Snail size as a determinant of prepatent period duration in Helisoma anceps infected with Halipegus occidualis (Digenea: Hemiuridae)

Pulmonate snails (Helisoma anceps) of varying sizes were exposed to 2, 4, 8, or 16 eggs of Halipegus occidualis and observed over several months to determine the duration of the prepatent period. Infection probability was positively correlated with number of eggs ingested. The number of eggs ingested did not, however, have a significant effect on the duration of the prepatent period, presumably because low infectivity of the eggs (15%) dictated that most patent infections arose from a single miracidium. There was a significant, positive correlation between snail length (measured at time of exposure) and duration of prepatent period, suggesting that density-dependent effects are, in part, responsible for the initiation of cercariae production in H. occidualis.     

On the morphology and life-history of Hemiurus luehei Odhner, 1905 (Digenea: Hemiuridae)

Previously undescribed cystophorous cercariae which develop in sporocyst germinal sacs in the tectibranch opisthobranch Philine denticulata (Adams) are shown to be cercariae of Hemiurus luehei Odhner, 1905 (Hemiuridae), a common stomach parasite of clupeid and salmonid fishes off the Atlantic coast of Europe, in the Baltic and the Mediterranean Sea. The free-swimming cercariae are seized by calanoid copepods. Temora longicornis (Müller) and Acartia tonsa Dana acted as suitable experimental intermediate hosts. Pressure by the copepod mouth appendages causes delivery tube eversion and the injection of the cercarcial body into the haemocoel. Sagitta sp. was found naturally infected with a metacercaria of H. luehei. The cystophorous cercaria and metacercaria are described using light and scanning electron microscopy. Adults from herring were examined under the stereoscan electron microscope.     

Halipegus eschi n. sp. (Digenea: Hemiuridae) in Rana vaillanti from Guanacaste Province, Costa Rica

Halipegus eschi n. sp. is described from the esophagus of Rana vaillanti from Guanacaste Province, Costa Rica. The new species differs from other known species of Halipegus on the basis of relative testis size, lateral extent of the uterus, vitelline follicle arrangement, egg size, and polar filament length.     

Electrophoretic differentiation of two Halipegus (Trematoda: Hemiuridae) congeners in an amphibian population

The presence of Halipegus eccentricus, which previously had been absent from a study area, was confirmed when marked genetic differences were observed within what had been thought to be a homogenous population of Halipegus occidualis from the buccal region of the green frog, Rana clamitans. Starch gel electrophoresis was used to determine levels of genetic variation in H. occidualis and H. eccentricus. The H. occidualis population had 12% polymorphic loci and average individual heterozygosity of 0.05. Endemism of snail hosts, limited potential for parasite immigration, monoecious adults, and asexual reproduction occurring in the molluscan intermediate host may be factors that minimize the extent of genetic variability in the parasite population.     

The Acanthocolpidae (Digenea) of fishes from the north-east Atlantic: the status of Neophasis Stafford, 1904 (Digenea) and a study of North Atlantic forms

The genus Neophasis is defined. The taxonomic status of the genus is discussed in the light of its life-cycle, hosts and cercarial and adult morphology. It is considered to be closest to the family Acanthocolpidae due to the utilisation of a fish second intermediate host, the form of the cercarial excretory system and, in the adult, the presence of a uterine seminal receptacle and the absence of an external seminal vesicle. The taxonomic value of morphological features are discussed and a key to the north Atlantic species given. The following species are described: Neophasis oculatus (Levinsen) from Myoxocephalus scorpius off West Greenland (type-material), Norway and Denmark, Lycodes esmarkii off NW Scotland and the Faroes, and L. vahli off Newfoundland; N. burti n. sp. (distinguished from N. oculatus by sucker-ratio and testicular configuration) from Myoxocephalus octodecemspinosus off Nova Scotia (type-locality) and New Brunswick, also? immature in Gadus morhua from the Gulf of St. Lawrence; N. anarrhichae (Nicoll) from Anarhichas lupus in the North Sea and off the Faroes; and N. pusilla Stafford from A. lupus off ‘eastern Canada’ (type-material) and Nova Scotia. The only other species in the genus is N. symmetrorchis Machida from the NW Pacific Ocean.     

Temperature effects on survival and excystment of cercariae of Halipegus occidualis (Trematoda)

Survival of the cystophorous cercariae of Halipegus occidualis (Trematoda: Hemiuridae) from naturally infected Helisoma anceps (Gastropoda: Planorbidae) was determined following constant temperature storage at 4, 16, 22, 26 and 30 degrees C. At 4 degrees C, 90% of cercariae survived for 14 weeks and 50% survived for 30 weeks. Survival varied inversely with temperature, and at 30 degrees, 90% survived for 1.5 weeks and 50% survived for 2.5 weeks. At the time of release from snails, cercariae excysted within 0.5 s of stimulation. Excystment times ultimately increased to 1-3 s at all temperatures as length of storage increased, but no connection was detected between excystment time and the activity of the cercaria. Excystment times began increasing while survival was still above 90%. Temperature specific mortality rates of cercariae were estimated, and used to simulate the survival of cercariae released naturally at various times of the year. The results of these simulations suggest that cercariae of H. occidualis are present in the environment at all times of the year, but that cercariae present in late winter (January to mid-March) may have an impaired excystment mechanism that reduces their infectivity.     

A review of the genus Parahemiurus Vaz & Pereira, 1930 (Digenea: Hemiuridae)

The genus Parahemiurus Vaz & Pereira, 1930 (syn.: Daniella Sahai & Srivastava, 1977) is defined, its major morphological characters discussed and a key to species given. The species P. merus (Linton, 1910) (syns: P. parahemiurus Vaz & Pereira, 1930, P. sardiniae Yamaguti, 1934, P. seriolae Yamaguti, 1934, P. platichthyi Lloyd, 1938, P. atherinae Yamaguti, 1938, P. harengulae Yamaguti, 1938, P. noblei King, 1962) and P. anchoviae Pereira & Vaz, 1930 are described. Other species recognized are P. clupeae Yamaguti, 1953, P. [originally Daniella] madrasensis (Sahai & Srivastava, 1977) n. comb. (syns: P. dussumieriai Hafeezullah, 1981, P. indicus Ahmad, 1981), P. ecuadori Manter, 1940, P. engraulisi Gupta & Jahan, 1977 (syns: P. cameroni Gupta & Ahmad, 1977, P. puriensis Ahmad, 1981, P. simhai Gupta & Gupta, 1978, P. tricanthusi Gupta & Puri, 1984) and P. yanamense Hafeezullah, 1980. Forms considered species inquirendae are P. arripidis Lebedev, 1971, P. clupeae of King (1964), P. dogieli Skrjabin & Guschanskaya, 1953, P. pseudosciaenae Shen, 1985 and P. trachichthodi Lebedev, 1968. Host and locality information is given in detail for all species. The complete life-cycle is not known, but metacercariae are reported in chaetognaths and teleosts. The definitive hosts of Parahemiurus spp. most frequently reported belong in the families Clupeidae and Carangidae and the genus is most commonly reported in temperate and subtropical waters.     

Reevaluation of the systematic status of Cotylotretus grandis (Digenea: Psilostomidae) with new morphological data

The digenetic trematode Cotylotretus grandis is redescribed on the basis of 130 specimens recovered from the body cavity and small intestine of 5 roseate spoonbills Ajaia ajaja (Aves, Ciconiiformes, Threskiornithidae) captured in the Pantanal do Mato Grosso, Brazil. The spoonbill A. ajaja is the only host recorded for specimens of C. grandis since 1819, when the species was proposed by Rudolphi as Distoma grande. The synonymy of Mesaulus Braun, 1902, with Cotylotretus Odhner, 1902, is discussed and confirmed after morphological analysis of the specimens.     

Hemiuridae (Digenea) from marine fishes of the Great Barrier Reef,Queensland, Australia

The following species are described, figured and/or recorded from the Great Barrier Reef at Heron Island or Lizard Island, Queensland, with comparative material reported from other areas of the ocean around Australia or New Guinea: Dinurus longisinus (new synonym: D. hippuri) from Seriola lalandi, Heron Island and Coryphaena hippurus, Papua New Guinea; Ectenurus trachuri from Caranx sexfasciatus, Diploprion bifasciatus, Pterocaesio marri, Seriola lalandi and Atherinomorus capricorniensis, Heron Island; Erilepturus hamati (with 25 new synonyms) from Lutjanus carponotatus, Lizard Island, Platycephalus bassensis, Coff's Harbour, NSW, P. fuscus, Coff's Harbour, NSW and Moreton Bay, Queensland, P. endrachtensis, Sillago analis, S. maculata, S. ciliata, Pseudorhombus arsius and Polydactylus sp. from Moreton Bay, Queensland and Lates calcarifer, Darwin, Northern Territory; Tubulovesicula angusticauda from Echeneis naucrates and Lethrinus miniatus, Heron Island and Anguilla reinhardtii, Moreton Bay and Bribie Island, Queensland; Elytrophalloides humerus from Trachinotus botla and T. coppingeri, Heron Island; Lecithochirium kawakawa from Euthynnus affinis Heron Island and Lizard Island: Lecithochirium cirrhiti (new synonyms: L. sammarae, L. nohu) from Sargocentron rubrum, Heron Island; Lecithochirium caesionis from Pterocaesio marri, heron Island; Plerurus digitatus (new synonyms: P. cynoglossi, P. atulis, P. scomberomori) from Plectropomus leopardus, Heron Island, Lutjanus erythropterus, Variola louti, Scomberomorus semifasciatus, Grammatorcynus bicarinatus and Carangoides embureyi, C. gymnostethoides, Lizard Island, Scomberomorus commerson, Heron Island, Lizard Island, New Britain, Papua New Guinea, Point Lookout, Queensland and Moreton Bay, Queensland, Euthynnus affinis Heron Island, Lizard Island and New Britain, Papua New Guinea, Sphyraena barracuda, Heron Island and Lizard Island, Scomberomorus munroi, S. queenslandicus and Saurida undosquamis, Moreton Bay, Queensland and Chirocentrus dorab, Bundaberg, Queensland. The Lecithochirium species-group Cirrhiti is considered indistiguishable from the species-group Lotellae.     

The Ultrastructure of the Daughter Sporocyst Tegument of Cercaria vaullegeardi Pelseneer (Digenea, Hemiuridae)

The body wall of the daughter sporocyst of Cercaria vaullegeardi Pelseneer, 1906 is unusual for a species without a birth pore in consisting of an outer anucleated, microvillous syncytial tegument and a cellular subtegument which lacks secretory cells. In contrast, most other known daughter sporocysts without birth pores have a nucleated region above the microvillous region of the tegument. The significance and functional aspects of this structure are discussed.     

Hemiuridae (Digenea) from marine fishes of the southern Indian Ocean: Dinurinae,Elytrophallinae, Glomericirrinae and Plerurinae

The following species are described, figured and/or recorded from regions of the southern Indian Ocean: Dinurus longisinus from Coryphaena hippurus, Cape Province and Western Australia and immature in Carangoides gymnostethoides, Western Australia; Ectenurus trachuri from Trachurus declivis, Tasmania, Erilepturus tiegsi from Platycephalus endrachtensis, Pomatomus saltator and Arripis georgianus, Western Australia and Sphyraena novaehollandiae, Tasmania; Tubulovesicula angusticauda (with eight new synonyms) from Trachichthodes gerrardi, South Australia and Gymnothorax woodwardi, Saurida undosquamis and Argyrops spinifer, Western Australia; Elytrophalloides oatesi from Meuschenia freycineti and Aldrichetta forsteri, South Australia; Elytrophalloides humerus n. sp. from Trachinotus botla, Natal; Lecithocladium angustiovum (with four new synonyms) from Rastrelliger kanagurta, Natal; Glomericirrus amadai from Hoplostethus atlanticus, Tasmania; Plerurus digitatus (with 17 new synonyms) from Euthynnus affinis, Scomberomorus commerson, Thunnus albacares, Scomberoides commersonianus, Alectis ciliaris, Pomatomus saltator, Saurida undosquamis, Rachycentron canadum, Natal and Lichia amia and Pomatomus saltator, Cape Province and Carangoides gymnostethoides, Western Australia; Synapto-bothrium sp. from Pomatomus saltator, Western Australia.     

Redescriptions of Aphanurus stossichii (Monticelli, 1891) and A. virgula Looss, 1907 (Digenea: Hemiuridae)

Aphanurus stossichii (Monticelli, 1891) is redescribed from Boops boops (L.) from various localities in the NE Atlantic region (off Spain), the Mediterranean (off Spain and Turkey) and the Black Sea (off Bulgaria). The material from the Atlantic coast of Spain showed differences in egg-size and shape and a characteristic ornamentation of the anterior third of the hermaphroditic duct. A comparison with samples from B. boops collected off Antalya (Turkey) and off the Bulgarian Black Sea coast revealed a considerable size variation and confirmed the presence of tubercles in the terminal part of the hermaphroditic duct. A. virgula Looss, 1907, previously considered a synonym of A. stossichii, is redescribed on the basis of voucher material from Engraulis encrasicolus ponticus Alexandrov collected off the Bulgarian Black Sea coast and previously identified as A. stossichii. A. virgula can be distinguished from A. stossichii by its smaller size and substantially lower range limits for all metrical features, the more posterior position of the base of sinus-sac, and the eggs being less numerous and larger in relation to the size of the body and gonads.     

Merlucciotrema praeclarum (Manter, 1934) (Digenea: Hemiuridae) redescribed from myctophiform and gadiform fishes of the North Atlantic

Merlucciotrema praeclarum (Manter, 1934) is redescribed from the deep-sea benthic fishes Bathysaurus mollis, B. ferox and Cataetyx laticeps from various localities in the northern Atlantic Ocean. The genus is close to Plerurus Looss, 1907, but is retained on the basis of the presence of a muscular sinus-organ and, possibly, a rudimentary or degenerate sinus-sac. The subfamily Plerurinae Gibson & Bray, 1979 and the genus Merlucciotrema are redefined.     

The role of habitat in structuring Halipegus occidualis metapopulations in the green frog

The transmission dynamics of the trematode Halipegus occidualis in its definitive host, Rana clamitans, have been examined over a 5-yr period in a North Carolina pond. The breeding season of green frogs coincides with the period of worm recruitment, during which time male frogs are territorial and females show strong site fidelity. This site fidelity allows inferences to be made regarding the suitability of a particular habitat for worm transmission based on frog infection intensities within that habitat. Four foci of infection were identified in the pond by plotting worm infrapopulation size against site of host capture. Sites within infection foci are characterized by shallow water and emergent vegetation, factors favorable for overlapping distributions of the 4 hosts in the life cycle of H. occidualis. Consistent year-to-year worm prevalences and intensities, despite fluctuations in frog population size, are thought to be the result of a relatively constant proportion of the frog population being present in infection foci each year. Removal of worms from heavily infected frogs in the fifth year resulted in further heavy worm recruitment by treated frogs suggesting that site selection can predispose a frog to heavy infection. Further, the sum of removed parasites and those recruited after parasite removal by treated frog hosts was higher than worm infrapopulations observed in previous years, indicating that worm density regulates parasite infrapopulation size in heavily infected frogs.     

Description of two new species of Glypthelmins Stafford, 1905 (Digenea: Macroderoididae) in Rana spp. from Mexico, based on morphology and mtDNA and rDNA sequences

Glypthelmins Stafford, 1905 includes 29 putative species commonly found in the intestine and liver of anurans from all over the world but mainly in the Americas. Partial sequences of the cytochrome c oxidase subunit 1 ( cox 1), ribosomal internal transcribed spacer region 2 (ITS2) and the large subunit 28S rDNA gene were obtained and analysed using pairwise distance matrices and parsimony methods in order to characterise the interrelationships between 14 isolates of four nominal species of Glypthelmins recognised on morphological grounds. The highest intra-specific sequence divergence occurred in the cox 1 (18.53%) sequence, followed by that of the ITS2 (5.44%) and 28S (4.63%). Genetic variability was detected between the three isolates originally identified as G. facioi Brenes et al., 1959 from two localities in Mexico and one locality in Costa Rica. Sequence divergence exhibited among these isolates ranged from 10.70 to 11.22%, from 0.48 to 0.97% and from 1.33 to 1.88% for cox 1, ITS2 and 28S, respectively. Phylogenetic analysis combining all three data-sets generated a single most parsimonious tree. The three isolates of G. facioi form a clade, with an isolate collected from frogs in Veracruz State as the sister group to an isolate from Tabasco State + G. facioi from Costa Rica. The information derived from pairwise distance of independent data-sets plus the phylogenetic information indicate that each of the two isolates from Mexico, identified a priori as G. facioi, represent separate species. A re-examination of specimens was carried out and a re-evaluation made of the morphological characters to find reliable differences that had been overlooked. As a consequence, G. brownorumae n. sp. from Tabasco and G. tuxtlasensis n. sp. from Veracruz are described based on molecular and morphological differences.     

The Lepocreadiidae (Digenea) of fishes from the north-east Atlantic: a review of the genusLepidapedon Stafford, 1904

The genusLepidapedon is subdivided into several species groups and subgroups of species based on the vitelline distribution and the length of the excretory vesicle. The species in each of the subgroups are listed and keys to the species in most subgroups are given. The following north-eastern Atlantic species are described or redescribed:Lepidapedon rachion fromMelanogrammus aeglefinus, Gadus morhua, Aspitrigla cuculus, Merlangius merlangus andPollachius pollachius; L. cambrensis fromEnchelyopus cimbrius; L. sommervillae n. sp. fromTrachyrincus scabrus, T. murrayi andCoryphaenoides guentheri; L. elongatum fromGadus morhua; L. gaevskayae fromCoryphaenoides (Nematonurus) armatus; L. discoveryi n. sp. fromCoryphaenoides (Nematonurus) armatus; L. arlenae n. sp. fromTrachyrincus scabrus andT. murrayi; L. mariannae n. sp. fromGaidropsarus argentatus; Lepidapedon spp. innom (Elongatum-group) fromCoryphaenoides guentheri andCoryphaenoides (Chalinura) leptolepis; L. desclersae n. sp. fromLepidion eques; L. beveridgei fromCoryphaenoides (Nematonurus) armatus andC. (Chalinura) mediterraneus; andL. zubchenkoi fromCoryphaenoides (Chalinura) leptolepis andC. (C.) profundicolus. The phylogeny, host-specificity and zoogeography of the genus are briefly discussed.     

The taxonomic status,host range and geographical distribution of Dollfuschella Vercammen-Grandjean, 1960 (Digenea: Halipeginae) from Xenopus spp. (Anura: Pipidae)

The taxonomy, host range and geographical distribution of halipegine digeneans from Xenopus spp. are reviewed. Dollfuschella Vercammen-Grandjean, 1960 is reinstated from synonymy with Halipegus Looss, 1899 because of the presence of a sinus-sac and weakly developed permanent sinus-organ. Only one representative of the genus, D. rodhaini Vercammen-Grandjean, 1960, is recognised, of which H. rhodesiensis Beverley-Burton, 1963 is considered a synonym. Based on previous literature records this parasite occurs in Xenopus laevis laevis from South Africa and Zimbabwe, X. l. victorianus from Zaire and Uganda, X. l. bunyoniensis from Rwanda, X. wittei from Zaire and Uganda, X. vestitus from Uganda and X. muelleri from Zaire (in some cases host identification has been revised). New host and/or locality records are from X. l. poweri in Zambia, X. l. victorianus in Rwanda, X. l. bunyoniensis in Uganda and X. clivii in Ethiopia. All known hosts belong to a clade characterised by multiples of 2n = 36 chromosomes. There are no records of halipegines from the other major lineage within Xenopus, X. tropicalis-like species with multiples of 2n = 20 chromosomes. This latter group occurs in lowland tropical rain forest from west Africa in contrast to the hosts of D. rodhaini which are found typically in grassland and wooded savanna and in montane forest biotypes. The distribution of D. rodhaini might, therefore, be limited by phylogenetic specificity to the definitive host group or by other ecological factors (e.g., availability of suitable molluscan hosts). Its wide geographical and host range, in common with some other parasite species from the 2n = 36 Xenopus lineage, may result from the lack of ecological or geographical barriers between different definitive host species and subspecies. However, significant geographical variation in egg-size occurs between northern D. rodhaini populations (north of about 15° S) and those from X. l. laevis in southern Africa. This is not considered sufficient for taxonomic recognition but it could reflect the operation of some isolating factor: parasite divergence concurs with evidence that X. l. laevis is evolutionarily relatively distant from the other (more northerly) members of the laevis Rassenkreis.     

The occurrence of the Japanese species Dinosoma lophiomi Toman, 1973 (Digenea: Hemiuridae) in Lophius piscatorius (Teleostei) from the western Mediterranean

Hemiurid worms from the stomach of the angler Lophius piscatorius off Corsica are described and considered to belong to a Japanese species, Dinosoma lophiomi Toman, 1973, which was originally described from an Indo-Pacific lophiid host. The apparent disjunctive distribution of this species and apparent differences in the terminal genitalia between the European and Japanese specimens are discussed. This is the first record of Dinosoma Manter, 1934 from the Mediterranean Sea. The validity of diagnostic features of Adinosoma Manter, 1947 is commented upon, and lists of records of species of Dinosoma and digeneans from L. piscatorius in the Western Mediterranean are included.     

The life-cycle of three species of the Mesometridae (Digenea) with comments on the taxonomic status of this family

The intra-molluscan stages of three species of the Mesometridae Poche, 1926 are described. The corresponding adult stages are intestinal parasites of herbivorous sparid teleosts. The cercariae develop in prosobranch gastropods. The larvae of Elstia stossichianum occur in Vermetus triqueter, those of Wardula capitellata are parasites of Barleeia rubra and those of Centroderma spinosissima are found in three related rissoid hosts species: Rissoa ventricosa, R. auriscalpium and R. similis. The phylogenetic status of the Mesometridae in the Digenea is discussed in relation to larval and life-cycle characters.