similarity patterns in helminth infracommunities within and among localities. We also investigated whether similarity in species composition of helminth component communities decreases with an increase of geographic distance between host populations. We found that the pattern of space use by the hosts rather than their social behaviour promotes differences in helminth species composition between host species as well as among host populations from different localities. The rate of distance decay of similarity in species composition of helminth component communities differed between the two hosts due to difference in the degree of environmental variation across their geographic ranges. We conclude that patterns of spatial variation in helminth species composition are driven mainly by host spatial behaviour and, to a lesser extent, by environment-associated factors.

We studied nestedness and its relationships with beta-diversity in flea communities harboured by three closely related rodent species (Rhabdomys pumilio, Rhabdomys intermedius, Rhabdomys dilectus) at two spatial scales (within and among host populations) in South Africa and asked (a) whether variation in species composition of flea communities within and among host populations follows a non-random pattern; if yes, (b) what are the contributions of nestedness and species turnover to dissimilarity (= beta-diversity) among flea communities at the two scales; and (c) do the degree of nestedness and its contribution to beta-diversity differ among host species (social vs solitary) and between scales. We found that nestedness in flea assemblages was more pronounced (a) in social than solitary host species and (b) at lower (among host individuals within populations) than at higher scale (among host populations). We also found that higher degree of nestedness was associated with its higher contribution to beta-diversity. Our findings support earlier ideas that parasite community structure results from the processes of parasite accumulation by hosts rather than from the processes acting within parasite communities.

We studied the structure of chigger mite (Trombiculidae) communities parasitic on a South African rodent, Rhabdomys pumilio. We aimed to determine whether: (a) different chigger species differ in preferences for certain body areas of a host and (b) chigger assemblages among body areas of the same host individual, are structured and if so, whether the structure of these assemblages is aggregative or segregative. Rhabdomys pumilio is parasitized by seven chigger species belonging to six genera. The three most abundant species (Leptotrombidium sp. nr. muridium, Schoutedenichia sp. and Neoschoengastia sp. A) displayed a non-random distribution across the host body, with the two most abundant species (L. sp. nr. muridium and Schoutedenichia sp.) significantly associated with the tail area. In addition, whenever non-randomness of chigger co-occurrence in the same body area was recorded, it indicated positive but not negative co-occurrences of different species. This might be due to similarity of chigger species in resource needs and strategies to avoid host defence efforts.

Laelaps giganteus and Laelaps muricola (Mesostigmata; Laelapidae) are widespread and locally abundant host generalists on small mammals in southern Africa. The large host range and complex life history of these ectoparasites may allude to possible intraspecific cryptic diversity in these taxa. To assess genetic and morphological diversity in L. giganteus and L. muricola, we sampled 228 rodents at eight localities in South Africa. This sample included nine previously recorded host species and on these, L. muricola was only recorded from Mastomys natalensis and Micaelamys namaquensis while L. giganteus was found on Rhabdomys dilectus and Lemniscomys rosalia. Phylogenetic analyses of partial mtDNA cytochrome oxidase subunit I (COI) and nuclear ITS1 data strongly supported the recognition of L. giganteus and L. muricola, a scenario partly supported by the Tropomyosin intron. Strong support for evolutionary distinct lineages within L. giganteus is found: L. giganteus lineage 1 is confined to R. dilectus and L. giganteus lineage 2 is confined to L. rosalia. These host specific monophyletic lineages were also separated by 9·84% mtDNA sequence divergence and 3·44% nuclear DNA sequence divergence. Since quantitative morphometric analyses were not congruent with these findings, these two lineages more than likely represent cryptic species.

Within southern Africa, the widely distributed four-striped mouse genus (Rhabdomys) is parasitized by, amongst others, the specific ectoparasitic sucking louse, Polyplax arvicanthis. Given the presence of significant geographically structured genetic divergence in Rhabdomys, and the propensity of parasites to harbour cryptic diversity, the molecular systematics of P. arvicanthis was investigated. Representatives of P. arvicanthis were sampled from Rhabdomys at 16 localities throughout southern Africa. Parsimony and Bayesian gene trees were constructed for the mitochondrial COI, 12S rRNA, 16S rRNA and nuclear CAD genes. Our findings support the existence of 2 genetic groups within P. arvicanthis separated by at least 25% COI sequence divergence, which is comparable to that observed among recognized Polyplax species. We therefore propose that these 2 genetic lineages probably represent distinct species and that the apparent absence of clear morphological differences may point to cryptic speciation. The 2 taxa have sympatric distributions throughout most of the sampled host range and also occasionally occur sympatrically on the same host individual. The co-occurrence of these genetically distinct lineages probably resulted from parasite duplication via host-associated allopatric divergence and subsequent reciprocal range expansions of the 2 parasite taxa throughout southern Africa.

Habitat fragmentation can adversely impact biodiversity, although where remnant fragments of natural vegetation provide favourable conditions the negative effects of fragmentation may be mitigated. Host-parasite systems in fragmented areas have only recently been examined, with parasites generally showing higher prevalence and richness in fragments, mediated by changes in host density. However, the effect of fragmentation on parasite body size and fecundity remains poorly investigated. Thus, here we compared the body size and condition of a generalist rodent host, Rhabdomys pumilio and the body size of 2 common flea species between pristine natural areas and remnant fragments within agriculture areas. Host body length, weight and body condition values were significantly larger in fragments than in pristine natural vegetation. Listropsylla agrippinae fleas showed the same pattern, being significantly larger in fragments, while Chiastopsylla rossi fleas did not differ in size between fragments and natural areas. The differential response of the 2 flea species may reflect the strength of association between the host and parasite, with the former spending a greater proportion of its lifespan on the host. Therefore, in this study agriculture fragments provide better conditions for both an opportunistic rodent and a closely associated flea species.

The gender-biased pattern of parasite infestation has been shown to be a complicated phenomenon that cannot be explained by a single mechanism but rather involves several different mechanisms. We asked what are the factors that affect the manifestation and extent of gender-biased parasitism and studied the relationship between parasite-related (mean abundance, mean species richness and total species richness of all parasites), host-related (rodent density and proportion of reproductive males and females both separately and together) and environment-related (mean daily maximal and minimal temperatures, rainfall and relative humidity) factors and the magnitude of gender-biased infestation of a South African rodent Rhabdomys pumilio by ixodid ticks, gamasid mites, lice and fleas. We found that spatial variation in gender differences in parasite infestation was affected by parasite-, host- and environment-related factors, although the set of factors affecting gender differences in infestation differed among higher taxa of ectoparasites. Gender differences in infestation by fleas and lice were affected mainly by parasite-related factors, whereas gender differences in infestation by ticks and, in part, by mites were affected mainly by host-related and environmental factors. In addition, spatial variation in most measures of gender difference in mite infestation remained unexplained.

We asked whether the occurrence and the extent of host gender-biased parasitism vary among higher parasite taxa, among individual species within these taxa and within parasite species among localities. To answer this question, we studied prevalence, abundance, species richness and the level of aggregation of ectoparasites (ticks, mites, lice and fleas) collected from male and female striped mice (Rhabdomys pumilio) in 9 localities of the Western Cape Province of South Africa. We used meta-analyses to compare parasitological variables between male and female hosts across localities for individual parasite species and higher taxa as well as across parasite species within a higher taxon. Whenever gender-biased parasitism was found, it indicated higher infestation of male than female hosts (except 1 low abundant mite species). We found that the occurrence and extent of gender-biased infestation varied mainly within a parasite species among localities and among parasite species within a higher taxon but not among parasite taxa. Our results suggest that the extent of a gender-biased pattern of parasite infestation of the same host may not only involve host-related mechanisms but also depends on biological features of a particular parasite species.