A new species of Monocotyle taschenberg, 1878 (Monogenea: Monocotylidae) from Dasyatis guttata (Dasyatidae)

Monocotyle guttatae n. sp. is described from the gills of the ray Dasyatis guttata (Bloch and Schneider, 1801) (Dasyatidae) off the coast of Brazil The new species can be readily differentiated from the other species of the genus in having a male copulatory organ with 2 loops and an accessory piece, 5-7 sclerites on the marginal haptoral papillae, and the absence of accessory sclerites on the dorsal surface of the posterior body. The present record confirms the presence of the genus in the subtropical waters of South America.     

Evolutionary expansion of the Monogenea

The evolutionary expansion of the monogeneans has taken place in parallel with the diversification of the fish-like vertebrates. In this article the main trends in monogenean evolution are traced from a hypothetical skin-parasitic ancestor on early vertebrates. Special consideration is given to the following topics: early divergence between skin feeders and blood feeders; diversification and specialization of the haptor for attachment to skin; transfer from host to host, viviparity and the success of the gyrodactylids; predation on skin parasites and camouflage; colonization of the buccal and branchial cavities; diversification and specialization of the haptor for attachment to the gills; phoresy in gill parasites; the development of endoparasitism and the origin of the cestodes; the success of dactylogyroidean gill parasites; the uniqueness of the polyopisthocotyleans; ovoviviparity and the colonization of the tetrapods. Host specificity has been the guiding force of coevolution between monogeneans and their vertebrate hosts, but the establishment of monogeneans on unrelated hosts sharing the same environment (host-switching) may have been underestimated. Host-switching has provided significant opportunities for evolutionary change of direction and is probably responsible for the establishment of monogeneans on cephalopod molluscs, on the hippopotamus and possibly on chelonians. There are indications that host-switching may be more common in monogeneans that spread by direct transfer of adults/juveniles from host to host. A limitation on the further expansion of monogeneans is the need for water for the dispersal of the infective larva (oncomiracidium).     

Host specificity of monogenean ectoparasites on fish skin and gills assessed by a metabarcoding approach

Monogeneans are highly diverse fish ectoparasites with a direct life cycle, widely distributed, and are known to generally display strict host specificity. Factors related to the hosts and the parasite have been suggested to explain this high specificity. Monogeneans have also been observed to colonise fish species not in their natural host range under experimental conditions. We developed a specific metabarcoding protocol and applied it on the Sparidae-Lamellodiscus host-parasite system, to assess parasite diversity on skin and gills of several sparid host species. We first demonstrated that the use of a metabarcoding approach provided a better understanding of the diversity of monogeneans associated with teleost skin and gills than traditional approaches based on morphological identification. We identified a high diversity of both expected and unexpected (never observed on this host species) Lamellodiscus spp. on each host species and on skin and gills. No significant difference in parasite diversity was found between skin and gills. These results suggest that the establishment of the observed host specificity in monogeneans relies on multiple levels of regulation, involving the survival capacity of the larvae and host recognition mechanisms.     

<Emphasis Type="Italic">Malalophus jensenae</Emphasis> n. g., n. sp. (Monogenea: Monocotylidae) from the gills of <Emphasis Type="Italic">Aetomylaeus vespertilio</Emphasis> (Myliobatidae) off northern Australia

Malalophus jensenae n. g., n. sp. is described from the gills of the ornate eagle ray Aetomylaeus vespertilio (Bleeker) collected off the eastern coast of the Gulf of Carpentaria, northern Queensland. The new genus is similar to Heliocotyle Neifar, Euzet & Ben Hassine, 1999, with which it shares a haptor bearing seven peripheral loculi and a single dorsal haptoral accessory structure. M. jensenae can be distinguished from species of Heliocotyle by the presence of numerous sclerotised sinuous ridges covering the ventral surfaces of the peripheral loculi of the haptor. It also lacks pseudosepta which are present on the haptor of Heliocotyle species. This is the first published record of a monogenean from an elasmobranch in the Gulf of Carpentaria, Australia.     

Furcohaptor cynoglossi n. g., n. sp., an ancyrocephaline monogenean gill parasite with a bifurcate haptor and a note on its adhesive attitude

An ancyrocephaline monogenean,Furcohaptor cynoglossi n. g., n. sp., is described from the gills of the marine teleost flatfishCynoglossus macrostomus caught off the coast of Kerala State in India. The parasite is also recorded onC. puncticeps. The haptor divides to form two elongated and slender haptoral arms, each bearing at the distal end a cluster of relatively small sclerites, comprising the following: two hamuli, one of which is rod-shaped and the other of typical hooked shape; a bar articulating with the typical hamulus; five marginal hooklets of typical shape, each with a domus; a sixth marginal hooklet which is shorter and more slender than the others and lacks a domus. This sixth marginal hooklet is hard to locate in some individuals. The haptoral arms embrace a primary gill lamella, their distal hook-bearing regions meeting or overlapping on the side of the lamella opposite to the parasite's body. Species of the genusBifurcohaptor Jain, 1958 differ in that two of the hamuli are greatly enlarged and support the haptoral arms.     

The surface topography of <Emphasis Type="Italic">Eudiplozoon nipponicum</Emphasis> (Monogenea) developmental stages parasitizing carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis> L.)

Using scanning electron microscopy (SEM), the external morphology of all developmental stages (egg, oncomiracidium, diporpa, just fused juvenile and adult) of the parasite, Eudiplozoon nipponicum (Monogenea, Diplozoidae), from the gills of carp was studied. During the ontogeny, the tegument, tegumentary and sensory structures are subsequently developed. The tegument of free swimming oncomiracidium occurs in two types — the ciliated and non-ciliated with numerous uniciliated sensory structures. An attachment apparatus starts to form during the oncomiracidium stage. Further developmental stages are adapted to the environment of the gills. Tegumentary folds become more apparent later in development and assist to the parasite’s attachment. In connection with its reproductive strategy, the two morphological structures of diporpa (ventral sucker and dorsal papilla) appear to play important role. On the gills, two individuals need to meet and these structures mediate the fusion between two diporpae. The hindbody of adult parasite is highly modified for attachment. The haptor, folds and lobular extensions are most developed. The forebody is flexible and able to interact with host gill tissue via the mouth and associated mouth structures. The process of food intake of the parasite was discussed.     

A new species of Dermopristis Kearn, Whittington & Evans-Gowing, 2010 (Monogenea: Microbothriidae), with observations on associations between the gut diverticula and reproductive system and on the presence of denticles in the nasal fossae of the host Glaucostegus typus (Bennett) (Elasmobranchii: Rhinobatidae)

Dermopristis cairae n. sp. (Monogenea: Microbothriidae) is described from the skin and possibly from the nasal fossae of the giant shovel-nosed ray Glaucostegus typus (Bennett). The new species is distinguished from D. paradoxus Kearn, Whittington & Evans-Gowing, 2010 by its larger size, body shape, lack of transverse ridges on the ventral surface and absence of a seminal receptacle. Extensive short gut branches lie dorsal to the testes and adjacent to the coiled region of the vas deferens and the o?type, possibly reflecting high metabolic demand in these areas. Denticles are present in the lining of the nasal fossae of G. typus, providing a firm substrate for the cement-based attachment of a microbothriid. However, confirmation that D. cairae inhabits the nasal fossae of G. typus is required.     

Origin and evolution of the hamuli in the monogeneans

The hypothesis of the origin and evolution of the hamuli in monogeneans is proposed. It is suggested that the hamuli originated as the adult attachment organs of protomonogeneans inhabited the gills of the first vertebrates. Primarily they were represented by two lateral pairs of large hooks disposed anterior to the larval haptor. The fundamental direction in the evolution of monogeneans was the concentration of all attachment structures on the growing haptor. It the course of this evolutionary process, the hamuli onchoblasts migrated to the haptor, in which they had reached the position in the hind part of the haptor. The neotenic evolution of the Dactylogyridea and Gyrodactyloidea resulted in the forming new hamuli pairs. The hooks of opposite sides of the haptor are joined in a single complex within each side by the transverse plates (bars). So the presence of 4 hamuli is plesiomorphy for all the monogeneans and the presence of the transverse bars and new hamuli pairs are apomorphy for the Dactylogyridea and Gyrodactyloidea, whose evolution was linked with that of the Teleostei. The origin of the new hamuli pairs and transverse bars in the Dactylogyridea and Gyrodactyloidea appears to be a convergence.     

Lamellodiscus sanfilippoi n. sp. (Monogenea, Diplectanidae) parasite from the gills of Diplodus sargus (Teleostei, Sparidae) in Mediterranean Sea

Lamellodiscus sanfilippoi n. sp. takes place, among the other species of Lamellodiscus, in the "ergensi" sub-group (Amine et Euzet, 2005) characterized by the morphology of the dorsal lateral bars of the haptor. This sub-group comprises, in the Mediterranean, L. ergensi Euzet and Oliver, 1966, L. kechemirae Amine and Euzet, 2005, L. tomentosus Amine and Euzet, 2005, all parasite of Diplodus sargus, and L. baeri Oliver, 1974 parasite of Pagrus pagrus. L. sanfilippoi can be distinguished from the previous species by the morphology and size of the dorsal lateral bars. The new species is close to Lamellodiscus furcillatus Kritsky, Jiménez-Ruiz and Sey, 2000, a parasite of Diplodus noct in the Persian Gulf, but differs by the size of the haptoral sclerotised pieces and the morphology of the male copulatory apparatus. Lamellodiscus gussevi Sanfilippo (1978) et Lamellodiscus abbreviatus Sanfilippo (1978) are considered as nomina nuda.     

Abnormalities of the attachment clamps of representatives of the family Diplozoidae

A comparative study has been made of the haptoral morphology of four species of diplozoon (Monogenea: Diplozoidae) from the gills of fish exposed to different levels of water pollution in two river systems in eastern Europe. An examination of the haptors of Paradiplozoon homoion (Bychowsky & Nagibina 1959), Paradiplozoon ergensi (Pejcoch 1968) and Paradiplozoon megan (Bychowsky & Nagibina 1959) from chub caught in the River Morava, Czech Republic and of Diplozoon paradoxum (Nordmann 1832) from bream recovered from the River Volga, Russia has revealed abnormalities to the attachment clamps. Two abnormal conditions were found: structural alterations to the attachment clamps and changes in the number of attachment clamps; these occurred both singly and in combination. A higher frequency of abnormal attachment clamps was found in diplozoons from fish caught in the more polluted localities of both rivers. The abnormalities have been recorded and their morphology compared in the light of conditions of environmental stress.     

Neotropical Monogenoidea. 54. Proposal of Aetheolabes n. g. (Dactylogyrinea: Diplectanidae), with the description of A. goeldiensis n. sp. from the gills of ‘pescada’ Plagioscion sp. (Teleostei: Sciaenidae) in Brazil

Aetheolabes goeldiensis n. g., n. sp. (Diplectanidae) is described from the gills of ‘pescada’ Plagioscion sp. (Sciaenidae) collected from the Baía de Marajó, about 30 km north of Belém, Pará, Brazil. The monotypic Aetheolabes n. g. is characterised, in part, by its type-species having the haptor and haptoral sclerites modified as a clasp for attachment to the gill tissue of its host, the copulatory complex situated far posterior to the intestinal bifurcation near the mid-length of the trunk, the vaginal pore apparently within the genital atrium, the tegument lacking scales, anchors atypical for diplectanids, and by lacking peduncular spines and squamodiscs. A. goeldiensis n. sp. closely resembles Diplectanum umbrinum Tripathi, 1957 from India and China by the haptoral sclerites forming a clasp, but differs from it primarily by the orientation of the reproductive organs and absence of squamodiscs.     

Septesinus gibsoni n. g., n. sp. (Monocotylidae: Heterocotylinae), from the gills of Himantura walga (Dasyatidae) off Sarawak, Borneo

Septesinus gibsoni n. g., n. sp. (Monocotylidae: Heterocotylinae) is described from the gills of the dwarf whipray Himantura walga (Müller & Henle) collected in marine waters off Sarawak (Borneo), Malaysia. Septesinus n. g. is distinguished from other genera in the Monocotylidae by a combination of characters, including a haptor with one central and seven peripheral loculi, the presence of a highly sinuous ridge surmounting all haptoral septa, four rounded accessory structures on the dorsal surface of the haptor, and the anterior region with two pairs of anteromedian and three pairs of anterolateral gland-duct openings. Septesinus n. g. is accommodated in the Heterocotylinae. Septesinus gibsoni n. sp. is described and fully illustrated, and a key to the genera of Heterocotylinae is provided. The composition of the ridges surrounding the mouth of a number of heterocotyline species and their usefulness as a taxonomic character are examined. The identity of four specimens of Monocotyle Taschenberg, 1878, also recovered from the gills of this host species, is discussed.     

The egg bundles of the monogenean Dionchus remorae and their attachment to the gills of the remora, Echeneis naucrates

Gills of three remoras, Echeneis naucrates L., from Heron Island, Queensland, Australia had few adults but many attached egg bundles of the monogenean, Dionchus remorae. Studies of fresh, and preserved and cleared, primary gill lamellae bearing egg bundles and investigations with scanning electron microscopy and serial wax sections reveal proliferated host epithelium surrounding and embedding part of a loop of egg-shell material to which eggs are tethered, but eggs in each bundle hang free of gill tissue. This hyperplasia appears to anchor egg bundles to the host's gills. However, hyperplasia will take time to develop and cannot play a part in tethering newly laid egg bundles. Possible advantages to the parasite by attaching eggs to the gills of its host include improved oxygenation of the eggs, and a reduced risk of egg predation. Egg attachment by D. remorae to their remora hosts seems well adapted to successful larval invasion of fish which exhibit phoresy.     

A new Neocalceostomatid (Monogenoidea) from the gills of the blackfin sea catfish, Arius jella (Siluriformes: Ariidae), in the Bay of Bengal, India

Thysanotohaptor n. gen. (Neocalceostomatidae) is proposed to accommodate Thysanotohaptor rex n. sp. collected from the gills of the blackfin sea catfish Arius jella Day (Siluriformes: Ariidae) from off the coast of Visakhapatnam, Bay of Bengal, Andhra Pradesh, India. Thysanotohaptor is differentiated from the other known neocalceostomatid genera by its species having multiple postgermarial testes (single testis in species of Neocalceostoma and Neocalceostomoides ), lacking a transverse bar associated with the ventral anchor pair (present in species of Neocalceostoma ), and possessing a disc-shaped haptor with a pleated marginal frill (frill absent in Neocalceostomoides spp.; Neocalceostoma spp. with delicate marginal membranes). The Neocalceostomatidae is considered valid within the Order Dactylogyridea based on its members having a haptor armed with 10 marginal and 4 ventral hooks and a germarium having a distal loop prior to uniting with the ootype; the family is not assigned to a suborder of Dactylogyridea because of uncertainty in part about the way in which the distribution of haptoral hooks evolved within the taxon.     

Dendromonocotyle lasti n. sp. from the skin and Monocotyle caseyae n. sp. (Monogenea: Monocotylidae) from the gills of Himantura sp. (Dasyatidae) in Moreton Bay, Queensland, Australia

Seven specimens of rays of the genus Himantura which could not be identified to species were collected from waters near Dunwich, Stradbroke Island, Moreton Bay, Queensland, Australia. The five smallest specimens of Himanturasp. (disc width 218-302 mm; four female, one male) had a banded tail and the dorsal surface was uniformly grey/brown. The two largest individuals of Himantura sp. (disc widths 460, 533 mm; female and male, respectively) also had a banded tail but the grey/brown dorsal surface had white spots. Two new monogenean species (Monocotylidae: Monocotylinae) are described from both the plain and white-spotted specimens of Himantura. Dendromonocotyle lastin. sp. is distinguished from other species in the genus by the number of papillae on the haptor, by the morphology of the male copulatory organ and by the morphology of the proximal portion of the vagina. The muscular sheath which surrounds the male copulatory organ is also unique having sclerotised spines at the distal end. Dendromonocotyle species are skin parasites, but a total of five juvenile specimens of D. lasti were found on the gills of four rays. Monocotyle caseyae n. sp. from the gills is characterised by the morphology of the male copulatory organ and its accessory piece. One specimen of M. spiremae Measures, Beverley-Burton & Williams, 1990, originally described from the gills of Himantura fai Jordan & Seale off Heron Island, Great Barrier Reef, Queensland, Australia, was also found on the gills of one Himantura specimen. The site and host-specificity of the parasites and the identity of the hosts are discussed.     

The attachment of the monogenean parasite Callorhynchicola multitesticulatus to the gills of its holocephalan host Callorhynchus milii

Llewellyn J. and Simmons J. E. 1984. The attachment of the monogenean parasite Callorhynchicola multitesticulatus to the gills of its holocephalan host Callorhynchus milii. International Journal for Parasitology14: 191–196. C. multitesticulatus when juvenile (up to about 10 mm long) attaches itself by four pairs of clamps to the secondary gill lamellae of its host the elephant fish Callorhynchus milii. However, in adult parasites the haptor invades the host tissue and the stalk connecting the haptor with the body-proper becomes surrounded by a sleeve of fibrous host inflammatory reaction tissue. The stalk is provided with annuli with anteriorly-directed flanges which are pressed into the walls of the sleeve of host tissue when the diameter of the stalk increases as a result of the contraction of well-developed sub-tegumentary longitudinal muscles.     

A new diplectanid (Monogenea) genus and species from the gills of the black snook, Centropomus nigrescens (Perciformes: Centropomidae) of the Pacific coast of Mexico

Cornutohaptor nigrescensi n. sp. (Diplectanidae) is described from the gills of the black snook, Centropomus nigrescens (Perciformes: Centropomidae) from the Pacific coast of Mexico. Cornutohaptor n. gen. is proposed for this new species and is characterized by possessing 2 intestinal ceca terminating blindly; a germarium looping right intestinal cecum; bilobed testis; 2 seminal vesicles; 7 pairs of hooks, each with protruding thumb; a grooved ventral bar and coiled male copulatory organ (MCO); an accessory piece comprising a "baglike structure" with an appendage; dorsal bars associated parallelly to body midline; and no adhesive accessory organs on the haptor. Cornutohaptor differs from all confamilial genera by including species with anchors with straight and deep root longest, hook pair 1 reduced in size, MCO with counterclockwise rings, and by the morphology of the accessory piece. Cornutohaptor nigrescensi most closely resembles species of Murraytrema Price, 1937, Lobotrema Tripathi, 1937, and Murraytrematoides Yamaguti, 1958, because of the absence of squamodiscs or lamellodiscs on the haptor and tegumental scales on the posterior portion of the body. Cornutohaptor differs from these genera in the position and number of haptoral bars (2 bars in Lobotrema spp., dorsal bars transversally associated in Murraytrema and Murraytrematoides spp.) and in having a coiled MCO (copulatory organ is a comparatively straight, poorly sclerotized tube in Murraytrematoides spp.). This is the first diplectanid described from a centropomid along the Pacific coast of Mexico.     

On the ontogeny of the haptor and the evolution of the Monogenea

Data on the ontogeny of the posterior haptor of monogeneans were obtained from more than 150 publications and summarised. These data were plotted into diagrams showing evolutionary capacity levels based on the theory of a progressive evolution of marginal hooks, anchors and other attachment components of the posterior haptor in the Monogenea (Malmberg, 1986). 5 + 5 unhinged marginal hooks are assumed to be the most primitive monogenean haptoral condition. Thus the diagrams were founded on a 5 + 5 unhinged marginal hook evolutionary capacity level, and the evolutionary capacity levels of anchors and other haptoral attachement components were arranged according to haptoral ontogenetical sequences. In the final plotting diagram data on hosts, type of spermatozoa, oncomiracidial ciliation, sensilla pattern and protonephridial systems were also included. In this way a number of correlations were revealed. Thus, for example, the number of 5 + 5 marginal hooks correlates with the most primitive monogenean type of spermatozoon and with few sensillae, many ciliated cells and a simple protonephridial system in the oncomiracidium. On the basis of the reviewed data it is concluded that the ancient monogeneans with 5 + 5 unhinged marginal hooks were divided into two main lines, one retaining unhinged marginal hooks and the other evolving hinged marginal hooks. Both main lines have recent representatives at different marginal hook evolutionary capacity levels, i.e. monogeneans retaining a haptor with only marginal hooks. For the main line with hinged marginal hooks the name Articulon-choinea n. subclass is proposed. Members with 8 + 8 hinged marginal hooks only are here called Proanchorea n. superord. Monogeneans with unhinged marginal hooks only are here called Ananchorea n. superord. and three new families are erected for its recent members: Anonchohapteridae n. fam., Acolpentronidae n. fam. and Anacanthoridae n. fam. (with 7 + 7, 8 + 8 and 9 + 9 unhinged marginal hooks, respectively). Except for the families of Articulonchoinea (e.g. Acanthocotylidae, Gyrodactylidae, Tetraonchoididae) Bychowsky's (1957) division of the Monogenea into the Oligonchoinea and Polyonchoinea fits the proposed scheme, i.e. monogeneans with unhinged marginal hooks form one old group, the Oligonchoinea, which have 5 + 5 unhinged marginal hooks, and the other group form the Polyonchoinea, which (with the exception of the Hexabothriidae) has a greater number (7 + 7, 8 + 8 or 9 + 9) of unhinged marginal hooks. It is proposed that both these names, Oligonchoinea (sensu mihi) and Polyonchoinea (sensu mihi), will be retained on one side and Articulonchoinea placed on the other side, which reflects the early monogenean evolution. Except for the members of Ananchorea [Polyonchoinea], all members of the Oligonchoinea and Polyonchoinea have anchors, which imply that they are further evolved, i.e. have passed the 5 + 5 marginal hook evolutionary capacity level (Malmberg, 1986). There are two main types of anchors in the Monogenea: haptoral anchors, with anlages appearing in the haptor, and peduncular anchors, with anlages in the peduncle. There are two types of haptoral anchors: peripheral haptoral anchors, ontogenetically the oldest, and central haptoral anchors. Peduncular anchors, in turn, are ontogenetically younger than peripheral haptoral anchors. There may be two pairs of peduncular anchors: medial peduncular anchors, ontogentically the oldest, and lateral peduncular anchors. Only peduncular (not haptoral) anchors have anchor bars. Monogeneans with haptoral anchors are here called Mediohaptanchorea n. superord. and Laterohaptanchorea n. superord. or haptanchoreans. All oligonchoineans and the oldest polyonchoineans are haptanchoreans. Certain members of Calceostomatidae [Polyonchoinea] are the only monogeneans with both (peripheral) haptoral and peduncular anchors (one pair). These monogeneans are here called Mixanchorea n. superord. Polyonchoineans with peduncular anchors and unhinged marginal hooks are here called the Pedunculanchorea n. superord. The most primitive pedunculanchoreans have only one pair of peduncular anchors with an anchor bar, while the most advanced have both medial and lateral peduncular anchors; each pair having an anchor bar. Certain families of the Articulonchoinea, the Anchorea n. superord., also have peduncular anchors (parallel evolution): only one family, the Sundanonchidae n. fam., has both medial and lateral peduncular anchors, each anchor pair with an anchor bar. Evolutionary lines from different monogenean evolutionary capacity levels are discussed and a new system of classification for the Monogenea is proposed.In agreeing to publish this article, I recognise that its contents are controversial and contrary to generally accepted views on monogenean systematics and evolution. I have anticipated a reaction to the article by inviting senior workers in the field to comment upon it: their views will be reported in a future issue of this journal. EditorIn agreeing to publish this article, I recognise that its contents are controversial and contrary to generally accepted views on monogenean systematics and evolution. I have anticipated a reaction to the article by inviting senior workers in the field to comment upon it: their views will be reported in a future issue of this journal. Editor