Helminth community structure of sympatric eastern American toad, Bufo americanus americanus, northern leopard frog, Rana pipiens, and blue-spotted salamander, Ambystoma laterale, from southeastern Wisconsin

One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.     

Helminth infracommunities of Rana vaillanti brocchi (Anura: Ranidae) in Los Tuxtlas, Veracruz, Mexico

A total of 76 adult individuals of Rana vaillanti were collected in Laguna Escondida, Los Tuxtlas, Veracruz, Mexico, and their helminth infracommunity structure was determined. Among the 21 helminth taxa collected (10 digeneans, 8 nematodes, and 3 acanthocephalans), the digenean Langeronia macrocirra reached the highest prevalence (64.4%), mean abundance (6.6), and mean intensity (10.4), as well as the highest total number of individuals (499). Only 2 frogs were uninfected, the remainder harbored between 1 and 7 helminth species and 1-102 individuals; mean species richness and abundance were 3.49 +/- 0.22 and 16.1 +/- 16.3, respectively. Langeronia macrocirra dominated in 50.6% of the infracommunities, with relatively low Berger-Parker index values (0.56); for this reason, the evenness was high (0.70 +/- 0.31), and consequently, diversity values are the highest recorded to date in species of Rana. However, patterns of helminth infracommunity richness and diversity were similar to those previously observed in amphibians. This structure is attributed to the feeding habits (between 66.7 and 81% of helminth species parasitizing R. vaillanti enter using the food web dynamics) and low vagility (the remainder species infect by host penetration).     

Nestedness in colonization-dominated systems: helminth infracommunities of Rana vaillanti Brocchi (Anura: Ranidae) in Los Tuxtlas, Veracruz, Mexico

Colonization probabilities of parasite species often are determined by the habitat preference and vagility of host individuals. Although extinction-based interpretations have been investigated for nested subset patterns of parasite infracommunities, the low relative frequency of nestedness in colonization-dominated systems makes the determination and interpretation of nested infracommunities of broad ecological importance. In these systems, ontogenetic shifts in habitat preference or diet of the host have the potential to produce nested subset patterns of parasite infracommunities. Helminth infracommunity structure was investigated for 76 Rana vaillanti individuals collected from Laguna Escondida, Los Tuxtlas, Veracruz, Mexico, in 1998. Pooled helminth infracommunities were significantly nested, as were penetrating and ingested helminth infracommunities when considered separately. Richness, diversity, and evenness of the helminth infracommunities were not correlated with host size, and did not differ between host sexes, suggesting that the structure of infracommunities simply is a product of the interaction between host individuals and their landscape mediated by individual differences in vagility. It is hypothesized that individual differences in recruitment can produce nested subset infracommunity patterns when the habitats or habitat preferences of hosts are themselves nested.     

Patterns of association, nestedness, and species co-occurrence of helminth parasites in the greater kudu, Tragelaphus strepsiceros, in the Kruger National Park, South Africa, and the Etosha National Park, Namibia

The helminth parasites of the greater kudu from the Kruger National Park (KNP), South Africa, and the Etosha National Park (ENP), Namibia, were examined to determine the major patterns of spatial and demographic variation in community structure and to evaluate nonrandomness in parasite community assembly. Nonmetric multidimensional scaling ordination procedures were used to test for differences in parasite community composition between hosts of the 2 parks and between hosts of different demographic groups within KNP. Infracommunities within KNP were also examined for patterns of nonrandomness using 2 null models, i.e., nestedness and species co-occurrence. Infracommunities of KNP and ENP were significantly different from each other, as were infracommunities of different host demographic groups within KNP. Parasite species in the greater kudu from KNP displayed significant levels of nestedness and were found to co-occur less frequently than expected by chance; however, this lack of co-occurrence was significant only when all demographic groups were considered. When restricted to any particular age class, co-occurrence patterns could not be distinguished from random. Overall, these data suggest that biogeography and host demographics are important factors in determining community organization of helminth parasites in the greater kudu.     

Helminth communities of two green frogs (Rana perezi and Rana saharica) from both shores of the Alboran Sea

The helminth communities of two populations of green frogs from both shores of the Alborón Sea (Western Mediterranean) were studied. Of the 79 frogs examined for helminths, 39 individuals of the species Rano saharica were collected from Bob-Taza (Morocco), and 40 of the species Rona perezi were collected from the Natural Park of the Sierra de Grazalema (Spain). Although the species richness of helminths was identical in the two sampled areas, the differences observed in the structure of the helminth infracommunities were quite important. Statistically, significant differences were found between the species richness and the diversity of the infracommunities of R. perezi female population and the other three studied statistical populations. The helminth component communities of these two green frogs can be considered as depauperate, although their infracommunities present interactive features.     

Helminth communities of herons (Aves: Ardeidae) in southern Italy

The helminth communities of nine species of herons from southern Italy were studied and compared. Of 24 taxa found including seven digeneans, seven nematodes, six cestodes and four acanthocephalans, only five taxa were found in more than one heron species, and five of the 21 taxa that could be identified to species level were classified as ‘heron specialists’. The total number of helminth species per heron species ranged from 1 in Botaurus stellaris to 9 in Ixobrychus minutus with infection levels generally low. A statistical comparison was carried out for herons with a sample size > 5. At the infracommunity level, only I. minutus clearly differed from other heron species. Diversity parameters of heminth infracommunities did not significantly differ among heron species. Species richness ranged from just 0.3 to 2.3 helminth taxa per individual host, and the Brillouin index, from 0 to 0.3. Total helminth abundance did not exceed 40 worms per host except in a single case of Ardeola ralloides. Infracommunities clearly were dominated by single helminth species. The present study confirms a depauperate helminth community in herons from southern Italy. Comparison with data from Spain and the Czech Republic showed strong quantitative similarities with values obtained in the present study. Results also suggest that the composition of local helminth communities are strongly variable depending on geographical location as is demonstrated by comparison with data from other European areas. However, whether herons in Europe naturally host depauperate helminth communities or these communities are depauperate because of other factors is unknown.     

The structure of helminth communities

The existence of both interactive and isolationist communities of helminths, each of which may be produced by any one of several processes, necessitates a pluralistic view of helminth community structure. A scheme involving three hierarchical scales of communities of helminths is proposed. Interactions between species occur in infracommunities, and should be looked for there. Interactions are often clearer in smaller groups of species, either among core species or within guilds. Infracommunities are samples of helminth communities at two larger scales: the helminth community in the host population provides core species, largely specialists, and that in the host community provides the generalists and the satellite species. The richness of each of the two large-scale communities is affected by various ecological, historical and evolutionary factors. The different concepts which have been applied to parasite communities are based on these factors.     

Helminth communities of green frogs Rana clamitans Latreille, from southwestern Michigan.

A total of 239 green frogs Rana clamitans, collected between June 3 and August 27, 1998 from 6 locations in southwestern Michigan, was examined for helminths. Of the 26 helminth taxa found, the larval cestode Mesocestoides sp. had the highest mean intensity, followed by the larval trematode Fibricola sp. Of the helminths that mature in frogs, Haematoloechus varioplexus had the highest prevalence and Gorgodera amplicava had the highest mean intensity. Frogs from 118th Pond had the highest species richness (20), mean helminth species richness (5.2), and mean helminth abundance (153.7). Frogs from Constantine East had the highest mean helminth species diversity (0.8778) and evenness (0.6033), followed by frogs from 118th Pond. In all comparisons of mean helminth community species richness, abundance, diversity, and evenness, adult frogs had significantly higher or higher values than did juveniles at each location. Jaccard's coefficients of similarity for the helminth communities for location pairs ranged from 0.545 to 0.823. Nine and 2 core helminth taxa occurred at the local and regional levels, respectively. The differences in several helminth community measures in green frogs among locations stress the importance of local ecological conditions on helminth community structure.     

Habitat Selection and Movement of Pond-Breeding Amphibians in Experimentally Fragmented Pine Forests

ABSTRACT Population-level responses of amphibians to forest management regimes are partly dictated by individual behavioral responses to habitat alteration. We examined the short-term (i.e., 24-hr) habitat choices and movement patterns of 3 amphibian species—southern leopard frogs (Rana sphenocephala), marbled salamanders (Ambystoma opacum), and southern toads (Bufo terrestris)—released on edges between forest habitats and recent clear-cuts in the Upper Coastal Plain of South Carolina, USA. We predicted that adult frogs and salamanders would preferentially select forest using environmental cues as indicators of habitat suitability. We also predicted that movement patterns would differ in clear-cuts relative to forests, resulting in lower habitat permeability of clear-cuts for some or all of the species. Using fluorescent powder tracking, we determined that marbled salamanders selected habitat at random, southern toads preferred clear-cuts, and southern leopard frogs initially selected clear-cuts but ultimately preferred forests. Frogs exhibited long-distance, directional movement with few turns. In contrast, toads exhibited wandering behavior and salamanders moved relatively short distances before locating cover. Southern toads and southern leopard frogs moved farther in forests, and all 3 species made more turns in clear-cuts than in forests. Habitat selection by southern toads did not vary according to body size, sex, or the environmental cues we measured. However, marbled salamanders were more likely to enter clear-cuts when soil moisture was high, and southern leopard frogs were more likely to enter clear-cuts when relative humidity and air temperature were higher in the clear-cut than in adjacent forest. Although we found evidence of reduced habitat permeability of clear-cuts for southern leopard frogs and southern toads, none of the species exhibited strong behavioral avoidance of the small (4-ha) clear-cuts in our study. Further studies of long-term habitat use and the potential physiological and other costs to individuals in altered forests are needed to understand the effects of forest management on population persistence. To reduce potentially detrimental effects of clear-cutting on amphibians in the Southeast, wildlife managers should consider the vagility and behavior of species of concern, especially in relation to the size of planned harvests adjacent to breeding sites.     

Host-parasite relationships of longnose dace, Rhinichthys cataractae, from the Ford River, Michigan.

A total of 1,115 longnose dace, Rhinichthys cataractae (family Cyprinidae), were examined for parasites from May 1983 through October 1986 from 3 localities in the Ford River in Michigan's Upper Peninsula. Thirteen parasite species (1 Monogenea, 2 Digenea, 2 Cestoda, 4 Nematoda, 1 Acanthocephala, 3 Protozoa) infected dace. The parasite faunas of dace, taxonomically and in species number, were similar between localities. Posthodiplostomum minimum minimum, Neascus sp., and Rhabdochona canadensis were the most common helminths infecting dace from each locality. The first 2 species did not exhibit consistent seasonal infection patterns between years, whereas the prevalence and mean intensity of R. canadensis in dace from the downriver locality were higher in summer 1983, 1984, and 1985. The intensity of infection of each of these helminth species significantly increased with host length. The prevalences and mean intensities of P. m. minimum, Neascus sp., and R. canadensis as well as the helminth infracommunity diversity were highest in dace from the upriver locality. The major factors that influenced parasite intensity were environmental factors that occurred when and where a fish began its life, the sequence of events that occurred in each habitat the fish encountered during its life, and the length of exposure (age of fish). Dace have isolationist helminth infracommunities arising from factors including ectothermy, a simple enteric system, restricted vagility, and being gape-limited. Allogenic helminths with indirect life cycles predominate in the depauperate helminth fauna of dace.     

The helminth community of the wood mouse, Apodemus sylvaticus, in the Sierra Espuna, Murcia, Spain

The helminth community of the wood mouse, Apodemus sylvaticus, in the Sierra Espuna was characterized after a complete analysis of its helminth community component and infracommunity structure relative to host age, sex and year of capture. The helminth community comprised 13 species: one trematode, four cestodes and eight nematodes. The cestode Pseudocatenotaenia matovi and the nematode Syphacia frederici were the most prevalent and abundant helminth species, respectively. Sixty four percent of mice analysed presented helminths with a direct cycle and 42% presented helminths with an indirect cycle. The helminth community presents a low diversity with infracommunities usually made up of only one or two helminth species. Host age and year of capture seem to play a major role in determining species richness and helminth diversity, but not in determining the abundance of helminths. Host sex does not seem to affect the infection rate nor the diversity. Further studies on more samples of wood mice and other small mammal species in this regional park are needed to explore any possible interactions between helminth communities in the host populations.     

Helminth communities of two sympatric skinks (Mabuya agilis and Mabuya macrorhyncha) from two "restinga" habitats in southeastern Brazil

The helminth fauna of two sympatric congeneric skinks (Mabuya agilis and M. macrorhyncha) from two distinct "restinga" habitats (Praia das Neves and Grussaí) in southeastern Brazil were studied, totalling four data sets (sample sizes ranging from 11 to 28). A total of ten helminth species were associated with the skinks: Raillietiella sp., Paradistomum parvissimum, Pulchrosomoides elegans, Oochoristica ameivae, Hexametra boddaertii, Parapharyngodon sceleratus, Physalopteroides venancioi, Physaloptera sp., an unidentified acuariid nematode and an unidentified centrorhynchid acanthocephalan. Except for Hexametra boddaertii (found only in Grussaí) and Pulchrosomoides elegans (found only in Praia das Neves), all helminth species were present at both localities. Half of the helminth species were present only as larvae and, in most cases, appear to represent paratenic parasitism. Overall prevalences of infection were high for both host species in both localities. Mabuya agilis tended to have richer and more diverse infracommunities than M. macrorhyncha. Some parameters of infection by individual helminth species seem to be related to the ecology of each Mabuya species. The parasite faunas were qualitatively very similar among species and/or localities, but quantitative similarities were more varied, due to differential representativeness of individual helminth species among host populations. The helminth communities of both skink species can be classified as non-interactive, being composed of site-specialists and immature stages of non-lizard parasites.     

Coexistence of helminth species in Lysapsus limellus (Anura: Pseudidae) from an Argentinean subtropical area: influence of biotic and abiotic factors

The need for studies on helminth communities of South American amphibians was addressed by examining changes in composition and population dynamics of the helminth component and infracommunities in the frog Lysapsus limellus Cope, 1862, from 1994 to 1996. Two pond types were considered, one permanent and one semipermanent. The main goals of this study were (1) to investigate the relationships between pond type, season, study time, frog body size, and frog sex and the presence-absence and counts of helminth parasite species in the frog host, L. limellus, and (2) to examine the co-occurrence of the different parasite species in the frog host in terms of the structure, assembly, and dynamics of the helminth infracommunity. Parasite presence and absence were analyzed using logistic regression and parasite counts were analyzed using log-linear modeling. To examine the association between parasite species, a principal components analysis was carried out on the correlation matrix for the counts. The pattern of co-occurrences was also examined through a randomization test. The main results were as follows: (1) five parasite species were found in the infracommunity; (2) the pond type and the host size class were the principal factors related to the presence and absence of parasites for the three dominant species, while the year of study and the host sex were only important for one of the species; (3) for the parasite counts, many more factors were significant, with body size class and pond type always important factors for the three core species of the infracommunity, with season, year of study, and sex also sometimes important, and (4) strong associations were observed between some helminth species of the host from the permanent pond, but the same species did not co-occur in frogs in the semipermanent pond. The life histories of the five species can be considered as opportunistic or "r strategists."     

Patterns in metazoan parasite communities of some oyster species

Metazoan parasite communities of Crassostrea gigas and Ostrea edulis from Great Britain, Crassostrea virginica from Mexico, and Saccostrea commercialis from Australia are described and summarized in terms of species composition, species richness, total number of individuals and dominance. Metazoan parasite communities in all host species were composed of turbellarians and the metacercarial stage of digeneans, with the exception of S. commercialis where only metacercariae were found. Arthropods, including one copepod and one mite species, were present only in British oyster species. All metazoan parasite communities of oysters had few species and low density of individuals. Richest communities were found in C. virginica at both component and infracommunity level. The least diverse component community occurred in S. commercialis. Infracommunities in O. edulis and S. commercialis never exceeded one species per host. The host response against parasites is suggested as the principal factor responsible for depauperate parasite communities of oysters. Environmental factors characteristic of tropical latitudes are likely to have enhanced both the number of species and the densities of parasites per host in the infracommunities of C. virginica.     

The helminth infracommunities in three species of herons (Aves: Ardeidae) from Ukraine

This survey analysed the helminth communities associated with three Ukrainian heron species (Ardea cinerea, Ardea purpurea, Nycticorax nycticorax) collected throughout 1947–2019. The sample analysed included 128 specimens, and 55 helminth species were identified. Of these, the most diverse taxonomic group was trematodes with 29 species, followed by cestodes with 14 species, nematodes with nine, and acanthocephalans with three species. The helminth infracommunities were found to be similar across all three host species. Furthermore, the distance to the coastal regions impacts both diversity and species richness in infracommunities.     

Similarity in parasite communities of the teleost fish Pinguipes brasilianus in the southwestern Atlantic: Infracommunities as a tool to detect geographical patterns

Patterns of distance decay in similarity among communities of the fish Pinguipes brasilianus (Teleostei: Pinguipedidae) from five areas in the southwestern Atlantic were investigated to determine whether the rate of decay varied depending on the community level or the parasite guild analyzed (ectoparasites, adult endoparasites and larval endoparasites). Similarities in species composition were computed at both the component community and infracommunity levels. Similarity indices were calculated between all possible pairs of assemblages from different zones. Infracommunity similarity values between and within host populations were averaged. Significance of linear regressions for similarity values against distance was assessed using randomization tests. Different patterns were observed for each guild, and similarity among infracommunities within host populations varied accordingly. Decay in similarity over distance was recorded for most communities. The slopes differed significantly between infracommunities and component communities in all cases, and stronger decay was always observed for infracommunities. Different geographical patterns in parasite communities were a consequence of variability in parasite availability in the different regions, modulated by oceanographic conditions, as well as variation among species in terms of host specificity and life-cycles strategies. Infracommunities showed a stronger effect of distance than component communities, probably due to the influence of short term and local variability of oceanographic conditions.     

Helminth infracommunities of the frogs Rana catesbeiana and Rana clamitans from Turkey Marsh, Michigan

One hundred twenty-seven bullfrogs, Rana catesbeiana, and 120 green frogs, Rana clamitans, collected in July and August 1987 from Turkey Marsh, Michigan, were examined for helminths. Of the 16 helminth species found, Falcaustra catesbeianae had the highest prevalence and mean intensity in bullfrogs. In green frogs, Megalodiscus temperatus had the highest prevalence and Haematoloechus parviplexus had the highest mean intensity. Taxonomically, 1,030 (30.5%) trematodes and 2,348 (69.5%) nematodes occurred in bullfrogs; 2,874 (96.4%) trematodes, 2 (0.1%) cestodes, and 105 (3.5%) nematodes were found in green frogs. The larger and more aquatic of the 2 hosts, R. catesbeiana, had the highest mean number of helminth species, helminth intensity, diversity, and evenness. Adult frogs of both species had a significantly higher mean number of helminth species than did juveniles. Significant correlations existed between the number of helminth species and frog length. Although the helminth communities of Turkey Marsh frogs are isolationist in character, they are more diverse than other helminth communities studied in amphibians.     

Helminth community structure in an expanding white-winged dove (Zenaida asiatica asiatica) population

Helminth communities of 171 fledged white-winged doves (Zenaida asiatica asiatica) from the expanding eastern population in Texas (USA) were examined from hosts collected 11 June to 19 September 1997 within their historical range, new breeding periphery, and an intermediate area. Eleven helminth species, representing 435 individuals, were found. Helminths occurred in three microhabitats, of which the small intestine was the most commonly occupied. Nematodes dominated numerically (76% of total worms), followed by cestodes (17%), and trematodes (7%). Infracommunities were species-poor; the most complex infracommunity consisted of three helminth species, which occurred in three host individuals, followed by two species that occurred in 13 hosts. The remaining 155 doves had one (70) or no (85) species. The overall helminth component community was species-poor and was dominated by Ascaridia columbae which occurred in 26% of the white-winged doves and accounted for 65% of all helminth individuals. Prevalence and abundance of A. columbae varied by geographic region and host age, but not by host sex. Helminth component communities varied by geographic region, host age, and host sex. These differences were primarily attributable to unique occurrences of uncommon species within specific host subpopulations. Results suggest that the white-winged doves' multimodal regional abundance pattern, sympatry with other columbids, and granivorous diet may be more important in shaping helminth community structure than the influences often associated with geographic range expansion.     

Richness,nestedness, and randomness in parasite infracommunity structure

Within a host population, parasite infracommunities vary in both richness and species composition. If interspecific interactions among parasites are important in shaping infracommunities, the structure of these assemblages is expected to differ from the one predicted by null models, i.e. from the one that would result from chance alone. Using data from the literature, I tested for discrepancies between observed and random patterns in the richness and composition of gastrointestinal helminth infracommunities of birds and mammals. Both the Poisson distribution and a more sophisticated null model, derived from prevalence of the different parasite species in the host population, usually provided a good fit to the observed distributions of infracommunity richness among hosts. This suggests that parasite species do not co-occur more or less frequently than expected by chance. In mammals, the co-occurrence of all available parasite species in the same host individual, or maximum potential infracommunity richness, was less likely to be observed when several parasite species were available; this is also a phenomenon expected from the random assembly of parasite species. Finally, there was no evidence for a nested subset pattern among parasite species in a host population: rate species were distributed independently of common ones. The overall picture emerging from these results is one in which parasite assemblages are more likely to be the product of random events than of predictable and repeatable processes.     

The helminth community of the skink Chalcides sexlineatus from Gran Canaria (Canary Islands)

A survey of the gastrointestinal helminth communities of a population of Chalcides sexlineatus Steindachner, a small skink endemic to Gran Canaria island (Canary Archipelago, Spain), was conducted to determine the prevalence, abundance and species diversity of intestinal parasites in these reptiles. Only three parasite species were found, one cestode, Oochoristica agamae Baylis, 1919 and two nematodes, Parapharyngodon micipsae (Seurat, 1917) and Pharyngodonidae gen. sp. Helminth infracommunities of C. sexlineatus showed low values of abundance and species richness and diversity, being more similar to the helminth community of Tarentola boettgeri boettgeri (Steindachner) rather than those of Gallotia stehlini (Schenkel), both syntopic with the sampled host.