Are glial cells of the Digenea (Platyhelminthes) muscle cells?

Muscle cells of a digenean fish blood fluke, Aporocotyle simplex, aggregate along the periphery of the cerebral ganglia. Solitary myocytons and sarcoplasmic processes with muscle fibres give rise to long, narrow lamellate projections, which are visible along the periphery and within ganglia. These ultrastructural observations suggest a switching of glial functions to muscle cells and represent additional evidence of the phylogenetic lability of glial cells in bilaterians.

Interrelationships of vitelline and muscle cells within the vitelline follicles of the blood fluke Aporocotyle simplex (Digenea, Aporocotylidae) and morphological evidence for the modification of vitelline material for eggshell formation.

This study of the fish blood fluke Aporocotyle simplex represents the first detailed transmission electron microscopical (TEM) investigation of the vitellarium of an aporocotylid digenean blood fluke. It revealed some unusual characteristics in the cytoarchitecture of the vitelline follicles and demonstrated modifications of the vitelline granules for eggshell formation. The vitelline follicles consist of vitellocytes at different developmental stages surrounded by sarcoplasmic processes of myocytes which occur throughout each follicle. Sites of intimate contact occur between the vitellocytes and the myocytes. Individual vitelline globules (0.1-0.2 µm in diameter) accumulate in quite small clusters of 10-20 and have a dense, heterogeneous matrix possessing central and peripheral regions with a greater density. Modifications of the vitelline globules take place within the clusters and are first apparent when the vitellocytes reach the lumen of the vitelline duct and vitelline reservoir. Globules within the clusters become confluent, and, when the vitellocytes reach the lumen of the oviduct and proximal ootype, these consolidated clusters contain a shapeless, loosely packed, dense material which is released from the vitellocytes by exocytosis. This investigation has provided morphological evidence for shell formation from modified vitelline globules in the form of a discontinuous, thin layer (~ 0.07 µm in thickness) of electron-dense shell material around the fertilized ovum and associated vitellocytes in the proximal ootype. The eggshell of intra-uterine eggs acquires an additional thin, heterogeneous outer layer, increasing its thickness to ~ 0.1 µm. The cytoarchitecture of the vitellarium, modifications of the vitelline globules within the clusters and the structure of the eggshell of A. simplex may prove to be of value in studies examining relationships between the three distinct lineages of aporocotylid digeneans.

Molecular Phylogeny and Dating Reveal a Terrestrial Origin in the Early Carboniferous for Ascaridoid Nematodes.

Ascaridoids are among the commonest groups of zooparasitic nematodes (roundworms) and occur in the alimentary canal of all major vertebrate groups, including humans. They have an extremely high diversity and are of major socio-economic importance. However, their evolutionary history remains poorly known. Herein, we performed a comprehensive phylogenetic analysis of the Ascaridoidea. Our results divided the Ascaridoidea into six monophyletic major clades, i.e., the Heterocheilidae, Acanthocheilidae, Anisakidae, Ascarididae, Toxocaridae, and Raphidascarididae, among which the Heterocheilidae, rather than the Acanthocheilidae, represents the sister clade to the remaining ascaridoids. The phylogeny was calibrated using an approach that involves time priors from fossils of the co-evolving hosts, and dates the common ancestor of the Ascaridoidea back to the Early Carboniferous (approximately 360.47-325.27 Ma). The divergence dates and ancestral host types indicated by our study suggest that members of the Ascaridoidea first parasitized terrestrial tetrapods, and subsequently, extended their host range to elasmobranchs and teleosts. We also propose that the fundamental terrestrial-aquatic switches of these nematodes were affected by changes in sea-level during the Triassic to the Early Cretaceous.

Unique ultrastructural characteristics of the tegument of the digenean blood fluke Aporocotyle simplex Odhner, 1900 (Digenea: Aporocotylidae), a parasite of flatfishes.

This paper includes the first transmission electron microscopical (TEM) study of the tegument of a member of the basal digenean family Aporocotylidae. Scanning electron microscopical investigations of the fish blood fluke Aporocotyle simplex show that each boss on the lateral body surface bears 12-15 simple, uniform spines which extend from 0.5-2.7 µm above the surface of the boss. TEM observations revealed that these spines reach deep beneath the distal cytoplasm of the tegument for much of their length (9-12 µm) and are surrounded by a complex of diagonal muscles in each boss. This is the first record of any digenean with so-called 'sunken' spines. The results suggest that aporocotylid spines arise from within the sarcoplasm of the boss diagonal muscles. The sunken cell bodies (perikarya) of the tegument are connected to the distal cytoplasm via ducts (specialised processes lined by microtubules); this in contrast to other digeneans studied, where they are connected via non-specialised cytoplasmic processes. Within the distal cytoplasm, the tegumental ducts of A. simplex are surrounded by invaginations of the basal membrane and release their cytoplasmic inclusions into the distal cytoplasm. These apparently unique morphological features of the tegument, especially the deep origin of the spines, may represent useful characteristics for understanding aporocotylid relationships, especially in view of the known variation in the spine patterns of aporocotylids.

Global spread of helminth parasites at the human-domestic animal-wildlife interface.

Changes in species distributions open novel parasite transmission routes at the human-wildlife interface, yet the strength of biotic and biogeographical factors that prevent or facilitate parasite host shifting are not well understood. We investigated global patterns of helminth parasite (Nematoda, Cestoda, Trematoda) sharing between mammalian wildlife species and domestic mammal hosts (including humans) using >24,000 unique country-level records of host-parasite associations. We used hierarchical modelling and species trait data to determine possible drivers of the level of parasite sharing between wildlife species and either humans or domestic animal hosts. We found the diet of wildlife species to be a strong predictor of levels of helminth parasite sharing with humans and domestic animals, followed by a moderate effect of zoogeographical region and minor effects of species' habitat and climatic niches. Combining model predictions with the distribution and ecological profile data of wildlife species, we projected global risk maps that uncovered strikingly similar patterns of wildlife parasite sharing across geographical areas for the different domestic host species (including humans). These similarities are largely explained by the fact that widespread parasites are commonly recorded infecting several domestic species. If the dietary profile and position in the trophic chain of a wildlife species largely drives its level of helminth parasite sharing with humans/domestic animals, future range shifts of host species that result in novel trophic interactions may likely increase parasite host shifting and have important ramifications for human and animal health.

The first ultrastructural observations of the egg-forming complex of Calicotyle affinis, an endoparasitic monocotylid monogenean which lacks a uterus.

This ultrastructural study of the female reproductive system of Calicotyle affinis, a monogenean without a uterus, demonstrates, for the first time in a monogenean, the presence of accessory cells located in the peripheral region of the ovary and a single large cell whose surface is penetrated by deep invaginations which restrict the passage of oocytes through the lumen of the distal extremity of the ovary. The cytoarchitecture of the epithelial lining of the two vaginae is a syncytium formed by an anucleate epithelial lining with sunken epithelial perykaria. The wall of the seminal receptacle is an enlarged prolongation of the vaginal epithelial lining. Fertilization occurs in the fertilization chamber and fertilized oocytes retain cortical granules within their cytoplasm, a characteristic known to occur in free-living flatworms. Our study also highlights concentrations of two distinct groups of Mehlis' gland cell ducts on either side of the proximal end of the ootype distinguished by ultrastructural characteristics of their secretory granules. The epithelial wall of the ootype is formed by a single layer of regular, columnar, glandular epithelial cells; these cells are closely adjacent, conjoined towards their apical region by septate junctions and produce rounded, electron-dense granules which are discharged into the ootype lumen via a merocrine or holocrine mechanism. Released granules concentrate around the eggshell and form an additional fibrous coat. The morphological diversity observed in the female reproductive system of the Monogenea is commented on it relation to clarifying patterns in monogenean evolution and for understanding the phylogeny of the Neodermata.

Ultrastructural characteristics of the male ducts and terminal genitalia of an endoparasitic monogenean, Calicotyle affinis Scott, 1911 (Monopisthocotylea: Monocotylidae), with the first detailed description of a copulatory stylet in a monogenean.

This investigation of Calicotyle affinis is the first ultrastructural description of the male reproductive system of a monocotylid monogenean. It has revealed numerous characteristics which are either new or clarify those determined from previous light microscopical observations on the morphology of Calicotyle spp. These include numerous individual testes, each surrounded by its own basal lamina; an ejaculatory bulb delimited by a basal lamina surrounded by two independent internal and external bands of muscles; a bipartite, but not fully divided, muscular chamber (an internal seminal vesicle) in the proximal region of the ejaculatory bulb; well-developed unicellular prostatic glands localised outside the ejaculatory bulb, the cytoplasmic extensions of which pass into the ejaculatory bulb, to form the ducts which open into the lumen of ejaculatory duct; and a male copulatory organ in the form of a coiled penis tube containing throughout its lumen a long, hollow stylet. The penis tube appears to be a modified prolongation of the ejaculatory duct and is supported by a basal lamina and muscle layers. The stylet wall consists of a homogenous, dense material with a narrow, denser inner lining; it is funnel-shaped proximally but narrows imperceptibly towards the distal end of the penis tube, and appears to be a derivative of the basal laminae of the ejaculatory bulb and distal ejaculatory duct. A comparative analysis of the nature of the stylet in relation to those found in free-living platyhelminths forms part of a search for characters useful for understanding monocotylid origins and relationships.

The unique uterine structure of the basal monogenean Chimaericola leptogaster (Monogenea: Polyopisthocotylea), an ectoparasite of the relictual holocephalan fish Chimaera monstrosa.

Ultrastructural studies of the monogenean uterus are few in number and no non-polystomatid polyopisthocotyleans have been investigated. The uterus of Chimaericola leptogaster, a basal polyopisthocotylean monogenean, has several unusual features, including six reflexed loops comprising four ascending and three descending, longitudinally oriented, linear sections. At the ultrastructural level, three readily distinguishable uterine regions and other distinctive characteristics are apparent. One novel feature occurs in the proximal uterus, where the lining forms a so-called 'single-layered multi-rowed cellular epithelium', which includes two types of cells, tall (ca. 14-19 µm in height) and short (ca. 6-9 µm in height) cells, both lying on the basement membrane. Although known from other bilaterian groups, this is the first record of this type of epithelium in the Neodermata. The lining of the middle uterine region comprises a single regular layer of columnar glandular epithelial cells, which produce numerous rounded, electron-dense granules associated with Golgi complexes. The presence of the uterine glands in the middle region of the uterus is an unusual feature for a monogenean, having previously been described only for basal orders of the Cestoda, i.e. the Gyrocotylidea, Caryophyllidea and Spathebothriidea. Seen in cross-section, the epithelium of the distal uterus contains three areas of tall single-layered columnar epithelium (ca. 30 µm deep) interspersed by three areas of flattened epithelium (ca. 0.2-0.9 µm deep). Such a pattern is quite different from those reported for other monogeneans and, indeed, other neodermatan groups. The investigation has shown that the outer layer of the fully formed eggshell is assembled from epithelial secretions in the middle uterine lumen, but is modified in terms of its shape in the distal uterus. Possible phylogenetic implications arising from the unusual features described are discussed in relation to other neodermatan groups and recent molecular phylogenies of the Bilateria.

An investigation of the host-specificity of metacercariae of species of Apophallus (Digenea: Heterophyidae) in freshwater fishes using morphological, experimental and molecular methods.

Metacercariae of species of the genus Apophallus Lühe, 1909, infecting the fins and skin of freshwater fishes, frequently cause black spot disease. Two species, Apophallus muehlingi (Jägerskiöld, 1899) and A. donicus (Skrjabin & Lindtrop, 1919), are known to occur in Hungarian fishes. It has generally been thought that metacercariae of A. muehlingi infect cyprinid fishes, whereas those of A. donicus develop in percids. As part of a morphological, experimental and molecular study, metacercariae were collected from 99 infected specimens of five cyprinid hosts (Abramis brama, Blicca bjoerkna, Chondrostoma nasus, Squalius cephalus, Scardinius erythrophthalmus) and 18 infected specimens of two percid hosts (Gymnocephalus cernua, Perca fluviatilis) in Hungarian natural waters (Lake Balaton, River Danube). Moreover, 1024 common carp (Cyprinus carpio) specimens collected from Hungarian fish ponds were investigated for Apophallus infection, but without positive results. For reliable species identification, experimental infections of chicks were carried in order to produce adult specimens from metacercariae collected from the fins and skin of the cyprinid and percid hosts. Within 8 days, adult specimens of both A. muehlingi and A. donicus developed in chicks infected with metacercariae from the cyprinid common bream (Abramis brama) and the white bream (Blicca bjoerkna) and the ruffe (Gymnocephalus cernua), a percid, respectively. The morphology of the collected metacercariae and adult individuals developed in the feeding experiments was characterised. A molecular analysis was extended to cercarial samples from the snail Lithoglyphus naticoides and to a single adult specimen of Apophallus from a fox. Sequences of 28 specimens were analysed using molecular methods (sequencing the internal transcribed spacer region and the cytochrome oxidase I subunit). Phylogenetic analysis was executed, and the Apophallus samples clustered into three distinct branches using both genes, A. muehlingi from cyprinids, A. donicus from percids and, a third, previously unknown, Apophallus clade, also from cyprinids.

Surface ultrastructural characteristics of Dictyocotyle coeliaca Nybelin, 1941 (Monopisthocotylea: Monocotylidae), an endoparasitic monogenean of rays.

Scanning and transmission electron microscopical observations were made of the surface topography of Dictyocotyle coeliaca (Nybelin, 1941) (Monopisthocotylea: Monocotylidae), a unique endoparasitic monogenean from the body cavity of the ray Amblyraja radiata (Elasmobranchii: Rajidae). Scanning investigation show the presence on the ventral side of the anterior body of smooth areas with pit-like depressions and shallow ridges, whereas the tegument of the middle and posterior regions of the body is extensively folded, and on the dorsal surface the tegument is smooth but interrupted by deep depressions. Transmission observation revealed the presence of invaginations of varying irregularity and depth, which form the various depressions of the tegumental surface. Non-ciliated, dome-shaped papillae occur singly or in groups and are common around the mouth and both the genital and vaginal pores, but exhibit no particular orientation. The haptor is much reduced, compared with that of ectoparasitic monocotylids, and is covered ventrally with an irregular array of 40-70 shallow loculi. Internal differences occur in the thickness of the syncytial tegumentary layer of the haptoral loculi and septa and also in the number of cytoplasmic inclusions. The locular surface has distinct ultrastructural characteristics, the most obvious of which are a honeycomb arrangement of small pockets measuring 0.6 × 0.75 to 1.0 × 1.7 µm in diameter, a terminal web beneath the surface plasma membrane and large outgrowths on the surface of the loculi filled with lysosome-like bodies. The surface specializations of the body and haptoral tegument of D. coeliaca are discussed in relation to their being adaptations to an endoparasitic environment.

Ultrastructural observations of the attachment organs of the monogenean Rajonchocotyle emarginata (Olsson, 1876) (Polyopisthocotylea: Hexabothriidae), a gill parasite of rays.

The present study uses scanning and transmission electron microscopical observations to examine the haptoral features of the hexabothriid polyopisthocotylean monogenean Rajonchocotyle emarginata from the gills of the elasmobranch Amblyraja radiata. The haptor possesses six equal, pedunculate suckers, each armed with a large, curved, hooked sclerite, and an appendix bearing two small, terminal suckers and a pair of minute hamuli. The outer side of the distal region of the hooked sclerite forms a large bulge along the antero-lateral side of each sucker and the proximal region forms a small bulge on the opposite postero-lateral side; part of the large bulge, which covers the small sclerite hook, surmounts the sucker rim. The sucker sclerite is situated outside the main sucker muscle complex of the sucker wall and is surrounded by its own muscular envelope; close to the sclerite itself, long, tightly packed lamella-like processes of sarcoplasm are present. A muscular sphincter surrounds the distal region of the sclerite shaft. The sclerite consists of three heterogeneous layers, but its hook region is distinguished from the shaft by the presence of an additional homogeneous layer. Hamuli within the appendix consist of three distinct layers loosely surrounded by sarcoplasmic lamellae. The luminal surface of the appendicular suckers is covered with unusual, long, thin, interconnected surface protrusions. In all other regions of the haptor, the tegumental surface is smooth, with numerous shallow pores penetrating the "terminal web" of the syncytial cytoplasm. Unicellular gland cells are localized close to the base of the appendix, with their ducts, containing non-homogenous secretory bodies, opening onto the anterior haptoral surface. Two types of sense receptors are visible on the haptoral surface. The ultrastructural features of the haptor are discussed in relation to our understanding of their function and the evolutionary relationships of the basal polyopisthocotylean monogenean groups Hexabothriidae and Chimaericolidae.

An annotated catalogue of the ascaridoid nematode parasites of Chinese vertebrates.

A catalogue, based on both examined specimens and the published literature, of all the ascaridoid nematodes recorded in China is presented. A total of 95 recognised species, representing 26 genera in five families, are reported. Detailed information on the type-host, type-locality, original reference, synonyms, annotated subsequent references of taxonomic importance, other host records, site of infection, location of type-specimens and distribution are listed for each recognised species. Additional comments on the taxonomic status of some species are also given. Moreover, some nomenclatural changes are proposed: (i) Toxascaris selenarctis Wang, 1965 and T. ailuri Wu, He & Hu, 1987 are placed in synonymy with Baylisascaris transfuga (Rudolphi, 1819); (ii) Raphidascaris lophii Wang & Wu, 1991 is a secondary homonym of R. lophii (Wu, 1949) and a replacement name, R. wangi nom. nov., is proposed for the former species; (iii) Aliascaris aetoplatea Luo, 2001 is transferred to Terranova Leiper & Atkinson, 1914, as T. aetoplatea (Luo, 2001) n. comb., and should be considered a species inquirenda; (iv) Ophidascaris orientalis (Wang, 1965) is resurrected as a valid species; (v) Phocascaris longispiculum Wang & Wu, 1991 and Ophidascaris agkistrodontis Wang, 1979 are treated as incertae sedis; and (vi) Hysterothylacium sauridae Li, Xu & Zhang, 2008 is listed as a nomen nudum.

The Mediterranean: high discovery rates for a well-studied trematode fauna.

Our knowledge of trematode diversity in Mediterranean Sea fishes is based on many contributions since the early 1800s (e.g. by Rudolphi, Stossich, Looss, Bartoli, Bray and Gibson). We have updated data from the Natural History Museum Host-Parasite Database and listed 302 digenean trematode species allocated to 146 genera in 29 families from 192 fish species (27% of the known fish-fauna) belonging to 76 families. The most diverse (with 31-41 species) digenean families (Opecoelidae Ozaki, 1925, Didymozoidae Monticelli, 1888 and Hemiuridae Looss, 1899) represent more than a third of the total richness (36% of species) and have been most frequently recorded (43% of the records). The overall mean number of species per host is close to recent global estimates for digenean richness in teleosts (1.57 vs 2.04, respectively), indicating a high diversity in the Mediterranean. The most diverse host families are also the best studied. However, three speciose host families (Rajidae de Blainville, Gobiidae Cuvier and Myctophidae Gill) appear under-studied and no digenean reports exist for 94 of 169 fish families present in the Mediterranean. Thus, although Mediterranean fishes appear well studied, further efforts are needed. Nevertheless, the descriptions of a large number of new taxa since 2000 indicate that focused efforts have resulted in a high discovery rate (2.4 species per year). Many of these new (often cryptic) taxa are the result of combined morphological and molecular methods, which promise more reliable estimates of digenean diversity in this region. We provide host-parasite lists for 192 species of fish in the Mediterranean comprising 890 host-parasite associations.

Ultrastructural characteristics of the caeca of basal polyopisthocotylean monogeneans of the families Chimaericolidae and Hexabothriidae parasitic on cartilaginous fishes.

Ultrastructural differences are shown between the caecal organization in three blood-feeding polyopisthocotylean monogeneans, i.e., the chimaericolid Chimaericola leptogaster from the holocephalan Chimaera monstrosa and two hexabothriids, Callorhynchocotyle callorhynchi from the holocephalan Callorhynchus capensis and Rajonchocotyle emarginata from the elasmobranch Amblyraja radiata. In C. leptogaster, digestive cells and connecting syncytium, joined close to the luminal surface by septate junctions, are arranged alternately along the caecal epithelial wall; the nuclear regions of both cell types are sunk below the general level of the caecal epithelium; a concave depression on the apical margin of the digestive cells bears lamellae; and this depression is covered by a lamellate bubble formed by thin projections emanating from the connecting syncytium. The luminal surface of the connecting syncytium is covered with outgrowths terminating in the form of long, narrow processes. In R. emarginata and C. callorhynchi, the predominant digestive cells are at different stages of development and occur in groups, developing digestive cells bulge into the caecal lumen from the connecting syncytium with contact sites present close to the luminal surface, and the luminal surface structures of both the connecting syncytium and the digestive cells are short lamellae. In these two hexabothriids, a holocrine (or apocrine) process for the elimination of digestive product is assumed via the detachment of fully differentiated, bulging digestive cells. Free, apparently sloughed digestive cells and residual bodies are present within the caecal lumen, and replacement digestive cells are numerous in the connecting syncytium. In the chimaericolid, free bubbles containing residual bodies and portions of digestive cells filled with degenerating digestive vesicles occur in the caecal lumen along with large amounts of male and female reproductive material. The usefulness of characteristics of the caecal ultrastructure as taxonomic traits at the family level is discussed.

The surface topography of Callorhynchocotyle callorhynchi (Manter, 1955) (Monogenea: Hexabothriidae), a parasite of the holocephalan fish Callorhinchus capensis.

A scanning electron microscopical study, incorporating some transmission electron microscopical observations, was undertaken on the surface topography of the gill parasite Callorhynchocotyle callorhynchi (Manter, 1955) (Monogenea: Hexabothriidae) from the Cape elephant fish Callorhinchus capensis (Holocephali) off the western coast of South Africa. The study revealed the presence of several new characteristics for this species. These include the presence of regularly distributed, knob-shaped projections on the surface of the haptor, haptoral appendix and sucker peduncles measuring 0.2 µm and in concentrations of approximately 100 per 10 µm(2) and the existence of a ridge which bisects each sucker lumen, forming two different loculi. We also report, for the first time for any monogenean, the presence of 'true spines'; these occur on the luminal surface of the haptoral suckers and have all of the characteristics of the tegumental spines of digeneans, i.e. they are situated within the distal syncytial tegumental cytoplasm, rest on the basal plasma membrane, have a uniform structure and are covered apically by the tegumental surface plasma membrane. These spines are simple, straight and single-pointed. Under the scanning electron microscopy (SEM), within an area of 20 µm in diameter, 23 such spines were counted, but their concentration and arrangement varies in different regions of the sucker. At their base, they measure about 1.5 µm in width and reach approximately 2 µm in height above the general level of the tegument, but transmission electron microscopy (TEM) measurements of an entire spine indicate that they may reach 3 µm in total length. The presence of spines, possessing similar morphological characteristics in both basal polyopisthocotylean monogeneans and digeneans, represents another characteristic which may prove useful in understanding the evolutionary relationships within the Neodermata.

The occurrence of metacercariae of Petasiger (Digenea: Echinostomatidae) in an unusual site, within the lateral line scales of cyprinid fishes.

During a regular veterinary inspection of fishes from Lake Balaton, Hungary, echinostomatid metacercariae (Digenea), with collar spines characteristic of species of the genera Petasiger Dietz, 1909 and Paryphostomum Dietz, 1909, were found in the lateral line scales of a roach Rutilus rutilus (Linnaeus), an apparently unique site. In a subsequent examination of 586 fishes from 20 different species, similar infections were found in 11 species. The infection was virtually restricted to the lateral line scales, other scales being infected only incidentally. These encysted metacercariae had 27 collar spines, including eight larger angle spines and 19 smaller dorsal spines arranged in two rows. Two types of metacercarial cyst were found. One type had a cyst diameter of 138-171 µm × 105-120 µm and three central dorsal spines that were larger than the remainder and tended to resemble the angle spines. The second type of metacercarial cyst had a diameter of 128-157 µm × 105-115 µm and all 19 dorsal spines of the metacercaria were of a similar size. ITS sequences of the second type of metacercaria exhibited a 100% similarity to sequences of two adult Petasiger phalacrocoracis (Yamaguti, 1939) specimens collected from the gut of Phalacrocorax carbo (Linnaeus) in Hungary and to P. phalacrocoracis deposited in the GenBank database. Sequences obtained from two metacercariae of the first type showed a 2.8-2.9 % difference from sequences of the second type of metacercaria and from those of adult specimens of P. phalacrocoracis from cormorants. Based on these results, the second type metacercaria is considered to be a larval stage of P. phalacrocoracis, but the identity of the first type is uncertain. The unusual location of these metacercariae in the lateral line scales is discussed in relation to their transmission.

Clamp ultrastructure of the basal monogenean Chimaericola leptogaster (Leuckart, 1830) (Polyopisthocotylea: Chimaericolidae).

The ultrastructure of the haptoral clamps of the chimaericolid monogenean Chimaericola leptogaster, a basal polyopisthocotylean from the gills of a holocephalan fish, is described. These clamps are characterized by the presence of two muscle blocks interrupted mid-anteriorly and mid-posteriorly and different kinds of hard structures: a single median and paired lateral sclerites embedded in the clamp wall; six spine-like structures directed towards the clamp lumen; and electron dense surface structures along the internal surface of the anterior clamp lips and along the luminal surface of the tegument of the clamp lumen. The lateral sclerites are situated deep within muscular tissue and are closely bounded by radial myofibrils, possessing a uniform electron dense matrix within which are hollow areas of different sizes. The median sclerite occupies an area between the clamp wall myofibrils and the luminal epithelium, is surrounded by a basement lamina and is composed of a heterogeneous matrix comprising two different morphological layers related to variations in the type and concentration of fibrils. Four of the spine-like structures are extensions of the margins of the two spindle-like muscle blocks in the clamps, i.e. the two anterior and two posterior structures, and the two others are situated at the lateral constrictions of the left and right muscle blocks. The electron dense surface structures are derivations of the clamp tegument or, to be more precise, its outer, densely fibrous region. These results are discussed in relation to the evidence that the haptoral clamps of C. leptogaster are apparently ancient origin.

Synopsis of the species of the genus Myxobilatus Davis, 1944 (Myxozoa: Myxosporea: Myxobilatidae).

A synopsis of the species of Myxobilatus Davis, 1944 (Myxozoa: Myxosporea: Myxobilatidae) is presented. Thirty-nine nominal species are included. The major characteristics and an illustration are given for each species based on the original records.

Lamellodiscus aff. euzeti Diamanka, Boudaya, Toguebaye & Pariselle, 2011 (Monogenea: Diplectanidae) from the gills of Cheimerius nufar (Valenciennes) (Pisces: Sparidae) collected in the Arabian Sea, with comments on the distribution, specificity and historical biogeography of Lamellodiscus spp.

Specimens of Lamellodiscus Johnston & Tiegs, 1922 (Monogenea: Diplectanidae) were collected from the gills of Cheimerius nufar (Valenciennes) (Sparidae) in the Arabian Sea. All of these parasites belonged to one and the same species, which is morphologically very close to L. euzeti Diamanka, Boudaya, Toguebaye & Pariselle, 2011. A different host, distant locality and small morphological differences compared with the original description of L. euzeti acted as a stimulus for a detailed redescription. The specimens from the Arabian Sea differ slightly in the details of the male copulatory organ (MCO) from the type-specimens of L. euzeti, which were re-examined, and from the respective drawings in its original description. Such differences include a longer inner process of the large element of the accessory piece associated with the proximal part of the copulatory tube, a longer point on the small element of the accessory piece associated with the distal part of the copulatory tube, and the presence of a smooth or slightly folded inner margin of this element rather than structures resembling spines which occur in the type-specimens of L. euzeti. Therefore, the present specimens infecting C. nufar in the Indo-Pacific may represent a different, but morphologically very similar species to the Atlantic form L. euzeti; consequently, they are recognised here as Lamellodiscus aff. euzeti. This form belongs to the 'ignoratus s. str.' subgroup of the genus. The composition of this subgroup is redefined to comprise 17 species, including L. corallinus Paperna, 1965 but excluding L. acanthopagri Roubal, 1981, and the morphology of the MCO of representatives of this group is clarified. A link between the diversity of Lamellodiscus species and the ancestral origin of present-day sparid species in the Tethys Sea is suggested. It is shown that Lamellodiscus spp. exhibit rather high levels of specificity to their hosts, since half of them parasitise only a single host species and c.90% infect closely related host species. Comparison of the levels of host-specificity of the species of this genus with other narrowly specific genera of the Dactylogyridea revealed that their estimations are comparable. The possibility of intra-host speciation within Lamellodiscus is discussed. It is shown that a co-evolutionary model is more discernible if it includes data on the occurrence of morphologically similar species from different regions and host taxa.

Ultrastructural characteristics of the vaginae of the basal monogenean Chimaericola leptogaster (Leuckart, 1830).

In the first study of the vaginal ultrastructure of any monogenean, the paired vaginae of Chimaericola leptogaster, a basal polyopisthocotylean from the gills of a holocephalan, are described. Each vagina opens laterally. A unique feature of this relict parasite is a short vagino-intestinal connecting duct, the lining of which is separated by septate junctions from the linings of the vagina and the intestine. After giving rise to this vagino-intestinal duct, each vagina travels in a posterior direction and opens into the vitelline collecting duct. The lining of each vagina close to the vaginal pore resembles the body tegument, the syncytial lining of which measures about 2.5 µm in thickness and has a dense surface layer 0.2 µm in thickness and different apical projections. Proximal to this and before the entrance to the vaginal-intestinal duct, the vaginal lining is characterised by: uniformly distributed electron-dense, lanceolate, spine-like surface structures of about 1.7 µm in length; three kinds of surrounding glands with three kinds of secretory granules (abundant oval or rounded electron-dense granules 0.7-2.5 µm in length, much less numerous spheroidal, vesicular granules 0.3-0.4 µm in diameter, and tightly packed, spheroidal granules 0.8 µm in diameter and containing fine particulate material of low density); and four kinds of sensory receptors (three uniciliate and one aciliate). These features are absent from the rest of the vagina. The likely roles of the vagino-intestinal connection, the different gland cells, sensory structures, and the armament of the distal vaginal regions are discussed. Considering the suggested polyopisthocotylean ancestor for the Neodermata, the relict parasite fauna of holocephalans and the unique vagino-intestinal connection in C. leptogaster, which is also known in turbellarians, an ancient origin for chimaericolids is supported.