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R Poulin

Publications and source records attributed to R Poulin.

At least 37 records · Page 2Linked to original sources

Life history constraints on the evolution of abbreviated life cycles in parasitic trematodes.

Abbreviations of the complex life cycle of trematodes, from three to two hosts, have occurred repeatedly and independently among trematode lineages. This is usually facultative and achieved via progenesis: following encystment in the second intermediate host, the metacercaria develops precociously into an egg-producing adult, bypassing the need to reach a definitive host. Given that it provides relatively cheap insurance against a shortage of definitive hosts, it is not clear why facultative progenesis has only evolved in a few taxa. Here a comparative approach is used to test whether progenetic trematodes are characterized by larger body size and egg volumes, two traits that correlate with other key life history features, than other trematodes. These traits may constrain the evolution of progenesis, because precocious maturation might be impossible when the size difference between the metacercaria and a reproductive adult is too large. First, trematode species belonging to genera in which progenesis has been documented were found not to differ significantly from other trematode species. Second, using within-genus paired comparisons across 19 genera in which progenesis has been reported, progenetic species did not differ, with respect to body size or egg size, from their non-progenetic congeners. Third, using intraspecific paired comparisons in species where progenesis is facultative, no difference was observed in the sizes of eggs produced by worms in both the intermediate and definitive host, suggesting that opting for progenesis does not influence the size of a worm's eggs. Overall, the lack of obvious differences in body or egg size between trematodes with truncated life cycles and those with the normal three-host cycle indicates that basic life history characteristics are not acting as constraints on the evolution of progenesis; trematodes of all sizes can do it. Why facultative progenesis is not more widespread remains a mystery.

Animals↗

Combining phylogenetic and ecological information into a new index of host specificity.

Host specificity has 2 independent facets: the extent to which different host species are used by a parasite, and the phylogenetic distances among these hosts. Although the number of host species exploited by a parasite commonly is used as a measure of host specificity, it fails to capture ecological and phylogenetic differences among hosts. Here, a new index of host specificity, S(TD)*, is developed and illustrated. This index measures the average taxonomic distinctness among the host species used by a parasite, weighted for the parasite's prevalence in the different hosts. For a given number of host species, the index approaches its minimum value when a parasite achieves high prevalence in a few closely related host species, and the index approaches its highest value when a parasite reaches its highest prevalence values in distantly related host species. Simple hypothetical examples are used to demonstrate the index's computation and some of its properties. The new index is influenced independently both by the taxonomic (or phylogenetic) affinities of a set of host species and by the distribution of prevalence values among these hosts. A single value cannot truly capture all the nuances of a phenomenon as complex as host specificity; nevertheless, the proposed index incorporates the features of specificity that are most relevant to parasitologists and will be a useful tool for comparative studies.

Animals↗

Detection of interspecific competition in parasite communities.

Matrices of correlation coefficients between the abundances or intensities of all pairs of helminth species, across all individual hosts in a sample, are regularly used to detect possible cases of interspecific competition in parasite communities. In these matrices, however, the range of possible values that any correlation coefficient can take is not -1 to 1, contrary to what is generally assumed. The number and magnitude of other correlation coefficients in a matrix will constrain the values that any given correlation can achieve. This property of matrices, and of inter-related natural variables, is explained and illustrated with 2 examples from real helminth communities. As a rule, the presence of many negative correlations in a matrix raises the lower value that any of them can possibly achieve. This has important but previously overlooked implications for the interpretation of correlation coefficients, and the detection of competition in natural parasite communities.

Animals↗

Relationships between parasite abundance and the taxonomic distance among a parasite's host species: an example with fleas parasitic on small mammals.

Opportunistic parasite species, capable of exploiting several different host species, do not achieve the same abundance on all these hosts. Parasites achieve maximum abundance on their principal host species, and lower abundances on their auxiliary host species. Taxonomic relatedness between the principal and auxiliary host species may determine what abundance a parasite can achieve on its auxiliary hosts, as relatedness should reflect similarities among host species in ecological, physiological and/or immunological characters. We tested this hypothesis with fleas (Siphonaptera) parasitic on small Holarctic mammals. We determined whether the abundance of a flea in its auxiliary hosts decreases with increasing taxonomic distance of these hosts from the principal host. Using data on 106 flea species from 23 regions, for a total of 194 flea-locality combinations, we found consistent support for this relationship, both within and across regions, and even after controlling for the potentially confounding effect of flea phylogeny. These results are most likely explained by a decrease in the efficiency of the parasite's evasive mechanisms against the host's behavioural and immune defences with increasing taxonomic distance from the principal host. Our findings suggest that host switching over evolutionary time may be severely constrained by the coupling of parasite success with the relatedness between new hosts and the original host.

Animals↗

The impact of sexual selection on Corynosoma magdaleni (Acanthocephala) infrapopulations in Saimaa ringed seals (Phoca hispida saimensis).

In free-living animals sexual selection is a central force shaping the spatial distribution of individuals in a population as well as sexual size dimorphism. We studied the influence of sexual selection on spatial distribution, female-to-male body size ratio, and female mating success of acanthocephalans in a natural host population of Saimaa ringed seal (Phoca hispida saimensis) harbouring a single intestinal helminth species, Corynosoma magdaleni. The acanthocephalans were always found along the full length of the small intestine; however, the site selection varied among the individual seals according to the age of the infection. The distribution of male acanthocephalans was not random with respect to females, with larger males tending to aggregate around non-mated females. A higher proportion of C. magdaleni females had copulated in seals with relatively more male worms. Male-male competition for access to females can be intense in C. magdaleni infrapopulation and may select for large males. We found that the larger the infrapopulation size, the smaller the males compared to females. In addition, the greater the female bias in the infrapopulation, the smaller the testes of males. Our study shows that sexual selection may be an important determinant of spatial distribution, male body size and female mating success of C. magdaleni in Saimaa ringed seal.

Acanthocephala↗

Fish population size, and not density, as the determining factor of parasite infection: a case study.

The diversity and abundance of parasites vary widely among populations of the same host species. These infection parameters are, to some extent, determined by characteristics of the host population or of its habitat. Recent studies have supported predictions derived from epidemiological models regarding the influence of host population density: parasite abundance and parasite species richness are expected to increase with increasing host population density, at least for directly transmitted parasites. Here, we test this prediction using a natural system in which populations of the crucian carp, Carassius carassius (L.), occur alone, with no other fish species, in a series of 9 isolated ponds in Finland. The ectoparasite communities in these fish populations consist of only 4 species of monogeneans (Dactylogyrus formosus, D. wegeneri, D. intermedius and Gyrodactylus carassii); the total and relative abundance of these 4 species varies among ponds, with one or two of the species missing from certain ponds. Across ponds, only one factor, total fish population size, explained a significant portion of the variance in both the mean number of monogenean species per fish and the mean total abundance of monogenean individual per fish. In contrast, fish population density did not influence either monogenean abundance or species richness, and neither did any of the other variables investigated (mean fish length per pond, number (of fish examined per pond, distance to the nearest lake, and several water quality measures). In our system, proximity among fish individuals (i.e. host population density) may not be relevant to the proliferation of monogeneans; instead, the overall availability of host individuals in the host population appeared to be the main constraint limiting parasite population growth.

Animals↗

Parasite biodiversity and its determinants in coastal marine teleost fishes of Brazil.

Recent studies of the forces behind the diversification of parasite assemblages have shed light on many aspects of parasite biodiversity. By using only parasite species richness as their measure of diversity, however, previous investigations have ignored the relatedness among parasite species and the taxonomic structure of the assemblages, which contain much information about their evolutionary origins. Here, we performed a comparative analysis across 50 species of fish from the coast of Brazil; we evaluated the effects of several host traits (body size, social behaviour, feeding habits, preference for benthic vs. pelagic habitats, depth range, and ability to enter brackish waters) on the diversity of their assemblages of metazoan parasites. As measures of diversity, we used parasite species richness, as well as the average taxonomic distinctness of the assemblage and its variance; the latter measures are based on the average taxonomic distance between any two parasite species in an assemblage. Unlike parasite species richness, taxonomic distinctness was unaffected by the number of host individuals examined per species. Fish body length proved to be the main predictor of parasite species richness, even when controlling for the confounding influences of host phylogeny and sampling effort, although it did not correlate with measures of parasite taxonomic distinctness. Predatory fish also had higher parasite species richness than planktivores, but this trend could not be confirmed using phylogenetically independent contrasts between host taxa. The main host feature associated with the taxonomic diversity of parasites was schooling behaviour, with schooling fish having more taxonomically diverse parasite assemblages than those of their non-schooling relatives. When focusing on endoparasite species only, both predatory feeding habits and a broad depth range were associated with the taxonomic distinctness of parasites. Our results suggest that certain host traits (i.e. body size) determine how many parasite species a host can accumulate over evolutionary time, whereas different host features influence the processes causing the taxonomic diversification of parasite assemblages.

Animals↗

Stability of Corynosoma populations with fluctuating population densities of the seal definitive host.

In theory there should be a strong coupling between host and parasite population sizes. Here, we investigated population size and structure in 3 species of acanthocephalans, Corynosoma semerne, C. strumosum and C. magdaleni, in ringed seals (Phoca hispida) from the Bothnian Bay over a period of more than 20 years. During this period, seal numbers first decreased markedly and then increased steadily; at the same time, a paratenic fish host particularly important for C. strumosum has gradually disappeared from the bay due to decreasing salinity. We found no evidence that the mean abundance of any of the 3 acanthocephalan species changed significantly over time, nor was there any relationship between parasite abundance at any point in time and seal numbers in the corresponding year. Based on the proportion of sexually mature female worms per infrapopulation, and on relationships between the sex ratio of worms and infrapopulation size, both C. magdaleni and C. semerme appear to be doing well, independently of the population size of their seal definitive hosts. In contrast, perhaps because of the loss of its main paratenic host, C. strumosum appears more at risk in the Bothnian Bay. Our results show that in complex natural systems, there are not necessarily simple, direct links between definitive host population size or density, and parasite population dynamics.

Acanthocephala↗

Equal partnership: two trematode species, not one, manipulate the burrowing behaviour of the New Zealand cockle, Austrovenus stutchburyi.

Metacercariae of the trematode Curtuteria australis (Echinostomatidae) accumulate in the foot of the New Zealand cockle Austrovenus stutchburyi, severely impairing the cockle's ability to burrow under the sediments. This results in increased predation by birds on cockles, and thus enhanced transmission rates of the parasite to its bird definitive hosts. This host manipulation by the trematode is costly: fish regularly crop the tip of the foot of cockles stranded on the sediment surface, killing any metacercariae they ingest. A second, previously undetected trematode species (characterized by 23 collar spines) co-existing with C. australis, has been found in the foot of cockles in the Otago Harbour, South Island, New Zealand. The relative abundance of the two species varies among localities, with the identity of the numerically dominant species also changing from one locality to the next. Both C. australis and the new species have a strong preference for encysting in the tip of the cockle's foot, where their impact on the burrowing ability of the host is greatest, and where they both face the risk of cropping by fish. Results indicate that these two species are ecological equivalents, and their combined numbers determine how the cockle population is affected.

Animals↗

Parasite-induced surfacing in the cockle Austrovenus stuchburyi: adaptation or not?

Parasite manipulation of host behaviour is a compelling example of the extended phenotype. However, in many cases, such manipulation may be incorrectly assumed. Previous work has demonstrated that Austrovenus stuchburyi cockles stranded on mud-flat surfaces due to an inability to re-burrow both contain significantly more metacercariae of the trematode Curtuteria australis and are predated by the definitive host of this parasite at a faster rate than burrowed cockles. These results have been interpreted as strong evidence for a manipulation of cockle behaviour by the trematode to facilitate transmission to the definitive host. The model presented here, however, indicates that the selective advantage to the parasite of the altered host behaviour is currently of a negligible level at our study site that is highly unlikely to have been realized as an adaptation over evolutionary time. Hence, there are no grounds on which the more parsimonious explanation, that the altered host behaviour observed is simply an incidental side-effect of infection, can be rejected. We thus maintain that for any change in the behaviour of infected hosts to be confirmed as potentially a parasite trait that has evolved in response to selection, the adaptive benefit taking into account the entire parasite life cycle may need to be considered.

Adaptation, Physiological↗

Effects of conspecifics and heterospecifics on individual worm mass in four helminth species parasitic in fish.

Intraspecific and interspecific effects on the growth and body size of helminths are rarely studied in natural situations, yet knowing what determines helminth sizes and thus fecundity is crucial to our understanding of helminth ecology and epidemiology. The determinants of average individual worm mass were investigated in four common species of helminths parasitic in trout, Salmo trutta. In the acanthocephalan Echinorhynchus truttae, there was a negative relationship between the intensity of infection by conspecifics and average individual worm size. However, in the acanthocephalans Pomphorhynchus laevis and Acanthocephalus anguillae and in the cestode Cyathocephalus truncatus, the relationship was positive: individual worms were larger on average when co-occurring with many conspecifics than when co-occurring with very few. In addition, the average mass of individual C. truncatus in a host decreased as the total mass of other helminth species in the same host increased. This interspecific effect involves the whole helminth community, as the combined effect of all other helminth species is a better predictor of reduced mass in C. truncatus than the mass of any other species taken on its own. These results illustrate the importance of considering helminth interactions and helminth growth in a natural setting.

Acanthocephala↗

Parasite specialization from a phylogenetic perspective: a new index of host specificity.

The host specificity of a parasite is not merely a function of how many host species it can exploit, but also of how closely related these host species are to each other. Here, a new index of host specificity is proposed, one that takes into account the average taxonomic or phylogenetic distance between pairs of host species used by a parasite. The index is derived from measures of taxonomic distinctness used in biodiversity studies. It is easy to compute and interpret, ranging from a minimum value of 1 when all host species are members of the same genus, to a maximum of 5, when all host species belong to different classes. The variance of this measure can also be computed, and provides additional information on the taxonomic or phylogenetic structure of the host assemblage. Using data on helminth parasites of Canadian freshwater fishes, we show that the new index, unlike the mere number of known host species, is independent of study effort i.e. the number of published records of a parasite. Although the index and the number of known hosts are not entirely independent statistically, each captures a different aspect of host specificity. For instance, although acanthocephalans infect significantly more host species than trematodes, cestodes or nematodes, there is no difference in the average index value among these 4 helminth taxa, suggesting that the average taxonomic distances between the host species of a parasite do not vary among these higher taxa. We recommend the use of our new index in future comparative studies of host specificity, in particular when the focus is on the evolutionary history of parasites and of their past colonizations of host lineages.

Animals↗

Spatial and temporal repeatability in parasite community structure of tropical fish hosts.

An assessment is made of the repeatability of parasite community structure in space for a marine fish, and in space and time for a freshwater fish from south-eastern Mexico. The marine fish species was the red grouper, Epinephelus morio (collected from 9 localities), and the freshwater species was the cichlid, Cichlasoma urophthalmus (collected from 6 localities: including monthly at 2 localities for 1 year, and bimonthly at 1 locality in 1990 and 1999). Pairwise interspecific associations and analyses of nested patterns in the distributions of parasite species among hosts were used in both fish species, with comparisons over time made only with the cichlid. Positive interspecific associations, and nested patterns were noted in some localities for both fish species, and/or at some sampling times for the cichlid fish. However, non-random patterns in the structure of parasite communities in these 2 host species only were observed sporadically. When present, nestedness in both fish species was apparently linked with a positive association between total infection intensities and fish size. Additionally, adjacent localities were more likely to display similar parasite community structure than distant ones. This preliminary result suggests that distance between localities is an important determinant of predictability in parasite community structure.

Animals↗

Egg size variability in trematodes: test of the bet-hedging hypothesis.

The hypothesis according to which egg size variability in hermaphroditic parasites results from bet-hedging was investigated in a comparative analysis using trematodes as a model. We hypothesized that the species reproducing mainly by self-fertilization should produce smaller eggs than those species that regularly practice cross-fertilization. Indeed, because self-fertilization is usually associated with inbreeding depression, selection should favor individuals spreading the risk of genetically disturbed development across more but smaller eggs, instead of producing fewer eggs, each possessing a large resource supply, of which many may fail to develop because of genetic deficiencies. On the basis of earlier theoretical and empirical studies, we assumed that the ratio length of testis-length of ovary positively correlates with the mating group size and, hence, with opportunities for cross-fertilization. In accordance with the bet-hedging hypothesis, we found, across trematode species, a positive relationship between this ratio and the mean egg volume produced by adults. This result was, however, observed only for the trematodes infecting birds and not for the species infecting fishes and mammals. In addition, once the influence of trematode phylogeny was taken into account, there was no significant trend, suggesting that phylogenetic legacies played a large role in generating the previous signal. Experimental tests of the bet-hedging hypothesis will be necessary to clarify the matter.

Animals↗

A gastropod scavenger serving as paratenic host for larval helminth communities in shore crabs.

The whelk Cominella glandiformis is an important predator-scavenger of New Zealand intertidal ecosystems; a few whelks can quickly eat all the soft tissues of recently dead crabs. In this study, we demonstrate that whelks can also ingest and act as paratenic hosts for at least 4 helminth species that use crabs as intermediate hosts: metacercariae of the trematode Maritrema sp. and of another unidentified trematode, larval acuariid nematodes, and cystacanths of the acanthocephalans Profilicollis spp. Large whelks ingest disproportionately more helminth larvae than small whelks, but the survival of parasites during their short stay in the whelks is not affected by whelk size. The majority of metacercariae and nematodes are passed out in whelk feces within 3 days of ingestion, whereas the few cystacanths found did not leave whelks until after that time; no parasite was left in whelks 5 days postingestion. Survival of all 4 helminth species was generally very high, though it decreased day by day in 2 species. Given that the avian definitive hosts of all 4 helminths also eat whelks, our results indicate that alternative transmission pathways exist and that parasites can take routes through food webs that are too often ignored.

Acanthocephala↗

Parasitism, community structure and biodiversity in intertidal ecosystems.

There is mounting evidence that parasites can influence the composition and structure of natural animal communities. In spite of this, it is difficult to assess just how important parasitism is for community structure because very few studies have been designed specifically to address the role of parasites at the community level, no doubt because it is difficult to manipulate the abundance of parasites in field experiments. Here, we bring together a large amount of published information on parasitism in intertidal communities to highlight the potential influence of parasites on the structure and biodiversity of these communities. We first review the impact of metazoan parasites on the survival, reproduction, growth and behaviour of intertidal invertebrates, from both rocky shores and soft-sediment flats. Published evidence suggests that the impact of parasites on individuals is often severe, though their effects at the population level are dependent on prevalence and intensity of infection. We then put this information together in a discussion of the impact of parasitism at the community level. We emphasize two ways in which parasites can modify the structure of intertidal communities. First, the direct impact of parasites on the abundance of key host species can decrease the importance of these hosts in competition or predator-prey interactions with other species. Second, the indirect effects of parasites on the behaviour of their hosts, e.g. burrowing ability or spatial distribution within the intertidal zone, can cause changes to various features of the habitat for other intertidal species, leading to their greater settlement success or to their local disappearance. Our synthesis allows specific predictions to be made regarding the potential impact of parasites in certain intertidal systems, and suggests that parasites must be included in future community studies and food web models of intertidal ecosystems.

Animals↗

Intra- and interspecific density-dependent effects on growth in helminth parasites of the cormorant, Phalacrocorax carbo sinensis.

The action of intra- and interspecific competition, mediated by density-dependent effects on growth, was investigated among the 3 helminth species found in the alimentary tract of 104 cormorants, Phalacrocorax carbo sinensis. Intraspecific density-dependent effects on worm sizes were observed in the abundant nematode Contracaecum rudolphii, as shown by a negative correlation between mean worm size and intensity of infection. Higher intensities of infection by C. rudolphii were also associated with more variable worm sizes in the nematode Syncuaria squamata, suggesting a one-sided and density-dependent interspecific effect. There was also clear evidence of some form of negative interaction between the nematode S. squamata and the acanthocephalan Southwellina hispida from two fronts. First, there was a strong negative correlation between the intensities of infection of the 2 species across hosts. Second, sizes of worms of 1 species became more variable as the number of worms of the other species per host increased, and vice versa. This interspecific density-dependent effect on growth was thus apparently symmetrical. We also found evidence that worm size is a predictor of egg output in the 3 helminth species, indicating that intra- and interspecific density-dependent effects on growth can affect population dynamics in these worms. These results illustrate the complex nature of density dependence in helminth growth, and how its effects can act both within and among species.

Acanthocephala↗

Inequalities in size and intensity-dependent growth in a mermithid nematode parasitic in beach hoppers.

Inequality in body sizes is a common feature in populations of helminth parasites, with potential consequences for egg production and population genetics. Inequalities in body lengths and the effects of intraspecific competition on worm length were studied in a species of mermithid nematode parasitic in the crustacean Talorchestia quoyana (Amphipoda: Talitridae). The majority of the 753 worms recovered were relatively small, and an analysis using a Lorenz curve and Gini coefficient suggested that there were no marked inequalities in body lengths among the worms. Total worm length in the 356 infected amphipods (i.e. the sum of the lengths of all the worms in a host) increased steadily as a function of the number of worms per amphipod, whereas the length of the longest worm per amphipod peaked in amphipods harbouring intermediate numbers of worms. This last result was not significantly accounted for by the observed increase in host size with increasing intensity of infection, but resulted from a correlation between worm length and host size. As the number of worms per amphipod increased, the relative sizes of the second-, third-, and fourth-longest worms per host increased markedly. This means that relative inequalities in sizes become less pronounced, i.e. subordinate worms get closer in size to the longest worm, as the number of worms per host increases. The main consequence of this phenomenon is that worm sizes in the mermithid population are more homogeneous than they would be if intraspecific competition had stronger effects on worm growth.

Animals↗