Functional richness, functional evenness, and use of niche space in parasite communities

Most efforts aimed at elucidating the factors responsible for the variation in species richness among different parasite communities have focused on host characteristics such as body mass or diet. Independently of host features, however, the way in which parasites use resources within the host may also affect the species richness of the community. The distribution of parasite individuals or biomass in niche space determines whether host resources are used evenly, or whether there are gaps in resource use, with some parts of the total niche being underutilized. Here, the concepts of functional richness and functional evenness are applied to parasite communities for the first time, using simple indices. Measurements of the distribution of species in niche space within communities, such as mapping the distribution of helminths along the length of the host's intestine, is standard practice in parasitology. In such cases, functional richness is simply the proportion of the total number of intestine sections available that are used by at least 1 worm, whereas functional evenness measures the evenness in the distribution of worm numbers or biomass across all niche sections that are occupied. Data on cestode communities of elasmobranchs are used to illustrate the use of these indices, and to show how important they can be in tests of ecological hypotheses. The indices presented here capture essential features of resource use in parasite communities, and can be useful tools for comparative analyses.     

Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.     

Body size and coexistence in gamasid mites parasitic on small mammals: null model analyses at three hierarchical scales

We studied body size ratio in gamasid mites (Acari: Mesostigmata) parasitic on Palearctic small mammals at 3 hierarchical scales, namely infracommunities (an assemblage of mites harboured by an individual host), component communities (an assemblage of mites harboured by a host population), and compound communities (an assemblage of mites harboured by a host community). We used null models and asked a) whether body size distributions in these communities demonstrate non‐random patterns; b) whether these patterns indicate segregation or aggregation of body sizes of coexisting species; and c) whether patterns of body size distribution are scale‐dependent, that is, differ among infracommunities, component communities, and compound communities. In most mite assemblages, the observed pattern of body size distribution did not differ from that expected by chance. However, meta‐analyses demonstrated that component and compound communities of gamasid mites consistently demonstrated a tendency to reduced body size overlap, while we did not find any clear trend in mite body size distribution across infracommunities. We discuss reasons for scale‐dependence of body size distribution pattern in parasite communities and propose ecological and evolutionary mechanisms that allowed the reduced body size overlap in component and compound communities of ectoparasites to arise.     

Latitudinal gradients of parasite richness: a review and new insights from helminths of cricetid rodents

The latitudinal diversity gradient (LDG), or the trend of higher species richness at lower latitudes, has been well documented in multiple groups of free‐living organisms. Investigations of the LDG in parasitic organisms are comparatively scarce. Here, I investigated latitudinal patterns of parasite diversity by reviewing published studies and by conducting a novel investigation of the LDG of helminths (parasitic nematodes, trematodes and cestodes) of cricetid rodents (Rodentia: Cricetidae). Using host–parasite records from 175 parasite communities and 60 host species, I tested for the presence and direction of a latitudinal pattern of helminth richness. Additionally, I examined four abiotic factors (mean annual temperature, annual precipitation, annual temperature range and annual precipitation range) and two biotic variables (host body mass and host diet) as potential correlates of parasite richness. The analyses were performed with and without phylogenetic comparative methods, as necessary. In this system, helminths followed the traditional LDG, with increasing species richness with decreasing latitude. Nematode richness appeared to drive this pattern, as cestodes and trematodes exhibited a reverse LDG and no latitudinal pattern, respectively. Overall helminth richness and nematode richness were higher in areas with higher mean annual temperatures, annual precipitation and annual precipitation ranges and lower annual temperature ranges, characteristics that often typify lower latitudes. Cestode richness was higher in areas of lower mean annual temperatures, annual precipitation and annual precipitation ranges and higher annual temperature ranges, while trematode richness showed no relationship with climate variables when phylogenetic comparative methods were used. Host diet was significantly correlated with cestode and trematode species richness, while host body mass was significantly correlated with nematode species richness. Results of this study support a complex association between parasite richness and latitude, and indicate that researchers should carefully consider other factors when trying to understand diversity gradients in parasitic organisms.     

Phylogenetic overdispersion in Floridian oak communities

Closely related species that occur together in communities and experience similar environmental conditions are likely to share phenotypic traits because of the process of environmental filtering. At the same time, species that are too similar are unlikely to co-occur because of competitive exclusion. In an effort to explain the coexistence of 17 oak species within forest communities in North Central Florida, we examined correlations between the phylogenetic relatedness of oak species, their degree of co-occurrence within communities and niche overlap across environmental gradients, and their similarity in ecophysiological and life-history traits. We show that the oaks are phylogenetically overdispersed because co-occurring species are more distantly related than expected by chance, and oaks within the same clade show less niche overlap than expected. Hence, communities are more likely to include members of both the red oak and the white + live oak clades than only members of one clade. This pattern of phylogenetic overdispersion arises because traits important for habitat specialization show evolutionary convergence. We hypothesize further that certain conserved traits permit coexistence of distantly related congeners. These results provide an explanation for how oak diversity is maintained at the community level in North Central Florida.     

Echinobothrium harfordi sp. nov. (Cestoda: Diphyllidae) from Raja naevus in the North Sea and English Channel.

Echinobothrium harfordi sp. nov. from the spiral intestine of Raja naevus in British waters is described and illustrated. The species most closely resembles E. affine as both possess two groups of eleven large apical hooks, but can be distinguished by fewer peduncle spines and by characteristics of the eggs. The intensity of infection in individual hosts was generally low, and although the incidence of infection was high in rays less than 35 cm in length, this fell towards zero in larger rays. The cestode showed a marked affinity for the first two tiers of the spiral intestine.     

Pollination niche overlap between a parasitic plant and its host

Niche theory predicts that species which share resources should evolve strategies to minimise competition for those resources, or the less competitive species would be extirpated. Some plant species are constrained to co-occur, for example parasitic plants and their hosts, and may overlap in their pollination niche if they flower at the same time and attract the same pollinators. Using field observations and experiments between 1996 and 2006, we tested a series of hypotheses regarding pollination niche overlap between a specialist parasitic plant Orobanche elatior (Orobanchaceae) and its host Centaurea scabiosa (Asteraceae). These species flower more or less at the same time, with some year-to-year variation. The host is pollinated by a diverse range of insects, which vary in their effectiveness, whilst the parasite is pollinated by a single species of bumblebee, Bombus pascuorum, which is also an effective pollinator of the host plant. The two species therefore have partially overlapping pollination niches. These niches are not finely subdivided by differential pollen placement, or by diurnal segregation of the niches. We therefore found no evidence of character displacement within the pollination niches of these species, possibly because pollinators are not a limiting resource for these plants. Direct observation of pollinator movements, coupled with experimental manipulations of host plant inflorescence density, showed that Bombus pascuorum only rarely moves between inflorescences of the host and the parasite and therefore the presence of one plant is unlikely to be facilitating pollination in the other. This is the first detailed examination of pollination niche overlap in a plant parasite system and we suggest avenues for future research in relation to pollination and other shared interactions between parasitic plants and their hosts.     

Morphology and coexistence of congeneric ectoparasite species: reinforcement of reproductive isolation?

Assuming that differences or similarities in morphology among congeneric parasite species living in the same habitat are not a random pattern, several hypotheses explaining morphological differences were tested: (i) reproductive isolation, (ii) niche restriction resulting from competition, and (iii) niche specialization. Congeneric monogenean (platyhelminth) ectoparasites parasitizing the gills of one host species were used as an ecological model. Morphometric distances of the attachment organ and morphometric distances of the copulatory organ between species pairs were calculated, Levin's niche size and Renkonen niche overlap indices were applied. Our results support the prediction that the function of niche segregation is to achieve reproductive isolation of related species in order to prevent hybridization (reinforcement of reproductive barriers). Parasite species living in the same niche differ greatly in the size of copulatory organ. Moreover, species coexistence is facilitated by an increase in morphometric distances of copulatory organ and niche centre distances. Our results also show that species living in overlapping niches have similar attachment organs, which supports the prediction that morphologically similar species have the same ecological requirements within one host and suggests small effects of interspecific competition for the evolution of morphological diversity of attachment organs. Specialist adaptations also seem to facilitate species coexistence and affect the niche distribution within host species. Parasite species that can colonize more than one host species, i.e. generalists, occupy more distant niches within host species than strictly host-specific parasites. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76, 125–135.     

Coexistence theory in the Cape Floristic Region: revisiting an example of leaf niches in the Proteaceae

Competition in plant communities is often a contentious issue because the mechanisms of competitive interactions are not obvious. We sought evidence that Proteaceae communities are competing along two leaf niche axes as observed in a previous study. Two functional characters, leaf size and leaf shape were measured on numerous individuals per species per plot of six communities from two different regions. Patterns of overdispersion along these leaf trait axes between species were observed, similar to the earlier study. The observed results were compared with the patterns expected under a null model using standard and novel indices to test the significance of trait dispersion between species within a plot. Competition and niche differentiation in the observed plots were not supported as the observed trait overlaps were not significantly different from the null expectation. Our results do not support the theory that Proteaceae communities compete along the two proposed functional leaf traits.     

Structure of the parasite communities of a coral reef fish assemblage (Labridae): testing ecological and phylogenetic host factors

The role of ecological and phylogenetic processes is fundamental to understanding how parasite communities are structured. However, for coral reef fishes, such information is almost nonexistent. In this study, we analyzed the structure of the parasite communities based on composition, richness, abundance, and biovolume of ecto- and endoparasites of 14 wrasse species (Labridae) from Lizard Island, Great Barrier Reef, Australia. We determine whether the structure of the parasite communities from these fishes was related to ecological characteristics (body size, abundance, swimming ability, and diet) and/or the phylogenetic relatedness of the hosts. We examined 264 fishes from which almost 37,000 individual parasites and 98 parasite categories (types and species) were recorded. Gnathiid and cestode larvae were the most prevalent and abundant parasites in most fishes. Mean richness, abundance, and biovolume of ectoparasites per fish species were positively correlated with host body size only after controlling for the host phylogeny, whereas no such correlation was found for endoparasites with any host variable. Because most ectoparasites have direct transmission, one possible explanation for this pattern is that increased space (host body size) may increase the colonization and recruitment of ectoparasites. However, endoparasites generally have indirect transmission that can be affected by many other variables, such as number of prey infected and rate of parasite transmission.     

No evidence of strong host resource segregation by phorid parasitoids of leaf-cutting ants

Resource segregation by species is a cornerstone ecological concept that may result from several processes such as interspecific competition, and can help structuring communities, in particular parasitoid communities. Phorid parasitoid flies that use ants as hosts usually employ one host per individual parasitoid, and thus the pressure for segregating the host resource should be high. At a particular community, these parasitoids might segregate resources by temporal differences in activity patterns, using different host species or nests from those available. Even if parasitoid species coexist on the same nest, they can take advantage of worker polymorphism and task division, searching for ants performing different tasks at different microsites of the same nest. Here we evaluated the segregation of parasitoid species in these hypothesized axes using leaf-cutting ant phorid parasitoids as a model system. We analyzed temporal data collected at two localities with contrasting host species richness; and compared parasitoid co-occurrence at the different niche axis. For most of the hypothesized niche axes tested we found either no departures from random expectations or significantly more niche overlap than expected by chance, ruling out the existence of biologically relevant host resource segregation in this system. However, there was evidence of segregation for some species, since one parasitoid species was only found in winter and another species showed a negative correlation of its abundance over nests with other two species. Furthermore, we found that several species were flexible in host use; Atta phorids varied in average host sizes preferred, whereas Acromyrmex phorids that were generalists were able to use different host species or microsites for host location. From an applied perspective, these results are encouraging when selecting species for the control of leaf-cutting ants because parasitoids coexistence seems to be unaffected by their overlap in niche dimensions.     

Niche shifts in Greater Antillean Anolis communities: effects of niche metric and biological resolution on null model tests

Summary Using published data, a Monte Carlo approach was used to determine if niche shifts in communities of Anolis lizards have occurred within two islands in the Greater Antilles. Communities at 4 sites on Puerto Rico and 4 sites on Jamaica were randomly assembled using resource-use data from any of the 4 sites with the constraint that the species list of the random community was identical to that observed for each site. The observed niche structure and that of the randomly assembled community were compared at each site. The null hypothesis tested was that one of several functions of niche overlap did not produce lower values in the observed communities compared to those randomly assembled. The sensitivity of this method to details of the analysis was investigated by using 5 variations on the procedure for quantifying the niche structure of the communities. I investigated the effects of (1) using Pianka's versus Levins' metric of niche overlap, (2) using complete resource-use data versus the marginals of the distributions, (3) asymmetric responses to overlap due to size differences between overlapping species, (4) using estimated electivities versus the observed proportional use of resource states, and (5) methods of niche analysis based on different degrees of biological resolution (e.g., community-wide statistics versus individual species responses). The results of the simulations and statistical comparisons did not provide a clear, unambiguous answer to the question of the occurrence of niche shifts. Statistical evidence for niche shifts was more frequently detected using Levins' metric compared to Pianka's, and shifts were easier to detect using high biological resolution (examination of individual species) compared to community-wide measures of niche structure response.     

Infracommunity structure of parasites of Hemigymnus melapterus (Pisces: Labridae) from Lizard Island, Australia: the importance of habitat and parasite body size

This study describes the community of all metazoan parasites from 14 individuals of thicklip wrasse, Hemigymnus melapterus, from Lizard Island, Australia. All fish were parasitized, and 4,649 parasite individuals were found. Twenty-six parasite species were identified although only 6 species were abundant and prevalent: gnathiid isopods, the copepod Hatschekia hemigymni, the digenean Callohelmis pichelinae, and 3 morphotypes of tetraphyllidean cestode larvae. We analyzed whether the body size and microhabitat of the parasites and size of the host affected understanding of the structure of the parasite community. We related the abundance, biovolume, and density of parasites with the host body size and analyzed the abundances and volumetric densities of some parasite species within microhabitats. Although the 2 most abundant species comprised 75% of all parasite individuals, 4 species, each in similar proportion, comprised 85% of the total biovolume. Although larger host individuals had higher richness, abundance, and biovolume of parasites than smaller individuals, overall parasite volumetric density actually decreased with the host body size. Moreover, parasites exhibited abundances and densities significantly different among microhabitats; some parasite species depended on the area available, whereas others selected a specific microhabitat. Parasite and habitat size exhibited interesting relationships that should be considered more frequently. Considerations of these parameters improve understanding of parasite community structure and how the parasites use their habitats.     

Community structure of helminth parasites of the tuna, Euthynnus affinis, from the Visakhapatnam coast, Bay of Bengal

An analysis is made of the community structure of the helminth parasites of the tuna Euthynnus affinis collected off the coast of Visakhapatnam, Bay of Bengal. The helminth fauna comprised 23 species, consisting of two monogeneans, 18 digeneans, one larval cestode, one nematode and one acanthocephalan. Didymozoids, represented by 14 species, were the dominant members of the parasite spectrum. The parasite assemblage is characterized by species richness as well as a high diversity. Most of the parasites found, the didymozoids in particular, were host specialists, exhibiting a high degree of host specificity. The fauna comprised three core species, four satellite species and many secondary species, with no potential for interaction among them but the degree of predictability is high in that each infracommunity comprised two or three core species superimposed by a few secondary species. Many host factors such as the varied diet, high vagility, long life span and endothermy appear to have contributed to the development of the species rich and diverse parasite communities in the tuna. There was evidence for a decrease in parasite density and an increase in diversity with increase in host size, indicating that host size has a profound influence on the component community structure.     

Bryophytes and ecological niche theory

Classical niche theory, particularly in terms of competitive exclusion, does not appear to apply as well to bryophytes as to other organisms. Bryophyte communities, as well as those of other plants and of animals, can be thought of in terms of individual species each utilizing particular portions of various resource or habitat continua. Quantitative studies carried out since 1981, particularly those involving niche breadth and overlap, are reviewed. Special attention is given to niche diversification in Sphagnum , Splachnaceae, bryophyte communities in streams, and to ephemeral bryophyte communities. Some bryophyte communities appear to have equilibrium characteristics and to contain species with relatively narrow niche breadths and with no or only partial niche overlap. In many habitats, however, bryophyte communites have non-equilibrium characteristics and diversification of species in microhabitats is opportunistic. Do any bryophyte communities persist long enough for complete saturation by species which have realized niches determined by competitive interactions? Recent studies indicate that this is the case for at least some Sphagnum communities, but that it is the exception not the rule for bryophytes.     

Cestodiasis in the red grouse in Scotland

Little is known about the cestode parasites of red grouse Lagopus lagopus scoticus. The carcasses of 71 free-living and grouse collected from northeastern Scotland (UK) between June 1991 and September 1992 were examined for the cestode Paroniella urogalli. Over 70% of young and old birds were infected in summer and winter. Cestode abundance ranged from 0 to 29 (mean +/- SE = 5.54 +/- 0.81) worms per host and did not differ significantly between young and old grouse, or between seasons. Cestode intensities did vary significantly between the sexes, but in opposite directions at two sites. A significantly greater total biomass (dry weight) of cestodes was found in grouse in the summer. Cestode biomass also was greater in young than in old grouse, although there was no significant difference after controlling for the effects of site, year and season. However, in the summer total cestode biomass in young grouse was significantly greater per unit bodyweight than in old grouse. No evidence was found for an effect of cestodiasis on grouse condition but a significant positive correlation existed between grouse comb condition and cestode abundance. Estimates of mean per capita worm mass suggested an intensity dependent reduction in the weight of individual cestodes in birds in their first summer. Per capita cestode mass was significantly lower in winter than summer, which may suggest that the production of mature proglottids varies with season. There was also a significant negative correlation between numbers of the nematode Trichostrongylus tenuis and the total biomass of P. urogalli in grouse.     

Parasite-associated growth enhancement in a fish-cestode system

Parasites impose an energetic cost upon their hosts, yet, paradoxically instances have been reported in which infection is associated with enhanced, rather than diminished, host growth rates. Field studies of these parasite effects are problematic, since the pre-infection condition of the hosts is generally unknown. Here, we describe a laboratory experiment in which the growth rate and body condition of 76 laboratory-reared three-spined stickleback fishes were examined before, during and after each fish was fed the infective stage of the parasitic cestode Schistocephalus solidus. Twenty-one of these fishes went on to become infected by the cestode. Fishes were individually housed and provided with an abundant food supply to eliminate the potentially masking effects of variable competitive ability. Infection occurred independently of fish gender, size, body condition or pre-exposure growth rate. After exposure to the cestode, infected fishes grew faster (excluding parasite weight) and maintained a similar or better body condition compared with uninfected fishes, despite developing enlarged spleens. The accelerated growth could not be explained by reduced gonadal development. This result, one of few demonstrations of parasite-associated growth enhancement in fishes, is discussed with respect to other such parasite systems.     

Big plants—do they limit species coexistence?

Aims According to conventional theory, larger plant species are likely to inflict more intense competition on other (smaller) species. We tested a deducible prediction from this: that a larger species should generally be expected to impose greater limits on the number of species that can coexist with it.Methods Species richness was sampled under plant canopies for a selection of woody species ('host' species) that display a wide range of adult sizes (from small shrubs to large trees), growing within natural vegetation of the Interior Douglas-fir zone of southern British Columbia, Canada. These data were compared with species richness levels sampled within randomly placed plots within the host species habitat.Important findings A prominent host species size effect on species richness was detected but only narrowly at the small end of the species size range. Across most (90%) of the increasing size range of host species, the number of species residing under the host canopy showed no significant decrease relative to the number expected by random assembly, based on species richness within randomly defined equivalent areas within the habitat of the host species. This apparent 'null effect', we suggest, is explained not because these larger species have no effect on community assembly. We postulate that larger species are indeed likely to be more effective in causing competitive exclusion of some smaller species (as expected from conventional theory), but that any potential limitation effect of this on resident species richness is offset for two reasons: (i) larger species also generate niche spaces that they cannot exploit under their own canopies and so have minimal impact (as competitors) on smaller species that can occupy these niches and (ii) certain other small species—despite small size—have effective competitive abilities under the severe competition that occurs within host neighbourhoods of larger species. These and other recent studies call for re-evaluation of traditional views on the role of plant size in affecting competitive ability and community assembly.     

Spiral valve parasites of blue and common thresher sharks as indicators of shark feeding behaviour and ecology

This study documented the parasite faunas of the spiral valves of blue sharks Prionace glauca (L. 1758) and common thresher sharks Alopias vulpinus (Bonnaterre, 1788) caught in the California Current Large Marine Ecosystem (CCLME) north of the Mexican border. The spiral valves of 18 blue and 19 thresher sharks caught in the CCLME from 2009 to 2013 were examined for parasites. Seven parasite taxa were found in blue sharks and nine in threshers. The tetraphyllidean cestode Anthobothrium sp. (78% prevalence) was the most common parasite in blue sharks, and the phyllobothriid cestode Paraorygmatobothrium sp. (90% prevalence) was the most common in threshers. An adult nematode of the genus Piscicapillaria was found in threshers for the first time and may be a new species. Adult individuals of Hysterothylacium sp. were found in both shark species. The adult acanthocephalan Rhadinorhynchus cololabis and remains of the parasitic copepod Pennella sp. – both parasites of Pacific saury, Cololabis saira – were found in the intestines of threshers, indicating recent feeding on saury. This study paves the way for a more comprehensive examination, including more samples and a wider variety of shark species, to provide a greater understanding of shark feeding behaviour and possibly provide information on shark population biology.     

Determinants and consequences of interspecific body size variation in tetraphyllidean tapeworms

Tetraphyllidean cestodes are cosmopolitan, remarkably host specific, and form the most speciose and diverse group of helminths infecting elasmobranchs (sharks, skates and rays). They show substantial interspecific variation in a variety of morphological traits, including body size. Tetraphyllideans represent therefore, an ideal group in which to examine the relationship between parasite body size and abundance. The individual and combined effects of host size, environmental temperature, host habitat, host environment, host physiology, and host type (all likely correlates of parasite body size) on parasite length were assessed using general linear model analyses using data from 515 tetraphyllidean cestode species (182 species were included in analyses). The relationships between tetraphyllidean cestode length and intensity and abundance of infection were assessed using simple linear regression analyses. Due to the contrasting morphologies between shark and batoid hosts, and contrasting physiologies between sharks of the Lamnidae family and other sharks, analyses were repeated in different subsets based on host morphology and physiologies (“sharks” vs. batoids) to determine the influence of these variables on adult tetraphyllidean tapeworm body size. Results presented herein indicate that host body size, environmental temperature and host habitat are relatively important variables in models explaining interspecific variations in tetraphyllidean tapeworm length. In addition, a negative relationship between tetraphyllidean body size and intensity of infection was apparent. These results suggest that space constraints and ambient temperature, via their effects on metabolism and growth, determine adult tetraphyllidean cestode size. Consequently, a trade-off between size and numbers is possibly imposed by external forces influencing host size, hence limiting physical space or other resources available to the parasites.