Genetic structure and invasion history of the house mouse (Mus musculus domesticus) in Senegal, West Africa: a legacy of colonial and contemporary times.

Knowledge of the genetic make-up and demographic history of invasive populations is critical to understand invasion mechanisms. Commensal rodents are ideal models to study whether complex invasion histories are typical of introductions involving human activities. The house mouse Mus musculus domesticus is a major invasive synanthropic rodent originating from South-West Asia. It has been largely studied in Europe and on several remote islands, but the genetic structure and invasion history of this taxon have been little investigated in several continental areas, including West Africa. In this study, we focussed on invasive populations of M. m. domesticus in Senegal. In this focal area for European settlers, the distribution area and invasion spread of the house mouse is documented by decades of data on commensal rodent communities. Genetic variation at one mitochondrial locus and 16 nuclear microsatellite markers was analysed from individuals sampled in 36 sites distributed across the country. A combination of phylogeographic and population genetics methods showed that there was a single introduction event on the northern coast of Senegal, from an exogenous (probably West European) source, followed by a secondary introduction from northern Senegal into a coastal site further south. The geographic locations of these introduction sites were consistent with the colonial history of Senegal. Overall, the marked microsatellite genetic structure observed in Senegal, even between sites located close together, revealed a complex interplay of different demographic processes occurring during house mouse spatial expansion, including sequential founder effects and stratified dispersal due to human transport along major roads.

Invasion genetics of a human commensal rodent: the black rat Rattus rattus in Madagascar.

Studies focusing on geographical genetic patterns of commensal species and on human history complement each other and provide proxies to trace common colonization events. On Madagascar, the unintentional introduction and spread of the commensal species Rattus rattus by people may have left a living clue of human colonization patterns and history. In this study, we addressed this question by characterizing the genetic structure of natural populations of R. rattus using both microsatellites and mitochondrial sequences, on an extensive sampling across the island. Such data sets were analysed by a combination of methods using population genetics, phylogeography and approximate Bayesian computation. Our results indicated two introduction events to Madagascar from the same ancestral source of R. rattus, one in the extreme north of the island and the other further south. The latter was the source of a large spatial expansion, which may have initially started from an original point located on the southern coast. The inferred timing of introduction events-several centuries ago-is temporally congruent with the Arabian trade network in the Indian Ocean, which was flourishing from the middle of the first millennium.

Beyond an AFLP genome scan towards the identification of immune genes involved in plague resistance in Rattus rattus from Madagascar.

Genome scans using amplified fragment length polymorphism (AFLP) markers became popular in nonmodel species within the last 10 years, but few studies have tried to characterize the anonymous outliers identified. This study follows on from an AFLP genome scan in the black rat (Rattus rattus), the reservoir of plague (Yersinia pestis infection) in Madagascar. We successfully sequenced 17 of the 22 markers previously shown to be potentially affected by plague-mediated selection and associated with a plague resistance phenotype. Searching these sequences in the genome of the closely related species Rattus norvegicus assigned them to 14 genomic regions, revealing a random distribution of outliers in the genome (no clustering). We compared these results with those of an in silico AFLP study of the R. norvegicus genome, which showed that outlier sequences could not have been inferred by this method in R. rattus (only four of the 15 sequences were predicted). However, in silico analysis allowed the prediction of AFLP markers distribution and the estimation of homoplasy rates, confirming its potential utility for designing AFLP studies in nonmodel species. The 14 genomic regions surrounding AFLP outliers (less than 300 kb from the marker) contained 75 genes encoding proteins of known function, including nine involved in immune function and pathogen defence. We identified the two interleukin 1 genes (Il1a and Il1b) that share homology with an antigen of Y. pestis, as the best candidates for genes subject to plague-mediated natural selection. At least six other genes known to be involved in proinflammatory pathways may also be affected by plague-mediated selection.

Development of Trichosomoides nasalis (Nematoda: Trichinelloidea) in the murid host: evidence for larval growth in striated muscle fibres.

Trichosomoides nasalis (Trichinelloidea) is a parasite of Arvicanthis niloticus (Muridae) in Senegal. Female worms that harbour dwarf males in their uteri, occur in the epithelium of the nasal mucosa. Young laboratory-bred A. niloticus were either fed females containing larvated eggs or intraperitoneally injected with motile first-stage larvae recovered from female uteri. Both resulted in successful infection. Organs examined during rodent necropsy were blood and lymphatic circulatory systems (heart, large vessels, lymphnodes), lungs, liver, kidneys, thoracic and abdominal cavities, thoracic and abdominal muscular walls, diaphragm, tongue, and nasal mucosa. Development to adult nasal stages took three weeks. Recovery of newly hatched larvae from the peritoneal fluid at four-eight hours after oral infection suggests a direct passage from the stomach or intestinal wall to the musculature. However, dissemination through the blood, as observed with Trichinella spiralis, cannot be excluded even though newly hatched larvae of T. nasalis are twice as thick (15 µm). Developing larvae were found in histological sections of the striated muscle of the abdominal and thoracic walls, and larvae in fourth moult were dissected from these sites. Adult females were found in the deep nasal mucosa where mating occurred prior to worms settling in the nasal epithelium. The present study shows a remarkable similarity between T. nasalis and Trichinella species regarding muscle tropism, but the development of T. nasalis is not arrested at the late first-larval stage and does not induce transformation of infected fibres into nurse cells. T. nasalis seems a potential model to study molecular relations between trichinelloid larvae and infected muscle fibres.

AFLP genome scan in the black rat (Rattus rattus) from Madagascar: detecting genetic markers undergoing plague-mediated selection.

The black rat (Rattus rattus) is the main reservoir of plague (Yersinia pestis infection) in Madagascar's rural zones. Black rats are highly resistant to plague within the plague focus (central highland), whereas they are susceptible where the disease is absent (low altitude zone). To better understand plague wildlife circulation and host evolution in response to a highly virulent pathogen, we attempted to determine genetic markers associated with plague resistance in this species. To this purpose, we combined a population genomics approach and an association study, both performed on 249 AFLP markers, in Malagasy R. rattus. Simulated distributions of genetic differentiation were compared to observed data in four independent pairs, each consisting of one population from the plague focus and one from the plague-free zone. We found 22 loci (9% of 249) with higher differentiation in at least two independent population pairs or with combining P-values over the four pairs significant. Among the 22 outlier loci, 16 presented significant association with plague zone (plague focus vs. plague-free zone). Population genetic structure inferred from outlier loci was structured by plague zone, whereas the neutral loci dataset revealed structure by geography (eastern vs. western populations). A phenotype association study revealed that two of the 22 loci were significantly associated with differentiation between dying and surviving rats following experimental plague challenge. The 22 outlier loci identified in this study may undergo plague selective pressure either directly or more probably indirectly due to hitchhiking with selected loci.

Plio-Pleistocene history of West African Sudanian savanna and the phylogeography of the Praomys daltoni complex (Rodentia): the environment/geography/genetic interplay.

Rodents of the Praomys daltoni complex are typical inhabitants of the Sudanian savanna ecosystem in western Africa and represent a suitable model for testing the effects of Quaternary climatic oscillations on extant genetic variation patterns. Phylogeographical analyses of mitochondrial DNA sequences (cytochrome b) across the distribution range of the complex revealed several well-defined clades that do not support the division of the clade into the two species currently recognized on the basis of morphology, i.e. P. daltoni (Thomas, 1892) and Praomys derooi (Van der Straeten & Verheyen 1978). The observed genetic structure fits the refuge hypothesis, suggesting that only a small number of populations repeatedly survived in distinct forest-savanna mosaic blocks during the arid phases of the Pleistocene, and then expanded again during moister periods. West African rivers may also have contributed to genetic differentiation, especially by forming barriers after secondary contact of expanding populations. The combination of three types of genetic markers (mtDNA sequences, microsatellite loci, cytogenetic data) provides evidence for the presence of up to three lineages, which most probably represent distinct biological species. Furthermore, incongruence between nuclear and mtDNA markers in some individuals unambiguously points towards a past introgression event. Our results highlight the importance of combining different molecular markers for an accurate interpretation of genetic data.

Geographic patterns of inversion polymorphisms in a wild African rodent, Mastomys erythroleucus.

By suppressing recombination and reducing gene flow, chromosome inversions favor the capture and protection of advantageous allelic combinations, leading to adaptive polymorphisms. However, studies in non-model species remain scarce. Here we investigate the distribution of inversion polymorphisms in the multimammate rat Mastomys erythroleucus in West Africa. More than 270 individuals from 52 localities were karyotyped using G-bands and showed widespread polymorphisms involving four chromosome pairs. No significant deviations from Hardy-Weinberg equilibrium were observed either through space or time, nor were differences retrieved in viability or sex contribution between cytotypes. The distribution of chromosomal variation, however, showed perfect congruence with that of mtDNA-based phylogeographic clades. Thus, inversion diversity patterns in M. erythroleucus appeared more related to historical and/or demographic processes than to climate-based adaptive features. Using cross-species chromosome painting and G-banding analyses to identify homologous chromosomes in related out-group species, we proposed a phylogenetic scenario that involves ancestral-shared polymorphisms and subsequent lineage sorting during expansion/contraction of West African savannas. Our data suggest that long-standing inversion polymorphisms may act as regions in which adaptation genes may accumulate (nucleation model).

CCR5 polymorphism and plague resistance in natural populations of the black rat in Madagascar.

Madagascar remains one of the world's largest plague foci. The black rat, Rattus rattus, is the main reservoir of plague in rural areas. This species is highly susceptible to plague in plague-free areas (low-altitude regions), whereas rats from the plague focus areas (central highlands) have evolved a disease-resistance polymorphism. We used the candidate gene CCR5 to investigate the genetic basis of plague resistance in R. rattus. We found a unique non-synonymous substitution (H184R) in a functionally important region of the gene. We then compared (i) CCR5 genotypes of dying and surviving plague-challenged rats and (ii) CCR5 allelic frequencies in plague focus and plague-free populations. Our results suggested a higher prevalence of the substitution in resistant animals compared to susceptible individuals, and a tendency for higher frequencies in plague focus areas compared to plague-free areas. Therefore, the CCR5 polymorphism may be involved in Malagasy black rat plague resistance. CCR5 and other undetermined plague resistance markers may provide useful biological information about host evolution and disease dynamics.

Description of two new species of Nippostrongylinae (Nematoda: Heligmonellidae) coparasites in three sympatric species of Mastomys spp. (Rodentia: Muridae) from Senegal.

Two new species of heligmosomoid Trichostrongylina nematodes belonging to the genera Neoheligmonella Durette-Desset, 1970 and Heligmonina Baylis, 1928 are described. They are parasitic in the small intestine of three species of Mastomys from Senegal living in sympatry: M. natalensis (Smith, 1834), M. erythroleucus (Temminck, 1853) and M. huberti (Wroughton, 1909). Neoheligmonella granjoni n. sp. is closely related to three species from Senegal. They concern: N. bai Diouf & Durette-Desset, 2002 and N. dielmensis Diouf, Bâ & Durette-Desset, 1998, both parasitic in Arvicanthis niloticus Geoffroy, 1903 and N. mastomysi Diouf et al., 1998, a parasite of M. erythroleucus. N. granjoni n. sp. differs from these species by having 15 cuticular ridges at mid-body versus 13, a large carene and spicules taking up 10-15% of body length versus 5.3-7.1%. Heligmonina kanei n. sp. differs from the most related species H. kotoensis Diouf, Daouda & Durette-Desset 2005, a parasite of M. natalensis from Benin in the following features: spicules taking up 11.6% of body length on average versus 16.8%; a female tail three times longer than the distance anus-vulva versus a tail of equivalent size to this distance. In N. granjoni n. sp., where the material is abundant in all three hosts, the infra-specific variations observed (morphological or morphometrical) were not related to the host species. This is the first report of the genera Neoheligmonella and Heligmonina in M. huberti. The relevance of the phenomenon of host capture concerning the evolution of these two genera is confirmed.

Description of two new species of Nippostrongylinae (Nematoda, Trichostrongyuna, Heligmosomoidea) coparasites in Arvicanthis ansorgei (Muridae) from Cameroon.

In the small intestine of a single Arvicanthis ansorgei from Cameroon, two new species of Nippostrongylinae were found: Neoheligmonella zero n. sp. and Heligmonina comerounensis n. sp. N. zero belongs to the Neoheligmonella species in which the right dorsal ridge is poorly developed. Among these species, N. bainae (Durette-Desset, 1970), a parasite of Steatomys opinus from Burkina Fasso, is a closely related species. It is differentiated by the presence of vulvar alae, the ratio uterus length/body length and the separation of rays 2 and 3 at two thirds of their length. N. zero is also closely related to N. kenyae (Yeh, 1958) a parasite of Rattus rattus kijabius from Kenya by the pattern of the caudal bursa and the ratio spicules length/body length. The synlophe of N. kenyae was not described in detail but it is differentiated from N. zero by the position of the excretory pore, situated just posteriorly to the nerve ring and in the female by the ratio ovejector length/body length which is smaller. H. camerounensis n. sp. is characterised by the ratio dorsal ridges/ventral ridges which is 4/7. It is differentiated from the species of which the synlophe has not been described by the pattern of the caudal bursa (type 1-4 with tendancy 1-3-1). It is the first report of Nippostrongylinae species in Cameroon and the first record of a species of the genus Heligmonina in an Arvicanthis.

A longitudinal survey of Borrelia crocidurae prevalence in rodents and insectivores in Senegal.

We report results of a longitudinal survey designed to determine the importance and the dynamics of Borrelia crocidurae, the spirochete responsible for tick-borne relapsing fever in West Africa in rodents and insectivores in a rural area of northern Senegal. A total of 954 animals were caught during bimonthly capture sessions over a two-year period. Positive thick blood smears were recorded in 17.6% of the 740 rodents and 7.3% of the 55 musk shrews tested. Variations of prevalence were analyzed in Arvicanthis niloticus and Mastomys huberti, which represented 62.7% and 28.3%, respectively. of the animals captured, and 65.7% and 27.6%, respectively, of the animals found infected. Borrelia crocidurae prevalence was significantly different between captures and fluctuated separately for each species. Age-specific prevalence of B. crocidurae showed distinct patterns, decreasing with age from 50% in younger juveniles to 3% in older adults for A. niloticus, while increasing with age from 8% to 23% for M. huberti. No relationship was observed with animal abundance or with the season of the year for either species. These findings suggest that the diversity of the population dynamics of host-vector-parasite associations in the Sahel region of Senegal may be a key factor for the relative stability of the borreliosis reservoir.

The spread of tick-borne borreliosis in West Africa and its relationship to sub-Saharan drought.

In West Africa, tick-borne relapsing fever is due to the spirochete Borrelia crocidurae and its geographic distribution is classically limited to the Sahel and Saharan regions where the vector tick Alectorobius sonrai is distributed. We report results of epidemiologic investigations carried out in the Sudan savanna of Senegal where the existence of the disease was unknown. A two-year prospective investigation of a rural community indicated that 10% of the study population developed an infection during the study period. Transmission patterns of B. crocidurae to humans and the small wild mammals who act as reservoirs for infection were similar to those previously described in the Sahel region. Examination of 1,197 burrows and blood samples from 2,531 small mammals indicated a considerable spread of the known area of distribution of A. sonrai and B. crocidurae. The actual spread of the vector and the disease has affected those regions where the average rainfall, before the start of the extended drought in West Africa, reached up to 1,000 mm and corresponds to the movement of the 750-mm isohyet toward the south from 1970 to 1992. Our findings suggest that the persistence of sub-Saharan drought, allowing the vector to colonize new areas in the Sudan savanna of West Africa, is probably responsible for a considerable spread of tick-borne borreliosis in this part of Africa.

A comparative study of three methods of detection of Borrelia crocidurae in wild rodents in Senegal.

In a rural area in Sénégal with a high incidence of tick-borne relapsing fever in humans, Borrelia crocidurae was studied in the blood and brain of wild rodents (Mastomys erythroleucus, Arvicanthis niloticus and Rattus rattus) using 3 methods: (i) direct examination of thick blood films; (ii) intraperitoneal inoculation of blood into white mice; (iii) intraperitoneal inoculation of homogenized cerebral tissue into white mice. Of the 82 rodents examined, the proportion of infected animals was respectively 2.4%, 7.3% and 14.6% for each method, and 18.3% for all 3 methods combined. Of the 12 animals with infected cerebral tissue, only 3 were found to have infected blood. These results suggest that isolated infections of the brain occur frequently in Senegalese wild rodents. Measurement of the real prevalence of B. crocidurae should therefore take into account these infections in addition to blood infections.

Fruit characters as a basis of fruit choice and seed dispersal in a tropical forest vertebrate community.

Interactions between a large community of vertebrate frugivore-granivores (including 7 species of large canopy birds, 19 species of rodents, 7 species of ruminants, and 6 species of monkeys), and 122 fruit species they consume, were studied for a year in a tropical rainforest in Gabon.The results show how morphological characters of fruits are involved in the choice and partitioning of the available fruit spectrum among consumer taxa. Despite an outstanding lack of specificity between fruit and consumer species, consideration of simple morphological traits of fruits reveals broad character syndromes associated with different consumer taxa. Competition between distantly related taxa that feed at the same height is far more important than has been previously supposed. The results also suggest how fruit characters could have evolved under consumer pressure as a result of consumer roles as dispersers or seed predators. Our analyses of dispersal syndromes show that fruit species partitioning occurs more between mammal taxa than between mammals and birds. There is thus a bird-monkey syndrome and a ruminant-rodent-elephant syndrome. The bird-monkey syndrome includes fruit species on which there is no pre-dispersal seed predation. These fruits (berries and drupes) are brightly colored, have a succulent pulp or arillate seeds, and no protective seed cover. The ruminant-rodent-elephant syndrome includes species for which there is pre-dispersal predation. These fruits (all drupes) are large, dull-colored, and have a dry fibrous flesh and well-protected seeds.

Susceptibility of rodents to infection with Schistosoma mansoni in Richard-Toll (Senegal).

The susceptibility of Arvicanthis niloticus, Mastomys huberti, Mastomys erythroleucus and Mus musculus was studied to assess the capacity of these rodents to transmit Schistosoma mansoni. The susceptibility was determined by the percentage of adult schistosomes recovered, the number of eggs per gramme of faeces, the viability of these eggs and the capacity of the rodents to maintain the life cycle of Schistosoma mansoni. The percentages of adult worms recovered were respectively 18%, 11.5%, 8.4% and 20.5% in A. niloticus, M. huberti, M. erythroleucus and M. musculus. After infection, they liberate in the environment viable eggs whose miracidia are infectious for the intermediate host (Biomphalaria pfeifferi). The mean egg load was 300 +/- 327.8 in A. niloticus; 664 +/- 673.5 in M. huberti; 240 +/- 304.8 in M. erythroleucus; 400 +/- 361.5 in M. musculus.

Histological study of Trichosomoides nasalis (Nematoda: Trichinelloidea) in the nasal cavities of the murid Arvicanthis niloticus, with associated pathology.

Histological study of the nasal cavities and upper maxillae of Arvicanthis niloticus naturally infected with Trichosomoides nasalis shows that the female worms reside in the epithelial monolayer of the nasal mucosa of the posterior and median cavities. Eggs laid by T. nasalis were infiltrated between the female body wall and the epithelial lining. Small groups of eggs, mixed with mucus and polymorphonuclear cells, were found in the nasal lumen, freed by rupture of the stretched epithelium. Two females and a few eggs were also found in the connective tissues. One male was found in a female uterus and two were apparently in the lumen of the nasal cavity but the surrounding tissues were disrupted. No male was identified in the lamina propria of the mucosa. However, significant inflammatory lesions occurred in the lamina propria, similar to those induced by the males of Anatrichosoma spp. which live in this part of the mucosa. In rodents, the lesions resulted in rhinosinusitis characterised by a lymphocytic infiltration leading to nasal obstruction.

[Resurgence of the plague in the Ikongo district of Madagascar in 1998. 2. Reservoirs and vectors implicated]. / Résurgence de la peste dans le district d'Ikongo à Madagascar en 1998. 2. Réservoirs et vecteurs impliqués.

Our survey of mammals and fleas arose as a result of an outbreak of bubonic plague at an usually low altitude in the Ikongo district (Madagascar), while a previous study had found anti-F1 antibodies in an endemic hedgehog. Animals were sampled with live traps in two hamlets (Antanambao-Vohidrotra, 540 m alt. and Ambalagoavy, 265 m alt.) and with pitfall traps in a neighbouring forest (750 m alt.). Rat fleas were collected by brushing the fur and free-living fleas by use of light traps. The introduced shrew Suncus murinus was found only in the village of Ambalagoavy while the black rat (Rattus rattus) was found in all three sites and the only seropositive rat was caught at Antanambao-Vohidrotra. In contrast, among the Tenrecidae (endemic shrews and hedgehogs) found in the forest near the first village, four animals were found seropositive for anti-F1 antibodies. One of them was carrying the endemic flea Paractenopsyllus pauliani, not yet reported as a vector of plague. The endemic vector of plague, Synopsyllus fonquerniei, was found only in the first village of Antanambao-Vohidrotra, and the cosmopolite flea Xenopsylla cheopis only in Ambalagoavy. Although no Yersinia pestis could be isolated and no F1-antigen could be detected in these animals, we found evidence of the recent transmission of plague in Antanambao-Vohidrotra and the nearby forest, but not in Ambalagoavy. These data corroborate with the sylvatic plague cycle hypothesis in Madagascar and its involvement in the outcome of the bubonic plague outbreak in this district.

[Resurgence of the plague in the Ikongo district of Madagascar in 1998. 1. Epidemiological aspects in the human population]. / Résurgence de la peste dans le district d'Ikongo à Madagascar en 1998. 1. Aspects épidémiologiques dans la population humaine.

Between the 20th October and the 18th November 1998, an outbreak of bubonic plague was declared in a hamlet in the Ikongo district of Madagascar. We conducted an epidemiological survey because of the re-emergence of the disease in this area (the last cases had been notified in 1965) and because of the low altitude compared to the classical Malagasy foci. The outbreak had been preceded by an important rat epizootics during September. A total of 21 cases were registered with an attack rate of 16.7% (21/126) and a lethality rate of 33% (7/21). The disease was more prevalent in males (66% of cases) and children aged < 15 years, as observed in general throughout the country. The anti-F1 seroprevalence among the contact population was 13.5% (13/96), probably attributable to subclinical infection by Yersinia pestis. No rodent was trapped during the survey, but an endemic hedgehog (Tenrec ecaudatus) was highly seropositive, suggesting a recent transmission of the plague bacillus among this species. The small mammals and vectors possibly involved in these new foci were investigated in May 1999.

[Update on plague in Madagascar]. / Actualités sur la peste à Madagascar.

After a thirty year period of successful control, bubonic plague showed the first signs of return in Madagascar where a fatal outbreak occurred in Antananarivo in 1978. A second outbreak was observed in Mahajanga in 1991 after more than a half century. In 1997, 459 confirmed or presumptive cases were reported, as compared to 150 to 250 cases during the last years. However the actual extent of this recrudescence must be placed in the perspective of a more efficient control program that has led to better reporting of suspected cases and availability of more accurate diagnostic techniques. Recent research has led to the development of highly effective immunological diagnostic tools (detection of antibodies and F1 antigen) allowing not only better surveillance of the disease in man and animals but also renewed study of the epidemiological cycle in the current environment. In this regard the capacity of several endemic fleas as vectors and the role of the rat Rattus norvegicus and the musk shrew Suncus murinus are currently under investigation. Genetic study of strains collected from 1936 to 1996 has demonstrated the appearance of 3 new ribotypes of Yersinia pestis since 1982 in the zones of strongest plague activity in Madagascar. A strain showing multiresistance to standard therapeutic antibiotic agents was isolated in 1995. Bubonic plaque is a priority health problem in Madagascar but remains a major concern for the rest of the world.

Comparative population genetics of a parasitic nematode and its host community: the trichostrongylid Neoheligmonella granjoni and Mastomys rodents in southeastern Senegal.

Contrasting host and parasite population genetic structures can provide information about the population ecology of each species and the potential for local adaptation. Here, we examined the population genetic structure of the nematode Neoheligmonella granjoni at a regional scale in southeastern Senegal, using 11 microsatellite markers. Using the results previously obtained for the two main rodent species of the host community, Mastomys natalensis and Mastomys erythroleucus, we tested the hypothesis that the parasite population structure was mediated by dispersal levels of the most vagile host. The results showed similar genetic diversity levels between host and parasite populations, and consistently lower levels of genetic differentiation in N. granjoni, with the exception of one outlying locus with a high F(ST). The aberrant pattern at this locus was primarily due to two alleles occurring at markedly different frequencies in one locality, suggesting selection at this locus, or a closely linked one. Genetic differentiation levels and isolation by distance analyses suggested that gene flow was high and random in N. granjoni at the spatial scale examined. The correlation between pair-wise genetic differentiation levels in the parasite and its main host was consistent with the hypothesis tested. Models of local adaptation as a function of the dispersal rates of hosts and parasites suggest that opportunities for local adaptation would be low in this biological system.