Co‐evolutionary patterns in congeneric monogeneans: a review of Dactylogyrus species and their cyprinid hosts

Patterns associated with the evolution of parasite diversity, speciation and diversification were analysed using Dactylogyrus species (gill monogeneans) and their cyprinid hosts as a model. The aim of this study was to use this highly specific host–parasite systems to review: (1) the diversity and distribution of Dactylogyrus species, (2) the patterns of organization and structure of Dactylogyrus communities, (3) the evolution and determinants of host specificity and (4) the mode of Dactylogyrus speciation and co‐evolutionary patterns in this Dactylogyrus–cyprinid systems. Dactylogyrus are a highly diverse group of parasites, with their biogeography and distribution clearly linked to the evolutionary history of their cyprinid hosts. The coexistence of several Dactylogyrus species on one host is facilitated by increasing niche distances and the differing morphology of their reproductive organs. The positive interspecific and intraspecific interactions seem to be the most important factors determining the structure of Dactylogyrus communities. Host specificity is partially constrained by parasite phylogeny. Being a strict specialist is an ancestral character for Dactylogyrus, being the intermediate specialists or generalists are the derived characters. The evolution of attachment organ morphology is associated with both parasite phylogeny and host specificity. Considering larger and long‐lived hosts or hosts with several ecological characters as the measures of resource predictability, specialists with larger anchors occurred on larger or longer‐living fish species. Intra‐host speciation, a mode of speciation not often recorded in parasites, was observed in Dactylogyrus infecting sympatric cyprinids. Sister parasite species coexisting on the same host occupied niches that differed in at least one niche variable. Intra‐host speciation, however, was not observed in Dactylogyrus species of congeneric hosts from geographically isolated areas, which suggested association by descent and host‐switching events.     

Trophic morphology features allow Astyanax endemic species coexistence in a Neotropical river system

Resource partitioning has an essential role in interspecific relations, especially in congener species, which share many morphological traits. In some places, small characids coexist through resource partitioning, which may reduce their interspecific competition. Astyanax species (e.g., Astyanax minor, Astyanax gymnodontus and Astyanax bifasciatus), for example, coexist in different water bodies from the Iguaçu River basin. These species have high phenotypic plasticity and many morphologic specializations that allow them to live in different habitats. Based on evidences that these species modified their feeding habits because of changes in resource availability in Iguaçu River, this study tested two hypotheses: (a) there are differences in head morphology, number of teeth and number of gill rakers among the species of Astyanax; and (b) there are differences in gill arch and gill raker morphology among the species of Astyanax, which may favour their trophic resource partitioning in Iguaçu River. The head morphological traits and quantitative morphological characters were summarized in a principal coordinates analysis (PCoA), and the analysis of similarities (ANOSIM) showed significant differences among species. Gill morphological measurements were analysed through analysis of covariance (ANCOVA), and it also showed significant differences in gill arch and gill raker morphology among species. Therefore, the analysis of ecomorphological traits related to trophic habits revealed some differences that may suggest a tendency of reducing competition for trophic resources in the Iguaçu River basin.     

Cophylogenetic relationships between Dactylogyrus (Monogenea) ectoparasites and endemic cyprinoids of the north-eastern European peri-Mediterranean region

The study of host–parasite coevolution is one of the cornerstones of evolutionary biology. The majority of fish ectoparasites belonging to the genus Dactylogyrus (Monogenea) exhibit a high degree of host specificity. Therefore, it is expected that their evolutionary history is primarily linked with the evolutionary history of their cyprinoid fish hosts and the historical formation of the landmasses. In the present study, we used a cophylogenetic approach to investigate coevolutionary relationships between endemic Cyprinoidea (Cyprinidae and Leuciscidae) from selected regions in southern Europe and their respective Dactylogyrus species. A total of 49 Dactylogyrus species including endemic and non-endemic species were collected from 62 endemic cyprinoid species in the Balkan and Apennine Peninsulas. However, 21 morphologically identified Dactylogyrus species exhibited different genetic variants (ranging from 2 to 28 variants per species) and some of them were recognized as cryptic species on the basis of phylogenetic reconstruction. Phylogenetic analyses revealed several lineages of endemic and non-endemic Dactylogyrus species reflecting some morphological similarities or host affinities. Using distance-based and event-based cophylogenetic methods, we found a significant coevolutionary signal between the phylogenies of parasites and their hosts. In particular, statistically significant links were revealed between Dactylogyrus species of Barbini (Cyprinidae) and their hosts belonging to the genera Aulopyge, Barbus and Luciobarbus. Additionally, a strong coevolutionary link was found between the generalist parasites D. alatus, D. sphyrna, D. vistulae, and their hosts, and between Dactylogyrus species of Pachychilon (Leuciscidae) and their hosts. Cophylogenetic analyses suggest that host switching played an important role in the evolutionary history of Dactylogyrus parasitizing endemic cyprinoids in southern Europe. We propose that the high diversification of phylogenetically related cyprinoid species in the Mediterranean area is a process facilitating the host switching of specific parasites among highly diverse congeneric cyprinoids.     

Predicting the similarity of parasite communities in freshwater fishes using the phylogeny,ecology and proximity of hosts

Parasite communities tend to be dissimilar in hosts that are geographically, phylogenetically, ecologically and developmentally distant from one another. The decay of community similarity is a powerful and increasingly common method of studying parasite beta diversity, but most studies have examined only a single type of distance. Here, we evaluate distances based on the phylogeny, ecology, spatial proximity and size of hosts, as predictors of the similarity of parasite communities in individual hosts, host populations and host species. We surveyed parasites in six species of fish collected simultaneously from six localities in the St. Lawrence River, Canada, and species in a common group of larval parasites were discriminated using DNA sequences from barcode region of cytochrome c oxidase I. Distances based on the habitat use patterns of host species were good predictors of short‐term, ecological similarity of parasite communities, such as that operating at the scale of the individual host. The genetic distance between host species was associated with almost all types of similarity at all scales, particularly qualitative and phylogenetic similarity of parasite communities at the level of populations and meta‐populations of hosts. The trophic level, diet, spatial proximity and size of hosts were poor predictors of parasite community similarity. The increased taxonomic resolution provided by molecular data increased the explanatory power of regression models, and different factors were implicated when parasite species were distinguished with DNA barcodes than when larval parasites were lumped into morphospecies, as is commonly practiced.     

Effect of habitat degradation on competition,carrying capacity,and species assemblage stability

Changes in species’ trophic niches due to habitat degradation can affect intra‐ and interspecific competition, with implications for biodiversity persistence. Difficulties of measuring species’ interactions in the field limit our comprehension of competition outcomes along disturbance gradients. Thus, information on how habitat degradation can destabilize food webs is scarce, hindering predictions regarding responses of multispecies systems to environmental changes. Seagrass ecosystems are undergoing degradation. We address effects of Posidonia oceanica coverage reduction on the trophic organization of a macroinvertebrate community in the Tyrrhenian Sea (Italy), hypothesizing increased trophic generalism, niche overlap among species and thus competition and decreased community stability due to degraded conditions. Census data, isotopic analysis, and Bayesian mixing models were used to quantify the trophic niches of three abundant invertebrate species, and intra‐ and interspecific isotopic and resource‐use similarity across locations differing in seagrass coverage. This allowed the computation of (1) competition strength, with respect to each other and remaining less abundant species and (2) habitat carrying capacity. To explore effects of the spatial scale on the interactions, we considered both individual locations and the entire study area (“‘meadow scale”). We observed that community stability and habitat carrying capacity decreased as P. oceanica coverage declined, whereas niche width, similarity of resource use and interspecific competition strength between species increased. Competition was stronger, and stability lower, at the meadow scale than at the location scale. Indirect effects of competition and the spatial compartmentalization of species interactions increased stability. Results emphasized the importance of trophic niche modifications for understanding effects of habitat loss on biodiversity persistence. Calculation of competition coefficients based on isotopic distances is a promising tool for describing competitive interactions in real communities, potentially extendible to any subset of ecological niche axes for which specimens’ positions and pairwise distances can be obtained.     

Distance decay of similarity among parasite communities of three marine invertebrate hosts

The similarity in species composition between two communities generally decays as a function of increasing distance between them. Parasite communities in vertebrate definitive hosts follow this pattern but the respective relationship in intermediate invertebrate hosts of parasites with complex life cycles is unknown. In intermediate hosts, parasite communities are affected not only by the varying vagility of their definitive hosts (dispersing infective propagules) but also by the necessary coincidence of all their hosts in environmentally suitable localities. As intermediate hosts often hardly move they do not contribute to parasite dispersal. Hence, their parasite assemblages may decrease faster in similarity with increasing distance than those in highly mobile vertebrate definitive hosts. We use published field survey data to investigate distance decay of similarity in trematode communities from three prominent coastal molluscs of the Eastern North-Atlantic: the gastropods Littorina littorea and Hydrobia ulvae, and the bivalve Cerastoderma edule. We found that the similarity of trematode communities in all three hosts decayed with distance, independently of local sampling effort, and whether or not the parasites used the mollusc as first or second intermediate host in their life cycle. In H. ulvae, the halving distance (i.e. the distance that halves the similarity from its initial similarity at 1 km distance) for the trematode species using birds as definitive hosts was approximately two to three times larger than for species using fish. The initial similarities (estimated at 1 km distance) among trematode communities were relatively higher, whereas mean halving distances were lower, compared to published values for parasite communities in vertebrate hosts. We conclude that the vagility of definitive hosts accounts for a high similarity at the local scale, while the strong decay of similarity across regions is a consequence of the low probability that all necessary hosts and suitable environmental conditions coincide on a large scale.     

Co-existence of nine gill ectoparasites (Dactylogyrus: monogenea) parasitising the roach (Rutilus rutilus l.): history and present ecology

Co-existence among potentially competing species can be favoured by niche specialisation and/or by reducing the overall intensity of competition via aggregated utilisation of fragmented resources. We investigated the respective roles of niche specialisation and aggregation in the case of nine congeneric monogenean parasites on the gills of Roach (Rutilus rutilus L.) belonging to the genus Dactylogyrus. The position of each individual parasite of the nine Dactylogyrus species was recorded. Niche breadth and niche overlap of parasite species were estimated. Comparative methods, which take into account phylogenetic information of the analysed species, were used. We reconstructed a phylogeny of the nine Dactylogyrus species based on morphological characters. We used the 'aggregation model of co-existence' in the model to test if species co-existence is facilitated when intraspecific aggregation exceeds interspecific aggregation. We observed a lack of negative correlation in abundance between pairs of parasites, and a negative correlation between niche size and parasite aggregation, for both intraspecific and interspecific aggregation. Our comparative analysis showed that parasite abundance is positively correlated with niche breadth. Then parasite abundance, and not interactions between Dactylogyrus species, seems to be the most important factor determining niche size This result gives some support to niche segregation by specialisation. Niche size was negatively correlated with both intraspecific and interspecific aggregation. No relationship was found between an increase of interspecific aggregation with an increase of niche overlapping, which suggests that competition may play little role. A lack of competition could be also confirmed by the lack of negative correlation in abundance between species pairs. A parsimony analysis of the evolution of gill distribution indicates a change in one parameter of the niche (arch, segment and/or area) at each branching event.     

福建琅岐岛台湾相思群落灌木层主要物种生态位与种间联结

为探究福建琅岐岛台湾相思(Acacia confusa)群落种间关系和群落演替动态,对其灌木层主要物种的生态位和种间联结特征进行了分析。结果表明,重要值和生态位宽度最大的物种为雀梅藤(Sageretia thea),其竞争优势、环境适应能力最强;灌木层主要物种的生态位相似性和生态位重叠较小,表明种间对环境资源的利用和要求存在较大差异,表现为较稳定的种间关系;灌木层主要物种的总体联结性为显著正联结,种对间正负联结比为1.72,检验显著率为7.35%,表明种对间联结程度较弱,独立性相对较强,物种竞争不激烈,群落处于相对后期且稳定的演替阶段;红锥(Castanopsis hystrix)与许多物种间存在垂直方向上对生境要求的互补性,生态位重叠较小,种间呈正联结关系;外来物种马缨丹(Lantanacamara)与福建胡颓子(Elaeagnus oldhamii)存在一定的竞争,与灌木层其他主要植物之间的资源竞争不激烈,其对该群落的影响相对较小。琅岐岛台湾相思群落目前已发展为地带性的顶级群落,物种竞争不激烈,种间关系较稳定;而海岛上恶劣的环境和外来物种的扩散会导致植物生境资源短缺,迫使物种聚集在...     

The biogeography of parasitism in sticklebacks: distance,habitat differences and the similarity in parasite occurrence and abundance

Similarity in parasite community composition often decreases with both increasing geographic distance and environmental dissimilarity between localities, though it is unknown whether similarity in local abundance of selected parasite species follows similar rules. We tested this using data on metazoan parasites in 126 stickleback (Gasterosteus aculeatus) populations, with locations from Eurasia, eastern North America, and western North America treated separately. Similarity values were regressed against pairwise distances between localities; after correcting for distance, the effect of environmental dissimilarity was assessed by splitting similarity values into those between pairs of localities with either similar, moderately different or very different salinity (freshwater, marine or brackish). For selected parasite species, pairwise similarity in abundance (mean no. parasites per host) were computed across all localities, and treated as above. Similarity in parasite community composition decreased with increasing distance between localities in all three geographic regions. A significant effect of environmental difference was found in all regions: for a given distance between two sites, their parasite communities were more similar if they were of the same salinity. Slopes for distance decay in similarity were consistently higher for eastern North America than for Eurasia. Among the 12 parasite species for which sufficient data were available, only 4 showed the expected relationship, i.e. the greater the geographical separation between host populations, the greater the difference in parasite abundance; also, significant effects of environmental differences in salinity were only found for 3 of these species. Our findings show that parasite communities of sticklebacks are structured by geographical distance and local salinity conditions. The results indicate that strong effects at the community level do not translate into corresponding effects at the population level, suggesting that parasite dispersal and population dynamics are controlled by different processes.     

Microhabitats of gill parasites (Monogenea and copepoda) of teleosts (Scomberoides spp.)

Ramasamy P., Ramalingam K., Hanna R. E. B. and Halton D. W. 1985. Microhabitat of gill parasites (Monogenea and Copepoda) of teleosts (Scomberoides spp.). International Journal for Parasitology15: 385–397. The parasites Vallisia indica, Allodiscocotyla chorinemi, Heterapta chorinemi, and Dionchus remorae exhibited site specificity on the gills of Scomberoides commersonianus, S. tol, S. lysan and S. tala, whereas the copepod Caligus sp. did not. Observations on site preference revealed microhabitat differences along the length of gill filaments, along the anterio-posterior axis of the gill, between external and internal gill filaments and on different gill arches. Site preference varied with parasite density on each gill and host. An interspecific association test between pairs of species revealed, in some cases, a positive association, and in other cases a negative association. There were apparently no association? between certain pairs of species. A comparison of intensity within pairs of parasite species infecting the same host revealed either an inverse or a direct correlation. The numerical dominance and prevalence of parasites differed on each host species. This study indicates that intra- and interspecific competition may occur among the gill parasites. The direction and speed of ventilation water-currents and certain intrinsic factors of the parasite themselves may determine their microhabitat restriction on the gills.     

Limiting similarity and the intensity of competitive effects on the mottled sculpin,Cottus bairdi,in experimental stream communities

The identification of potential competitors has been driven by the concept of limiting similarity. Lacking are explicit tests of interaction strength among morphologically similar and dissimilar species. I used the mottled sculpin, Cottus bairdi, as a focal species in an artificial stream experiment designed to compare the effect of intraspecific competition to interspecific cometition from two very different species: a congener, the Kanawha sculpin (C. carolinae ssp.), and an unrelated species, the fantail darter (Etheostoma flabellare). The differences in morphology between these two species generate specific predictions under limiting similarity regarding the likelihood of competition and its relative strength: the congener should be a more important potential competitor. Increased fish density had a strong effect on the multivariate response of survival and growth, and on the relative condition of C. bairdi, indicating competition. The effect of additional C. bairdi or Kanawha sculpins were roughly equal, but the effect of E. flabellare was significantly greater. The most important potential impact on C. bairdi came from interspecific competition by a species that is smaller and very different in morphology, contrary to predictions based on limiting similarity.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Ecological correlates of the distribution limits of two poeciliid species along a salinity gradient

Identifying the environmental factors responsible for the formation of a species' distribution limit is challenging because organisms interact in complex ways with their environments. However, the use of statistical niche models in combination with the analysis of phenotypic variation along environmental gradients can help to reduce such complexity and identify a subset of candidate factors. In the present study, we used such approaches to describe and identify factors responsible for the parapatric distribution of two closely‐related livebearer fish species along a salinity gradient in the lowlands of Trinidad, West Indies. The downstream distribution limits of Poecilia reticulata were strongly correlated with the brackish–freshwater interface. We did not observe significant phenotypic variation in life‐history traits for this species when comparing marginal with more central populations, suggesting that abrupt changes in conditions at the brackish–freshwater interface limit its distribution. By contrast, Poecilia picta was present across a wide range of salinities, although it gradually disappeared from upstream freshwater localities. In addition, P. picta populations exhibited an increase in offspring size in localities where they coexist with P. reticulata, suggesting a role for interspecific competition. The parapatric distribution of these two species, suggests that P. reticulata distributions are limited by an abiotic factor (salinity), whereas P. picta is limited by a biotic factor (interspecific competition). Similar parapatric patterns have been previously described in other systems, suggesting they might be a common pattern in nature. © 2013 The Linnean Society of London     

Competition from sea to mountain: Interactions and aggregation in low‐diversity monogenean and endohelminth communities in twospot livebearer Pseudoxiphophorus bimaculatus (Teleostei: Poeciliidae) populations in a neotropical river

1. The role of interspecific interactions in structuring low‐diversity helminth communities is a controversial topic in parasite ecology research. Most parasitic communities of fish are species‐poor; thus, interspecific interactions are believed to be unimportant in structuring these communities.
2. We explored the factors that might contribute to the richness and coexistence of helminth parasites of a poeciliid fish in a neotropical river.
3. Repeatability of community structure was examined in parasitic communities among 11 populations of twospot livebearer Pseudoxiphophorus bimaculatus in the La Antigua River basin, Veracruz, Mexico. We examined the species saturation of parasitic communities and explored the patterns of species co‐occurrence. We also quantified the associations between parasitic species pairs and analyzed the correlations between helminth species abundance to look for repeated patterns among the study populations.
4. Our results suggest that interspecific competition could occur in species‐poor communities, aggregation plays a role in determining local richness, and intraspecific aggregation allows the coexistence of species by reducing the overall intensity of interspecific competition.

     

Competition for food between an introduced crayfish and two fishes endemic to the Colorado River basin

Synopsis Crayfish are not native to the Colorado River basin (CRB), however they are now established in portions of the mainstem and in many tributaries. I used density manipulation experiments in a laboratory setting to determine intra- and interspecific competition for food between Orconectes virilis, an aggressive polytrophic crayfish now common in the CRB, and two native fishes: Gila chub, Gila intermedia, and flannelmouth sucker, Catostomus latipinnis. I tested each fish species in separate trials. Growth of Gila chub decreased when animal densities increased, however they were more affected by intraspecific competition than by crayfish presence. In contrast, growth of flannelmouth suckers was more affected by crayfish than by intraspecific competition. Crayfish growth was not significantly altered by presence of either fish. Crayfish thus reduced fish growth by competition for food, but the effect differed markedly between the two species. An erratum to this article can be found at .     

The evolution of competing species of terrestrial salamanders: niche partitioning or interference?

Summary One of the central assumptions of evolutionary ecology is that interspecific competition is a potent evolutionary force acting on coexisting species. There are few animal species that provide an opportunity for an experimental analysis of the evolutionary consequences of the phenomenon. We have taken advantage of the fact that two species of terrestrial salamander,Plethodon glutinosus andP. jordani, have different altitudinal distributions on two mountain ranges in North Carolina. Field removal experiments showed that interspecfic competition was much stronger in the Great Smoky Mountains than in the Balsam Mountains, and transplant experiments between the two mountain ranges showed that neither species from the Balsam Mountains had a measurable effect on its congener in the Smokies, although both species from the Smokies had strong negative effects on the Balsam congeners. Other experiments were conducted on the behavioral and ecological changes that have (or have not) evolved in the two areas. Our studies show that increased interspecific interference was the major evolutionary response of these largePlethodon species to interspecific competition, and that partitioning of food or microhabitat was not involved.     

Aggregation and species coexistence of ectoparasites of marine fishes.

Interspecific interaction may lead to species exclusion but there are several ways in which species can coexist. One way is by reducing the overall intensity of competition via aggregated utilisation of fragmented resources. Known as the 'aggregation model of coexistence', this system assumes saturation and an equilibrium number of species per community. In this study we tested the effects of interspecific aggregation on the level of intraspecific aggregation among ectoparasites of marine fishes (36 communities of gill and head ectoparasite species). If parasite species are distributed in a way that interspecific aggregation is reduced relative to intraspecific aggregation then species coexistence is facilitated. We found a positive relationship between parasite species richness and fish body size, controlling for host phylogeny. A positive relationship between infracommunity species richness and total parasite species richness was also found, providing no evidence for saturation. This result supports the view that infracommunities of parasites are not saturated by local parasite residents. The observed lack of saturation implies that we are far from a full exploitation of the fish resource by parasites. Ectoparasites were aggregated at both population and species levels. However, only half of the ectoparasite communities were dominated by negative interspecific aggregation. We found that infracommunity parasite species richness was positively correlated with the level of intraspecific aggregation versus interspecific aggregation. This means that intraspecific aggregation increases compared with interspecific aggregation when total parasite species richness increases, controlling fish size and phylogeny. This supports one assumption of the 'aggregation model of coexistence', which predicts that interspecific interactions are reduced relative to intraspecific interactions, facilitating species coexistence.     

Do parasites confer a disadvantage to hybrids?

Summary This paper is the second of three studies of a natural hybrid,Rutilus rubilioxAlburnus alburnus found in Lake Mikri Prespa in Northern Greece. It compares the parasite load of the hybrid to that of its parents, focussing onDactylogyrus spp.,Diplozoon sp.,Bolbophorus confusus andPomphorhynchus bosniacus. The hybrid's gill tissue is parasitised almost exclusively by the parentalDactylogyrus. It follows that theDactylogyrus group constitutes an excellent biological marker for hybridisation. Furthermore, the hybrid is characterised by the following two features: 1. high susceptibility to parasitic infections, 2. important role of ecological components in the recruitment of parasites: for monoxenous parasites (Dactylogyrus spp.), the spatial position of the hybrid and for heteroxenous parasites (P. bosniacus), the trophic position. The parasitology of this particular hybrid may be useful as a model for the properties and fitness of hybrids in general.     

Genetic structures of allopatric and sympatric populations in Pseudorasbora pumila pumila and Pseudorasbora parva

Pseudorasbora pumila pumila was collected to investigate its genetic relationships from 11 localities in Hokkaido and eastern Honshu of Japan with Pseudorasbora parva from 6 localities in Hokkaido, eastern Honshu, and Polovynka River (a tributary of the Amur River in eastern Russia) in 1996–1998. In Hokkaido and Akita prefectures (Honshu), three P. pumila pumila populations were distributed with P. parva sympatrically. Samples of P. pumila pumila and P. parva were analyzed based on 27 allozyme loci. Genetic differentiation between P. pumila pumila and P. parva (D = 0.522–0.622, average 0.601) was almost equal to or greater than that between other congeneric fish species previously reported. In the three sympatric populations of two species, which had arisen from artificial transplantation of the latter into localities of the former, the individual frequency composition of genetic types (P. pumila pumila, P. parva, F₁ hybrid, and later filial generation hybrid) were significantly different from each other. Differing from the previous studies, F₁ hybrids were fertile and coexisting populations of the two species included the F₁ hybrid and the later filial generation hybrid. These results suggest that genetic population dynamics of sympatric populations of the two species would be influenced not only by hybridization but also by ecological and environmental factors. Alternatively, the existence of populations that possessed both the alleles specific to P. pumila pumila and those specific to P. parva may indicate that introgressive hybridization, instead of species displacement, has occurred between the two species under a certain condition.     

Habitat segregation in a salt marsh among adult sticklebacks (Gasterosteidae)

Synopsis The spatial distribution, seasonal abundance and diel activity of four sticklebacks,Gasterosteus aculeatus, G. wheatlandi, Pungitius pungitius, andApeltes quadracus. coexisting in a St. Lawrence salt marsh were examined to see how these closely related species share their habitat. While all four species breed in the Riviè des Vases, a tidal creek, only three species are found in the adjacent salt marsh pools,A. quadracus being absent. Results are interpreted in terms of avoidance of interspecific competition for space during the relatively short breeding season at this latitude.