High Predation Pressure by an Introduced Flatworm on Land Snails on the Oceanic Ogasawara Islands1

The introduction of a predatory flatworm, Platydemus manokwari, has been considered a cause of the decline of endemic land snails on the tropical oceanic islands. To clarify the effect of P. manokwari on land snail survival in the field, we examined survival rates of snails experimentally placed in areas where snails are absent (i.e., P. manokwari is present) on Chichijima, Ogasawara (Bonin) Islands. We found that over 50 and 90 percent of the snails were dead after 3 and 11 d, respectively, and that the main cause of mortality was predation by P. manokwari.     

The cause of mollusk decline on the Ogasawara Islands

Decline of land snails on the Ogasawara Islands was studied. In Hahajima, major alien predators such as Euglandina rosea and Platydemus manokwari are not present, but some small endemic snails, for example, Hirasea spp. and Ogasawarana spp., are already rare and more common endemic snails, for example, Mandarina spp., are also declining in the northern mountains. The decline cannot be directly explained by forest deforestation and by its subsequent regeneration. Three species of flatworms were found to eat small snails under captive conditions. The distribution of these flatworms is restricted to the northern mountains of Hahajima where Mandarina is declining and its survival is low. These predators are plausible candidates as a cause of the decline of the endemic snails.     

Long‐term stasis and short‐term divergence in the phenotypes of microsnails on oceanic islands

Phenotypic divergence is often unrelated to genotypic divergence. An extreme example is rapid phenotypic differentiation despite genetic similarity. Another extreme is morphological stasis despite substantial genetic divergence. These opposite patterns have been viewed as reflecting opposite properties of the lineages. In this study, phenotypic radiation accompanied by both rapid divergence and long‐term conservatism is documented in the inferred molecular phylogeny of the micro land snails Cavernacmella (Assimineidae) on the Ogasawara Islands. The populations of Cavernacmella on the Sekimon limestone outcrop of Hahajima Island showed marked divergence in shell morphology. Within this area, one lineage diversified into types with elongated turret shells, conical shells and flat disc‐like shells without substantial genetic differentiation. Additionally, a co‐occurring species with these types developed a much larger shell size. Moreover, a lineage adapted to live inside caves in this area. In contrast, populations in the other areas exhibited no morphological differences despite high genetic divergence among populations. Accordingly, the phenotypic evolution of Cavernacmella in Ogasawara is characterized by a pattern of long‐term stasis and periodic bursts of change. This pattern suggests that even lineages with phenotypic conservatism could shift to an alternative state allowing rapid phenotypic divergence.     

Potential impacts of the invasive flatworm Platydemus manokwari on arboreal snails

The introduction of the snail-eating flatworm Platydemus manokwari (Tricladida: Rhynchodemidae) has been considered a cause of the extinction of native land snails on several Pacific islands. Although P. manokwari is known to attack land snails on the ground, whether P. manokwari attacks snails on trees remains unclear. To clarify the effect of P. manokwari on arboreal snails, we examined survival rates of land snails experimentally placed on tree trunks (0.5–2.0 m above the ground) in a forest on Chichijima, Ogasawara (Bonin) Islands, in the northwestern Pacific Ocean. The survival of snails experimentally placed on tree trunks with artificially created snail scent trails rapidly decreased for 7 days, and the mortality was caused by P. manokwari predation. However, snails placed on tree trunks without snail scent trails were not attacked by P. manokwari. Therefore, P. manokwari climbed tree trunks, likely tracking the snail scent. We found that over 40% of the snails placed on tree trunks with snail scent trails were eaten by P. manokwari within 7 days. This experiment supports the hypothesis that P. manokwari predation is an important cause of the rapid decline or extinction of native arboreal snails on Pacific islands.     

The recent history and population structure of five Mandarina snail species from subtropical Ogasawara (Bonin Islands, Japan)

The effect of Pleistocene climate change on the organisms of tropical and subtropical regions is rather poorly understood. We therefore studied the land snail genus Mandarina (Bradybaenidae) of oceanic Ogasawara (Bonin Islands, Japan), with the aim of using population genetic data to understand their recent history. Our analysis of a mitochondrial 16S ribosomal RNA region from more than 600 snails in five ground-living species suggests that populations on the small islands of Mukoujima, Anejima, Imotojima and Meijima, as well as on the low-lying southern and central parts of Hahajima, have probably undergone recent bottlenecks followed by subsequent expansions. Except between the main island of Hahajima and Mukouijima, there is almost no evidence for gene flow among islands even though the islands were connected repeatedly by land bridges through the Pleistocene. Within islands the population structure is severe, suggestive of a long-term, low level of gene flow (F(ST) is frequently greater than 0.5 among geographically close populations). Finally, there is a marked genetic patchiness, meaning that genetically close populations are sometimes separated by genetically distant populations. These patterns could be a consequence of expansion from bottlenecks, low active dispersal and founder effects caused by rare long-distance migrants. Unfortunately, the exact nature of the refugia and bottlenecks remains unknown because the palaeoclimate of this region is poorly understood. Dating the population size changes is also challenging because the molecular clock is uncertain. We suggest, however, that arid conditions or deforestation induced by decreased atmospheric CO(2) may have been the main factor in determining population size.     

Effects of habitat history and extinction selectivity on species-richness patterns of an island land snail fauna

Aim Local‐scale diversity patterns are not necessarily regulated by contemporary processes, but may be the result of historical events such as habitat changes and selective extinctions that occurred in the past. We test this hypothesis by examining species‐richness patterns of the land snail fauna on an oceanic island where forest was once destroyed but subsequently recovered. Location Hahajima Island of the Ogasawara Islands in the western Pacific. Methods Species richness of land snails was examined in 217 0.25 × 0.25 km squares during 1990–91 and 2005–07. Associations of species richness with elevation, current habitat quality (proportion of habitat composed of indigenous trees and uncultivated areas), number of alien snail species, and proportion of forest loss before 1945 in each area were examined using a randomization test and simultaneous autoregressive (SAR) models. Extinctions in each area and on the entire island were detected by comparing 2005–07 records with 1990–91 records and previously published records from surveys in 1987–91 and 1901–07. The association of species extinction with snail ecotype and the above environmental factors was examined using a spatial generalized linear mixed model (GLMM). Results The level of habitat loss before 1945 explained the greatest proportion of variation in the geographical patterns of species richness. Current species richness was positively correlated with elevation in the arboreal species, whereas it was negatively correlated with elevation in the ground‐dwelling species. However, no or a positive correlation was found between elevation and richness of the ground‐dwelling species in 1987–91. The change of the association with elevation in the ground‐dwelling species was caused by greater recent extinction at higher elevation, possibly as a result of predation by malacophagous flatworms. In contrast, very minor extinction levels have occurred in arboreal species since 1987–91, and their original patterns have remained unaltered, mainly because flatworms do not climb trees. Main conclusions The species‐richness patterns of the land snails on Hahajima Island are mosaics shaped by extinction resulting from habitat loss more than 60 years ago, recent selective extinction, and original faunal patterns. The effects of habitat destruction have remained long after habitat recovery. Different factors have operated during different periods and at different time‐scales. These findings suggest that historical processes should be taken into account when considering local‐scale diversity patterns.     

Isolation and characterization of Ligustrum micranthum (Oleaceae) microsatellite loci using paired‐end Illumina reads

Twenty‐six microsatellite loci were developed and characterized for Ligustrum micranthum, a species endemic to the Ogasawara Islands, Japan. The genetic structure of this species must be clarified in order to restore the island's ecosystem. A total of 8511 primer pairs were designed from de novo sequencing. Of the 48 primer pairs selected, amplification and polymorphisms were tested using one population each from the Chichijima and Hahajima Islands of the Ogasawara Islands. Twenty‐six microsatellite loci were successfully amplified and the number of alleles for these loci ranged from five to 31 per locus, and the mean expected heterozygosities were 0.858 and 0.849, respectively. No significant deviation from the Hardy–Weinberg equilibrium was observed in either population, and no significant linkage disequilibrium was detected between any locus pair. The microsatellite loci reported in this study can be used in future studies to evaluate the genetic structure and mating system of L. micranthum.     

Prey preference and gregarious attacks by the invasive flatworm Platydemus manokwari

The flatworm Platydemus manokwari (Tricladida: Rhynchodemidae) preys on various species of land snail, and its introduction to areas outside of its native range is thought to have caused the extinction of native land snails on several Pacific islands. Platydemus manokwari occurs in areas where land snails have been absent since its invasion, suggesting that the flatworm can prey on animals other than land snails. To identify the alternative prey and prey preferences of P. manokwari, I examined the feeding ecology of P. manokwari under field and laboratory conditions. Individuals were observed attacking live earthworms in a forest where land snails are already extinct, on Chichijima, Ogasawara (Bonin) Islands, Japan. I also observed them attacking earthworms and a species of isopod and land snail in the lab. To prey on the worm, similar to other earthworm-eating flatworms, P. manokwari wrapped itself around the prey and fed on it by inserting its pharynx into the earthworm’s body. Large earthworms were able to escape P. manokwari attacks by autotomy, but the autotomized body parts were eaten. Several P. manokwari individuals together attacked earthworms that were larger than themselves. The laboratory experiments showed that such gregarious attacks increased predation success on both large earthworms and land snails. The flatworms also attacked the isopods, although the predation rate was low. These results demonstrate that P. manokwari is a polyphagous predator of slow-moving soil invertebrates (land snails, slugs, earthworms, and isopods) and that invasion by P. manokwari may directly and indirectly impact native soil fauna.     

Carnivore mollusks as natural enemies of invasive land flatworms

There are several records of the carnivorous behaviour of land flatworms, considered to be top‐predators in their micro‐habitats, by preying upon various species of invertebrates. However, there is little knowledge of predators on land‐flatworms. The possible impact of invasive land flatworms on prey populations has caused widespread concern, when considering their predatory behaviour, combined with recent human influence on the distribution of certain species. This work is the first record of predation on land flatworms by a carnivorous snail. Various‐sized land flatworms of 10 native species of the subfamily Geoplaninae, as well as the exotic species Bipalium kewense (subfamily Bipaliinae), were offered to Rectartemon depressus (Gastropoda, Streptaxidae), which accepted all. The predator also fed on the snail Bradybaena similaris. The snails were maintained in laboratory for an average period of 12 months based on a mixed diet of flatworms and B. similaris, suggesting that the snail is a polyphagous predator. Because certain land‐flatworm species have been described as invasive species which may have a potential impact on prey populations in native and man‐made ecosystems, it is proposed that carnivorous snails of other native species, as potential predators of flatworms, should be tested for possible use in biological control programmes of these invasive planarians.     

How did this snail get here? Several dispersal vectors inferred for an aquatic invasive species

1. How species reach and persist in isolated habitats remains an open question in many cases, especially for rapidly spreading invasive species. This is particularly true for temporary freshwater ponds, which can be remote and may dry out annually, but may still harbour high biodiversity. Persistence in such habitats depends on recurrent colonisation or species survival capacity, and ponds therefore provide an ideal system to investigate dispersal and connectivity. 2. Here, we test the hypothesis that the wide distributions and invasive potential of aquatic snails is due to their ability to exploit several dispersal vectors in different landscapes. We explored the population structure of Physa acuta (recent synonyms: Haitia acuta, Physella acuta, Pulmonata: Gastropoda), an invasive aquatic snail originating from North America, but established in temporary ponds in Doñana National Park, southern Spain. In this area, snails face land barriers when attempting to colonise other suitable habitat. 3. Genetic analyses using six microsatellite loci from 271 snails in 21 sites indicated that (i) geographically and hydrologically isolated snail populations in the park were genetically similar to a large snail population in rice fields more than 15 km away; (ii) these isolated ponds showed an isolation‐by‐distance pattern. This pattern broke down, however, for those ponds visited frequently by large mammals such as cattle, deer and wild boar; (iii) snail populations were panmictic in flooded and hydrologically connected rice fields. 4. These results support the notion that aquatic snails disperse readily by direct water connections in the flooded rice fields, can be carried by waterbirds flying between the rice fields and the park and may disperse between ponds within the park by attaching to large mammals. 5. The potential for aquatic snails such as Physa acuta to exploit several dispersal vectors may contribute to their wide distribution on various continents and their success as invasive species. We suggest that the interaction between different dispersal vectors, their relation to specific habitats and consequences at different geographic scales should be considered both when attempting to control invasive freshwater species and when protecting endangered species.     

Seashore in the mountain: limestone‐associated land snail fauna on the oceanic Hahajima Island (Ogasawara Islands,Western Pacific)

Because land snails inhabiting the seashore are most likely to be carried by ocean currents or by attaching to seabirds, land snail fauna on oceanic islands include species derived from the mainland ancestors inhabiting the seashore. If habitat use of the island descendants is constrained by the ecology of the mainland ancestor, the island species that moved from the coastal habitat to the inland habitat may still be restricted to relatively exposed microhabitats with high pH, calcium carbonate‐rich substrates, and poor litter cover. We tested this hypothesis by investigating the association between environmental conditions and species diversity of seashore‐derived species of the endemic land snails on the oceanic Hahajima Island (Ogasawara Islands). Seashore‐derived species showed higher species richness on limestone outcrops than non‐limestone areas, whereas the other species showed no significant increase in species richness in limestone outcrops. There was a higher proportion of seashore‐derived species on the limestone ridges than on the soil of dolines, even in the limestone area. Accordingly, the species derived from the seashore of the mainland are restricted to microhabitats with poor vegetation cover, poor litter cover, high pH, and calcium carbonate‐rich substrates, which supports the hypothesis that the inland species on an island derived from the mainland seashore still prefer environments similar to the seashore. In addition, the seashore‐derived species on the limestone outcrop include cave‐dwellers lacking functional eyes. This suggests that the probability of colonizing a cave environment is restricted to seashore‐derived species. The findings obtained in the present study suggest that habitat use of the ancestral lineages can constrain habitat use of the descendants, even in the oceanic islands with depauperate fauna. This bias in the species composition on the limestone outcrop constrains lineages that can colonize and adapt to the inside of caves, and therefore, habitat use of the ancestral lineages affects the ability of descendant lineages to colonize novel habitats. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 686–693.     

The demographics of destruction: isolated populations of arboreal snails and sustained predation by rats on the island of Moloka’i 1982–2006

Populations of Partulina redfieldi, an achatinelline tree snail studied in four isolated trees, grew 100–900% between 1983 and 1995. Beginning in 1995, populations declined by 85%, and shells of rat-killed snails accumulated beneath the trees. While rat-marked shells were always present in the study area, numbers increased significantly. Despite a rat-abatement program begun in 1995, the snails continued to disappear, which we conclude was due to continued rat migration into the study area, despite baiting, and a switch in rat-food preference toward the snails. In neighboring forest where tree canopies are more continuous, snail density is lower and rat predation is not apparent. Captive-bred snails were successfully introduced to a small unoccupied tree in the same area in 1989, and this population suffered the same fate as the natural snail populations. Since 2000, P. redfieldi populations have remained low and rat predation continues.     

Distribution,movement, and microhabitat use of the introduced predatory snail Euglandina rosea in Hawaii: implications for management

The purposeful introduction of the land snail Euglandina rosea, which feeds exclusively on snails and slugs, has been implicated as a major factor in the decline of diverse Pacific island land snail faunas. We report on the distribution, movement patterns, and microhabitat preferences of E. rosea in a gulch in the Waianae Mountains, Oahu, Hawaii, because such data will help focus management actions at a local scale to protect native snail populations in areas where E. rosea is established. The Waianae Mountains harbor many endangered or threatened snails, most currently found in isolated habitat patches near the ridges. Conversely, most living individuals (28/29) and shells (46/56) of E. rosea were collected within the gulch, which supported higher densities of other native and non‐native snails, and was cooler and more moist than the ridges. Thirteen individuals of E. rosea were tracked (eight directly using a bobbin and thread method, and five indirectly by mark–recapture); most (10/13) moved on average <2.5 m per week (range 0.1–25.21 m), and all stayed within the gulch. Members of E. rosea preferred leaf litter over open, fern/shrub, or wood microhabitats. There were large differences in the population density of E. rosea over small spatial scales, indicating that there may be places where native snail populations could persist even in areas where populations of E. rosea are established. Identifying areas with differing population densities of E. rosea is critical for not only understanding why some native snail species may be more vulnerable to extinction, but also to locate areas where predation pressure is low and conservation efforts will be most likely to succeed.     

Application of harmonic radar technology to monitor tree snail dispersal

Abstract. Planned conservation efforts for tree snails of the endangered genus Achatinella, endemic to the island of O'ahu, Hawai'i, will include translocations among the remaining wild and captive‐bred populations. In order to establish optimal levels of artificial migration among neighboring groups of snails within fragmented populations, efforts to determine natural dispersal rates through direct observation were initiated. Capture–mark–recapture (CMR) efforts have proved inadequate for obtaining the requisite dispersal estimates, due to low recapture probabilities. In addition, snail dispersal beyond the boundaries of a finite CMR study site was indistinguishable from mortality. In the preliminary study reported here, both the low recapture probability and dispersal detection problems of past CMR efforts were addressed by using harmonic radar tracking. This approach yielded rough dispersal estimates that were unattainable using CMR alone by providing 100% recapture rates even beyond the normal survey area boundaries. Extensive snail movements within clusters of connected trees were frequently observed after tracking for merely a few hours, although movements between unconnected trees were rare and recorded only after monthly survey intervals. Just 11 out of 40 tracked snails made between‐tree movements (average distance of 4.94±1.52 m) during the entire 7‐month study, and provided the only data utilizable for inferring gene flow in and out of subpopulations. Meteorological data loggers were deployed when tracking began to look for an association between such snail movement and weather fluctuations. The resultant data indicate that increases in both wind gusts and humidity facilitate dispersal (R²=0.77, p‐value <0.001), and that passive wind dispersal alone may be responsible for many snail movements (R²=0.59, p‐value=0.0014). Despite having provided coarse estimates of short‐term dispersal and corresponding wind influences, the limitations of the radar method can be substantial.     

Leech predation on juvenile freshwater snails: effects of size,species and substrate

Summary The leechGlossiphonia complanata does not appear to have substantial impact on snail populations, but this may be due to most studies focusing on adult snails rather than juvenile snails. In this study I investigated how predation rates ofG. complanata feeding on newly-hatched and juvenile snails was affected by snail species, snail size, snail density and substrate, in a laboratory experiment. Number of snails eaten increased with increasing density resulting in a type II functional response curve. Predation rates were higher when leeches were feeding onLymnaea emarginata than onPhysa gyrina, whereas there was no significant difference in predation rates when they were feeding onL. emarginata andHelisoma anceps. Sandy substrates and greater snail size resulted in decreased predation rates. Sand reduced movement speed ofG. complanata, which probably reduced encounter rates. Thus, there was a comparatively large effect of leech predation on newly-hatched snails, due to a high probability of encounter and high predation rates, but spatial and temporal refuges probably reduce the importance of leech predation as a structuring force in freshwater snail assemblages.     

Ecological and morphological diversification within single species and character displacement in Mandarina,endemic land snails of the Bonin Islands

The endemic land snail species Mandarina hahajimana has undergone extensive habitat and morphological diversification within the Hahajima islands in the Bonin archipelago. This species has diversified into populations with ground, arboreal and semi-arboreal life histories. In addition, arboreal populations and semi-arboreal populations show diversification in preferences of species and positions of the tree on which they are found. Shell morphologies of M. hahajimana exhibit remarkable geographical variation, and they have a clear relationship with their life histories. The morphological variation of M. hahajimana results from adaptation to different lifestyles. The habitats of these populations influence the relationships with other species of Mandarina coexisting with M. hahajimana. This suggests that the morphological and ecological divergence within M. hahajimana has been induced by competitive interaction with other species of Mandarina. Character displacement may have played an important role in promoting adaptive radiation of Mandarina in the Bonin Islands.     

Influence of rearing and experimental temperatures on predator avoidance behaviour in a freshwater pulmonate snail

1. Predicted increases in the temperature of freshwaters is likely to affect how prey species respond to predators. We investigated how the predator avoidance behaviour of the freshwater pulmonate snail Lymnaea stagnalis is influenced by the temperature at which it was reared and that at which behavioural trials were carried out. 2. Crawl‐out behaviour of juvenile snails from two populations (high predation risk versus low predation risk) reared at either 15 or 20 °C was assessed in response to predation cues (predatory fish kairomones and conspecific alarm cues) in behavioural trials at both 15 and 20 °C. 3. Trial temperature had a significant effect on the time that snails spent in avoidance, regardless of their population of origin. Crawl‐out behaviour was greater during behavioural trials at 15 °C, but there was no effect of trial temperature on the speed with which animals showed avoidance behaviour. 4. There was no interactive effect of rearing temperature (RT) and trial temperature, but the effect of RT on avoidance behaviour did differ between populations. For an RT of 15 °C, snails from the South Drain (high risk) population showed a more rapid and longer avoidance response than those from the Chilton Moor (low risk) population. In contrast, for snails reared at 20 °C, there was no difference between populations for the duration of the avoidance response and snails from Chilton Moor crawled out faster than those from South Drain. 5. Hence, whilst (predictable) differences relative to natural predation threat in crawl‐out behaviour were apparent at 15 °C, raising the developmental temperature to 20 °C eliminated or, in the case of latency, reversed these differences. This suggests that L. stagnalis populations that cohabit with predatory fish and experience high developmental temperatures may have a reduced ability to respond to fish predation risk.     

Genetic structure of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens and its implications for the management of threatened island populations

The Red‐headed Wood Pigeon Columba janthina nitens is endemic to the Ogasawara Islands, an oceanic island chain located 1000 km south of the main islands of Japan. The subspecies is at high risk of extinction because of its small population size and restricted habitat range. We undertook genetic analyses of this pigeon using sequences of a portion of the mitochondrial control region and five microsatellite markers to estimate the genetic characteristics of two wild populations from the Bonin and Volcano Islands, as well as one captive breeding population. The genetic diversity of the wild individuals was exceptionally low in both the mitochondria (nucleotide diversity = 0.00105) and at the microsatellite (3.2 alleles per locus and H_E = 0.12) loci. Higher numbers of microsatellite genotypes were observed in the Volcano Islands population than in the Bonin Islands population, which may be because of the relatively low impact of human disturbance. The most common mitochondrial haplotypes and microsatellite alleles observed in the two wild populations were completely fixed in the captive population. Our results suggest that the genetic diversity of the captive population needs to be increased. However, introduction of a wild individual into a captive population can lead to a decreased genetic diversity in the wild population and therefore should be done with caution. The genetic differentiation between the Bonin and the Volcano island groups was low, and the populations of the two island groups should be regarded as a single evolutionarily significant unit. However, special consideration is required for habitat conservation in the Volcano Islands, which may be functioning as a sanctuary for the Red‐headed Wood Pigeon. For the long‐term conservation of threatened bird species that live on remote oceanic islands, determination of management units considering gene flow caused by their flying capacity and maintenance of genetically suitable wild and captive populations are essential.     

Factors affecting the distribution and abundance of the commensal <Emphasis Type="Italic">Temnocephala iheringi</Emphasis> (Platyhelminthes: Temnocephalidae) among the southernmost populations of the apple snail <Emphasis Type="Italic">Pomacea canaliculata </Emphasis>(Mollusca: Ampullariidae)

Temnocephala iheringi is the most common temnocephalan inhabiting the mantle cavity of the apple snail Pomacea canaliculata, a freshwater neotropical gastropod that has become a serious rice pest in Southeastern Asia. T. iheringi has been recorded from Mato Grosso (Brazil) to water bodies associated with the Río de la Plata river (Argentina). During an extensive survey in the southern limit of the native area of P.␣canaliculata the presence of T. iheringi eggs was recorded in several apple snail populations, extending the known distribution of the commensal more than 400 km southwards. The aim of this study was to understand the factors affecting the distribution and abundance of T. iheringi among populations of P.␣canaliculata. Only 23% of the apple snail populations inhabiting streams harboured temnocephalans while the occurrence among lentic ones was 71%. T. iheringi was found mostly in populations of apple snails living in non-alkaline sites and where snails attaining sizes larger than 4 cm were very common. The prevalence of the temnocephalans in lentic populations was higher than 90%. The number of eggs on the shell (not including the umbilicus) ranged between 0 and 470 and was different among populations of P.␣canaliculata. The prevalence and number of eggs were lower in the lotic populations, except for a stream population immediately downstream of a lake with commensals. There was no difference between males and females of P. canaliculata neither in the prevalence nor in the number of eggs on the shell. The southernmost population of the world of P. canaliculata harbours commensals that tolerate cold winter water temperatures (4–5 °C) as well as its host. On the other hand, T. iheringi was found only in sites with bicarbonate concentrations lower than 6.6 meq l⁻¹, suggesting that the tolerance of the commensal is very much lower than that of the apple snail (up to 9.95 meq l⁻¹). The number of worms inside each snail or the life history variation of P. canaliculata could explain the influence of the size of the snails on the occurrence of T. iheringi. In the big-sized snails, where the number of commensals is higher, the probability of survival of at least one worm is also higher, specially during the hibernation period, when crawling and feeding are null and snails remain buried. On the other hand, P. canaliculata snails from lentic populations are generally bigger and mostly iteroparous, while those inhabiting streams are smaller and semelparous. In these populations the snails have access to mate only with snails of their same cohort, while in iteroparous populations they can copulate with individuals of other cohorts, allowing the inter-generation transmission of worms and the long term persistence of the population of commensals.     

Evolutionary aspects of the metapopulation dynamics of Biomphalaria pfeifferi,the intermediate host of Schistosoma mansoni

Combining genetic and demographic data is a powerful approach to study adaptation process and evolutionary forces acting in natural populations. We focus on the freshwater snail Biomphalaria pfeifferi, the intermediate host of Schistosoma mansoni. Twenty‐one populations sampled in the south of Madagascar were genotyped at six microsatellite loci. Demographic parameters and parasitic prevalence were estimated monthly over the year preceding the genetic sampling. Our results indicate that populations experience recurrent bottlenecks and size fluctuations, which strongly depresses the genetic diversity within population. The recolonization of depleted sites involves genetically differentiated immigrants. We detected frequent migration events along rivers and rare migration events between watersheds. This explains the high level of differentiation observed among populations. The negative regression observed between the prevalence of S. mansoni and the genetic diversity of B. pfeifferi populations indicates that host consanguinity may affect prevalence through the genetic mechanisms involved in resistance. Coevolutionary outcomes are also influenced by the relative migration rates of snails and flukes, but the parasite local adaptation may be prevented by rare long distance dispersal in snails and the phylogeographical patterns of colonization of both hosts and snails.