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J N Caira

Publications and source records attributed to J N Caira.

At least 19 recordsLinked to original sources

Orectolobicestus n. g. (Cestoda: Tetraphyllidea), with the description of five new species and the transfer of Phyllobothrium chiloscyllii to the new genus.

ORECTOLOBICESTUS N. G. (CESTODA: Tetraphyllidea) is erected for six cestode species parasitising bamboo sharks (Orectolobiformes: Chiloscyllium). Members of this genus differ from all other phyllobothriid genera in possessing modified maisiform spinitriches on their distal bothridial surfaces. In addition, they are easily recognised in that they share the following unique combination of characters: their vitelline fields are interrupted by the ovary, their necks are scutellate, and their bothridia bear an apical sucker and marginal loculi. Five new species of Orectolobicestus are described, including O. tyleri n. sp. from Chiloscyllium punctatum off Borneo, O. lorettae n. sp. from C. cf. punctatum off Australia, O. mukahensis n. sp. and O. kelleyae n. sp. from C. indicum off Borneo, and O. randyi n.sp. from C. hasselti also from off Borneo. In addition, Phyllobothrium chiloscyllii Subhapradha, 1955 is transferred to the new genus. O. chiloscyllii (Subhapradha, 1955) n. comb. is readily distinguished from all five new species in its greater total length. In addition to a number of proglottid features, O. kelleyae n. sp. and O. randyi n. sp. clearly differ from the other three new species in their possession of trifid, rather than fully serrate, spinitriches on their proximal bothridial surfaces. The latter two species conspicuously differ from one another in total number of proglottids (11-21 vs 27-38). O. tyleri n. sp. generally has fewer proglottids than O. lorrettae n. sp. (7-17 vs 13-23) and, like O. mukahensis n. sp., possesses scutes that are spathate rather than elongate. O. tyleri n. sp. is readily distinguished from O. mukahensis n. sp. in its possession of fewer proglottids (7-17 vs. 19-29). The five new species of Orectolobicestus share derived bothridial microthrix features with Phyllobothrium squali Yamaguti, 1952, Thysanocephalum sp., Orygmatobothrium sp., Ruhnkecestus Caira & Durkin, 2006 and species of Paraorygmatobothrium Ruhnke, 1994. Among these taxa, Orectolobicestus most closely resembles Paraorygmatobothrium and Ruhnkecestus in its vitelline fields being interrupted by the ovary and the possession of a scutellate neck.

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Mode of attachment and lesions associated with trypanorhynch cestodes in the gastrointestinal tracts of two species of sharks collected from coastal waters of Borneo.

Lesions associated with two species of tapeworms within the digestive tract of wild-caught specimens of the bull shark, Carcharhinus leucas, and the sicklefin weasel shark, Hemigaleus microstoma, from Malaysian Borneo are described. Portions of the glandular stomach and pyloric gut with parasites were removed and fixed in 10% formalin buffered in sea water. Whole mounts, histological sections of tissues with and without worms in situ, and scanning electron microscopy images of detached worms were examined. Both species of cestodes belonged to the trypanorhynch family Tentaculariidae. Heteronybelinia estigmena was found in large numbers parasitizing the pyloric gut of C. leucas; an unidentified tentaculariid was found in relatively small numbers in both the glandular stomach and pyloric gut of H. microstoma. Both species burrowed their scoleces deeply in the mucosa and attached via hooked tentacles and unciniform microtriches of the scolex. The lesions induced by the parasites were marked in both sharks and ranged from acute necrotizing to chronic granulomatous gastroenteritis. Regenerative hyperplasia and intestinal metaplasia of gastric epithelium were also present. The severity and character of pathology was causally linked to the intensity of infection, the attachment mode of the parasites, and to the anatomophysiological relationships within the gut of the host shark.

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Five new species of Acanthobothrium (Tetraphyllidea: Onchobothriidae) from the freshwater stingray Himantura chaophraya (Batoidea: Dasyatidae) in Malaysian Borneo.

Five new species of Acanthobothrium (Tetraphyllidea: Onchobothriidae) are described from the spiral intestine of the Freshwater whipray, Himantura chaophraya, in the Kinabatangan River in Malaysian Borneo. Based on criteria set forth in a previous categorization scheme for species of Acanthobothrium, these consist of 3 Category 1 species, Acanthobothrium asnihae n. sp., Acanthobothrium saliki n. sp., and Acanthobothrium zainali n. sp.; a Category 8 species, Acanthobothrium etini n. sp.; and a Category 2 species, Acanthobothrium masnihae n. sp.. Acanthobothrium asnihae n. sp. differs from all Category 1 species in its possession of a horizontal band of weak musculature that divides the posterior loculus in half. Among Category 1 species, A. saliki n. sp. differs from all but Acanthobothrium southwelli in its possession of postovarian testes. It differs from A. southwelli in its possession of fewer testes and a greater number of proglottids. Acanthobothrium zainali n. sp. differs from the 25 other Category 1 species in a combination of overall size, muscular pad and hook shape, arrangement and number of testes, ovary configuration in cross section, position of ovarian isthmus, and genital pore position. Acanthobothrium etini n. sp. is distinguished from all 5 other Category 8 species in its lack of testes from the proglottid antiporal and postporal regions and in testis number. Acanthobothrium masnihae n. sp. differs from the 35 other Category 2 species in its possession of fewer testes, postporal testes, or a greater number of proglottids. A key to Acanthobothrium species parasitizing H. chayophraya is presented. This represents the first report of Acanthobothrium from freshwater stingrays belonging to a family other than the Potamotrygonidae.

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An unusual blood sequestering tapeworm (Sanguilevator yearsleyi n. gen., n. sp.) from Borneo with description of Cathetocephalus resendezi n. sp. from Mexico and molecular support for the recognition of the order Cathetocephalidea (Platyhelminthes: Eucestoda).

Sanguilevator yearsleyi n. gen., n. sp. and Cathetocephalus resendezi n. sp. are described from the Broadfin shark, Lamiopsis temmincki, in Malaysian Borneo and Carcharhinus leucas in Mexico, respectively. The new genus is unusual in its possession of internal chambers and channels in its scolex that appear to house extensive quantities of host white and red blood cells, respectively. Histology reveals an extremely intimate association between host tissue and the surface of the apical pad of the scolex. Positive staining with periodic acid-Schiff of the surface of the pad of the scolex and the linings of the chambers and channels suggests that an adhesive substance may be produced in these regions. However, explanations for how and why host blood cells come to reside within the scolex remain elusive. Cathetocephalus resendezi n. sp. is distinctive in the form of the papillae in the papillate band of the scolex and also in the inconspicuous nature of the rugose base of the scolex. Scanning electron microscopy of both new taxa as well as Cathetocephalus thatcheri, Cathetocephalus australis and an undescribed species of Cathetocephalus collected from Carcharhinus amboinensis in Australia, suggests that the papillae surrounding the pad of the scolex are of significant taxonomic utility in distinguishing among species in these groups. Parsimony and Bayesian analyses of sequence data (766 bases of 18S rDNA and 405 bases of 28S rDNA) generated from ethanol preserved specimens of C. thatcheri and S. yearsleyi, when compared with equivalent data available for 40 cestode species in GenBank, resulted in trees that support previous propositions that Cathetocephalus should be placed in the order Cathetocephalidea. The results suggest that Sanguilevator should also be considered to belong to this order.

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A revision of Phoreiobothrium (Tetraphyllidea: Onchobothriidae) with descriptions of five new species.

Five new species of Phoreiobothrium (Cestoda: Tetraphyllidea) are described from carcharhiniform sharks collected from a diversity of localities throughout the world. These are Phoreiobothrium perilocrocodilus n. sp. from Negaprion acutidens, Phoreiobothrium anticaporum n. sp. from Negaprion brevirostris, Phoreiobothrium lewinense n. sp. from Sphyrna lewini, Phoreiobothrium robertsoni n. sp. from Carcharhinus brachyurus, and Phoreiobothrium blissorum n. sp. from C. plumbeus, This brings the number of described species of Phoreiobothrium to 17. The new species from the lemon sharks, P. perilocrocodilus n. sp. and P. anticaporum n. sp., both possess more bothridial subloculi than do all species from hammerhead sharks, and fewer subloculi than do all species from sharks of Carcharhinus spp. Phoreiobothrium anticaporum n. sp. is unique among Phoreiobothrium species in that its genital pore is located in the extreme anterior portion of the proglottid. Phoreiobothrium lewinense n. sp. is unique in its possession of a combination of 8-11 subloculi and basal hook prongs that lack internal channels. Phoreiobothrium robertsoni n. sp. possesses 25-29 subloculi, triangular bothridia that flare posteriorly, and hooks that extend well beyond the middle of the bothridia. Phoreiobothrium blissorum n. sp. is most similar to Phoreiobothrium lasium in that it possesses rectangular bothridia with greater than 22 subloculi and long filitriches on its scolex. However, it differs from the latter species in that rather than articulating with one another, the bases of the medial and lateral hooks are widely separated; it also possesses 5-6 (vs.) 7-8 columns of testes. Phoreiobothrium lasium and Phoreiobothrium tiburonis are redescribed. The host associations of P. lasium are reevaluated and a neotype is designated for this species. The mature proglottid of P. tiburonis is described for the first time from voucher material collected from the type host and locality. These data suggest that the proglottid information provided in the original description, which was obtained from detached gravid proglottids, may have come from a tapeworm species other than P. tiburonis. All 7 species treated here in detail were examined with scanning electron microscopy; most were found to exhibit a microthrix pattern similar to that described previously for other members of the genus, e.g., Phoreiobothrium manirei. Bladelike spinitriches were present on the strobila, cephalic peduncle, and proximal bothridial surfaces, but were lacking from the distal bothridial surfaces. These spinitriches varied in length and density among species. Filitriches were present on all of these surfaces, but varied somewhat in length among species. Phoreiobothrium lewinense n. sp. is exceptional in that spinitriches were not seen on the proximal bothridial surfaces. A key to Phoreiobothrium species is provided. This work brings the number of carcharhinid shark species known to host this onchobothriid genus to 6, and the number of sphyrnid species to 5. Like many other onchobothriid tapeworms, species of Phoreiobothrium appear to exhibit oioxenous specificity for their hosts. Given this, and the fact that a total of 9 genera and 46 species of carcharhinid and sphyrnid sharks have yet to be examined for Phoreiobothrium, the diversity of the genus globally is likely to greatly exceed that described to date.

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Anthobothrium lesteri n. sp. (Cestoda: Tetraphyllidea) in Carcharhinus melanopterus from Heron Island, Australia, with comments on its site, mode of attachment, reproductive strategy and membership of the genus.

Anthobothrium lesteri n. sp. in Carcharhinus melanopterus from Heron Island, Australia is described and figured. A combination of characters, namely the number of testes, a weakly laciniate strobila and mature proglottis which is conspicuously longer than wide distinguishes it from other species that appear to be consistent with van Beneden's original concept of Anthobothrium . The scolex and the peduncle are relatively small and frail when compared with the rest of the strobila and with the detached proglottis. A myzorhynchus is absent and the stalked, undivided, simple bothridia have thickened margins. Two circular areas of muscles, which resemble accessory suckers, occur in the centre of each bothridium. There is a distinct cephalic peduncle with spiniform blade-like microtriches followed by craspedote proglottides which have a posterior four-lobed velum. Apolysis is well developed and the oncosphere has a polar filament and a dense covering of spines. Comments are made on the site, the mode of attachment, an aspect of the reproductive strategy of the new species in relation to intestinal eversion in some sharks, and membership of the genus.

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Five new species of Pedibothrium (Tetraphyllidea: Onchobothriidae) from the tawny nurse shark, Nebrius ferrugineus, in the Pacific Ocean.

The spiral intestines of 7 Tawny nurse sharks (Nebrius ferrugineus; Rhincodontidae) from Australia and French Polynesia were examined for tapeworms. These sharks hosted 5 new species of Pedibothrium. The 5 species were prepared for light and scanning electron microscopy. Whereas 3 of the species exhibit the bipronged hooks typical of members of Pedibothrium, 1 species was found to possess tiny unipronged hooks or to lack some, or all, hooks, and all individuals of the fifth species appeared to lack bothridial hooks entirely. Nonetheless, the otherwise remarkable similarities between these species and the species of Pedibothrium bearing bipronged hooks were considered to be sufficient to justify the placement of these 2 species in this genus. All 5 species possess proglottid features typical of Pedibothrium, including a uterus that extends anteriorly only to the level of the cirrus sac and a cirrus sac that is bent anteriorly and is crossed by the vagina. All 5 species also exhibit uniloculated bothridia. Of particular note was the fact that all 5 species exhibit bladelike spinitrichs with elongated distal tips, a form of microthrix currently known only from species of Pedibothrium. As a consequence, these species were treated as possessing modified hooks or as having lost hooks. The diagnoses of Pedibothrium and Onchobothriidae were emended to include these species. Each of the 5 new species bears a strong resemblance to 1 or more species of Pedibothrium hosted by a species of rhincodontid shark other than N. ferrugineus. Much of Southwell's type and voucher material of Pedibothrium was located and used to verify the identity of Pedibothrium kerkhami and many of the hosts from which Southwell's material of Pedibothrium was collected. This material suggests that P. kerkhami parasitizes Stegostoma fasciatum and not N. ferrugineus.

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Two new species of Yorkeria (Tetraphyllidea: Onchobothriidae) from Chiloscyllium punctatum (Elasmobranchii: Hemiscylliidae) in Thailand.

Two new species of the tapeworm genus Yorkeria are described from the spiral intestine of a specimen of the Brownbanded bambooshark, Chiloscyllium punctatum, collected from a fish market in Bangsarai, Thailand in 1996. The larger of the 2 new tapeworm species, Y. hilli, possesses fewer proglottids, fewer testes, and is smaller than most of its congeners. The smaller of the 2 new species, Y. kelleyae, conspicuously differs from all of its congeners in its possession of medial hooks that are only slightly larger than the lateral hooks (rather than conspicuously larger). Scanning electron microscopy shows both species to possess elongate filitriches on their distal prehook locular surfaces and elongate filitrichs and spinitrichs on their proximal bothridial, distal posthook locular surfaces, pedicels, and cephalic peduncles. In both species, the spinitrichs on the proximal bothridial surfaces, pedicels, and peduncles are very large and readily visible with light microscopy. Whereas the spinitrichs on these 4 surfaces are bluntly rounded in Y. hilli, they are pointed in Y. kelleyae. Examination of specimens identified as Y. parva, collected by several previous workers from a diversity of hosts and localities, calls into question the conspecificity of at least some of this material. The unusual scolex form suggests that the configuration of the cerebral ganglionic mass in species of Yorkeria may differ from that seen in most other tetraphyllidean taxa.

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An investigation of the co-evolutionary relationships between onchobothriid tapeworms and their elasmobranch hosts.

There is general consensus that the living elasmobranchs comprise a monophyletic taxon. There is evidence that, among tetraphyllidean tapeworms, the approximately 201 hooked species (Onchobothriidae) may also comprise a monophyletic group. Determinations of host specificity are contingent upon correct specific identifications. Since 1960, over 200 new elasmobranch species and over 100 new onchobothriid species have been described. Some confidence can be placed in host and parasite identifications of recent studies, but specific identifications provided in older literature in many cases are suspect. There is some consensus among published works on the phylogenetic relationships among elasmobranchs. Phylogenetic relationships among onchobothriids remain largely unresolved. Elasmobranchs have been poorly sampled for onchobothriids; records exist for approximately 20% of the 911 species and approximately 44% of the 170 elasmobranch genera. Onchobothriids are remarkably host specific, exhibiting essentially oioxenous specificity for their definitive hosts. Multiple onchobothriid species commonly parasitise the same host species; in some cases these are congeners, in other cases these are members of two different onchobothriid genera. There is substantial incongruence between available host and parasite phylogenies. For example, Acanthobothrium is by far the most ubiquitous onchobothriid genus, parasitising almost all orders of elasmobranchs known to host onchobothriids, yet, there is no evidence of major clades of Acanthobothrium corresponding to postulated major subgroupings of elasmobranchs (e.g. Galea and Squalea or sharks and rays). Potamotrygonocestus appears to be among the most basal onchobothriid groups, yet it parasitises one of the most derived elasmobranch groups (the freshwater stingray genus Potamotrygon). It appears that congeners parasitising the same host species are not necessarily each other's closest relatives. At this point the preliminary and limited available data suggest that, at least in this system, strict host specificity is not necessarily indicative of strict co-evolution. This study was extremely limited by the lack of available robust phylogenies for onchobothriids and elasmobranchs.

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Two new species of Litobothrium Dailey, 1969 (Cestoda: Litobothriidea) from thresher sharks in the Gulf of California, Mexico, with redescriptions of two species in the genus.

As part of a survey of the metazoan parasites of elasmobranchs of the Gulf of California, Mexico, the spiral intestines of 10 pelagic thresher sharks Alopias pelagicus and one bigeye thresher shark A. superciliosus were examined for tapeworms. Eight of the A. pelagicus specimens examined were found to host Litobothrium amplifica and L. daileyi. Both tapeworm species are redescribed based on examination of this new material with light and scanning electron microscopy, and the ranges of most of the measurements for these species are expanded; scanning electron micrographs and detailed illustrations and measurements of their segment anatomy are presented for the first time. An argument is made that the identification of the original host specimens of these species was in error and that A. pelagicus is likely to be the correct original host. In addition, L. nickoli n. sp., a third species in the genus hosted by A. pelagicus, was found in three of the 10 individual hosts examined. This species differs from all six known Litobothrium species in the form of the pseudosegments of the scolex, the anterior two being essentially non-cruciform, while the latter three are distinctly cruciform. All other species possess either no non-cruciform or at most one non-cruciform segment anteriorly. The single specimen of A. superciliosus examined was found to host the new species, L. janovyi. This species differs from L. coniformis, L. gracile and L. amsichensis in its possession of four rather than three, three and five cruciform pseudosegments, respectively. It lacks the modificiations of the fourth pseudosegment seen in L. amplifica and lacks the anterior non-cruciform fifth pseudosegment found in L. daileyi. It most closely resembles L. alopias but differs among other features in its greater total length, greater number of segments and longer mature segments. SEM of the four species collected from the Gulf of California as well as material of L. amsichensis from the goblin shark Mitsukurina owstoni that hosted the type-specimens of this species show that all surfaces of the body of all five species bear a dense covering of long filiform microtriches. L. amplifica bears a single row of large spine-like structures throughout most of the posterior margins of the first and second cruciform pseudosegment only. L. daileyi possesses one to two rows of overlapping spine-like structures on the posterior margins of the first four pseudosegments with the exception of the medial projections. The fifth pseudosegment lacks these structures. L. janovyi n. sp. bears spine-like structures on the lateral margins of only the third and fourth pseudosegments. L. nickoli n. sp. bears spine-like structures throughout the posterior margins of the first and second pseudosegments, and throughout the posterior margins of the third and fourth pseudosegments with the exception of the medial projections; the fifth pseudosegment lacks these structures. L. amsichensis bears no structures that could be considered to be spine-like on any of its pseuosegments, but possesses a border of densely arranged larger microtriches on the posterior margin of all five pseudosegments. A key to the species is included.

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Chaetotaxy and ultrastructure of sensory receptors in the cercaria of a species of Allassogonoporus Olivier, 1938. (Digenea:Lecithdendriidae).

A standard procedure that combines chaetotaxic, ultrastructural and neuromorphological observations has recently provided a new perspective to the study of cercarial sensory systems. In the present work, we aimed to extend the use of this combination of techniques to investigate the chaetotaxy of Allassogonoporus sp. in conjunction with the ultrastructure of sensory receptors and neuromorphology. Five nerve regions were distinguished. A conspicuous bilobed cerebral ganglion was observed at the level of the pharynx. The chaetotaxic pattern was generally consistent with that of other lecithodendriids. Four types of receptors were distinguished with scanning electron microscopy. These types differed in cilium length (short, moderately long or long) and tegumentary collar length (moderately low or high). Internal ultrastructure of receptor type IIAL revealed an unsheathed cilium, a closed basal body, septate extracellular junctional complexes and thickened nerve collars. Some receptor types were site-specific. Long uniciliated receptors were found mainly on the dorsal surface, whereas short uniciliated receptors were widespread across the tegument. Ultrastructure and site-specificity observations suggest that most sensory receptors are mechanoreceptors, probably reflecting the important role mechanoreception plays in host finding.

Acetylthiocholine↗

Two new species of Acanthobothrium Beneden, 1849 (Tetraphyllidea: Onchobothriidae) from horn sharks in the Gulf of California, Mexico.

Between 1993 and 1996, 26 individuals of two species of horn sharks were collected from the upper Gulf of California and their spiral intestines examined for onchobothriid tapeworms. Heterodontus francisci was found to host Acanthobothrium puertecitense n. sp. Based on the criteria of Ghoshroy & Caira (2001), this is a Category 4 species (with >15 mm total length, >50 proglottids, >80 testes and a symmetrical ovary). It differs from the four Category 4 species previously reported from either the western Atlantic or eastern Pacific Oceans, including A. bajaense (emend.) which was described from H. francisci, in hook-shape, total length, number of proglottids, number of testes and number of columns of vitelline follicles. H. mexicanus was found to host Acanthobothrium santarosaliense n. sp., which, based on the criteria of Ghoshroy & Caira (2001) is a Category 3 species (with >15 mm total length, >50 proglottids, >80 testes and an asymmetrical ovary). A. santarosaliense differs from the five Category 3 species previously reported from either the western Atlantic or eastern Pacific Oceans in its size, euapolytic nature, lack of dark matrix from the bases of the hooks, hook-shape, and distribution and number of testes. In addition to the difference in ovarian symmetry, A. puertecitense possesses fewer testes, fewer proglottids, is a shorter worm, possesses a more anterior genital pore and vitelline follicles which extend further forward anteriorly than in A. santarosaliense. This represents the first report of onchobothriids from H. mexicanus and the first report of onchobothriids from H. francisci in the Gulf of California. The apparent host-specificity of both new species should be viewed with caution until a greater sample of host individuals of both species can be examined. The fact that both new tapeworm species are known only from the Gulf is also considered to be a preliminary result, given the small sample of host individuals examined from these regions to date. This brings the total number of species of Heterodontus known to host Acanthobothrium to four and the total number of Acanthobothrium species described from heterodontiform sharks to five.

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Ultrastructure and chaetotaxy of sensory receptors in the cercariae of a species of Crepidostomum Braun, 1900 and Bunodera Railliet, 1896 (Digenea: Allocreadiidae).

Previous investigations of cercarial sensory systems have focused on chaetotaxy and ultrastructure of sensory receptors and have revealed chaetotaxic patterns within families, genera, and species as well as different types of sensory receptors. However, chaetotaxic and ultrastructural observations have rarely been combined. We investigated the ultrastructure of cercarial sensory receptors in conjunction with the chaetotaxy and neuromorphology in 2 allocreadiid species belonging to the genera Crepidostomum and Bunodera. Cercariae were treated with acetylthiocholine iodide and silver nitrate, and for scanning electron microscopy (SEM) and transmission electron microscopy (TEM). Similar cholinergic nerve networks were revealed. Chaetotaxy was consistent with that of other allocreadiids. Seven and 6 types of receptors were distinguished with SEM in Crepidostomum sp. and Bunodera sp., respectively. Types differed in number of cilia (1 or 2), cilium length (short, moderately long, or long), presence or absence of a tegumentary collar and a domelike base, and tegumentary collar length (low, moderately low, or high), TEM of some types revealed unsheathed cilia, basal body, and thickened nerve collars. Some receptor types were site specific. Thus, long uniciliated receptors were concentrated on the dorsal surface. Other types, such as short uniciliated receptors, were widespread throughout most regions. Ultrastructure and site-specificity observations suggest that most receptors are mechanoreceptors.

Acetylthiocholine↗

Four new species of Acanthobothrium (Cestoda: Tetraphyllidea) from the whiptail stingray Dasyatis brevis in the Gulf of California, Mexico.

Examination of the spiral intestines of 29 whiptail stingrays, Dasyatis brevis, from 6 sites in the Gulf of California, Mexico in 1993 and 1996 resulted in the discovery of 4 new species of Acanthobothrium: Acanthobothrium bullardi, Acanthobothrium dasi, Acanthobothrium rajivi, and Acanthobothrium soberoni. This brings the total number of species of Acanthobothrium known from the eastern Pacific Ocean to 34. With 22 additional species reported from elasmobranchs from the western Atlantic Ocean, and over 100 species globally, the number of interspecific comparisons required to justify the designation of a new species is rather unwieldy. To facilitate these and future comparisons, and in the absence of a phylogenetic hypothesis for this genus, the 56 species of Acanthobothrium from these 2 geographic regions were categorized for 4 characters: total length (< or = or > 15 mm), number of segments (< or = or > 50 segments), number of testes (< or = or > 80), and symmetry of poral and aporal ovarian lobes. These 56 taxa and their categories are presented in tabular form. Based on these characters, A. bullardi, A. dasi, and A. rajivi are category 2 species (they are relatively small, possess few segments, relatively few testes, and exhibit asymmetrical ovaries); A. soberoni is a category 6 species (it is a relatively longer worm with a larger number of segments, but with fewer testes and an asymmetrical ovary). All 4 species differ from 1 another and from species in other geographic regions in further subtleties of these 4 characters as well as hook size and relative length of hook prongs, cirrus sac size, genital pore position, number of columns of testes anterior to the cirrus sac, and number of postvaginal testes. Five specimens that appear to represent a new species of Acanthobothroides were also collected. This species is figured and some details of the morphology are described, but the material was considered to be insufficient to allow for formal description of this species at this time. This is the first record of either of these genera of onchobothriid tapeworms from the Gulf of California.

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Erudituncus n. gen. (Tetraphyllidea: Onchobothriidae) with a redescription of E. musteli (Yamaguti, 1952) n. comb. and comments on its hook homologies.

Examination of type and new material of Platybothrium musteli Yamaguti, 1952, resulted in the erection of a new genus, Erudituncus, the new combination Erudituncus musteli (Yamaguti, 1952), and the redescription of the newly combined species. This species exhibits novel hook morphology. Each bothridium bears 2 pairs of hooks, the axial pair comprising 1 medial and 1 lateral bipronged hook and the abaxial pair comprising a lateral unipronged hook and a medial, simple hook element. Based on the position of the hook talons, we suggest that the inner prong of the axial hooks of Erudituncus is homologous to the single prong of the axial hooks of Calliobothrium and the axial prongs of Dicranobothrium, Platybothrium, Phoreiobothrium, and Onchobothrium; the outer prong of the axial hooks of Erudituncus is homologous to the abaxial prongs of Dicranobothrium, Platybothrium, Phoreiobothrium, and Onchobothrium; and the abaxial hooks of Erudituncus and Calliobothrium are homologous. Erudituncus musteli was lacking from 31 individuals of the type host of record, the triakid shark Mustelus manazo Bleeker, 1854, examined for this study, but was obtained from 2 individuals of the triakid shark Hemitriakis japanica (Muller and Henle, 1839). Therefore, H. japanica is proposed as the probable type host of E. musteli.

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On the phylogenetic relationships among tetraphyllidean, lecanicephalidean and diphyllidean tapeworm genera.

This study had two main objectives: (1) to construct an extensive, explicit list of characters and character states that might serve as a starting point, and perhaps even a model, for the compilation of a more complete list of characters for all cestode taxa; and (2) to use this character list to generate a hypothesis of the phylogenetic relationships among species representing most of the tetraphyllidean, lecanicephalidean and diphyllidean genera. Specimens of one species in each of 48 genera of tetraphyllideans, eight genera of lecanicephalideans, the three genera of diphyllideans, two genera of proteocephalideans and two genera of trypanorhynchs, were examined as whole-mounts and sections, with light and scanning electron microscopy. A list of 120 morphological characters was compiled. Four phylogenetic analyses were conducted using PAUP* and/or NONA. The first was a comprehensive analysis with the 56 tetraphyllidean and lecanicephalidean species as ingroups and the remaining seven species as outgroups. The second was an analysis of the three diphyllidean species as ingroups and the two proteocephalidean and the two trypanorhynch species as outgroups. The third was an analysis of the eight lecanicephalidean species and the "tetraphyllideans" Echeneibothrium sp. and Pseudanthobothrium n. sp. as ingroups and an outgroup consisting of the seven species used as outgroups in the first analysis. In the fourth analysis, the ingroup consisted of the 14 hooked tetraphyllideans (onchobothriids), and the outgroup consisted of the seven species used as outgroups in the first analysis. The results of these analyses support the following phylogenetic hypotheses: The diphyllideans are monophyletic and Echinobothrium n. sp. and Macrobothridium sp. are more closely related to one another than either is to Ditrachybothridium macrocephalum. The tetraphyllideans, lecanicephalideans and proteocephalideans are more closely related to each other than they are to the diphyllideans or the trypanorhynchs. The ordinal status of the lecanicephalideans is dubious. The lecanicephalidean species are more closely related to some of the tetraphyllidean taxa than these tetraphyllidean taxa are to the remainder of the tetraphyllidean taxa. The proteocephalideans appear to belong within the tetraphyllidean clade. The "tetraphyllidean" species Echeneibothrium sp. and Pseudanthobothrium n. sp. are members of the lecanicephalidean clade. The position of "Discobothrium" n. sp. within the lecanicephalideans is dubious. Within the tetraphyllideans, the non-acetabulate species Litobothrium daileyi, Disculiceps galapagoensis and Cathetocephalus sp. are the most basal members of the group. The family Onchobothriidae is monophyletic, as it is currently defined. Within the onchobothriids, the uniloculate species are basal to the multiloculate species; the species with unipronged hooks are basal to the species with multipronged hooks. Although relationships among the phyllobothriids, as they are currently defined, remain poorly resolved, the family Phyllobothriidae is not monophyletic. These results suggest that some aspects of the classification of the lecanicephalidean and tetraphyllidean taxa require revision. However, such revision should be based on further analyses including a broader representation of the genera and species in these groups.

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Two new species of Echinobothrium (Cestoidea: Diphyllidea) from myliobatiform elasmobranchs in the Gulf of California, México.

Two new species of diphyllidean cestodes, Echinobothrium mexicanum n. sp., and Echinobothrium fautleyae n. sp., are described, the former from the spiral intestines of the bat rays Myliobatis californicus and Myliobatis longirostris and the latter from the spiral intestine of the cownose ray Rhinoptera steindachneri, all caught in the Gulf of California, México. Echinobothrium mexicanum is most easily distinguished from the majority of the species of the genus by the number of large apical hooks in each dorsoventral group (23) and from the remainder in the number of lateral hooklets in each group on the scolex. Echinobothrium fautleyae differs conspicuously from all other members of the genus in that the first and last lateral hooklets in each group are distinctly larger than the other hooklets in each group. Serial sections of the scoleces of these 2 new species confirm that the attachment structures on the scolex are bothria, rather than bothridia, as has been described in other species in the genus, suggesting that the attachment structures of the scolex should be examined in more detail in the genus in general. Palmate microtriches of various forms cover the proximal bothrial surfaces of both new species and are also found on the dorsal bothrial surfaces of E. fautleyae. The distal bothrial surfaces of E. mexicanum are covered with slender spiniform and short filiform microtriches. The cephalic peduncles of both species possess short filiform microtriches only. Comparison of these 2 new species with the other 3 species in this genus that have been examined with SEM suggests that the various forms and distributions of these microtriches is unique in each of these 2 species. We believe these features will prove to be taxonomically useful.

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Evolution of the major lineages of tapeworms (Platyhelminthes: Cestoidea) inferred from 18S ribosomal DNA and elongation factor-1alpha.

The interrelationships of the tapeworms (Platyhelminthes: Cestoidea) were inferred by analysis of complete gene sequences (approximately 2,200 bp) of 18S small subunit ribosomal DNA (18S) and partial gene sequences (approximately 900 bp) of elongation factor-1alpha (Ef-1alpha). New collections were made of 23 species representing each of the 14 currently recognized orders of tapeworms, including the Amphilinidea, Gyrocotylidea, and the 12 orders of the Eucestoda. Sequences were determined directly from polymerase chain reaction (PCR) products by either manual or automated methods. Nucleotide sequences of platyhelminth species outside of the Cestoidea were obtained for rooting the resulting trees. The 18S sequences were aligned with reference to the secondary structural features of the gene and the Ef-1alpha sequences were aligned with reference to their corresponding amino acid residues. Significant length variation among taxa was observed in the V2, V4, and V7 variable regions of the 18S gene. Such positions where sequences could not be aligned confidently were excluded from the analyses. Third codon positions of the Ef-1alpha gene were inferred to be saturated at an ordinal level of comparison. In addition, a short (approximately 35 bp) intron region of the Ef-1alpha gene was found to be shared only among the eucestode taxa, with the exception of Spathebothrium simplex (Spathebothriidea), which lacked the intron. Complete alignments showing structural features of the genes and sites excluded from analysis are provided as appendices. The sequence data were partitioned into 7 data sets in order to examine the effects of analyses on different subsets of the data. Analyses were conducted on the 2 genes independently, different codon positions of Ef-1alpha, amino acid sequences of Ef-1alpha, and combinations thereof. All subsets of the data were analyzed under the criterion of maximum parsimony as well as minimum evolution using both maximum-likelihood estimated, and LogDet-transformed distances. Results varied among the different data partitions and methods of analysis. Nodes with strong character support, however, were consistently recovered, and a general pattern of evolution was observed. Monophyly of the Cestoidea (Amphilinidea + Gyrocotylidea + Eucestoda) and Eucestoda and the traditionally accepted positions of the Amphilinidea and Gyrocotylidea as sister lineages to the Eucestoda were supported. Within the Eucestoda, the Spathebothriidea was found to be the sister of all other eucestodes. The remaining orders generally formed a diphyletic pattern of evolution consisting of separate difossate and tetrafossate lineages. This pattern was not universally observed among the analyses, primarily because the trypanorhynch and diphyllidean taxa showed instability in their phylogenetic position. Additional relationships that showed high levels of nodal support included a sister relationship between the Pseudophyllidea and Haplobothriidea and a clade uniting the Cyclophyllidea, Nippotaeniidea, and Tetrabothriidea. The Tetraphyllidea, as currently defined, was found to be paraphyletic without the inclusion of the orders Proteocephalidea and, possibly, Lecanicephalidea. Ordinal status of a monophyletic Litobothriidea, currently classified within the Tetraphyllidea, was found to be supported from a phylogenetic perspective.

Animals↗