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Robert Poulin

Publications and source records attributed to Robert Poulin.

At least 19 recordsLinked to original sources

Clonal diversity of the marine trematode Maritrema novaezealandensis within intermediate hosts: the molecular ecology of parasite life cycles.

We quantified the clonal diversity of the New Zealand marine trematode Maritrema novaezealandensis (n = 1250) within Zeacumantus subcarinatus snail (n = 25) and Macrophthalmus hirtipes crab (n = 25) intermediate hosts using four to six microsatellite loci, and investigated the potential biological and physical factors responsible for the observed genetic patterns. Individual snails harboured one to five trematode genotypes and 48% of snails were infected by multiple parasite genotypes. Overall, the number of parasite genotypes did not increase with snail size, but was highest in intermediate-sized snails. Significantly larger numbers of parasite genotypes were detected in crabs (relative to snails; P < 0.001), with 16-25 genotypes recovered from individual crabs. Although crabs are typically infected by small numbers of cercariae sourced from many snails, they are occasionally infected by large numbers of cercariae sourced from single snails. The latter cases explain the significant genetic differentiation of trematode populations detected among their crab hosts (F(ST) = 0.009, P < 0.001). Our results suggest that the timing of infection and/or intraspecific competition among parasite clones within snails determine(s) the diversity of parasite clones that snails harbour. The presence of a large number of infected snails and tidal mixing of cercariae prior to infection results in crabs potentially harbouring hundreds of parasite genotypes despite the crabs' territorial behaviour.

Animals↗

The scaling of total parasite biomass with host body mass.

The selective pressure exerted by parasites on their hosts will to a large extent be influenced by the abundance or biomass of parasites supported by the hosts. Predicting how much parasite biomass can be supported by host individuals or populations should be straightforward: ultimately, parasite biomass must be controlled by resource supply, which is a direct function of host metabolism. Using comparative data sets on the biomass of metazoan parasites in vertebrate hosts, we determined how parasite biomass scales with host body mass. If the rate at which host resources are converted into parasite biomass is the same as that at which host resources are channelled toward host growth, then on a log-log plot parasite biomass should increase with host mass with a slope of 0.75 when corrected for operating temperature. Average parasite biomass per host scaled with host body mass at a lower rate than expected (across 131 vertebrate species, slope=0.54); this was true independently of phylogenetic influences and also within the major vertebrate groups separately. Since most host individuals in a population harbour a parasite load well below that allowed by their metabolic rate, because of the stochastic nature of infection, it is maximum parasite biomass, and not average biomass, that is predicted to scale with metabolic rate among host species. We found that maximum parasite biomass scaled isometrically (i.e., slope=1) with host body mass. Thus, larger host species can potentially support the same parasite biomass per gram of host tissues as small host species. The relationship found between maximum parasite biomass and host body mass, with its slope greater than 0.75, suggests that parasites are not like host tissues: they are able to appropriate more host resources than expected from metabolically derived host growth rates.

Animals↗

Diversity of trematode genetic clones within amphipods and the timing of same-clone infections.

The genetic diversity of trematodes within second intermediate hosts has important implications for the evolution of trematode populations as these hosts are utilized after the parasites reproduce asexually within first intermediate hosts and before sexual reproduction within definitive hosts. We characterised the genetic clonal diversity of the marine trematode Maritrema novaezealandensis within amphipod (Paracalliope novizealandiae) second intermediate hosts using four to six microsatellite loci to determine if multiple copies of identical trematode clones existed within naturally infected amphipods. To determine the relative timing of infections by identical clones within hosts, trematode metacercariae were assigned to six developmental stages and the stages of identical clones were compared. The genotypes of 306 trematodes were determined from 44 amphipods each containing more than one trematode. Six pairs of identical trematode clones were recovered in total (representing five amphipods: 11% of amphipods with greater than one trematode) and all pairs of clones belonged to the same developmental stage. This suggests that identical clone infections are effectively synchronous. A general decrease in the number of metacercariae recovered, prevalence, and mean intensity of infection for each subsequent developmental stage coupled with large numbers of metacercariae (>9) only being recovered from recent infections, supports the occurrence of post-infection amphipod mortality and/or within-host trematode mortality. Taken together, our results indicate that natural infections are characterised by high genetic diversity, but that amphipods also periodically encounter "batches" of genetically identical clones, potentially setting the stage for interactions within and between clonal groups inside the host.

Amphipoda↗

Is abundance a species attribute? An example with haematophagous ectoparasites.

Population density is a fundamental property of a species and yet it varies among populations of the same species. The variation comes from the interplay between intrinsic features of a species that tend to produce repeatable density values across all populations of the same species and extrinsic environmental factors that differ among localities and thus tend to produce spatial variation in density. Is inter-population variation in density too large for density to be considered a true species character? We addressed this question using data on abundance (number of parasites per individual host, i.e. equivalent to density) of fleas ectoparasitic on small mammals. The data included samples of 548 flea populations, representing 145 flea species and obtained from 48 different geographical regions. Abundances of the same flea species on the same host species, but in different regions, were more similar to each other than expected by chance, and varied significantly among flea species, with 46% of the variation among samples accounted by differences between flea species. Thus, estimates of abundance are repeatable within the same flea species. The same repeatability was also observed, but to a lesser extent, across flea genera, tribes and subfamilies. Independently of the identity of the flea species, abundance values recorded on the same host species, or in the same geographical region, also showed significant statistical repeatability, though not nearly as strong as that associated with abundance values from the same flea species. There were also no strong indications that regional differences in abiotic variables were an important determinant of variation in abundance of a given flea species on a given host species. Abundance thus appears to be a true species trait in fleas, although it varies somewhat within bounds set by species-specific life history traits.

Animals↗

Ecological characteristics of flea species relate to their suitability as plague vectors.

The ability of vector-borne diseases to persist and spread is closely linked to the ecological characteristics of the vector species they use. Yet there have been no investigations of how species used as vectors by pathogens such as the plague bacterium differ from closely related species that are not used as vectors. The plague bacterium uses mammals as reservoir hosts and fleas as vectors. The ability of different fleas to serve as vectors is assumed to depend on how likely they are to experience gut blockage following bacterial multiplication; the blockage causes fleas to regurgitate blood into a wound and thus inject bacteria into new hosts. Beyond these physiological differences, it is unclear whether there exist fundamental ecological differences between fleas that are effective vectors and those that are not. Here, using a comparative analysis, we identify clear associations between the ability of flea species to transmit plague and their ecological characteristics. First, there is a positive relationship between the abundance of flea species on their hosts and their potential as vectors. Second, although the number of host species exploited by a flea is not associated with its potential as a vector, there is a negative relationship between the ability of fleas to transmit plague and the taxonomic diversity of their host spectrum. This suggests a correlation between some ecological characteristics of fleas and their ability to develop the plague blockage. The plague pathogen thus uses mainly abundant fleas specialized on a narrow taxonomic range of mammals, features that should maximize the persistence of the disease in the face of high flea mortality, and its transmission to suitable hosts only. This previously unrecognized pattern of vector use is of importance for the persistence and transmission of the disease.

Animals↗

Variation in infection parameters among populations within parasite species: intrinsic properties versus local factors.

Within any parasite species, variation among populations in standard infection parameters (prevalence, intensity and abundance) is an accepted fact. The proportion of hosts infected and the mean number of parasites per host are not fixed values across the entire geographic range of any parasite species. The question is whether this inter-population variation occurs within a narrow, species-specific range and is thus driven mainly by the biological features of the parasite, or whether it is substantial and unpredictable, leaving population parameters at the mercy of local conditions. Here, the repeatability of estimates of prevalence, intensity and abundance of infection was assessed across populations of the same parasite species, for 77 metazoan parasite species of Canadian freshwater fishes. Overall, parameter values from different populations of the same parasite species were more similar to each other and more different from those of other species, than expected by chance alone. Much of the variation in parameter values in the dataset was associated with differences between parasite species, rather than with differences among populations within species. This was particularly true for intensity and abundance of infection; in contrast, prevalence values, while somewhat repeatable among populations of the same species, still showed considerable variation. Among the higher taxa investigated (monogeneans, trematodes, cestodes, nematodes, acanthocephalans, copepods), there was no evidence that species of one taxon display intrinsically greater variation in population parameters than species of other taxa. Overall, the results suggest that intensity and abundance of infection are real species characters, though somewhat variable. This conclusion supports the view that the biological features of parasite species can potentially override local environmental conditions in driving parasite population dynamics.

Animals↗

A new cercaria and metacercaria of Acanthoparyphium (Echinostomatidae) found in an intertidal snail Zeacumantus subcarinatus (Batillaridae) from New Zealand.

A new 23-collar-spined cercaria and metacercaria are described from intertidal molluscs of the coast of New Zealand. The new cercaria found emerging from the mud snails Zeacumantus subcarinatus (Sowerby) (Prosobranchia: Batillariidae) is characterized mainly by the number and arrangement of the cephalic glands, the size of the suckers, and the size and number of the collar spines. The cercaria encysts in the cockle Austrovenus stutchburyi (Wood) (Bivalvia: Veneridae) which lives in sympatry with the first intermediate host throughout New Zealand. Laboratory infections of cockles by cercariae from naturally infected snails resulted in metacercariae identical to those found in naturally infected cockles. The main features of the cercaria and metacercaria are the presence of a reniform collar with 23 spines, two pairs of small cephalic glands at the oral sucker level, another two pairs of much longer ones posterior to the pharynx, and the excretory vesicle Y-shaped with the main collecting canals extending to the anterior level of the ventral sucker. Each of the main collecting canals had 10-11 pairs of bilateral diverticula between the anterior edge of the ventral sucker and anterior body end. A brief discussion of its possible life cycle and ecology is also provided.

Animals↗

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

The number of species coexisting in a community may be regulated by local factors (e.g., competitive interactions), or by regional processes (e.g., dispersal from a regional species pool). The relative importance of local and regional processes can be inferred from the shape of the relationship between local and regional species richness. We investigated this relationship in communities of fleas parasitic on small mammals at two spatial scales: between the richness of fleas on individual hosts (infracommunities) and that of fleas on host populations (component communities), and between the richness of component communities and that of the entire regional species pool. We tested linearity (proportional sampling) versus curvilinearity with an asymptote (species saturation) by plotting "local" against "regional" species richness of fleas either among host species or within host species among populations. At the two spatial scales, we found consistent curvilinear relationships between species richness of the more "local" communities and richness of the more "regional" communities. This was true across all host species in the data set and for geographic subsets, even after controlling for the influence of sampling effort on estimates of species richness, and that of host phylogeny in interspecific analyses. We also tested for density compensation in species-poor communities. There was no strong evidence for density compensation at the infracommunity level, although its existence at the component community level appeared likely. Our results suggest that identical patterns in local-versus-regional species richness observed on two different spatial scales arise via different mechanisms: infracommunities appear saturated with flea species most likely because of local processes, such as host immune defenses, whereas component communities are saturated with species through interspecific competition, possibly among larval stages.

Animals↗

Parasite species coexistence and limiting similarity: a multiscale look at phylogenetic, functional and reproductive distances.

The factors that control biodiversity have been the focus of numerous recent investigations; these include species interactions, speciation, environmental gradients and heterogeneity, all of these operating differently at each observation scale. We used a null model to examine the influence of two forces shaping the community structure of Dactylogyrus species parasitic on roach (Rutilus rutilus) gills: interspecific competition that might prevent the coexistence of the most similar species and environmental filters that might result in the most similar species coexisting together. The study was carried out on two sets of fish from two different localities in the Morava river basin (Czech Republic) to evaluate the consistency of the results across host populations, and at three different scales of observation to test for the scale dependence of assembly rules. In decreasing order, from largest to smallest, the three spatial scales investigated were: the individual fish, the individual gill arch and individual sections of each gill arch. The similarity between pairs of parasite species was measured using three different criteria: (1) phylogeny, (2) quantitative functional traits consisting of parasite size and morphometric measurements of the attachment organ's sclerotized parts and (3) qualitative attributes of reproductive organs. First, our study reveals a strong conservatism of ecological characters for the nine Dactylogyrus parasite species, in particular regarding the attributes of their copulatory organs. Second, our study did not find any limitation of similarity among coexisting Dactylogyrus species due to interspecific competition, irrespective of the scale and the similarity measures considered. Conversely, our results support the niche filtering hypothesis, preventing the co-occurrence of species too dissimilar from one another. This process is particularly apparent at the scale of the individual fish and for functional traits associated with the hard parts of the parasites' attachment organs. In both localities, the Dactylogyrus species that occur on the same fish individuals tend to have similar values for the haptor dimensions. Our study supports previous studies on monogenean parasite communities indicating a weak influence of competition as a structuring force, but it goes a step further by identifying environmental filtering as a key process shaping these communities.

Animals↗

Climate warming may cause a parasite-induced collapse in coastal amphipod populations.

Besides the direct impact on the general performance of individual organisms, the ecological consequences of climate change in terrestrial and marine ecosystems are expected to be determined by complex cascading effects arising from modified trophic interactions and competitive relationships. Recently, the synergistic effect of parasitism and climate change has been emphasised as potentially important to host population dynamics and community structure, but robust empirical evidence is generally lacking. The amphipod Corophium volutator is an ecologically important species in coastal soft-bottom habitats of the temperate North Atlantic, and commonly serves as host to microphallid trematodes that cause intensity-dependent and temperature-dependent mortality in the amphipod population. Using a simulation model parameterised with experimental and field data, we demonstrate that a 3.8 degrees C increase in ambient temperature will likely result in a parasite-induced collapse of the amphipod population. This temperature increase is well within the range predicted to prevail by the year 2075 in the International Wadden Sea region from where the model data are obtained. Due to the amphipods' ecological importance, their population decline may impact the coastal ecosystem as a whole.

Amphipoda↗

Evolution of host specificity in fleas: is it directional and irreversible?

Evolutionary trends in the evolution of host specificity have been the focus of much discussion but little rigorous empirical testing. On the one hand, specialization is often presumed to lead irreversibly into evolutionary dead ends and little diversification; this would mean that generalists might evolve into specialists, but not vice versa. On the other hand, low host specificity may limit the risk of extinction and provide more immediate fitness benefits to parasites, such that selection may favour evolution toward a generalist strategy. Here, we test for directionality in the evolution of host specificity using a large data set and phylogenetic information on 297 species of fleas parasitic on small mammals. The analyses determined whether host specificity, measured both as the number of host species exploited and their taxonomic diversity, was related to clade rank of the flea species, or the number of branching events between an extant species and the root of the phylogenetic tree (i.e., the total path length from the root of the tree to the species). Based on regression analyses, we found positive relationships between the number of host species used and clade rank across all 297 species, as well as within one (Hystrichopsyllidae) of four large families and one of seven large genera investigated separately; in addition, we found a positive relationship between the taxonomic diversity of host species used and clade rank in another of the seven genera. These results suggest a slight evolutionary trend of decreasing host specificity. Using a much more conservative likelihood ratio test, however, a random walk, or null model, of evolution could not be discarded in favour of the directional trends in all cases mentioned above. Still, these results suggest that host specificity may have tended to decrease in many flea lineages, a process that could have been driven by the benefits of exploiting a wide range of host species.

Adaptation, Biological↗

What are the factors determining the probability of discovering a flea species (Siphonaptera)?

Our aim was to determine which of four variables (number of host species exploited by the parasite, taxonomic distinctness of these hosts, geographic range of the principal host, and year of description of this host) was the best predictor of description date of fleas. The study used previously published data on 297 flea species parasitic on 197 species of small mammals from 34 different regions of the Holarctic and one region from the Neotropics. We used both simple linear and multiple regressions to evaluate the relationships between the four predictor variables and the year of flea description, on species values as well as on phylogenetically independent contrasts. Whether or not the analyses controlled for flea phylogeny, all predictor variables correlated significantly with year of flea description when tested separately. In multiple regressions, however, the number of exploited host species was the best predictor of the date of flea description, with the geographic range of the principal host species as well as the date of its description having a lesser, though significant, influence. Overall, our results indicate that a flea species is more likely to be discovered and described early if its biological characteristics (exploitation of many host species) and those of its hosts (long-known to science, broad geographic distributions) increase its chances of being included in a collection. Because the variables we investigated only explained about 10-11% of the variation in year of description among flea species, other factors must be important, such as temporal variability in the activity of flea taxonomists.

Animals↗

Impact of a microphallid trematode on the behaviour and survival of its isopod intermediate host: phylogenetic inheritance?

The extent to which the ability of parasites to alter host behaviour is phylogenetically inherited as opposed to independently evolved has received little attention. We investigated the impact of an undescribed species of Microphallus on the behaviour and survival of its host, the freshwater isopod Austridotea annectens, to determine if it produced effects comparable to those induced by other trematodes of this genus. There was no difference between the vertical distribution and responses to light of infected isopods and those of uninfected isopods. In contrast, we found that infected isopods were more active swimmers than uninfected isopods, and that they failed to show the evasive responses shown by uninfected isopods when exposed to a simulated predator. There was no detectable effect of Microphallus infection on isopod survival, however. Overall, our results provide no strong evidence that the behavioural manipulation shown by our Microphallus species and that displayed by the well-studied Microphallus papillorobustus are inherited from a common ancestor rather than independently derived.

Animals↗

Causes of inter-individual variation in reproductive strategies of the parasitic nematode Graphidioides subterraneus.

Several factors capable of affecting the amount of resources available to an individual parasite, such as the number of other parasites in the host or host quality, may cause variability in reproductive success among parasites. Variation in egg output and mean egg volume was investigated among adult females of the nematode Graphidioides subterraneus, parasitic in the herbivorous subterranean rodent Ctenomys talarum (Octodontidae). Female nematode body size correlated strongly with the number of eggs produced. However, neither host body mass nor the number of other nematodes per host had any influence on the number or volume of eggs produced by the parasites. There was also no evidence for a trade-off between the number of eggs produced and mean egg volume among female nematodes. All these results suggest that resource supply to individual worms is not limited by host size or by the number of conspecific parasites vying for the same resources, despite the 30-fold variation in intensity of infection and the twofold variation in host body mass observed in the present study. Instead, resource availability does not appear to constrain reproduction in G. subterraneus, with its host providing a stable, predictable environment.

Animals↗

Speciation in parasites: a population genetics approach.

Parasite speciation and host-parasite coevolution should be studied at both macroevolutionary and microevolutionary levels. Studies on a macroevolutionary scale provide an essential framework for understanding the origins of parasite lineages and the patterns of diversification. However, because coevolutionary interactions can be highly divergent across time and space, it is important to quantify and compare the phylogeographic variation in both the host and the parasite throughout their geographical range. Furthermore, to evaluate demographic parameters that are relevant to population genetics structure, such as effective population size and parasite transmission, parasite populations must be studied using neutral genetic markers. Previous emphasis on larger-scale studies means that the connection between microevolutionary and macroevolutionary events is poorly explored. In this article, we focus on the spatial fragmentation of parasites and the population genetics processes behind their diversification in an effort to bridge the micro- and macro-scales.

Animals↗

Molecular ecology of parasites: elucidating ecological and microevolutionary processes.

We review studies that have used molecular markers to address ecological and microevolutionary processes in parasites. Our goal is to highlight areas of research that may be of particular interest in relation to the parasitic lifestyle, and to draw attention to areas that require additional study. Topics include species identification, phylogeography, host specificity and speciation, population genetic structure, modes of reproduction and transmission patterns, and searching for loci under selection.

Animals↗

Alternative reproductive strategies in the progenetic trematode Coitocaecum parvum: comparison of selfing and mating worms.

The progenetic opecoelid trematode Coitocaecum parvum can reproduce either precociously by selfing in its second intermediate amphipod host or by mating in its normal definitive fish host. In this study, we describe and compare the infection parameters and some life history traits of both egg-producing worms and non-egg producing worms in both their second intermediate and definitive hosts. We showed that 58% of worms start to produce eggs while still in the amphipod. The relative abundance of progenetic worms increased with amphipod size, and egg-producing worms achieved greater size in amphipods than in fish. These 2 findings support the reproductive insurance hypothesis. No difference in size was revealed between eggs produced in the amphipods and those produced in the fish. Although more information is needed to thoroughly assess the respective costs and benefits of selfing and mating in this species, our conclusion is that adopting progenesis may have few, if any, long-term negative consequences for the parasite.

Amphipoda↗