The taxonomic and phylogenetic status of digeneans from the genus Timoniella (Digenea: Cryptogonimidae) in the Black and Baltic seas.

Timoniella spp. are cryptogonimid flukes (Digenea: Cryptogonimidae) that parasitize the guts of fish in brackish waters. Timoniella imbutiforme, a species from the Mediterranean Sea, is recorded in the Black Sea, while T. balthica has been described from the Baltic Sea. In this paper, we clarify the taxonomic status of Timoniella populations in the Baltic and Black seas. Adults and metacercariae of Timoniella spp. were sampled from localities in the Mediterranean Sea (France), Black Sea (Ukraine) and Baltic Sea (Germany) and subjected to molecular and morphological analysis, including Bayesian phylogenetic reconstruction based on concatenated sequences of ITS1-ITS2-28S. This allowed us to construct a new key to species of the genus Timoniella. Our results suggest that T. balthica forms part of the Boreal-Atlantic relict fauna of the Black Sea and should now be considered a junior synonym of T. imbutiforme.

Did biogeographical processes shape the monogenean community of butterflyfishes in the tropical Indo-west Pacific region?

Geographical distribution of parasite species can provide insights into the evolution and diversity of parasitic communities. Biogeography of marine parasites is poorly known, especially because it requires an understanding of host-parasite interactions, information that is rare, especially over large spatial scales. Here, we have studied the biogeographical patterns of dactylogyrid parasites of chaetodontids, one of the most well-studied fish families, in the tropical Indo-west Pacific region. Dactylogyrid parasites were collected from gills of 34 butterflyfish species (n=560) at nine localities within an approximate area of 62millionkm2. Thirteen dactylogyrid species were identified, with richness ranging from 6 to 12 species at individual localities. Most dactylogyrid communities were dominated by Haliotrema angelopterum or Haliotrema aurigae, for which relative abundance was negatively correlated (ρ=-0.59). Parasite richness and diversity were highest in French Polynesia and the Great Barrier Reef (Australia) and lowest in Palau. Three biogeographic regions were identified based on dactylogyrid dissimilarities: French Polynesia, characterised by the dominance of H. angelopterum, the western Pacific region dominated by H. aurigae, and Ningaloo Reef (Australia), dominated by Euryhaliotrema berenguelae. Structure of host assemblages was the main factor explaining the dissimilarity (turnover and nestedness components of the Bray-Curtis dissimilarity and overall Bray-Curtis dissimilarity) of parasite communities between localities, while environment was only significant in the turnover of parasite communities and overall dissimilarity. Spatial structure of localities explained only 10% of the turnover of parasite communities. The interaction of the three factors (host assemblages, environment and spatial structure), however, explained the highest amounts of variance of the dactylogyrid communities, indicating a strong colinearity between the factors. Our findings show that spatial arrangement of chaetodontid dactylogyrids in the tropical Indo-west Pacific is primarily characterised by the turnover of the main Haliotrema spp., which is mainly explained by the structure of host assemblages.

Pelagic to demersal transition in a coral-reef fish, the orbicular batfish Platax orbicularis.

Behavioural and ecological observations were made on young, reared Platax orbicularis in Opunohu Bay, Moorea, French Polynesia, during their transition from the pelagic, dispersive stage to the reef-orientated demersal stage. Seventy-two young P. orbicularis (17-75 mm standard length, LS ) were released in the pelagic zone and 20 (40-70 mm LS ) adjacent to the reefs. Swimming speed was slow (mean 5·2 cm s(-1) ) and independent of size. An ontogenetic descent was observed: the smallest P. orbicularis swam at the surface, medium-sized P. orbicularis swam in midwater (mean 5-13 m) and the largest P. orbicularis swam to the bottom, where many lay on their sides. Platax orbicularis swam southerly on average, away from the ocean and into the bay. Smaller P. orbicularis were more likely to swim directionally than larger individuals. Young P. orbicularis released near reef edges swam at similar, but more variable speeds (mean 6·6 cm s(-1) ). About half of those released near reefs swam away, but fewer swam away from an inshore fringing reef than from a patch reef near the bay mouth. Many P. orbicularis swam up the slope onto the reef top, but the little settlement observed was near the reef base. Average, near-reef swimming direction was also southerly. Some reef residents, in particular the triggerfish Balistapus undulatus, harassed young P. orbicularis.

Parasite acquisition by larval coral-reef fishes.

Of 164 fish larvae belonging to 11 species sampled from the island of Moorea, French Polynesia, 30% had at least one parasite individual and parasite prevalence ranged between 0 for Chaetodon citrinellus and 80% for Parupeneus barberinus. Parasites were present only in larvae that had fed and were present in the gut but absent from gills, body cavity and muscle. Parasites appeared to be acquired by ingestion of intermediate hosts when the larvae fed on the outer slope, prior to reef colonization.

Monorchiids (Platyhelminthes: Digenea) of chaetodontid fishes (Perciformes): biogeographical patterns in the tropical Indo-West Pacific.

Species richness and biogeography of the monorchiid genus Hurleytrematoides was studied by the examination of 2834 individuals of 45 species of Chaetodontidae at six major sites in the tropical Indo-West Pacific: Heron Island, Lizard Island, Ningaloo (Western Australia), Palau, New Caledonia and Moorea (French Polynesia). In total, 18 species were distributed among six sites; descriptions are provided for eight new species: H. boucheti n. sp., H. combesi n. sp., H. deblocki n. sp., H. dollfusi n. sp., H. euzeti n. sp., H. kulbickii n. sp., H. pasteuri n. sp., and H. planesi n. sp. Overall richness ranged from zero to five Hurleytrematoides species per chaetodontid species. Seven Hurleytrematoides species were found at only one locality and eleven were found at multiple localities. Only one species, H. morandi, was found at all localities. Individual localities had between six (Moorea) and 10 (Heron Island) species; we attribute Moorea's depauperate parasite fauna to its isolation and distance from the Indo-Philippine centre of biological diversity. Using cluster analysis of 18 species of Hurleytrematoides and 45 species of chaetodontids sampled in the Indo-West Pacific, we show that the localities on the Great Barrier Reef (Heron Island and Lizard Island) and New Caledonia have the most similar chaetodontid and parasite fauna of any locality pairs. Cluster analysis results also show that the similarity of the chaetodontid assemblages at five of the six localities is relatively high and that Ningaloo has the most distinct fauna. Similarity values based on sharing of species of Hurleytrematoides are generally lower than those for their hosts; Moorea, Ningaloo and Palau all have low similarity to New Caledonia and Great Barrier Reef sites. We attribute these distinctions to the differential dispersal capability of the fish and their parasites. Chaetodontids have long-lived mobile pelagic larvae, the dispersal of which would be most affected by prominent biogeographical barriers, such as that between the Indian and Pacific Oceans. In contrast, monorchiids have no obvious dispersal stage, and vast distances have the capacity to act as effective barriers to dispersal. We conclude that the present distributions of species of Hurleytrematoides in the Indo-Pacific are driven by historical opportunity and capacity to disperse, and that some disjunct distributions are sculpted by stochasticity.

Regulation of Anguillicola crassus (Nematoda) infrapopulations in their definitive host, the European eel, Anguilla anguilla.

The parasitic nematode Anguillicola crassus was recently introduced into populations of the European eel, Anguilla anguilla. We investigated, under experimental conditions, the regulation of A. crassus infrapopulations. We tested the effects of (1) the resource-limited habitat of the parasite and (2) the coexistence of several developmental stages in its niche (the swim-bladder) on the composition of the infrapopulations. The results revealed that the respective effects of these factors differed substantially during the course of the infection. Third-stage larvae (L3s) establishment would not be constrained by the size of the swim-bladder. Their moult to fourth-stage larvae (L4s) would be accelerated as the number of L3s increased. The moulting time of L4s to adults would be reduced by males and would be constrained by the size of the swim-bladder. However, the moult of L4s to adults and their further development would be synchronized with those of the opposite sex. At the time of mating, the number of males and the body weight of adults would depend on the size of the swim-bladder. Soon after the laying of eggs, the developmental constraint on the late L3s would decrease. When adults die, constraints would cease and late larval stages would moult to become adults.

Differential gene expression analysis in European eels (Anguilla anguilla, L. 1758) naturally infected by macroparasites.

We analyzed the relationships between the macroparasite community of the European eel and the expression of genes involved in the host physiology during its continental life. The genes studied are implicated in (1) host response to environmental stress, i.e., heat shock protein 70 (HSP70) and metallothionein (MT); (2) osmoregulation, i.e., beta thyroid hormone receptor (betaTHR) and Na+/K+ATPase; and (3) silvering, i.e., betaTHR, freshwater rod opsin (FWO), and deep-sea rod opsin (DSO). All were enumerated by quantitative reverse-transcription polymerase chain reaction. The epizootiological results for 93 yellow eels caught in the Salses-Leucate Lagoon (France) included 11 species: 1 nematode, 2 acanthocephalans, 1 monogenean, and 7 digeneans. The molecular results revealed (1) a significant negative relationship between digenean abundance and the expression level of all the tested genes, except FWO; (2) a significant negative relationship between the abundance of the nematode Anguillicola crassus and the expression level of the Na+/K+ATPase gene; and (3) a significant positive relationship between the A. crassus abundance and the expression level of the MT gene. Eels infected with digeneans had, on average, a lower level of expressed genes. We hypothesize that the parasites may disturb the eel's ability to withstand environmental stress and delay their migration to the Sargasso Sea because of degeneration of the gut. We further propose that the effect of the invasive species, A. crassus, on the gene expression was mainly linked to an increased trophic activity of infected eels. Moreover, it is possible that the parasite may have an effect on the fish's migratory behavior, which is tied to reproductive purposes. Additional work, including an experimental approach, is required to confirm our hypotheses.

Biased sex ratio in the European eel (Anguilla anguilla) swim-bladder parasite Anguillicola crassus, experimentally induced by 11-ketotestosterone.

Parasites are intimately connected to the host in which they live, and some may be affected by the polluted environment of their host. The present study describes the effect of a steroid hormone (11-ketotestosterone) on the sex ratio of the invasive hematophagous nematode Anguillicola crassus Kuwahara, Niimi & Itagaki, 1974, when experimentally injected to European eels, Anguilla anguilla. Our results showed that this steroid induced a significant male-biased ratio in the nematode A. crassus infrapopulations, suggesting that the presence of endocrine disruptors in the environment may lead to skewed sex ratios among parasites.

Acanthocephalus reunionensis n sp (Acanthocephala: Echinorhynchidae), a parasite of Anguilla species (Anguillidae) from Reunion Island.

In a survey of 118 eels Anguilla bicolor, A. marmorata and A. mossambica, (Anguillidae) indigenous to Reunion Island in the Mascarene island group, western Indian Ocean, a new species of acanthocephalan, Acanthocepholus reunionensis n. sp., was found. With a proboscis hook formula of 19 rows of 4-5 hooks, and elongated cement glands arranged in three pairs, this species differs from all other species in the genus. This is the first record of the genus Acanthocephalus occurring in eels from the African Region.

Infracommunity level aggregation in the monogenean communities of crucian carp (Carassius carassius).

Aggregation is one of the distinctive features in parasite-host relationships, which has generally been studied at the level of host communities. Parasite aggregation at the infracommunity level may nevertheless be important for intraspecific interactions such as parasite mating success and opportunities for cross-fertilization. In the present paper, we studied the infracommunity aggregation of 3 highly abundant Dactylogyrus (Monogenea) species occurring on the gills of crucian carp (Carassius carassius). In line with the previous work on monogenean communities, we observed no competition between the species. At the species level, parasites were distributed unevenly on the gills showing aggregation in the majority of infracommunities. However, aggregation decreased with increasing parasite abundance, which supports the hypothesis that less aggregation may be needed to ensure successful mating when the distance to a potential mate decreases with increasing number of conspecifics. Lack of interspecific interactions, species specific site selection and the importance of study scale for aggregation patterns in dactylogyrids are discussed.

Sex discriminatory effect of the acanthocephalan Acanthocephaloides propinquus on a gobiid fish Gobius bucchichii.

Most studies which aim at detecting effects of parasites on fish show that intermediate stages of parasites affect their host while adult parasites are usually less virulent in the final host. We studied the effect of the acanthocephalan Acanthocephaloides propinquus on one of its final hosts, the fish Gobius bucchichii. This study showed that the adult parasites affect host fitness. Moreover, our results showed that at the same level of infection in male and female gobiids, the female's reproductive success was negatively correlated to parasite abundance while the male's reproductive success was not. The negative effects on females includes reductions in gonado-somatic index and egg production. We hypothesized that it might be more difficult for females to compensate the cost of parasitism because of the very high cost of egg production. We discussed these results in terms of host population regulation.

Aggregation of congeneric parasites (Monogenea: Dactylogyrus) among gill microhabitats within one host species (Rutilus rutilus L.).

Parasite aggregation within the gill arches of 1 host species (Rutilus rutilus L.) was studied in the case of 9 congeneric monogeneans belonging to the genus Dactylogyrus. Both intra- and interspecific aggregation were followed by considering gill arches as independent patches. Parasite species were found to be aggregated both intra- and interspecifically. We showed that the intensity of parasite infection was positively related to intraspecific aggregation only for the more abundant species. No relationship was found between intensity of parasite infection and niche size when correcting for total parasite intensity. This may suggest that when parasite intensities are low, intraspecific competition may not have a strong effect on parasites. Conversely, when looking at the evolution of niche size and intraspecific aggregation for each species separately, intraspecific aggregation decreased for the most abundant species (D. crucifer) suggesting that this species is more competitive than others when total parasite intensity increases. When considering interspecific aggregation, Dactylogyrus species were found to be positively and negatively aggregated. Following the prediction of morphological and ecological similarity for congeneric species, Dactylogyrus species similar in morphometry of attachment parts tend to be more positively aggregated among their gill microhabitats than Dactylogyrus species with morphometric differences in attachment parts.

Water temperature influencing dactylogyrid species communities in roach, Rutilus rutilus, in the Czech Republic.

Dactylogyrid species (Monogenea) communities were studied in roach, Rutilus rutilus, collected from two localities in the basin of Morava river, Czech Republic, during the period from April to November 1997 and March to September 1998 to determine the effect of water temperature on parasite abundance, species richness and diversity. Dactylogyrid species were found to co-occur on the gills of roach with up to six species found on the same host individual. Nine dactylogyrid species were identified with the abundance of each reaching a very low level. Niche size was considered to increase with species abundance even when water temperature was high. There was a strong effect of water temperature on abundance of the common dactylogyrid species (D. crucifer, D. nanus, D. rutili and D. suecicus) as well as of the rare species D. rarissimus. The temporary occurrence of the rare species was found without any temperature effect. Water temperature did not affect the relationship between abundance and niche size. Niche size increased with abundance, even when the water temperature was high, which suggests that negative interspecific interactions are not important within dactylogyrid communities.

Endoparasite species richness of new caledonian butterfly fishes: host density and diet matter.

Ecological factors may influence the number of parasites encountered and, thus, parasite species richness. These factors include diet, gregarity, conspecific and total host density, habitat, body size, vagility, and migration. One means of examining the influence of these factors on parasite species richness is through a comparative analysis of the parasites of different, but related, host species. In contrast to most comparative studies of parasite species richness of fish, which have been conducted by using data from the literature, the present study uses data obtained by the investigators. Coral reef fishes vary widely in the above ecological factors and are frequently parasitized by a diverse array of parasites. We, therefore, chose to investigate how the above ecological factors influence parasite species richness in coral reef fishes. We investigated the endoparasite species richness of 21 species of butterfly fishes (Chaetodontidae) of New Caledonia. We mapped the diet characters on the existing butterfly fish phylogeny and found that omnivory appears to be ancestral. We also mapped the estimated endoparasite species richness, coded from low to high parasite species richness, on the existing butterfly fish phylogeny and found that low parasite species richness appears to be associated with the ancestral state of omnivory. Different dietary and social strategies appear to have evolved more than once, with the exception of obligate coralivory, which appears to have evolved only once. Finally, after controlling for phylogenetic relationships, we found that only the percentage of plankton in the diet and conspecific host density were positively correlated with endoparasite species richness.

Sexual competition in an acanthocephalan parasite of fish.

Acanthocephalans are polygamous parasites of vertebrates and some species are known to aggregate in sexual congress to mate. Such a reproductive behaviour could lead to male-male competition for access to females and could have consequences for sexual selection. We dissected 87 gobiid fish, Gobius bucchichii, harbouring 891 acanthocephalans, Acanthocephaloides propinquus. The parasites were sexed and their body sizes were measured. Testicular volume was also evaluated in 82 males in order to establish their phenotypic sexual investment in relation to the estimated sex ratio. We found that parasite intensity (i.e. the number of individuals/fish) was not correlated with fish size, but that parasite size was significantly related to host size. Our results showed that there was a significant relationship between the mean female body size and their number within one host. We found that when the percentage of male parasites in a host increased, presumably increasing male-male competition for access to females, males had a larger testicular volume. We discuss these results in terms of energy allocation, sexual and sperm competition. We conclude that competition for space should be less important for males than competition for access to females. Moreover, increasing testis size should confer advantages to males especially for their reproductive success when sperm competition occurs.

Parasites of recruiting coral reef fish larvae in New Caledonia.

Recruiting coral reef fish larvae from 38 species and 19 families from New Caledonia were examined for parasites. We found 13 parasite species (Platyhelminthes: Monogenea, Cestoda and Trematoda) but no acanthocephalan, crustacean or nematode parasites. Over 23% of individual fish were infected. Didymozoid metacercariae were the most abundant parasites. We conclude that most of the parasites are pelagic species that become 'lost' once the fish larvae have recruited to the reef. Larval coral reef fish probably contribute little to the dispersal of the parasites of the adult fish so that parasite dispersal is more difficult than that of the fish themselves.

Community structure of digenean parasites of sparid and labrid fishes of the Mediterranean sea: a new approach.

The aim of this work was to study the structure of the parasite communities of Digeneans of 2 families of Teleost fishes (Sparidae and Labridae) of the Mediterranean sea. We tried to quantify the importance of both the microhabitat requirements of the parasite species and the effect of host biological factors on the parasite communities. We applied, for the first time in parasite community studies, the Canonical Correspondence Analysis (CCA) to analyse (i) the spatial distribution of parasite species within the digestive tract of the hosts; (ii) the host's biological factors (such as diet, host length, gregariousness and abundance) that may influence this spatial distribution of parasite species. Our results showed that potential microhabitats were vacant in the 2 host families studied revealing a lack of niche saturation because either there was little inter- and/or intraspecific competition or there were enough available space and resources within the host. Our results also indicated that the position of the parasite in the digestive tract is much more important than host biological factors for the structure of parasite community. Finally, we highlight the potential use of the CCA method for controlling for phylogenetic constraints in multi-species analyses.

Comparative analysis: a tool for studying monogenean ecology and evolution.

The comparative approach has been used widely in evolutionary ecology during the last 10 years. In the first part of this study, taking monogeneans of Mediterranean marine fishes as an example, we highlight the pitfalls that may arise in interspecific studies. In the second part, the independent contrast method is used to control for phylogenetic effects in order to evaluate the determinants of monogenean species richness and their specificity. Our results show the importance of controlling data for both host sample size and phylogeny to avoid confounding effects. Thereafter, taking into account both sampling effort and phylogenetic effects, our results show that host size is the main factor affecting monogenean species richness and specificity.

Parasitism of Gobius bucchichii Steindachner, 1870 (Teleostei, Gobiidae) in protected and unprotected marine environments.

We collected 396 Gobius bucchichii, Steindachner, 1870 (Teleostei, Gobiidae) in and around the marine reserve of Cerbère-Banyuls, in the southeast of France, between March and July 1994. Five species of adult parasites were found: one acanthocephalan, Acanthocephaloides propinquus Dujardin, 1845 (Acanthocephala, Arhythmacanthidae); one nematode, Cucullanus sp. (Nematoda, Cucullanidae); and three species of digenetic trematodes, Helicometra sp. (Digenea, Opecoelidae), Derogenes sp. (Digenea, Hemiuridae) and Deretrema scorpaenicola Bartoli, 1990 (Digenea, Zoogonidae). Fishes collected in a protected area were on average, larger, older, had a higher percentage of regenerated scales, and harbored more parasites.