Accessory Sclerites Research Articles

A new Acrulia species from Danish amber (Coleoptera: Staphylinidae: Omaliinae: Omaliini)

ABSTRACT Acrulia danica Shavrin sp. n. from late Eocene Danish amber is described and illustrated based on a female holotype. The new species, embedded within a small piece of amber, was scanned using X-ray micro-computed tomography (μCT), to enable the detailed study of its morphology including the taxonomically important structure of the accessory sclerite. Previously, only three extant species of the genus Acrulia Thomson, 1858 were known from the Palaearctic Region. The new species is compared with extant congeners and the importance of a detailed study of the fossil entomofauna of the Danish amber is discussed in view of the hypotheses about its different origin from other European ambers. urn:lsid:zoobank.org:act:67972F3A-DF6A-48D5-A4BD-96EF8AA87DCF

Chromosomal Relationships of Two New Species in the Simulium vernum Macquart Species-Group (Diptera: Simuliidae) from Nepal

The polytene chromosomes and larvae of two species of black flies, Simulium gandakiense new species and S. magnacentrum new species, are described from the Himalaya Mountains of Nepal. Twenty-three chromosomal rearrangements were discovered in the two species, of which eight are shared. The species differ by four fixed inversions and by having unique sex chromosomes. Additionally, S. gandakiense has B chromosomes and S. magnacentrum has a chromocenter. Both species have a set of four fixed chromosomal inversions and accessory sclerites in the larvae, all of which are shared with members of the S. vernum species-group in the Oriental Region, suggesting a biogeographic connection between the Himalayas and the Oriental Region for the S. vernum species-group.

Two species of Dioncopseudobenedenia Yamaguti, 1965 (Monogenea: Capsalidae) from Naso unicornis (Forsskål) (Acanthuridae) and Coryphaena hippurus Linnaeus (Coryphaenidae) in western Japan, with a description of D. elongata n. sp. from N. unicornis.

Three species of Dioncopseudobenedenia Yamaguti, 1965 (Monogenea: Capsalidae: Benedeniinae) have been described, all found parasitising Naso spp. (Acanthuridae) and Siganus lineatus (Siganidae), in Hawaii, the Great Barrier Reef, and New Caledonia in the Pacific. In this paper, D. elongata n. sp. is described, found parasitising the gills of N. unicornis (Forsskål) caught off Okinawa-jima Island. Dioncopseudobenedenia kala Yamaguti, 1965 is also reported, found parasitising the gills of N. unicornis, and those of a new host, the common dolphinfish Coryphaena hippurus Linnaeus (Coryphaenidae), off Okinawa and Kochi prefectures, Japan. The new species differed from known species in both nuclear (28S rDNA and ITS1-5.8S-ITS2) and mitochondrial DNA (cox1) sequences. Morphologically, the new species differed from other species of Dioncopseudobenedenia in possessing a longer accessory sclerite and a thick, short penis. No species of Dioncopseudobenedenia have been detected during past examinations of C. hippurus; this instance was probably an accidental infection. A key for identification of the species of Dioncopseudobenedenia is also provided herein.

Light and transmission electron microscopy of the haptoral sclerites of the monogenean gill parasite Macrogyrodactylus clarii.

The present study represents the first detailed description of the haptoral sclerites of Macrogyrodactylus clarii Gussev 1961. Light microscopy reveals outgrowths of the hamuli roots; two lateral spine-like extensions of the hood-like accessory sclerites; a ridged, fan-shaped distal end of an accessory sclerite; and two thread-like accessory sclerites with biforked ends associated with the pointed hooked region of each hamulus. Transmission electron microscopy (TEM) reveals the presence of parallel tubules in the root of the hamuli and structural differences along the length of the hamulus including the root, shaft, and pointed hooked region. The root consists of two layers, the shaft four layers and pointed hooked region only one layer with dense outer serrations. Characteristic features of the hamulus root are the presence of longitudinally orientated parallel tubules in its central core and parallel electron-dense ridges in the outer layer of its middle region; features not observed in either the shaft or the pointed hooked region. Each hamulus blade of M. clarii is associated with haptoral gland cells producing electron-dense secretory bodies. The 16 marginal hooklets each consist of a blade (sickle) articulating with a handle at the guard region and a domus. TEM revealed structural differences between the handle, the blade at the articulation region, and the distal hooked region. The domus, a filamentous thread-like sclerite at the light microscope level, consists of two electron-dense, fibrous thickenings connected to each other by a cytoplasmic process. Each marginal hooklet is associated with a small cavity and a large reservoir of homogeneous particles and secretory bodies. The possible functions of these structures are discussed in relation to equivalent features in other monogeneans.

Muscular Arrangement and Sclerite Morphology in the Haptor of Tetraonchus monenteron (Monogenea, Dactylogyridea).

Haptoral musculature of T. monenteron was studied using phalloidin staining and confocal microscopy, with sclerites visualized in the reflection confocal mode. Haptoral armature of T. monenteron consists of ventral and dorsal pairs of anchors, a ventral bar, eight pairs of marginal hooks and at least three pairs of accessory sclerites. Anchors are operated by 14 muscles, of which the most prominent are extrinsic muscles, the transverse muscle interconnecting the ventral anchors, three muscles connecting the ventral anchor to the ventral bar, and four muscles of the dorsal and ventral anchors inserting on the haptoral wall. The extrinsic muscles are attached to the braceshaped sclerites, which in turn are connected to the dorsal anchors via muscle bundles. The gaffing action of the dorsal anchors is likely to be achieved by the extrinsic muscles and the transverse muscle that clamps the extrinsic muscles against the body wall. The ventral anchors are probably held in attached position by the transverse muscle and four muscles inserting on the ventral bar and haptoral wall. The haptoral musculature may have potential utility for tetraonchid taxonomy.

Three new species of Gyrodactylus von Nordmann, 1832 described from Goodea atripinnis (Pisces: Goodeidae), an endemic freshwater fish from the central highlands of Mexico.

Goodea atripinnis Jordan, 1880 has a broad range of habitats and is the most widespread species of the endemic goodeid fishes, which inhabit the central highlands of Mexico. This species is known to be host to a high diversity of helminth parasites from which only three belong to the genus Gyrodactylus von Nordmann, 1832: G. lamothei Mendoza-Palmero, Sereno-Uribe et Salgado-Maldonado, 2009, G. mexicanus Mendoza-Palmero, Sereno-Uribe et Salgado-Maldonado, 2009, and G. tomahuac Rubio-Godoy, Razo-Mendivil, García-Vásquez, Freeman, Shinn et Paladini, 2016. Here, we describe three new species of Gyrodactylus collected from G. atripinnis, which were characterised morphologically (sclerites of the attachment apparatus) and molecularly (sequences of the internal transcribed spacer region of the rDNA): Gyrodactylus iunuri n. sp., Gyrodactylus katamba n. sp. and Gyrodactylus tepari n. sp. These new species were collected in three different states in the Mexican Highlands: Guanajuato, Jalisco and Querétaro. Both morphological and molecular data support the hypothesis that two distinct groups of gyrodactylids infect goodeid fishes: G. iunuri n. sp., G. tepari n. sp. and G. tomahuac possess robust hamuli and are closely related phylogenetically; while G. katamba n. sp. resembles G. lamothei in having slender hamuli with accessory sclerites adjacent to the hamuli root, and apparently shares a common ancestor with gyrodactylids infecting poeciliid fishes. New locality records of G. tomahuac are presented. The addition of the three new species of Gyrodactylus as parasites of G. atripinnis makes monogeneans the second most abundant parasite group known to infect this host.

Redescription ofAllobenedenia dischizosepta(Suriano, 1975) n. comb. (Monogenoidea: Capsalidae: Trochopodinae) from the Gills of Argentine Sea Basses (Acanthistius) in the Southwestern Atlantic Ocean

We herein redescribe Allobenedenia dischizosepta (Suriano, 1975) n. comb. (Monogenoidea: Capsalidae) based on light microscopic analysis of the holotype and 18 paratypes from the gill of Acanthistius brasilianus (Cuvier, 1828) (Perciformes: Serranidae) along with light and scanning electron microscopy of 25 newly collected specimens from the gill of Acanthistius patachonicus (Jenyns, 1842) (new host record) in the southwestern Atlantic Ocean off Argentina. Contradicting the original description of this species and its generic diagnosis, all specimens had 5 loculi (2 anterior plus 3 posterior loculi) plus 5 corresponding haptoral septa only (central loculus absent). The original description of this species erroneously specified 6 loculi, including 2 small anterolateral loculi. Because A. dischizosepta was the type species for Tetrasepta Suriano, 1975 and because it fits the diagnosis of Allobenedenia Yamaguti, 1963, the genus therefore is regarded a junior synonym of Allobenedenia. Allobenedenia dischizosepta differs from its congeners by lacking a haptoral peduncle as well as by having 5 haptoral loculi, accessory sclerites that lack a conspicuous dorsal protuberance and that are weakly bifid, by posssesing robust anterior hamuli that are strongly recurved for half of their total length, by having a male accessory gland reservoir in the extreme posterior portion of the penis sac and lateral to the vitelline reservoir abutting the ovary, and by the presence of a straight (not coiled) proximal portion of the ejaculatory duct within the penis sac.

A new species of Tereancistrum (Monogenea, Dactylogyridae) from the gills of three Leporinus species (Characiformes, Anostomidae) and a revised description of Tereancistrum parvus

The present study describes Tereancistrum flabellum n. sp. (Dactylogyridae, Ancyrocephalinae) from the gills of the anostomid fishes Leporinus friderici, Leporinus amblyrhynchus and Leporinus elongatus from two freshwater ecosystems in the south east of Brazil. This new species is mainly characterized by the morphology of the copulatory complex (such as the MCO base formed by two fan-shaped structures, and accessory piece flattened, curved, rigid and channeled), a dorsal anchor with a well-developed superficial and inconspicuous deep root, and the shape of the accessory anchor sclerite with small spathulate termination. Tereancistrum flabellum n. sp. is the first record of a dactylogyrid from L. amblyrhynchus. The description of Tereancistrum parvus is also emended to correct and complement previous descriptions and the species is reported for the first time in Schizodon nasutus.

Monocotyle luquei n. sp. (Monogenea: Monocotylidae), from the gills of diamond stingray Dasyatis dipterura (Jordan and Gilbert, 1880) (Myliobatiformes: Dasyatidae), in the South Pacific.

Monocotyle luquei n. sp. (Monogenea: Monocotylidae) was described from gills of diamond stingray Dasyatis dipterura (Jordan and Gilbert, 1880) (Dasyatidae) off Peru. The new species can be differentiated from the other species of the genus by the combination of the following characteristics: (1) accessory sclerites on the dorsal posterior surface of the body absent, (2) only one testis is present, (3) 1-2 loops in the copulatory organ, (4) the male copulatory organ with a sclerotized accessory piece, (5) shape of five sclerites in marginal papillae, (6) size of anchor and (7) posterolateral septa bifurcated. This is the first record of species of Monocotyle Taschenberg, 1878 from the southern Pacific.

Trophic structure and function in the larva of predatory muscid flies (Diptera, Muscidae)

Assessed multiple times over a 100-year period, yet poorly understood, we provide a new view of trophic structure and function in predatory muscid larvae based on Phaonia goberti (Mik) and Phaonia subventa (Harris) (Diptera, Muscidae). Trophic structure and function were investigated by morphological analysis, direct observation and filming. Larvae search for prey using a compartmentalised body. The rear compartment grips the substrate, while the middle one turns to the sides and the front one grabs prey. Two feeding mechanisms were recorded, sucking and lunging. Sucking occurs when the head is anchored inside the prey, and fluids are ingested using the pump in the head skeleton. Lunging is the head moving forwards and backwards, coordinated with lowering and raising of the mandibles during which fluids and tissue are gathered into the cup-shaped, oral cavity prior to sucking in. These mechanisms rely on prey being pierced, and this is achieved by a remarkable structural and functional partnership between the pseudocephalon, mandibles and accessory sclerites. The partnership involves tightening the integument of the prey between the oral bars and disengaging the mandible hooks to pierce it. Due to connections between the oral bars and pseudocephalon, the hooks do not extend from their sheaths except when piercing, an unusual feature in larvae of the Cyclorrhapha (Diptera).

New Records of Species of Macrovalvitrematidae Yamaguti, 1963 (Monogenea) from Puerto Rico Including One New Genus and Two New Species.

New records of Macrovalvitrematidae Yamaguti, 1963 are reported from Puerto Rico including Buccamagna archosarga n. gen., n. sp. from the western Atlantic sea bream Archosargus rhomboidalis, Pseudotagia pomadasys n. sp. from the roughneck grunt Pomadasys corvinaeformis, and new locality records for the species Neopterinotrematoides avaginata Suriano, 1975 and Macrovalvitrema argentinensis (Suriano, 1975) n. comb. from the whitemouth croaker Micropogonias furnieri . Buccamagna n. gen. is based on the anterior end and the oral disc being much larger and wider than width of the body proper, by having a pair of rectangular and sclerotized oral suckers, by lacking a pharynx, by having an excessive narrowness in the proximal area of both valves, and by having both valves of the 3 pairs of large clamps armed with several accessory sclerites. Pseudotagia pomadasys n. sp. is distinguished by the absence of serration of lateral sclerites and the relative size of the clamps, having the 2 larger pairs positioned centrally in the haptor and the 2 smaller pairs in the outer positions. Macrovalvitrematoides and Neomacrovalvitrema are synonymized with Macrovalvitrema, which is amended to include Macrovalvitrema micropogoni (Pearse, 1949) n. comb. and Macrovalvitrema argentinensis (Suriano, 1975) n. comb. based on recognition of identical clamp morphologies. In this study the clamp is emphasized as the key taxonomic character for macrovalvitrematid genera. Except for Pseudotagia, the Macrovalvitrematidae have distinctively longer than wide clamps (at least twice as long as wide); with the dorsal valve larger and with a more-complex skeletal structure (g1, g2, i, k, and f sclerites) than ventral valve (c and d sclerites); and 2 clamp morphologies. Characteristics not previously described but present in all macrovalvitrematid monogeneans except Pseudotagia included: (1) asymmetrical plate (plate b) along the ventral valve with a thin fissure on one side, and (2) mirror image arrangement of the pairs of clamps.

Sperm transfer in monogenean (platyhelminth) parasites.

There are three major groups of parasitic platyhelminths (flatworms). The digeneans and cestodes are endoparasites, while the monogeneans are ectoparasites mostly on the gills or skin of fishes. Monogeneans are hermaphrodite and, with the exception of the gyrodactylids, mostly protandrous, the male reproductive system maturing before the female system. Their ectoparasitic life-style provides unique opportunities to observe the reproductive biology of living platyhelminths, opportunities restricted in digeneans and cestodes by their endoparasitic habits. Moreover, the male copulatory organs (MCOs) of monogeneans are of special interest because of their perplexing diversity, ranging from sclerotised penis tubes, many with accessory sclerites, to cirruses and genital atrium armature (hooks and spines). The relatively few accounts in the literature of mating in monogeneans are reproduced in this review, together with consideration of the following aspects of sperm transfer: structure and function of MCOs; self-insemination; spermatophores and pseudospermatophores; "hypodermic" and transtegumental insemination; tissue fusion; glands associated with MCOs and vaginae; finding a mating partner.

Histopathology associated with haptor attachment of the ectoparasitic monogenean Neobenedenia sp. (Capsalidae) to barramundi, Lates calcarifer (Bloch).

Journal of Fish DiseasesVolume 38, Issue 12 p. 1063-1067 Short Communication Histopathology associated with haptor attachment of the ectoparasitic monogenean Neobenedenia sp. (Capsalidae) to barramundi, Lates calcarifer (Bloch) A Trujillo-González, Corresponding Author A Trujillo-González James Cook University, Centre for Sustainable Tropical Fisheries and Aquaculture, College of Marine and Environmental Sciences, Townsville, Qld, AustraliaCorrespondence A Trujillo-González, Marine Parasitology Laboratory, Centre for Sustainable Tropical Fisheries and Aquaculture and the School of Marine and Tropical Biology, James Cook University, Townsville, Qld 4811, Australia (e-mail: alejandro.trujillogonzalez@my.jcu.edu.au)Search for more papers by this authorL K Johnson, L K Johnson James Cook University, Centre for Biosecurity in Tropical Infectious Diseases, College of Public Health, Medical & Vet Sciences, Townsville, Qld, AustraliaSearch for more papers by this authorC C Constantinoiu, C C Constantinoiu James Cook University, Centre for Biosecurity in Tropical Infectious Diseases, College of Public Health, Medical & Vet Sciences, Townsville, Qld, AustraliaSearch for more papers by this authorK S Hutson, K S Hutson James Cook University, Centre for Sustainable Tropical Fisheries and Aquaculture, College of Marine and Environmental Sciences, Townsville, Qld, AustraliaSearch for more papers by this author A Trujillo-González, Corresponding Author A Trujillo-González James Cook University, Centre for Sustainable Tropical Fisheries and Aquaculture, College of Marine and Environmental Sciences, Townsville, Qld, AustraliaCorrespondence A Trujillo-González, Marine Parasitology Laboratory, Centre for Sustainable Tropical Fisheries and Aquaculture and the School of Marine and Tropical Biology, James Cook University, Townsville, Qld 4811, Australia (e-mail: alejandro.trujillogonzalez@my.jcu.edu.au)Search for more papers by this authorL K Johnson, L K Johnson James Cook University, Centre for Biosecurity in Tropical Infectious Diseases, College of Public Health, Medical & Vet Sciences, Townsville, Qld, AustraliaSearch for more papers by this authorC C Constantinoiu, C C Constantinoiu James Cook University, Centre for Biosecurity in Tropical Infectious Diseases, College of Public Health, Medical & Vet Sciences, Townsville, Qld, AustraliaSearch for more papers by this authorK S Hutson, K S Hutson James Cook University, Centre for Sustainable Tropical Fisheries and Aquaculture, College of Marine and Environmental Sciences, Townsville, Qld, AustraliaSearch for more papers by this author First published: 17 October 2014 https://doi.org/10.1111/jfd.12320Citations: 9Read the full textAboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Citing Literature Volume38, Issue12December 2015Pages 1063-1067 RelatedInformation

Simulium (Nevermannia) khunklangense, a new species of black fly (Diptera: Simuliidae) from Chiang Mai, Thailand.

Simulium (Nevermannia) khunklangense sp. nov. is described from females, males, pupae and larvae collected in Doi Inthanon National Park, Chiang Mai, Thailand. This new species is placed in the vernum species-group of the subgenus Nevermannia and is similar to S. (N.) chomthongense Takaoka & Srisuka described from Doi Inthanon National Park, Thailand, but is distinguished in the male by the number of enlarged upper-eye facets and the relative width of the hind basitarsus against the hind tibia and femur, and in the pupa by the short common basal stalk of the gill and the cocoon with an anterodorsal bulge or a short anterodorsal projection. Taxonomic notes are provided to separate this new species from five other known species of the vernum species-group, which share an accessory sclerite on the larval abdomen, a rare characteristics in this species-group.

A new species of Araucnephia (Diptera: Simuliidae) from the Northeast Region of Brazil

We describe and illustrate the pupa, larva, and both adults of a new species in the genus Araucnephia Wygodzinsky &Coscarón, 1973 (A. cearensis nov. sp.). The species was found in a highland area in Ceará State, Northeast Brazil. The pupa canbe separated from those of other Araucnephia pupae based on the number of gill filaments, which range from 25 to 35 branches(versus 9 or 14), and by it being completely encased by the cocoon. The larva can be separated from those of other Araucnephiabased on the presence of a median accessory anal sclerite. Adults are typical of the genus. This species extends the northern range of the genus Araucnephia, which is postulated to be Gondwanan in origin.

Redescription and New Host Record of Capsala laevis (Monogenoidea: Capsalidae: Capsalinae) from Gill of Roundscale Spearfish, Tetrapturus georgii (Perciformes: Istiophoridae) in the Northwestern Atlantic Ocean

Specimens of a capsalid collected from the gill arches of 2 roundscale spearfish, Tetrapturus georgii Lowe, 1840, (Perciformes: Istiophoridae), captured in the northwestern Atlantic Ocean were identified as Capsala laevis (Verrill, 1875) Johnston, 1929 by having the combination of papillae on the ventral surface of haptor, dorsomarginal body sclerites in a single column extending the entire body length, haptoral accessory sclerites, conical papillae distributing over the ventral body surface, and an anterior attachment organ with a fimbriated posterior margin. The new specimens plus the holotype were used to conduct a taxonomic redescription of C. laevis using light and scanning electron microscopy. We documented that the holotype (USNPC No. 7179) and the new specimens of C. laevis from roundscale spearfish each had papillae on the ventral surface of the anterior attachment organs and sensory papillae on the dorsal body surface. Although data are insufficient at this time to justify proposal of a new species, the new specimens differed from the holotype and published accounts of C. laevis by having a sinistral dorsomarginal patch comprising 27-35 sclerites whereas the holotype has a dorsomarginal patch comprising 60 sclerites. Capsala laevis morphologically most closely resembles Capsala ovalis (Goto, 1894) Price, 1938 , but can be most easily differentiated from it by having dorsomarginal body sclerites. This represents the first record of any parasite from the recently taxonomically resurrected roundscale spearfish, long considered by some as a junior subjective synonym of white marlin, Tetrapturus albidus Poey, 1860 and, concomitantly, a new host record for Capsalidae Baird, 1853. An updated list of host records for C. laevis is provided. A perusal of that literature reveals that the identity of the type host for C. laevis is indeterminate beyond Istiophoridae species and that subsequent reports of the type host as ' T. albidus ' are presumptuous (originally reported in 1875 by Verrill as "bill-fish" only). Our results indicated that 2 records of C. laevis from the swordfish, Xiphias gladius Linnaeus, 1758, (Perciformes: Xiphiidae) are dubious, i.e., study of the museum voucher USNPC No. 8154 indicates that Linton's 1940 record from the northwestern Atlantic Ocean likely represents a new species of Capsala Bosc, 1811 and that the Kayiş et al. 2010 record from the Aegean Sea likely depicts a species of Capsaloides Price, 1938.

Revision of Dioncopseudobenedenia Yamaguti, 1965 including the description of D. ancoralis sp. n. (Monogenea: Capsalidae) from Pacific acanthuroid teleosts

Dioncopseudobenedenia Yamaguti, 1965 (Monogenea: Capsalidae) is redefined. Dioncopseudobenedenia kala Yamaguti, 1965 (type species) is redescribed from type material from Hawaii and from new specimens from Heron Island, Queensland, Australia and New Caledonia. We made detailed observations on D. macracantha Yamaguti, 1968 from type material from Hawaii, and from new material from Heron Island and New Caledonia. Dioncopseudobenedenia ancoralis sp. n. is described from the gill chamber of Siganus lineatus (Valenciennes) from Green Island and Heron Island, Australia and from New Caledonia. This study confirms that only one pair of large central sclerites is present on the haptor in Dioncopseudobenedenia species. The male copulatory organ in species of Dioncopseudobenedenia is a penis contained in a fluid-filled space (= penis canal) with weakly muscular walls. Dioncopseudobenedenia kala and D. ancoralis bear a sclerite at the tip of the penis. In D. macracantha, the structure of the penis, which has no terminal sclerite, indicates it may combine the functions of a penis and a cirrus. Dioncopseudobenedenia is compared with Calicobenedenia Kritsky et Fennessy, 1999, the other capsalid genus with a single pair of large sclerites on the haptor. The large haptoral sclerites in species of Dioncopseudobenedenia resemble accessory sclerites, whereas those of C. polyprioni Kritsky et Fennessy, 1999 resemble hamuli. Observations of oncomiracidia confirmed that the large haptoral sclerites in D. kala are accessory sclerites. Haptoral morphology suggests that different Dioncopseudobenedenia spp. employ different means of attachment. Mating behaviour was observed twice between two different pairs of D. kala specimens from Heron Island. Two preserved specimens from Nouméa, New Caledonia had structures near the dorsal vaginal pore that we interpret as spermatophores. This is the first report of spermatophores in a capsalid inhabiting the gill chamber. The geographic distribution of Dioncopseudobenedenia spp. is discussed.

Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963 is redescribed from new material collected from ventral skin surfaces of the Australian cownose ray, Rhinoptera neglecta Ogilby (Elasmobranchii: Myliobatidae) from waters in Moreton Bay and from captive rays in a public display aquarium in Townsville, from R. cf. neglecta and Rhinoptera sp. from Weipa, Queensland, Australia and from R. javanica Müller & Henle caught in the Arabian Gulf north of Dubai, United Arab Emirates. Benedeniella posterocolpa (Hargis, 1955) Yamaguti, 1963 is redescribed from new specimens collected from the cownose ray, R. bonasus (Mitchill) (Myliobatidae) from several localities in the Gulf of Mexico and in the Chesapeake Bay region of the south-eastern and eastern USA, respectively. Detailed anatomical redescriptions demonstrate that each Benedeniella species share several dorsal structures (paired anterior horns, excretory papillae and posterior conical structures at the junction of the body with the haptor), anterolateral grooves on the ventral surface of each anterior attachment organ, a similar route for tendons associated mostly with the accessory sclerites and the anterior hamuli and a long vagina. The different path of the vagina and the position of the vaginal pore are the simplest characters to discriminate the two species. Benedeniella unnithani Gupta & Chanana, 1976 from the gills of a Caranx sp. (Teleostei: Perciformes: Carangidae) off Kavaratti, Laccadive Islands, Arabian Sea, India is considered a species incertae sedis. Morphological parallels are identified, in particular concerning the anterior attachment organs of species in Benedeniella, Branchobdella Kearn, Whittington & Evans-Gowing, 2007, Entobdella Blainville in Lamarck, 1818, Neoentobdella Kearn & Whittington, 2005 and Pseudoentobdella Yamaguti, 1963. Of these entobdelline genera, species in three of them parasitise principally the skin of elasmobranch rays (Dasyatidae; Myliobatidae; Rhinobatidae); only species in Branchobdella and Entobdella parasitise flatfish teleosts (Paralichthyidae; Pleuronectidae; Soleidae). On the basis of the redescriptions presented here plus morphological similarities, host associations and recently published molecular genetic data, Benedeniella is moved from the Benedeniinae Johnston, 1931 and the diagnosis for the Entobdellinae Bychowsky, 1957 is amended to unite Benedeniella, Branchobdella, Entobdella, Listrocephalos Bullard, Payne & Braswell, 2004, Neoentobdella and Pseudoentobdella. The affinities of species in a seventh capsalid genus, Trimusculotrema Whittington & Barton, 1990 (currently in Benedeniinae), are considered briefly based on morphology and host association and reassigned to the Entobdellinae.

A revision of Entobdella Blainville in Lamarck, 1818, with special emphasis on the nominal (type) species “Entobdella hippoglossi (Müller, 1776) Blainville, 1818” (Monogenea: Capsalidae: Entobdellinae) from teleost flatfishes, with descriptions of three new species and a new genus

Species of Entobdella Blainville in Lamarck, 1818, from skin (less commonly gills) of teleost flatfishes and occasionally from round-bodied teleosts are reviewed, with special reference to the nominal (type) species “Entobdella hippoglossi (Müller, 1776) Blainville, 1818” (Monogenea: Capsalidae: Entobdellinae) and to E. pugetensis Robinson, 1961. In the late 20th century the following 5 species were synonymised with E. hippoglossi: E. brattstroemi Brinkmann, 1952; E. curvunca Ronald, 1957; E. rosaceus Crane, 1972; E. squamula (Heath, 1902) Johnston, 1929; E. steingroeveri (Cohn, 1916) Johnston, 1929. We have shown that E. brattstroemi and E. squamula are valid, E. curvunca is a synonym of E. hippoglossi and E. rosaceus and E. steingroeveri are species inquirendae. Specimens of E. steingroeveri, first described in 1916 by Cohn, have been re-examined. Parasites from the Pacific Ocean on Hippoglossus stenolepis Schmidt, 1904 (Pacific halibut) and Eopsetta jordani (Lockington, 1879) (Petrale sole) (Pleuronectiformes: Pleuronectidae) and parasites geographically isolated in the Atlantic Ocean on H. hippoglossus (Linnaeus, 1758) (Atlantic halibut) and Reinhardtius hippoglossoides (Walbaum, 1792) (Greenland halibut) (Pleuronectiformes: Pleuronectidae) have all been regarded as E. hippoglossi. We confirm that E. hippoglossi sensu stricto is a parasite of the Atlantic halibut, but is accompanied on the same host by a second entobdelline species, E. vanbenedeni n. sp. It differs from the type species, E. hippoglossi in the anatomy of the distal region of the vagina and in the number, distribution and shape of papillae on the ventral surface of the haptor. Parasites from H. stenolepis are morphologically similar to E. hippoglossi on H. hippoglossus (no second species has been found on H. stenolepis), but differences in the number and shapes of haptor papillae support proposal of the parasites from H. stenolepis as a distinct species, E. stenolepis n. sp. Our anatomical studies of these parasites, with the exception of E. pugetensis, have revealed a cluster of several seminal receptacles opening into the ovo-vitelline duct. Previous reports of a single seminal receptacle in E. stenolepis (as E. hippoglossi) and E. steingroeveri are erroneous. Furthermore, reports of the vagina opening proximally into the alleged single seminal receptacle are also incorrect; direct and indirect evidence suggests that in all these parasite species, including E. pugetensis, the vagina opens proximally into the vitelline reservoir. Entobdella squamula from Paralichthys californicus (Ayers, 1859) (California flounder) (Pleuronectiformes: Paralichthyidae) differs from E. stenolepis, E. hippoglossi and E. vanbenedeni in the relative lengths of the accessory sclerites and anterior hamuli. Parasites originally identified as E. squamula from Hippoglossina macrops Steindachner, 1876 (Bigeye flounder) (Pleuronectiformes: Paralichthyidae), caught off the coast of Chile by the Lund University Chile Expedition (1948 – 49), differ from E. squamula in the shapes of the accessory sclerites and are regarded as a new species, E. brinkmanni n. sp. E. aegyptiacus Amer, 1990 must be regarded as a species inquirenda. A key to the 7 valid species of Entobdella recognised by us is included. Entobdella pugetensis differs in the following ways from all other species of Entobdella: the parasite is found on gills (less commonly on skin) of Atheresthes stomias (Jordan & Gilbert, 1880) (Arrowtooth flounder) (Pleuronectiformes: Pleuronectidae); the penis sac has a thick, possibly muscular, wall; the vagina is a relatively wide, short, straight tube with a conspicuous opening; no seminal receptacles have been observed; there are no papillae on the ventral surface of the haptor. We regard these distinctions as sufficient to erect a new genus, Branchobdella n. gen. to accommodate B. pugetensis n. comb. Assimilation of spermatophores in skin-parasitic Entobdella spp. and in the gill parasite B. pugetensis is discussed.

Review of the Capsalinae (Monogenea: Capsalidae)

The Capsalinae Baird, 1853 (Monogenea: Capsalidae) is revised based on a thorough review of original descriptions and examination of type museum material, where available, to validate species. A total of 262 type and voucher specimens was studied representing apparently 42 of the 60 currently described capsaline species. A combination of characters that should be independent of variation due to specimen preparation techniques was chosen to discriminate species. These characters include the presence/absence of papillae on the ventral surface of the haptor, the presence/absence and the morphology of haptoral accessory sclerites and the presence/absence of dorsomarginal body sclerites and their morphology and distribution. We consider that only 36 of the 60 nominal capsaline species are valid. We could find no support for Caballerocotyla Price, 1960 and therefore we synonymise it with Capsala Bosc, 1811. Under the current concept we recognise 22 species of Capsala, 7 species of Capsaloides Price, 1938, 3 species of Nasicola Yamaguti, 1968 and 4 species of Tristoma Cuvier, 1817. The following Capsala species are considered valid: C. albsmithi (Dollfus, 1962) n. comb.; C. biparasitica (Goto, 1894) Price, 1938; C. caballeroi Winter, 1955; C. foliacea (Goto, 1894) Price, 1938; C. gouri Chauhan, 1951; C. gregalis (Wagner & Carter, 1967) n. comb.; C. interrupta (Monticelli, 1891) Johnston, 1929; C. katsuwoni (Ishii, 1936) Price, 1938; C. laevis (Verrill, 1875) Johnston, 1929; C. maccallumi Price, 1939; C. magronum (Ishii, 1936) Price, 1938; C. manteri Price, 1951; C. manteriaffinis (Mamaev, 1968) n. comb.; C. martinierei Bosc, 1811; C. notosinense (Mamaev, 1968) n. comb.; C. nozawae (Goto, 1894) Price, 1938; C. onchidiocotyle (Setti, 1899) Johnston, 1929; C. ovalis (Goto, 1894) Price, 1938; C. paucispinosa (Mamaev, 1968) n. comb.; C. pelamydis (Taschenberg, 1878) Price, 1938; C. poeyi (Pérez-Vigueras, 1935) Price, 1938; C. pricei Hildago-Escalente, 1950. We consider the following Capsaloides species valid: C. cornutus (Verrill, 1875) Price, 1938; C. cristatus Yamaguti, 1968; C. hoffmannae Lamothe-Argumedo, 1996; C. magnaspinosus Price, 1939; C. nairagi Yamaguti, 1968; C. perugiai (Setti, 1898) Price, 1938; C. sinuatus (Goto, 1894) Price, 1938. The following Nasicola species are deemed valid: N. brasiliensis Kohn, Baptista-Farias, Santos & Gibson, 2004; N. hogansi Wheeler & Beverley-Burton, 1987; N. klawei Stunkard, 1962. Presently, we consider the following Tristoma species valid: T. adcoccineum Yamaguti, 1968; T. adintegrum Yamaguti, 1968; T. coccineum Cuvier, 1817; T. integrum Diesing, 1850. A list of new and re-established synonyms is provided. The status of each species is discussed in detail and a key to all capsaline species that we consider valid is presented. The following 5 capsaline species are considered to be species inquirendae: Caballerocotyla phillippina Velasquez, 1982; Capsala megacotyle (Linstow, 1906) Johnston, 1929; Tristoma fuhrmanni Guiart, 1938; T. levinsenii Monticelli, 1891; T. uncinatum Monticelli, 1889. The importance of careful character selection to discriminate between capsaline species and the need for studies of live parasites to obtain additional characters based on reproductive structures is addressed. Hostspecificity in the Capsalinae is also discussed.