Infrapopulation Size Research Articles

High within-host genetic variation of the nematode Spirocerca lupi in a high-density urban dog population

The nematode worm Spirocerca lupi has a cosmopolitan distribution and can cause the death of its final canid host, typically dogs. While its life cycle, which involves a coprophagous beetle intermediate host, a number of non-obligatory vertebrate paratenic hosts and a canid final host, is well understood, surprisingly little is known about its transmission dynamics and population genetic structure. Here we sequenced cox1 to quantify genetic variation and the factors that limit gene flow in a 300km2 area in South Africa. Three quarters of the genetic variation, was explained by differences between worms from the same host, whereas a quarter of the variation was explained by differences between worms from different hosts. With the help of a newly derived model we conclude that while the offspring from different infrapopulations mixes fairly frequently in new hosts, the level of admixture is not enough to homogenize the parasite populations among dogs. Small infrapopulation sizes along with clumped transmission may also result in members of infrapopulations being closely related.

Infrapopulations ofProcamallanus elatensisFusco & Overstreet, (Nematoda: Camallanidae) in the rabbitfishSiganus rivulatus(Teleostei, Siganidae) from the Saudi coast of the Red Sea

Factors regulating gastrointestinal nematode infrapopulations are to a great extent density-dependent. Here, 23 natural infrapopulations (41-281 individuals) of the viviparous nematode Procamallanus elatensis (Camallanidae) from the fish Siganus rivulatus were found distributed in a well-defined fundamental niche (posterior 55% of the intestine). Immature worms were mostly found in the anterior third of this niche, while mature worms were found in aggregations posterior to them and followed by gravid females in the posterior 20% of the intestine. This distribution strongly suggests that worms migrate towards the posterior intestine while they mature, copulate and reproduce. In small infrapopulations, the sex ratios were distinctly female-biased and the number of gravid females was low. In large infrapopulations, the sex ratios were distinctly male-biased and the number of gravid females was high. However, the mean lengths of both immature males and females and mature females decreased dramatically as the infrapopulation size increased, while those of mature males increased significantly. These results strongly suggest intraspecific competition and density-dependent regulation of mean worm length, and the increase in the mean length of mature males strongly suggests intense sexual selection and competition between mature males. Production of larvae by female worms decreased significantly as the infrapopulation size increased, suggesting a density-dependent reduction in female worm fertility. The results are statistically significant and strongly suggest that infrapopulation self-regulation is through density-dependent mechanisms, in which development of immature worms, infrapopulation size, sex ratio, sexual selection and carrying capacity of the fundamental niche play essential roles in shaping and regulating the infrapopulations.

Genetic and morphological variation in Echinorhynchus gadi Zoega in Müller, 1776 (Acanthocephala: Echinorhynchidae) from Atlantic cod Gadus morhua L.

Previous studies have shown considerable variability in morphological features and the existence of genetically distinct sibling species in the acanthocephalan Echinorhynchus gadi Zoega in Müller, 1776. The aim of the present study was to follow up and extend those earlier studies by using a combination of DNA analysis and morphometrics to investigate differences between samples of E. gadi from Atlantic cod Gadus morhua L. caught at five fishing grounds in the Baltic Sea and three in different parts of the North Atlantic. Twelve morphological features were measured in 431 specimens of E. gadi, 99 individuals were studied by polymerase chain reaction-restriction fragment length polymorphosm (PCR-RFLP), and selected PCR products were sequenced. The molecular analyses showed the nucleotide sequences of E. gadi rDNA from cod caught at all the sampling sites to be identical. The comparative morphological study, in contrast, revealed significant differences between samples of E. gadi from different sampling sites and showed the separation of E. gadi into two groups corresponding approximately to the systematic classification of cod into the two subspecies, Atlantic G. morhua morhua and Baltic G. morhua callarias. The E. gadi infrapopulation size had a significant effect on some of the morphological features. The results are discussed in relation to cod population biology, the hydrography of the study area and the history of the Baltic Sea formation.

Population Biology of Contracaecum rudolphii Sensu Lato (Nematoda) in the Great Cormorant (Phalacrocorax carbo) from Northeastern Poland

Parameters related to the occurrence, aggregation, and population structure of the nematode Contracaecum rudolphii Hartwich, 1964 , in the great cormorant ( Phalacrocorax carbo ) from northeastern Poland were analyzed. A total of 491 birds of different ages (adults, immatures, and nestlings) was examined; the cormorants studied were taken from both fresh- and brackish water habitats. Contracaecum rudolphii were found in stomachs of 454 birds (92.5%); the 46,244 nematodes included third- (L3) and fourth-stage larvae (L4), and sub-adult and adult females and males. The distribution of nematodes in the host population were highly aggregated. The occurrence of C. rudolphii was significantly dependent on the host's age and habitat, as well as on season; the proportion of development stages differed significantly depending on birds' age and season (the latter only in adult birds). The infrapopulations of C. rudolphii in the adult cormorants showed distinct density-dependent correlations: that is, as the infrapopulation size increased, the number of adult females C. rudolphii decreased, and the proportion of larvae and sub-adult females increased. A higher proportion of larvae and sub-adult females characterized the component population structure of the nematodes in the cormorant nestlings, compared with adult birds, probably because of immune system deficiency in the immature birds, coupled with the development of the nematode population. Seasonal changes in the C. rudolphii population, observed in the adult cormorants, were not related to seasonality of the L3 occurrence in food; instead, the changes are believed to have resulted from independent processes of elimination of the oldest nematodes and their replacement by larval stages that subsequently matured.

The Comparison of Mean Crowding Between Two Groups

The infrapopulation size, or crowding, is an important measure in parasitology because of its impact on many facets of parasite biology. However, crowding values are difficult to handle statistically because of dependencies. Therefore, a bootstrap procedure was proposed in order to compare crowding between 2 groups or samples of hosts. Here, we propose Lepage's location-scale test based on intensities as an alternative to test whether there is a difference in mean crowding. Simulation results indicate that Lepage's test is superior to the bootstrap, even when sample sizes are large as needed for the bootstrap procedure. Lepage's test, however, can also be applied with moderate or small sample sizes.

Is metal accumulation in Pomphorhynchus laevis dependent on parasite sex or infrapopulation size?

Concentrations of the elements As, Cd, Co, Cu, Fe, Mn, Mo, Ni, Pb, Sn, V, Zn were analysed by inductively coupled plasma mass spectrometry (ICP-MS) in the acanthocephalan Pomphorhynchus laevis and its fish host Barbus barbus. A total of 27 barbels were collected from the Danube River in autumn 2006 close to the town Kozloduy (685 river kilometer) on the Bulgarian river bank. Fish were divided into 3 groups. According to their P. laevis infrapopulation size hosts were considered as heavily infected (>100 worms per fish) and lightly infected (<20 worms per fish). The third group was used to compare heavy metal concentrations between male and female P. laevis. The 5 elements As, Cd, Cu, Pb and Zn were detected in significantly higher concentrations in parasites compared to host tissues (muscle, intestine, liver). According to the calculated mean bioconcentration factors, 3 more elements (Co, Mn, V) showed usually higher concentrations in P. laevis. Comparisons between heavily and lightly infected fish revealed significant differences only for V with higher concentrations for the heavily infected group. Concerning sex-specific metal accumulation V and Zn showed significant differences (V, at P<0.05; Zn, at P=0.05), with higher levels of both metals in females of P. laevis. Our results suggest that - for the metals analysed - the size of the parasite infrapopulation plays no role in the degree of metal accumulation. Similarly, parasite sex seems not to be a crucial factor for metal accumulation in the parasites. Thus, for metal monitoring purposes there is no need to take these aspects into account, which makes the use of parasites as bioindicators more applicable.

Sex ratio and morphological polymorphism in an isolated, endemicTeladorsagia circumcinctapopulation

Teladorsagia circumcincta is a polygamous nematode that exhibits morphological polymorphism. Sex ratio is typically female biased and the male nematodes occur in association with the genetically similar, minor morphotypes Teladorsagia davtiani and Teladorsagia trifurcata. In experimental infections, sex ratio (proportion male) and the proportion of minor male morphs observed have been shown to be influenced by both host and nematode-related factors. As similar investigations from natural systems are rare, this study examined whether sex ratio and minor male morph frequency were associated with host age and sex and nematode infra-population size in the isolated Soay sheep population on St Kilda. Count data for Teladorsagia were analysed for sheep of all age classes and both sexes from the winters of three consecutive population crashes (1999, 2002 and 2005). Generally, the intensity of Teladorsagia nematodes increased with host age until the age of 2 years before decreasing. In 2005, abundance of nematodes was generally higher than in the previous crashes, nematode sex ratio was negatively associated with host age and tended to be negatively associated with nematode intensity. Within the male nematode subpopulation, T. circumcincta always predominated, followed by T. davtiani and then T. trifurcata, with little variation in the relative proportions between hosts. The presence of each minor morph was primarily associated with the intensity of male T. circumcincta and, in those hosts where all three male morphs were detected, intensity of each minor morph was most associated with intensity of Teladorsagia females. Therefore, in a year when the nematode was generally more abundant, sex ratio appeared to be influenced by both host and nematode-related factors, whereas in all years examined, the frequency of morphological polymorphism was primarily density dependent.

Sex ratio in flea infrapopulations: number of fleas, host gender and host age do not have an effect

This study set out to determine whether the sex ratio of fleas collected from host bodies is a reliable indicator of sex ratio in the entire flea population. To answer this question, previously published data on 18 flea species was used and it was tested to see whether a correlation exists between the sex ratio of fleas collected from host bodies and the sex ratio of fleas collected from host burrows. Across species, the female:male ratio of fleas on hosts correlated strongly with the female:male ratio of fleas in their burrows, with the slope of the regression overlapping 1. Controlling for flea phylogeny by independent contrasts produced similar results. It was also ascertained whether a host individual is a proportional random sampler of male and female fleas and whether the sex ratio in flea infrapopulations depends on the size of infrapopulations and on the gender and age of a host. Using field data, the sex ratio in infrapopulations of 7 flea species parasitic on 4 rodent species was analysed. Populations of 3 species (Nosopsyllus iranus, Parapulex chephrenis and Xenopsylla conformis) were significantly female-biased, whereas male bias was found in 1 species (Synosternus cleopatrae). In general, the sex ratio of fleas collected from an individual rodent did not differ significantly from the sex ratio in the entire flea population. Neither host gender, and age nor number of fleas co-occurring on a host affected (a) the sex ratio in flea infrapopulations and (b) the probability of an infrapopulation to be either female- or male-biased.

Intraspecific density-dependent effects on growth and fecundity of Diplosentis nudus (Harada, 1938) Pichelin et Cribb, 2001 (Acanthocephala, Cavisomidae)

AbstractDuring June and July of 2007, a total of 130 specimens of the fish Rastrelliger kanagurta Cuvier (Teleostei, Scombridae), ranging between 19–31 cm in total length, were caught in the Red Sea off the coast of Sharm El-Sheikh, South Sinai, Egypt, and examined for infections by acanthocephalans (65 fish/month). Of this number, 29 (22.30%) were slightly or heavily parasitized by the acanthocephalan Diplosentis nudus (Harada, 1938) Pichelin et Cribb, 2001 (Cavisomidae); no other helminth parasites were found in the intestine of R. kanagurta. Twenty-nine infrapopulations of D. nudus, ranging from 23–218 individuals were collected from the infected fishes. These infrapopulations were distributed in a well-defined fundamental niche along the intestine of R. kanagurta, where the distribution of male worms was not random with respect to female worms size and position and suggests that the male-male competition for access to female may be intense and may select for large males. No correlation between fish size and infrapopulation size was observed. Correlations between female-to-male sex ratio and infrapopulation size, numbers of females and their mean lengths, numbers of males and their mean lengths, mean female length and mean male length within infrapopulation were very strong, and clearly suggest that as the infrapopulation size increased, the number of females and their mean lengths decreased and the number of males and their mean lengths increased. Combination of these results strongly suggests density-dependent effects and competition between male worms. The relationship between the mean female length or size and the number of eggs within its pseudocoel was strongly positive; egg production by female worm significantly decreases as the infrapopulation size increases, suggesting density-dependent reduction in female worm fecundity. Tendency for the variability in male testes size was not significant in infrapopulations of D. nudus. All of these results are discussed.

Aspectos quantitativos e morfométricos das infrapopulações de platynosomum illiciens (trematoda: dicrocoeliidae) em paroaria dominicana (passeriformes: emberizidae) do estado da Bahia, Brasil

During the study of the metazoan parasites of Paroaria dominicana Linnaeus, 1758 (Passeriformes), four infrapopulations of Platynosomum illiciens (Brown, 1901) (Trematoda), parasitic in the liver ducts were collected, with intensity of infection 10, 11, 18 and 36 specimens respectively. Differences between the means of morphometrical values and the total number of eggs were statistically tested. The liver weigh were not significantly correlated with avian total weigh and length, and with the total parasitic abundance. Positive correlations were observed between the size of infrapopulations and the total number of eggs, and between parasite body size and the eggs. size. Negative correlations were observed between parasitic intensity and liver weigh; between the size of infrapopulations and the parasite body size; between total number and the size of eggs and between parasite body size and total number of eggs. These results suggests that host with smaller liver had greater infrapopulations. These greater infrapopulations showed smaller parasite body size and higher size and number of eggs.

DENSITY-DEPENDENT TOPOGRAPHICAL SPECIALIZATION IN GYRODACTYLUS ANISOPHARYNX (MONOGENOIDEA, GYRODACTYLIDAE): BOOSTING TRANSMISSION OR EVADING COMPETITION?

Viviparous gyrodactylids are remarkable monogenoid ectoparasites, not only because of their speciousness, but also due to their unusually wide range of hosts. Although many factors have been proposed to determine the location where gyrodactylids attach to their hosts, little is known about how their preference for specific host body regions changes over the course of infection. In this study, we investigate the dynamics of topographical specialization of the parasite Gyrodactylus anisopharynx on 2 of its natural freshwater fish hosts (Corydoras paleatus and C. ehrhardti), as well as a naïve host (C. schwartzi). We recorded the spatial location of this parasite from the foundation of the infrapopulation to its extinction to assess how topographical specialization is affected by host species, the size and the age of the infrapopulation, and the possibility of transmission among hosts. Our results indicate that topographical specialization is negatively correlated with infrapopulation size and only marginally affected by infrapopulation age. Also, the degree of specialization was different among host species, but seemed unaffected by the possibility of transmission among hosts. Therefore, observed changes in spatial specialization of G. anisopharynx do not appear to represent adaptive responses to maximize their transmission. Rather, mechanisms such as increased competition and/ or local immune responses might cause parasites to occupy less favorable regions of the body of their hosts with increasing density.

Stability ofCorynosomapopulations 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.

POPULATION STRUCTURE OF ANISAKIS SIMPLEX (NEMATODA) IN HARBOR PORPOISES PHOCOENA PHOCOENA OFF DENMARK

The population structure and habitat selection of Anisakis simplex in 35 harbor porpoises off Denmark are described. The nematodes were collected from the stomach and duodenal ampulla and were categorized as third-stage larvae, fourth-stage larvae, subadults, and adults. The porpoises harbored 8,043 specimens of A. simplex. The proportion of adults and subadults increased with infrapopulation size. The number of development stages across infrapopulations covaried significantly (Kendall's test of concordance). Concordance was higher in hosts with the highest intensities than in those with low and medium intensities. All stages occurred mainly in the forestomach, but this trend was stronger for the adults. Adult and subadult sex ratios did not depart significantly from 1:1. Our data suggested that recruitment and duration of each stage were the main factors accounting for infrapopulation structure. The preference of A. simplex for the forestomach conformed with previous studies, but the narrower distribution of adults relative to other stages might indicate a strategy to enhance mating opportunities. Information on sex ratios of A. simplex is scarce and contradictory. We suggest that the discrepancies might partly reflect differences in categorization criteria and statistical methods.

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.

Infrapopulation sizes of co-occurring trematodes in the snail Ilyanassa obsoleta.

This study addresses the infrapopulation sizes of 2 larval trematode species Himasthla quissetensis and Zoogonus rubellus as they co-occur within their estuarine snail host Ilyanassa obsoleta. Rediae of H. quissetensis and sporocysts of Z rubellus were counted in snails singly infected with each parasite and in snails infected with both. Comparisons of the counts indicate that infrapopulations of H. quissetensis were unaffected by co-occurrence with Z rubellus. However, Z. rubellus infrapopulations were reduced when co-occurring with H. quissetensis. It is proposed that this situation does not result from an interspecific interaction between parasite species. Although this double infection is relatively frequent in certain snail populations, it is contended that these trematode species do not co-occur often enough to evolve responses to one another. However, the host environment must be encountered in each life cycle, and both trematode species must be adapted to use it. On this basis, whatever happens when these 2 species occupy the same host is based on adaptations of the parasites to the host. It is proposed that these parasites are adapted to self-limit their infrapopulations in the snail host. They can, thus, preserve and use the host for many years and thereby enhance total cercarial transmission (fitness). Infrapopulation sizes would be determined by host resource levels, which, among other factors, would be influenced by the presence of multiple parasite species. In single infections, by far the most common situation, host resource levels would be set by the nutritional status or age (size) of the host (or both). The reduced infrapopulation sizes of Z rubellus on co-occurrence suggest that this trematode is more sensitive to host resource levels than is H. quissetensis.

Infrapopulation Sizes of Co-Occurring Trematodes in the Snail Ilyanassa obsoleta

Infrapopulation Sizes of Co-Occurring Trematodes in the Snail Ilyanassa obsoleta

Sources of variability in aggregation and sex ratios of Crataerina melbae (Diptera: Hippoboscidae) among adult colonial alpine swifts.

Aggregation of Crataerina melbae flies on breeding adult alpine swifts (Apus melba) was low when compared with other host-parasite systems and varied with sampling date, year, and sex of the flies. Generalized linear models were performed to ascertain which factors, extrinsic and/or intrinsic to the host, explained variability in the number of louse flies present on a single host, i.e., abundance. Overall abundance was unrelated to any host characteristic but varied slightly among years. Abundance of female flies varied among years, but also with date of sampling, the number of females increasing as the breeding season advanced. In contrast, abundance of males decreased as the season progressed, independently of host characteristics. Despite these different patterns, the number of flies of each sex on a given host was strongly intercorrelated. These results suggest that mate attraction may explain aggregation patterns in this louse fly species. Overall sex ratio of louse flies did not differ from unity. However, the proportion of males decreased during the breeding season, as a consequence of the opposite sex-related seasonal patterns in parasite abundance. Sex-ratio variability was not related to host characteristics or to infrapopulation sizes.

The Role of Habitat in Structuring Halipegus occidualis Metapopulations in the Green Frog

The transmission dynamics of the trematode Halipegus occidualis in its definitive host, Rana clamitans, have been examined over a 5-yr period in a North Carolina pond. The breeding season of green frogs coincides with the period of worm recruitment, during which time male frogs are territorial and females show strong site fidelity. This site fidelity allows inferences to be made regarding the suitability of a particular habitat for worm transmission based on frog infection intensities within that habitat. Four foci of infection were identified in the pond by plotting worm infrapopulation size against site of host capture. Sites within infection foci are characterized by shallow water and emergent vegetation, factors favorable for overlapping distributions of the 4 hosts in the life cycle of H. occidualis. Consistent year-to-year worm prevalences and intensities, despite fluctuations in frog population size, are thought to be the result of a relatively constant proportion of the frog population being present in infection foci each year. Removal of worms from heavily infected frogs in the fifth year resulted in further heavy worm recruitment by treated frogs suggesting that site selection can predispose a frog to heavy infection. Further, the sum of removed parasites and those recruited after parasite removal by treated frog hosts was higher than worm infrapopulations observed in previous years, indicating that worm density regulates parasite infrapopulation size in heavily infected frogs.

Infrapopulation Dynamics of Halipegus occidualis and Halipegus eccentricus (Digenea: Hemiuridae): Temporal Changes within Individual Hosts

Individual infrapopulations of Halipegus occidualis and Halipegus eccentricus (Hemiuridae) in the buccal cavities of their amphibian host (Rana clamitans) were monitored for changes over time. Rates and patterns of parasite maturation, recruitment, and loss were estimated, and the probabilities of infection for definitive hosts in this system were calculated. Although variable, the time required for maturation of immature worms (once in the buccal cavity) was as little as 1 wk. Relatively rapid losses of worms from individual infrapopulations were observed, although this did not appear to inhibit additional recruitment of these trematodes. Also, it was clear that these species overwintered within their amphibian hosts. There was no statistical difference among the 3 sampling years in the probability of acquiring an initial or a subsequent infection with either parasite. However, we suggest that frogs with an existing infrapopulation may have a greater probability of adding worms if those hosts are within certain foci of infection within the pond. The probability of frogs becoming infected in this system increased in late spring and peaked in midsummer. By examining the temporal changes within individual infrapopulations, it is clear that dynamic changes in infrapopulation size and composition may occur, suggesting that rates of trematode transmission may be greater than previously estimated.

Seasonal Population Dynamics of Halipegus occidualis and Halipegus eccentricus (Digenea: Hemiuridae) in Their Amphibian Host, Rana clamitans

The seasonal population dynamics of Halipegus occidualis and Halipegus eccentricus (Hemiuridae) in their amphibian host, the green frog (Rana clamitans), were examined for 3 yr (weekly, April through October). Frogs were caught, marked, and examined for H. occidualis and H. eccentricus, both of which occur as immatures and adults in the buccal cavity. Frogs were then released and allowed to continue natural recruitment and loss of the parasites. It was thus possible to monitor individual infrapopulations over successive time periods. One-hundred and forty-nine frogs were caught and released, with a total of 328 observations. Overall, the levels of both parasites among male and female frogs were similar, as were levels of infection among adult and juvenile hosts. There was, however, no correlation between the total number of H. occidualis and the body size (snout-vent length) of R. clamitans. In contrast, there was a significant correlation between the total number of H. eccentricus and frog size. Recruitment of both species began in May, peaked in June/July, and ended in July (H. eccentricus) or August (H. occidualis). In general, both the prevalence and relative density of H. occidualis was greater than that of H. eccentricus and may be related to space constraints in the buccal cavity of R. clamitans. Adults of both species were observed from April through October. Following increases in parasite recruitment, infrapopulation sizes declined in September 1992 and in August of 1993 and 1994. Examination of variance to mean ratios indicated that both species were overdispersed in the frogs. Large declines in the variance to mean ratios for H. occidualis after periods of greatest recruitment are most likely associated with the loss of larger infrapopulations, suggesting that there may be density-dependent regulation of infrapopulation size. By monitoring individual hosts using the mark-release-recapture protocol, dynamic changes in parasite infrapopulations were observed, e.g., there were losses of immature worms and rapid changes in infrapopulation sizes, observations that would not be made with typical host-parasite systems.