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Scolex and tentacular armature of Pseudonybelinia odontacantha from marine plankton. a Capilliform microtriches (Cm) at the anterior end of scolex. b, d Unciniform microtriches (Um) on the distal bothrial surface. c Acerosate microtriches on the distal bothrial surface. e Capilliform microtriches on the pars postbulbosa (also on pars vaginalis and bulbosa). f, g. Appendix, short Wliform microtriches (Fm). h Appendix end. Note the absence of any capilliform microtriches
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The surface ultrastructure of two monotypic trypanorhynch genera is described based on new material of Grillotiella exilis (Linton, 1909) and type material of Pseudonybelinia odontacantha Dollfus 1966. In G. exilis, spiniform microtriches cover the bothrial surfaces and the anterior part of the pars vaginalis posterior to the bothria.
Bifurcate mic...
Citations
... Scanning electron microscopy (SEM) reveals different kinds of microtriches across the entire surface of the tegument of the different groups of cestodes (Chervy, 2009;Faliex et al., 2000;Caira et al., 1999). There are different forms of microtriches in the different groups of trypanorhynchs, such as capilliform, papilliform, palmate, filiform, and others ( Figure 2) (Whittaker, 1985;Palm, 2008;Caira et al., 2010;Menoret and Ivanov, 2015;Haseli et al., 2016). ...
... Internal organs, e.g., the shape and position of the testes are also important characters. The partial fusion of two or four bothria as well as the surface ultrastructure is difficult to observe in light microscopy (LM), whereas scanning electron microscopy (SEM) can illustrate this fusion, the ultrastructure and the hook patterns, especially in smaller worms (Palm 2008). Due to its high resolution of the surface ultrastructure and high magnification, SEM has been widely used to study trypanorhynch cestodes (e.g., Palm 1995, 1997, 2004, 2008, Morales-Ávila et al. 2019. ...
... The partial fusion of two or four bothria as well as the surface ultrastructure is difficult to observe in light microscopy (LM), whereas scanning electron microscopy (SEM) can illustrate this fusion, the ultrastructure and the hook patterns, especially in smaller worms (Palm 2008). Due to its high resolution of the surface ultrastructure and high magnification, SEM has been widely used to study trypanorhynch cestodes (e.g., Palm 1995, 1997, 2004, 2008, Morales-Ávila et al. 2019. ...
... A very high magnification and visualisation of, e.g., macromolecules of 1 nm size, is possible with SEM. Consequently, it has been applied to illustrate the surface ultrastructure of trypanorhynch cestodes, including the tentacular armature ( Fig. 2D) (Palm 1995, 2004, 2008, Morales-Ávila et al. 2019. One disadvantage is that the specimen used for SEM cannot be analysed for internal structures and cannot be used for any other microscopic technique. ...
... Internal organs, e.g., the shape and position of the testes are also important characters. The partial fusion of two or four bothria as well as the surface ultrastructure is difficult to observe in light microscopy (LM), whereas scanning electron microscopy (SEM) can illustrate this fusion, the ultrastructure and the hook patterns, especially in smaller worms (Palm 2008). Due to its high resolution of the surface ultrastructure and high magnification, SEM has been widely used to study trypanorhynch cestodes (e.g., Palm 1995, 1997, 2004, 2008, Morales-Ávila et al. 2019. ...
... The partial fusion of two or four bothria as well as the surface ultrastructure is difficult to observe in light microscopy (LM), whereas scanning electron microscopy (SEM) can illustrate this fusion, the ultrastructure and the hook patterns, especially in smaller worms (Palm 2008). Due to its high resolution of the surface ultrastructure and high magnification, SEM has been widely used to study trypanorhynch cestodes (e.g., Palm 1995, 1997, 2004, 2008, Morales-Ávila et al. 2019. ...
... A very high magnification and visualisation of, e.g., macromolecules of 1 nm size, is possible with SEM. Consequently, it has been applied to illustrate the surface ultrastructure of trypanorhynch cestodes, including the tentacular armature ( Fig. 2D) (Palm 1995, 2004, 2008, Morales-Ávila et al. 2019. One disadvantage is that the specimen used for SEM cannot be analysed for internal structures and cannot be used for any other microscopic technique. ...
Taxonomic issues within Trypanorhyncha, e.g., the inaccurate light microscopic visualisation of the hook patterns, are solvable by confocal laser scanning microscopy (CLSM). We applied CLSM imaging to study Trygonicola macropora (Shipley et Hornell, 1906) and Dollfusiella michiae (Southwell, 1929) from Neotrygon caeruliopunctata Last, White et Séret from Bali, Indonesia. To illustrate the strength and limitations of CLSM, images of Otobothrium cysticum (Mayer, 1842) and Symbothriorhynchus tigaminacantha Palm, 2004, both permanent mounts from a collection, were also processed. The CLSM created image stacks of many layers, and edited with IMARIS Software, these layers resulted in three-dimensional images of the armature patterns and internal organs of both species. BABB (benzylalcohol and benzylbenzuolate) clearing was applied to T. macropora. We conclude that trypanorhynch cestodes stained with Mayer-Schuberg's acetic carmine permanently mounted in Canada balsam are suitable for CLSM, allowing detailed analyses of museum type-material as well as freshly collected and processed worms. BABB resulted in imaging the testes in detail, suggesting other stains to be used for CLSM in trypanorhynch cestode research. Application of CLSM for studies of other cestode groups is highly recommended.
... The Tentacularioidea includes the families Paranybeliniidae Schmidt, 1970and Tentaculariidae Poche, 1926(see Palm, 2008Morales-Á vila et al., 2019), both of them with a homeoacanthous armature distinguishing them from the mainly heteroacanthous typical armature observed in the Eutetrarhynchoidea Guiart, 1927. However, the species of the Paranybeliniidae differ from those of the Tentaculariidae in having two, rather than four, bothria, a characteristic microthrix pattern and a metacestode with a blastocyst (Palm, 2008;Morales-Á vila et al., 2019). Although metacestodes of both families have so far been reported from marine holoplanktonic euphausiids, the plerocercoids of the Tentaculariidae occur in many marine invertebrates and a wide range of teleosts worldwide (Palm, 2004). ...
... All measurements are reported in micrometres and presented as the range followed by the mean, standard deviation (when n C 15), the number of the measured worms (N) and the total number of measurements for each character (n) in parentheses. The classification follows Palm (2004Palm ( , 2008. The type-specimens were deposited in the Museum für Naturkunde Berlin (ZMB), Germany and the Lawrence R. Penner Collection (LRP), Storrs, USA. ...
Two new tentaculariid species were found infecting carcharhiniform sharks from off the coasts of Malaysian Borneo and the southwestern coast of the Baja California Sur, Mexico. Both new species exhibit a homeoacanthous heteromorphous basal and a homeoacanthous homeomorphous metabasal armature. Since this hook arrangement is unique within the tentaculariids and the taxonomy in this group deeply depends on the tentacular armature, Reimeriella n. g. is erected to accommodate R. varioacantha n. sp. ex Carcharhinus sorrah (Müller & Henle) and R. mexicoensis n. sp. ex Sphyrna lewini (Griffith & Smith). Unlike R. mexicoensis n. sp., R. varioacantha n. sp. has a pars bothrialis not overlapping the pars bulbosa and the number of testes is higher. Reimeriella mexicoensis n. sp. possesses very large uncinate to falcate hooks in the basal armature, while in R. varioacantha n. sp. these hooks are almost the same in size as the remaining hooks in both the basal and metabasal armature. The latter species is the first tentaculariid species where the metabasal armature very closely resembles an eutetrarhynchid with a heteroacanthous typical homeomorphous metabasal armature and a high number of spiniform hooks per half spiral row (10–11 vs 6–7 in R. mexicoensis n. sp.) in the metabasal and apical armature. This pattern provides further morphological evidence for the close relationship of the Eutetrarhynchoidea and the Tentacularioidea. Reimeriella varioacantha n. sp. enriches the trypanorhynch fauna from off the coast of Malaysian Borneo while R. mexicoensis n. sp. is a novel record of a tentaculariid trypanorhynch from the Mexican Pacific.
... Cestodes of the order Trypanorhyncha are distributed worldwide. Over 300 species infect the stomach and intestine of their elasmobranch final hosts and their metacestodes infect marine invertebrates (mostly zooplanktonic) and teleost fish (Campbell and Beveridge, 1994;Palm, 1999Palm, , 20042008;Caira and Jensen, 2017). However, while most trypanorhynchs have been reported from teleost and elasmobranch fish (Palm, 2004), there are several reports of metacestodes infecting marine zooplankton (Shipley and Hornell, 1906;Anantamaran, 1963;Dollfus, 1966Dollfus, , 1967Grabda, 1968;Slankis and Shevchenko, 1974;Shimazu, 1975Shimazu, , 1982Shimazu, , 19992006;Reimer, 1977;Mauchline, 1980; https://doi.org/10.1016/j.ijppaw.2019.08.006 ...
... Cestodes of the order Trypanorhyncha are distributed worldwide. Over 300 species infect the stomach and intestine of their elasmobranch final hosts and their metacestodes infect marine invertebrates (mostly zooplanktonic) and teleost fish (Campbell and Beveridge, 1994;Palm, 1999Palm, , 20042008;Caira and Jensen, 2017). However, while most trypanorhynchs have been reported from teleost and elasmobranch fish (Palm, 2004), there are several reports of metacestodes infecting marine zooplankton (Shipley and Hornell, 1906;Anantamaran, 1963;Dollfus, 1966Dollfus, , 1967Grabda, 1968;Slankis and Shevchenko, 1974;Shimazu, 1975Shimazu, , 1982Shimazu, , 19992006;Reimer, 1977;Mauchline, 1980; https://doi.org/10.1016/j.ijppaw.2019.08.006 ...
... Over 300 species infect the stomach and intestine of their elasmobranch final hosts and their metacestodes infect marine invertebrates (mostly zooplanktonic) and teleost fish (Campbell and Beveridge, 1994;Palm, 1999Palm, , 20042008;Caira and Jensen, 2017). However, while most trypanorhynchs have been reported from teleost and elasmobranch fish (Palm, 2004), there are several reports of metacestodes infecting marine zooplankton (Shipley and Hornell, 1906;Anantamaran, 1963;Dollfus, 1966Dollfus, , 1967Grabda, 1968;Slankis and Shevchenko, 1974;Shimazu, 1975Shimazu, , 1982Shimazu, , 19992006;Reimer, 1977;Mauchline, 1980; https://doi.org/10.1016/j.ijppaw.2019.08.006 Mooney and Shirley, 2000;Gómez-Gutiérrez et al., 2010González-Solís et al., 2013). ...
Plerocerci of the monotypic Paranybelinia otobothrioides were found parasitizing the subtropical neritic krill Nyctiphanes simplex in the Gulf of California, Mexico. The plerocerci were recovered from two microhabitats of the intermediate host, typically embedded inside the digestive gland (hepatopancreas) or rarely in the hemocoel. The morphology of the simple, single-layered blastocyst surrounding the entire scolex is unique within the Trypanorhyncha by having four large funnel-like pori or openings possibly with feeding and/or excretory function. One of the openings is located anteriorly and three at the posterior end. Scolex surface ultrastructure shows hamulate and lineate spinitriches covering the bothrial surface, capilliform filitriches at the anterior scolex end and on the scolex peduncle, and short papilliform filitriches on the long appendix. This pattern resembles that of species of the Tentaculariidae; but differs in that the hamulate spinitriches, which appear lineate at the bothrial margins, densely cover the entire distal bothrial surface. Tegumental grooves are present on the posterior bothrial margin, lacking spinitriches. Paranybelinia otobothrioides and Pseudonybelinia odontacantha share the following unique combination of characters: two bothria with free lateral and posterior bothrial margins, homeoacanthous homeomorphous armature, tegumental grooves, the distribution of the hamulate spinitriches, and the absence of prebulbar organs. Both genera infect euphausiids as intermediate hosts. Sequence data of the partial ssrDNA gene place Pa. otobothrioides sister to the family Tentaculariidae, and the Kimura two-parameters (K2P) distance between Pa. otobothrioides and species of the family Tentaculariidae ranged from 0.027 to 0.039 (44-62 nucleotide differences). These data suggest both species be recognized in a family, the Paranybeliniidae, distinct from, albeit as sister taxon to, the Tentaculariidae. High prevalence of infection (
... The Otobothrioidea Dollfus, 1942 known as one of the five superfamilies of Palm's (2004) system, after transferring the Paranybeliniidae Schmidt, 1970 to the Tentacularioidea Poche, 1926 by Palm (2008), comprises two families, the Pseudotobothriidae Palm, 1995 andthe Otobothriidae Dollfus, 1942, which include three and nine genera, respectively (see Palm, 2004;Beveridge & Campbell, 2005;Schaeffner et al., 2011;Schaeffner & Beveridge, 2013a). Regardless of the discrepancy about the validity of the Pseudotobothriidae (see Schaeffner et al., 2011), one pseudotobothriid species, Pseudotobothrium dipsacum (Linton, 1897), and five otobothriid species, i.e. ...
... Whether the Otobothrioidea is classified into the Pseudotobothriidae Palm, 1995 and the Otobothriidae (sensu Palm, 1997Palm, , 2004 (see Palm, 1997Palm, , 2004Palm, , 2008 or this superfamily is considered as a monotypic taxon (see Beveridge et al., 2000) supported strongly by Schaeffner et al. (2011), the new species, as a member of the Otobothriidae (sensu Palm, 1997Palm, , 2004 or Otobothriidae (sensu Beveridge et al., 2000), exhibits the closest morphological similarity to Pristiorhynchus palmi Schaeffner & Beveridge, 2013, the only species of its genus. Both species possess a long and acraspedote scolex lacking a velum; a pars vaginalis which is longer than the pars bothrialis; a retractor muscle originating in the posterior third of the bulb; a distinctive basal armature; a heteroacanthous atypical metabasal armature with five heteromorphous hooks per principle row, three intercalary hooks in a single row; the long segments each with a postequatorial genital pore; a median tubular uterus and two longitudinal columns of the testes arranged in a single layer. ...
A survey on the cestode fauna of Paragaleus randalli Compagno, Krupp & Carpenter in the Persian Gulf resulted in the discovery of a new trypanorhynch species of the family Otobothriidae Dollfus, 1942, the second otobothrioid species hosted by the family Hemigaleidae Hasse. The new species exhibits the closest morphological similarity to Pristiorhynchus palmi Schaeffner & Beveridge, 2013, the type- and only species of its genus. However, the new species differs from P. palmi in the position of the bothrial pits, the morphology and oncotaxy of the basal armature, the commencement of the hook files on the internal surface, a wider scolex peduncle in the pars bulbosa than in the pars vaginalis, a long neck and the presence of a lateral bothrial groove connecting the two bothrial pits to each other. The latter character is a unique trait within the Otobothrioidea Dollfus, 1942. Considering such differences, a new genus, Olgaella n. g., was erected to accommodate O. elenae n. g., n. sp. within the Otobothriidae. The evolutionary relatedness of the bothrial pits of the Otobothrioidea and the bothrial grooves of the Lacistorhynchoidea Guiart, 1927 is discussed.
... Tentacles and bothria can contract together or singly and, thus, control by the nervous system is most likely. There are some data available about the sensory organs in the tegument of Trypanorhyncha (Davydov and Biserova 1985;Biserova 1987Biserova , 1991aPalm 1995Palm , 1997Palm , 2000Palm , 2004Palm , 2008Palm et al. 1998Palm et al. , 2000Casado et al. 1999;Jones 2000). Sensory organs of Grillotia erinaceus are located diffusely at the outer and inner surface of the bothria, and also regular ciliated nerve endings are situated at the marginal area of bothria (Biserova 1987(Biserova , 1991a. ...
... Tentacles and bothria can contract together or singly and, thus, control by the nervous system is most likely. There are some data available about the sensory organs in the tegument of Trypanorhyncha (Davydov and Biserova 1985;Biserova 1987Biserova , 1991aPalm 1995Palm , 1997Palm , 2000Palm , 2004Palm , 2008Palm et al. 1998Palm et al. , 2000Casado et al. 1999;Jones 2000). Sensory organs of Grillotia erinaceus are located diffusely at the outer and inner surface of the bothria, and also regular ciliated nerve endings are situated at the marginal area of bothria (Biserova 1987(Biserova , 1991a. ...
... The morphology and distribution of microtriches in Trypanorhyncha are considered to be of phylogenetic importance and represent a potentially significant taxonomic character (Palm 1995(Palm , 2004. The tegument of N. surmenicola has a number of ultrastructural features which make it significantly different from Parachristianella sp. and other Trypanorhyncha. ...
The sensory organs in the tegument of two trypanorhychean species – Nybelinia surmenicola and adult Parachristianella sp. (Cestoda, Trypanorhyncha) were studied with the aim of ultrastructural description and a comparative analysis. The Nybelinia surmenicola plerocercoid lacks papillae with sensory cilia on the bothria adhesive surface. We found the unciliated sensory organ within the median bothrial fold. This unciliated free nerve ending contains the central electron-dense disc, three dense supporting rings, and broad root. The nerve ending locates in the basal matrix under the tegument. The tegument of N. surmenicola has a number of ultrastructural features which makes it significantly different from other Trypanorhyncha: i) the tegumental cytoplasm has a plicated constitution in a form of high apical and deep basal folds; ii) numerous layers of the basal matrix are presented in subtegument; iii) the squamiform and bristle-like microtriches N. surmenicola lack the base and the basal plate. In contrast, numerous ciliated and unciliated sensory organs were found in Parachristianella sp.: six types on the bothria and one type in the strobila’ tegument. Ultrastructural constitution of ciliated free nerve endings and unciliated basal nerve endings of Parachristianella sp. has many common features with sensory organs of other cestodes. In comparison with other Trypanorhyncha, all Nybelinia species studied have less quantity of the bothrial sensory organs. This fact may reflect behavioral patterns as well as phylogenetic position of Nybelinia into Trypanorhyncha. Our observations of living animals conventionally demonstrate the ability of N. surmenicola plerocercoids to locomote in forward direction on the Petri dish surface. The participation of the bothrial microtriches in a parasite movement has been discussed.
Key words: Cestoda, fine structure, receptors, microtriches, tegument
... The surface ultrastructure of trypanorhynchs is diverse and includes a broad range of different microthrix forms and shapes. Over the last two decades many scientists have focused on this morphological feature and various trypanorhynch species have been observed with SEM (Richmond & Caira, 1991;Palm, 1995Palm, , 1997Palm, , 2000Palm, , 2004Palm, , 2008Palm et al., 1993Palm et al., , 1998Jones & Beveridge, 1998;Campbell et al., 1999;Jones, 2000;Beveridge & Campbell, 2001). Recently, Chervy (2009) proposed a unified terminology for the complex morphology of microtriches of cestodes. ...
Redescriptions are provided for five incompletely described species of Otobothrium Linton, 1890: Otobothrium alexanderi Palm, 2004 from two species of carcharhinid sharks, Carcharhinus cautus (Whitley) and C. melanopterus (Quoy & Gaimard) at three localities off northern Australia; O. australe Palm, 2004 based on material collected from the type-host and type-locality and from six additional myliobatid and carcharhinid host species off Western Australia, the Northern Territory and northern Queensland; O. insigne Linton, 1905 from Rhizoprionodon terraenovae (Richardson) and Sphyrna tudes (Valenciennes) in the Atlantic Ocean off Senegal and the Democratic Republic of the Congo; O. mugilis Hiscock, 1954, previously known only from larval stages, based on adults from five sphyrnid and carcharhinid definitive host species off northern Australia and Malaysian Borneo; and O. penetrans Linton, 1907 from material collected from two species of hammerhead sharks (Sphyrnidae) in the Red Sea off Jordan and the Indian Ocean off Western Australia. Additional host and locality records are added for the type-species, O. crenacolle Linton, 1890 and for O. carcharidis (Shipley & Hornell, 1906). Two descriptions are provided for Otobothrium spp. treated here as Otobothrium sp. 1 from C. melanopterus off northern Australia and Otobothrium sp. 2 from Sphyrna zygaena (Linnaeus) in the Gulf of California, Mexico.
... Taxonomical and ecological studies on fish parasites from Indonesia have been intensified in recent years (e.g. Palm et al., 2007Palm et al., , 2008Palm et al., , 2011Yuniar et al., 2007;Palm, 2008Palm, , 2011Bray & Palm, 2009;Kuchta et al., 2009;Rü ckert et al., 2009aRü ckert et al., , b, 2010Kleinertz, 2010, Kleinertz et al., 2012, Dewi & Palm, 2013Kuhn et al., 2013), taking into account the high parasite biodiversity at this tropical location. The purpose of the present study is an assessment of the fish parasite fauna of E. coioides, a widely distributed and rapidly developing mariculture species in Indonesia, from additional sampling sites. ...
A total of 195 Epinephelus coioides (Hamilton, 1822) were studied for fish parasites from Javanese (Segara Anakan lagoon) and Balinese waters. Up to 25 different parasite species belonging to the following taxa: one Ciliata, one Microsporea, five Digenea, one Monogenea, four Cestoda, four Nematoda, one Acanthocephala, one Hirudinea and seven Crustacea were identified with four new host and locality records. The dominant parasites included the monogenean Pseudorhabdosynochus lantauensis (53.3-97.1%), the nematode Spirophilometra endangae (23.3-42.9%), the digenean Didymodiclinus sp. (2.9-40.0%), the nematodes Philometra sp. (22.6-34.3%) and Raphidascaris sp. (2.9-28.6%), and the isopod Alcirona sp. (6.7-31.4%). Regional differences for E. coioides were found in terms of endoparasite diversity, total diversity according to Shannon-Wiener, Simpson index and Evenness. A comparison with published data from Sumatera revealed highest endoparasite diversity (Shannon-Wiener: 1.86/1.67-2.04) and lowest ectoparasite/endoparasite ratio (0.73/0.57-0.88) off the Balinese coast, followed by Lampung Bay, Sumatera (1.84; 0.67), off the coast of Segara Anakan lagoon (1.71; 0.71), and in the lagoon (0.30/0.19-0.66; 0.85/0.67-1.00). The presented data demonstrate the natural range of these parameters and parasite prevalences according to habitat and region, allowing adjustment of the scale that has been used in the visual integration of the parasite parameters into a star graph. The parasite fauna of E. coioides in Segara Anakan lagoon 'improved' from 2004 until 2008/09, possibly related to earlier oil spill events in 2002 and 2004. The use of grouper fish parasites as an early warning system for environmental change in Indonesian coastal ecosystems is discussed.
... 1g, 1h). Extremely large microtrichia (the length of the apical part of which reaches 13 µm and exceeds by ten times the length of common blade like microtrichia) and small microtrichia with a short CHARACTERIZATION OF CESTODA TISSUE ORGANIZATION 147 (Biserova, 1987;Palm, 2008); (f) giant microtrichia (Poddubnaya et al., 2007); (i) spine (Mount, 1970); (j), (k) epithelium of reproductive ducts with microvilli and cilia. round apical part were described in the Echinophalus wageneri in pseudophyllid cestodes (Poddubnaya et al., 2007) (Figs. ...
... 1a, 1f). In addition to microtrichia, hooks are localized on proboscises of the attachment apparatus (Mount, 1970;Thomas, 1983;dárská and Nebesá ová, 2003;Palm, 2008) (Fig. 1i). ...
Taking into account our own and published data, a comprehensive ultrastructural estimation of the main tissue systems (muscular, nervous, epithelial, and tissues of the internal environment) was conducted for the Cestoda class. The facts on the structure of the cambial system were summarized, and peculiarities of its organization (as compared with other classes of flatworms) were detected. Characterization of the tissue organization of cestodes was suggested and justified.
