Some aspects of the taxonomy and biology of dracunculoid nematodes parasitic in fishes: a review

The nematode superfamily Dracunculoidea includes 166 recognized species, of which 150 (90%) are parasitic in about 300 species of freshwater, brackish-water and marine fishes. Fish dracunculoids are placed in 31 genera (86% of all dracunculoid genera) belonging to eight of the nine dracunculoid families: Anguillicolidae, Daniconematidae, Guyanemidae, Lucionematidae, Micropleuridae, Philometridae, Skrjabillanidae, and Tetanonematidae; the genus Lockenloia is considered incertae sedis. Because of difficulties in studying fish dracunculoids, associated with their morphological and biological peculiarities, most species of these largely histozoic parasites are poorly known and males of the majority of species and of eight genera have not yet been discovered. It is apparent that the present classification system of dracunculoids as a whole does not reflect phylogenetic relationships and a taxonomic revision of this nematode group, based on detailed morphological (including SEM and TEM), life history and molecular studies of individual species, is quite necessary. Data on the biology of fish dracunculoids is scarce. In known cases, their life cycles involve copepods, ostracods or branchiurids as intermediate hosts and, sometimes, fish paratenic hosts are known to occur in dracunculoid species parasitizing as adults piscivorous definitive hosts. However, nothing is known about the life cycles of representatives of 20 genera. Some species of dracunculoids, particularly of philometrids, are highly pathogenic and are known as agents of serious fish diseases. During recent years, especially the importance of Philometra spp. parasitizing the gonads of many species of marine fishes has increased due in particular to the rapid development of marine aquaculture, because they may significantly decrease fish reproduction or even cause full parasitic castration. Therefore, further detailed studies on fish dracunculoids are significant not only from the theoretical viewpoint, but they may also have practical implications.     

Some aspects of the taxonomy and biology of adult spirurine nematodes parasitic in fishes: a review

About 300 species belonging to four superfamilies (Gnathostomatoidea, Habronematoidea, Physalopteroidea and Thelazioidea) of the nematode suborder Spirurina are known as the adult parasites of freshwater, brackish-water and marine fishes. They are placed in four families, of which the Gnathostomatidae, including Echinocephalus with a few species and the monotypic Metaleptus, are parasites of elasmobranchs, whereas Ancyracanthus contains one species in teleosts; the Physalopteridae is represented in fish by four genera, Bulbocephalus, Heliconema, Paraleptus and Proleptus, each with several species in both elasmobranchs and teleosts. The majority of fish spirurines belongs to the Rhabdochonidae, which includes 10 genera (Beaninema, Fellicola, Hepatinema, Heptochona, Johnstonmawsonia, Megachona, Pancreatonema, Prosungulonema, Rhabdochona and Vasorhabdochona) of species parasitizing mainly teleosts, rarely elasmobranchs, and the Cystidicolidae with about 23 genera (Ascarophis, Caballeronema, Capillospirura, Comephoronema, Crenatobronema, Cristitectus, Ctenascarophis, Cyclozone, Cystidicola, Cystidicoloides, Johnstonmawsonoides, Metabronema, Moravecnema, Neoascarophis, Parascarophis, Prospinitectus, Pseudascarophis, Pseudoproleptus, Salvelinema, Similascarophis, Spinitectoides, Spinitectus, Sterliadochona), with many species parasitic in teleosts only. Because of difficulties in studying fish spirurines, associated with their morphological and biological peculiarities, most species of these parasites are poorly known. It is apparent that their present classification system does not reflect phylogenetic relationships and a taxonomic revision of this nematode group, based on detailed morphological (including SEM and TEM), life history and molecular studies of individual species, is quite necessary. In Cystidicolidae, several genera have been based on details in the cephalic structures visible only with the aid of SEM, but it will be evident whether or not these tiny features are of generic importance only when more cystidicolids are described using SEM and comparative molecular data become available. Data on the biology of fish spirurines are scarce. In known cases, their life cycles involve aquatic arthropods (crustaceans or insects) as intermediate hosts, in which, sometimes, the larvae undergo a precocious development and may even attain adulthood and become gravid in these invertebrates; sometimes, fish paratenic hosts are known to occur in cystidicolids parasitizing as adults piscivorous definitive hosts. Some spirurine species are pathogenic and are known as causative agents of serious fish diseases. Consequently, further detailed studies on fish spirurines are significant not only from the theoretical viewpoint, but they may also have practical implications.     

The biology of nematodes of the family Capillariidae Neveu-Lemaire, 1936

The present knowledge of the life cycles of nematodes of the family Capillariidae is reviewed and these data are considered from the viewpoint of a new system of the classification of genera in this family (Moravec 1982). An analysis of the relevant literature as also own studies have shown that, in this nematode group, there occur both direct (homoxenous) life cycles without an intermediate host (Baruscapillaria, Pseudocapillaria, Calodium, Pseudocapillaroides, partly also Capillaria, Eucoleus and Aonchotheca) and heteroxenous cycles with participation of obligate intermediate hosts that are usually oligochaetes and rarely fishes (Schulmanela, Pearsonema, partly also Capillaria, Eucoleus and Aonchotheca). A remarkable case is the species Aonchotheca philippinensis, an intestinal parasite of man, with alternative life cycles, i.e. either with participation of the intermediate host or without it (autoinfection), this being dependent on whether eggs or larvae are produced by the female parasites. The transmission of some capillariid species with a direct life cycle may include paratenic hosts (oligochaetes, fishes). Capillariids undergo four moults during their ontogenetic development, the first of which taking place inside the body of the intermediate host in case of heteroxenous cycles. The present knowledge of the biology of nematodes of the Capillariidae is very incomplete; their life cycles have hitherto been studied (in a different extent) in members of only 9 out of 22 presently valid genera (approximately in 7% of recognized capillariid species).     

Review of microsporidia-mosquito relationships: from the simple to the complex

Microsporidia in mosquitoes can be divided into two categories based on their life cycles and host-parasite relationships. Some species of microsporidia exhibit simple life cycles with one spore type responsible for oral (horizontal) transmission. They affect only one generation of the mosquito and are not usually host or tissue specific. Brachiola algerae and Vavraia culicis are examples of species isolated from mosquitoes with relatively straightforward life cycles (one spore type) and simple host-parasite relationships. B. algerae and a close relative of V. culicis have also been isolated from a vertebrate (human) host but sources for these infections are unknown. In contrast to B. algerae and V. culicis, polymorphic (heterosporous) microsporidia in mosquitoes are characterized by complex life cycles involving multiple spore types responsible for horizontal and vertical transmission. They affect two generations of the mosquito and some involve an obligate intermediate host. These microsporidia are generally very host and tissue specific with complex developmental sequences comprised of unique stages and events. The microsporidium Edhazardia aedis is a pathogen of Aedes aegypti and does not require an intermediate host. The developmental cycle of E. aedis is characterized by four sporulation sequences, two in the parental host and two in the filial generation. Recent speculation relative to the source of B. algerae human infection have implicated infected mosquitoes and raised concerns about the safety of mosquito microsporidia in general. The subject of this review is to compare and contrast three species of microsporidia from mosquitoes, two with broad host ranges (B. algerae and V. culicis) and one specific to mosquitoes (E. aedis). This review describes features that distinguish mosquito-parasitic microsporidia with simple life cycles and broad host ranges from truly mosquito-specific microsporidian parasites with complex life cycles.     

The role of mite pocket-like structures on Agama caudospinosa (Agamidae) infested by Pterygosoma livingstonei sp. n. (Acari: Prostigmata: Pterygosomatidae)

Pterygosoma livingstonei sp. n. collected from the Kenyan lizard Agama caudospinosa Meek shows morphological affinities with other South African congener species parasitizing lizards of the genus Agama, especially with P. triangulare Lawrence, 1936, but it differs in having glabrous genua II and III. P. livingstonei shows affinities with the Lawrence's hispida species group in the characters of genital and peripheral setae. This new species was found concentrated in a nuchal "mite pocket-like structure", a behaviour previously unreported among species belonging to the genus Pterygosoma. Mite pockets (or acarodomatia, acarinaria) of lizards typically house damaging chigger mites, and are usually interpreted as the evolutionary host's response to limit damage caused by parasites. Because scale mites are permanent ectoparasites and less damaging than seasonally occurring larval trombiculids, the heavy infestation by P. livingstonei in the nuchal skin folds of its host is interpreted as a consequence of the best utilisation of an available protected site by these mites that spend their entire life cycle on their host and whose primitive body shape prevents them from seeking shelter beneath the scales of their lizard host.     

A review of Dendromonocotyle (Monogenea: Monocotylidae) from the skin of stingrays and their control in public aquaria

Dendromonocotyle species (Monogenea: Monocotylidae) are the only monocotylids to parasitize the skin of chondrichthyan hosts. Currently 11 species are recorded from the skin of ray species in the Dasyatidae, Myliobatidae and Urolophidae. There have been increasing reports of Dendromonocotyle outbreaks on rays kept in public aquaria. This paper provides a broad review of Dendromonocotyle that should assist taxonomists and aquarists with species identification and help decisions on potential control methods for Dendromonocotyle infections. The taxonomy and host-specificity of Dendromonocotyle are discussed and a key to current species is provided. We summarise what little is known about the biology of Dendromonocotyle including egg embryonation and hatching, feeding, camouflage and reproduction. The efficacy of freshwater baths, chemical treatments and biological control measures such as the use of cleaner fish for Dendromonocotyle is also discussed. We demonstrate that effective control of Dendromonocotyle on captive rays is hampered by the lack of basic biological data on the life cycle of the parasites. A case history is provided outlining the success of a public aquarium (Underwater World, Mooloolaba, Queensland, Australia) in managing D. pipinna infections on captive Taeniura meyeni without chemical intervention simply by taking measures to reduce host stress.     

The interaction of fish trypanosome culture forms with some lectins

The interactions of fish trypanosome culture forms with 11 purified lectins were compared using the agglutination test in microwell plates. Altogether, ten stocks of ten different freshwater fish species were examined. Three basic types of cell-lectin interactions were observed on the microscopical level. The strong agglutination of all stocks regardless their original host was found in the presence of Con A, PSA, RCA60, and RCA120, which implies the presence of relatively high amounts of sugar residues of D-mannose and D-galactose in the surface of culture forms of these parasites. Weak agglutinations of some stocks were observed in the presence of LCA, PNA, SBA, and WGA lectins, but their low intensity makes them not sufficiently reliable for stock characterization. The lectins UEA I, HPA, and PHA caused no agglutination. In conclusion, in case of unequivocal results no remarkable differences in the interactions of various stocks of trypanosomes culture forms with used lectins were observed. These results imply the high degree of similarity of their main cell surface saccharide structures.     

New evidence on a cold case: trophic transmission, distribution and host-specificity in Hedruris spinigera (Nematoda: Hedruridae)

The life cycle of Hedruris spinigera Baylis, 1931 (Nematoda: Hedruridae) is determined here with the first formal identification of the parasite's intermediate host: the crustacean amphipod Paracorophium excavatum Thomson. Adult H. spinigera are redescribed from specimens collected from the stomach of fishes, Retropinna retropinna (Richardson) and Aldrichettaforsteri (Valenciennes), from Lake Waihola, New Zealand. Immature adults of the parasite collected from intermediate hosts (P. excavatum) are also described for the first time. The prevalence, abundance and intensity of infection of H. spinigera in several fish species are quantified along with the occurrence of P. excavatum, the parasite's intermediate host, in fish stomach contents. Although H. spinigera's transmission mode (trophic transmission) and fish diet potentially expose all fish species to infection, some level of host specificity must exist as parasite prevalence, abundance and intensity of infection vary greatly between potential definitive host species. We suggest here that the anatomy of the fish digestive tract and especially that of the stomach plays an important role in host suitability for H. spinigera. While P. excavatum is the only intermediate host in Lake Waihola, H. spinigera was found in six different fish species: Aldrichetta forsteri, Galaxias maculatus (Jenyns), Retropinna retropinna, Rhombosolea retiaria Hutton, Perca fluviatilis Linnaeus and Salmo trutta Linnaeus; although typical hedrurid attachment and mating positions were observed only in R. retropinna and A. forsteri. The limited distribution of H. spinigera is most likely due to that of its different host species (intermediate and definitive), all inhabitants of coastal fresh and brackish waters.     

Schistosomes causing cercarial dermatitis: a mini-review of current trends in systematics and of host specificity and pathogenicity

The human infection known under the names cercarial dermatitis or swimmers' itch is generally associated with swimming in lakes all over the world, however, a number of outbreaks of cercarial dermatitis developing in salt or brackish waters are also reported. The disease presents as allergic reaction which is able to trap and eliminate the parasites in the skin. However, the infection can be linked to more than skin symptoms under certain circumstances. Recent studies on bird schistosomes have shown that during primary infections of noncompatible hosts (mice) the parasites may migrate through visceral and nervous tissues of mammals. Up to date, cercarial dermatitis has been mostly associated with the cercariae of bird schistosomes of the genus Trichobilharzia. Recent findings of new genera and species indicate, however, broader spectrum of causative agents of the disease with different life cycles, host specificity and pathogenicity.     

Microsporidian xenomas in fish seen in wider perspective

The history of understanding xenoparasitic complexes or xenomas provoked in the host cell by various protists and especially by microsporidia is outlined. Microsporidia have been known to produce xenomas in oligochaetes (e.g., genera Bacillidium, Burkea, Hrabyeia, Jirovecia, species of the collective group Microsporidium), crustaceans (e.g., Abelspora, Mrazekia), insects (e.g., Polydispyrenia, Thelohania) and poikilothermic vertebrates, mostly fish (Alloglugea, Amazonspora, Glugea, Ichthyosporidium, Loma, Microfilum, Microgemma, Neonosemoides, Pseudoloma, Spraguea, Tetramicra). An overview of characters of xenomas caused by species of these genera is presented. The study of microsporidia causing xenomas in fish offers an insight into cell pathology and is of interest since many of these species are important agents of diseases in commercial fish. Xenomas produced from a few types of target cell display a complete change of organisation of the host cell and differ, according to the agent, in their structure. Recent data show that proliferation of the parasite may have already started in the cells transporting the parasites to the final site of xenoma formation. However, these are preliminary revelations and most of the facets of the life cycle are still to be clarified. Curiously, xenoma-forming microsporidia do not seem to be strictly host specific. The salient features of fish microsporidian xenomas are discussed, such as role of the xenoma, whether its features are host- or microsporidium-dependent, development and demise of the xenoma in the course of time, and host reaction phenomena. The need of further research is emphasised.     

Triaenophorus nodulosus and T. crassus in fish from northern Finland

10,288 fish specimens of 34 species from three water bodies in northern Finland were studied mainly in 1978-1982 Triaenophorus nodulosus and T. crassus parasites. Six of the 31 species from the Bothnian Bay were found to harbour T. nodulosus plerocercoids, four of the 11 from the cold, oligotrophic Lake Yli-Kitka in northeastern Finland and only one of the 6 from the small, eutrophic Lake Kuivasj?rvi. The ruff (Gymnocephalus cernuus) was most essential intermediate host in the lakes, as supported by stomach analysis of predator fish. Although the highest prevalences were found in adult burbot (Lota lota) in two of the areas, these fish are more likely a blind end in the life cycle of T. nodulosus in the present case. T. crassus larvae were found only in three coregonid species and once in the intestine of a lamprey (Lampetra fluviatilis) from the sea. Neither the prevalence nor the intensity of infection in most of the intermediate fish hosts of T. nodulosus was seen to increase with the length of the fish. The only exception was the ruff in Lake Yli-Kitka, where a sharp increase was encountered. No prominent seasonal variation in Triaenophorus larvae infections was found in any of the cases. The prevalence of T. nodulosus infection in its final host, the pike (Esox lucius), was highest in the sea (93%) and lowest in the eutrophic lake (47%). Seasonally, the lowest T. nodulosus infections were found in June-July in the two lakes. A clear maturation cycle was also found, with the youngest worms in the summer months in all cases. T. crassus was found in every third pike in the sea and its proportion in relation to T. nodulosus is higher in the smallest and largest fish. In only 12%t of the pikes did the number of T. crassus exceed that of T. nodulosus, however, the mean ratio being 1:13 to favour of T. nodulosus. The results give an indication that the composition of the fish fauna in each water body has a considerable influence on the T. nodulosus burden and its distribution among its potential intermediate fish hosts. With the most stable environmental conditions and the greatest variety of fish species, the parasite burden in the Bothnian Bay is most evenly distributed among the 6 intermediate hosts. The fewer fish species occur in the water body, as in the present eutrophic lake, the more the significance of a few or even one intermediate host will increase.     

Temporal and spatial distribution of glochidial larval stages of European unionid mussels (Mollusca: Unionidae) on host fishes

Glochidia are the larval stage of freshwater unionid mussels that parasitize the fins and gill apparatus of fish. A total of 22 fish species were examined for the presence of glochidia whose distribution on individual hosts was studied on three common fish species, the roach Rutilus rutilus (L.), perch Percafluviatilis L. and bitterling Rhodeus sericeus (Pallas). Between 1997 and 1999, the fish were obtained from the rivers Morava and Kyjovka and surrounding water pools in the Czech Republic. The glochidia of two genera, Unio and Anodonta, were found. Anodonta glochidia were observed on 10 fish species, Unio glochidia on 17 fish species. There was a difference in spatial distribution of glochidia on the body of the host fish. Unio glochidia were predominantly located on the gills, whereas most Anodonta glochidia were found on the fins, with the highest numbers of glochidia were observed on the margin of the pectoral fins. For the gill apparatus, Unio glochidia were found predominantly on the second and third arch. Anodonta glochidia were predominantly found during winter and spring (November-May), whereas Unio glochidia were more abundant during May and June. The number of glochidia was positively correlated with fish length in perch highly infected by Anodonta glochidia and perch infected by Unio glochidia. Of the three fish species, the highest occurrence of parasites was found on perch with fewer observed on roach. In spite of the close relationship between bitterling and unionid mussels, glochidiosis was rare on this fish species.     

Studies on transmission and life cycle of Enteromyxum scophthalmi (Myxozoa), an enteric parasite of turbot Scophthalmus maximus

In order to elucidate the transmission and dispersion routes used by the myxozoan parasite Enteromyxum scophthalmi Palenzuela, Redondo et Alvarez-Pellitero, 2002 within its host (Scophihalmus maximus L.), a detailed study of the course of natural and experimental infections was carried out. Purified stages obtained from infected fish were also used in in vitro assays with explants of uninfected intestinal epithelium. The parasites can contact and penetrate loci in the intestinal epithelium very quickly. From there, they proliferate and spread to the rest of the digestive system, generally in an antero-posterior pattern. The dispersion routes include both the detachment of epithelium containing proliferative stages to the intestinal lumen and the breaching of the subepithelial connective system and local capillary networks. The former mechanism is also responsible for the release of viable proliferative stages to the water, where they can reach new fish hosts. The finding of parasite stages in blood smears, haematopoietic organs, muscular tissue, heart and, less frequently, skin and gills, suggests the existence of additional infection routes in transmission, especially in spontaneous infections, and indicates the role of vascular system in parasite dispersion within the fish. The very high virulence of this species in turbot and the rare development of mature spores in this fish may suggest it is an accidental host for this parasite. This may also question the existence of a two-host life cycle involving an actinosporean stage in this species. Further studies are needed to clarify this open point of the life cycle.     

Activity levels and predator detection by amphipods infected with an acanthocephalan parasite, Pomphorhynchus laevis

The acanthocephalan parasite Pomphorhynchus laevis (Müller, 1776) uses freshwater amphipods as its intermediate host. In order to complete the life cycle, the infected amphipod must be consumed by a fish, where the acanthocephalan will mature and reproduce. Parasite transmission, and therefore fitness, could be enhanced if infected amphipods fail to detect or avoid predatory fish. We compared the activity levels of infected and non-infected amphipods, Echinogammarus stammeri (Karaman, 1931), in both the presence and absence of odours from its natural, definitive host, the fish Leuciscus cephalus (L.). Throughout the experiment, infected amphipods were more active than were non-infected individuals. The non-infected amphipods reduced their activity after the addition of fish odours, but the infected amphipods failed to show a significant decrease. The failure of infected amphipods to reduce activity levels in the presence of fish odour may reflect a parasite strategy to increase its chances of transmission by making its amphipod host more vulnerable to predation by fish.     

Differential host utilisation by different life history stages of the fish ectoparasite Argulus foliaceus (Crustacea: Branchiura)

In this study we examine differences in the occurrence of life history stages of the destructive fish ectoparasite Argulus foliaceus (L., 1758) on eight fish species (stickleback, rudd, roach, gudgeon, bream, tench, crucian carp and common carp) sampled from a mixed-species recreational fishing lake on nine occasions during late spring and summer. Total numbers ofA. foliaceus, as well as the number of larval, juvenile and adult parasite stages, from each fish were recorded along with the fish species. Lice generally exhibited an aggregated distribution approximating a negative binomial distribution. Significant differences in the prevalence, intensity and intensity frequency distribution were observed between life history stages and between host species. In general, all life history stages of A. foliaceus exhibited an over-dispersed distribution. However, larval lice did show some degree of aggregation particularly within the stickleback samples. Infection data for parasite larval stages suggested that sticklebacks are more likely to be infected than other host species. For adult lice, however, carp appeared to be the main host. We propose that A. foliaceus infection characteristics are predominantly determined by the level of host exposure to the parasite and its life history stages (larval, juvenile and adult) rather than by an innate difference in host susceptibility related to individual host factors such as immune responses. We conclude that host exposure is determined by the parasite-host behavioural interplay related to species-specific ecology and behavioural traits such as microhabitat preference and normal swimming speed.     

Gill parasites of Cephalopholis argus (Teleostei: Serranidae) from Moorea (French Polynesia): site selection and coexistence

The distribution and coexistence of gill ectoparasites of 121 specimens of Cephalopholis argus Bloch et Schneider, caught between October 1994 and October 1995, were investigated. Adults of the monogenean Benedenia sp. and copepod Hatschekia sp., the larval caligid copepod Caligus sp. (copepodite and chalimus stages), and praniza larvae of the isopod Gnathia sp. were found. All species were aggregated within the host population. Infracommunities were poor, with only 40.5% of fish infected by two parasite species. Only two individual fish harboured all the parasite species observed at the component community level. Prevalences were less than 50% and mean intensities were low (less than 6 parasites/host). No dominant parasite species were observed in the host population. The spatial distribution of each parasite species was studied on different partitions of the gill arches. Adult parasite stages that are mobile showed much overlap in their distribution, whereas temporarily attached larvae of Caligidae were more site specific. Copepodite and chalimus larvae showed niche restriction that is probably due to gregarious behaviour. Positive associations between caligid larvae reflected intraspecific interaction for site and/or resources. Each of the Caligus sp. larval stages prefers specific sites, as do the adults, which occur exclusively in the buccal cavity of the host. Infracommunities were too poor and too few to induce processes of interspecific competition.     

Parasite communities of eels Anguilla anguilla in freshwater and marine habitats in Iceland in comparison with other parasite communities of eels in Europe

Ninety-five eels from one marine and three freshwater localities in Iceland were examined for parasites. Twenty species were found, 12 from marine habitat, 12 from freshwater and 4 species were found in both habitats. These are: Eimeria anguillae, Chilodonella hexasticha, Trichodina fultoni, T. jadranica, Myxidium giardi, Myxobolus kotlani, two Zschokkella spp., Derogenes varicus, Deropristis inflata, Diplostonmum sp., Plagioporus angulatus, Podocotyle atomon, Anisakis simplex (larva), Eustrongylides sp. (larva), Hysterothylacium aduncum (larva), Raphidascaris acus (larval and adult stages), Bothriocephalus claviceps, Proteocephalus macrocephalus, and a pseudophyllidean larva. Thirteen of these species are new parasite records from Icelandic waters. The component community of marine eels was characterized by low diversity and a high dominance of a single species. Overall, seven species of helminths were observed, up to five different species occurring in an individual fish. The component community of the freshwater eels was species-poor with low diversity and relatively high dominance of single species. A between-sites difference in the freshwater eels was considerable; only Diplositonun sp. was found at more then one sampling site. Similar to previous studies, there is a total replacement of freshwater macroparasite species by marine ones in saline waters. But unlike research abroad in which species richness decreases with higher salinity, the marine eels in Iceland have considerably higher richness than the freshwater ones. The parasite communities of freshwater eels in Iceland are, in general species-poorer, less diverse and having higher Berger Parker (BP) dominance than other eel communities in Europe. Marine eels have on the other hand comparable species richness, are less diverse and with a high BP dominance.     

Importance of the life cycle in sympatric host race formation and speciation of pathogens

ABSTRACT Numerous morphological species of pathogenic fungi have been shown to actually encompass several genetically isolated lineages, often specialized on different hosts and, thus, constituting host races or sibling species. In this article, we explore theoretically the importance of some aspects of the life cycle on the conditions of sympatric divergence of host races, particularly in fungal plant pathogens. Because the life cycles classically modeled by theoreticians of sympatric speciation correspond to those of free-living animals, sympatric divergence of host races requires the evolution of active assortative mating or of active host preference if mating takes place on the hosts. With some particular life cycles with restricted dispersal between selection on the host and mating, we show that divergence can occur in sympatry and lead to host race formation, or even speciation, by a mere process of specialization, with strong divergent adaptive selection. Neither active assortative mating nor active habitat choice is required in these cases, and this may explain why the phylo-genetic species concept seems more appropriate than the biological species concept in these organisms.     

An annotated list of parasites (Isopoda, Copepoda, Monogenea, Digenea, Cestoda and Nematoda) collected in groupers (Serranidae, Epinephelinae) in New Caledonia emphasizes parasite biodiversity in coral reef fish

Abstract: Over a 7-year period, parasites have been collected from 28 species of groupers (Serranidae, Epinephelinae) in the waters off New Caledonia. Host-parasite and parasite-host lists are provided, with a total of 337 host-parasite combinations, including 146 parasite identifications at the species level. Results are included for isopods (5 species), copepods (19), monogeneans (56), digeneans (28), cestodes (12), and nematodes (12). When results are restricted to those 14 fish species for which more than five specimens were examined and to parasites identified at the species level, 109 host-parasite combinations were recorded, with 63 different species, of which monogeneans account for half (32 species), and an average of 4.5 parasite species per fish species. Digenean records were compared for 16 fish species shared with the study of Cribb et al. (2002); based on a total of 90 parasite records identified at the species level, New Caledonia has 17 new records and only seven species were already known from other locations. We hypothesize that the present results represent only a small part of the actual biodiversity, and we predict a biodiversity of 10 different parasite species and 30 host-parasite combinations per serranid. A comparison with a study on Heron Island (Queensland, Australia) by Lester and Sewell (1989) was attempted: of the four species of fish in common and in a total of 91 host-parasite combinations, only six parasites identified at the species level were shared. This suggests strongly that insufficient sampling impairs proper biogeographical or ecological comparisons. Probably only 3% of the parasite species of coral reef fish are already known in New Caledonia.