Cephalic Peduncle Research Articles

Two new species of Scyphophyllidium (Cestoda: Phyllobothriidea) from Chaenogaleus macrostoma (Bleeker) (Elasmobranchii: Carcharhiniformes) from the Persian Gulf, Iran.

Two new species of Scyphophyllidium Woodland, 1927 are described from the Hooktooth shark, Chaenogaleus macrostoma (Bleeker), from the Persian Gulf, Iran. Scyphophyllidium hormuziense n. sp. is assigned to morphological category 2 of its genus because it possesses bothridial marginal loculi and an apical sucker on each flat bothridium, and lacks bothridial facial loculi. Within category 2, it is distinguished from its congeners, except for S. janineae (Ruhnke, Healy and Shapero, 2006), by possessing, rather than lacking, a prominent cephalic peduncle. It differs from S. janineae in the distribution of the vitelline follicles. Scyphophyllidium iraniense n. sp. is assigned to morphological category 5 because its bothridia are essentially flat and lack proximal apertures, marginal loculi, facial loci and semi-circular muscle bands, and bears weakly serrate gladiate spinitriches on its distal bothridial surfaces. Within its morphological category, the presence of a long cephalic peduncle distinguishes S. iraniense n. sp. from its congeners lacking this structure or possessing a short cephalic peduncle. It further differs from S. arnoldi (Ruhnke and Thompson, 2006) and S. typicum (Subhapradha, 1955) in total length, from S. paulum (Linton, 1897) in lacking a conspicuous band of muscles along the locular periphery of the bothridia, and from S. kirstenae (Ruhnke, Healy and Shapero, 2006) in the distribution of the vitelline follicles. This study brings the number of the valid Scyphophyllidium species of the Persian Gulf to five.

NOVELTY AND PHYLOGENETIC RELATIONSHIPS WITHIN THE SERENDIPEIDAE (CESTODA: "TETRAPHYLLIDEA").

Nanoduplicibothrium n. gen. is erected for the subgroup containing the smallest members of the "tetraphyllidean" family Serendipeidae with bothridia fused lengthwise in 2 pairs that lack both a distinct row of posterior loculi and a cephalic peduncle. Two new species in this genus are described. These are Nanoduplicibothrium leanneae n. gen. n. sp. from Rhinoptera bonasus off South Carolina and Nanoduplicibothrium megaphallum n. sp. from Rhinoptera jayakari off Mozambique. Two species currently assigned to Duplicibothrium are transferred to the new genus as Nanoduplicibothrium paulum n. comb and Nanoduplicibothrium jillae n. comb. and the diagnosis of Duplicibothrium is emended so that it aligns with the revised membership of the group. Duplicibothrium bilai n. sp. is also described from R. jayakari off Mozambique. The description of these species provides formal names for 3 species included in previously published molecular phylogenetic work under the provisional names Duplicibothrium n. sp. 2, Duplicibothrium n. sp. 4, and Duplicibothrium n. sp. 5, respectively. Erection of the new genus substantially reduces the number of instances of congeners in the family parasitizing the same host species because in most instances the pairs of species now represent 1 species each in Nanoduplicibothrium and Duplicibothrium. Sequence data for the D1-D3 region of the 28S rDNA gene were generated for Serendip for the first time from an undescribed species from Aetomylaeus asperrimus collected off Panama. This finding also expands the known host associations of the Serendipeidae beyond the Rhinopteridae to include a species of Myliobatidae. A maximum-likelihood phylogenetic analysis of all species of serendipeids for which data for the D1-D3 region of the 28S rDNA gene are available confirms the reciprocal monophyly of Nanoduplicibothrium, Duplicibothrium, and Serendip. The phylogenetic placement of the fourth genus in the family-the monotypic Glyphobothrium-remains to be determined.

Two new species of Acanthobothrium Blanchard, 1848 (Cestoda, Onchoproteocephalidea) from Urobatisjamaicensis (Cuvier, 1816) (Elasmobranchii, Batoidea, Myliobatiformes) of the Mexican Caribbean.

Eight specimens of Urobatisjamaicensis were collected from four localities of the Yucatan Peninsula, Mexico, of which four specimens were infected with cestodes of two new species of Acanthobothrium. Acanthobothriumgarciaprietoisp. nov. differs from congeners by a combination of characters including the size of the scolex and bothridia, length of the cephalic peduncle, length of the axial and abaxial prongs and total length of the abaxial prongs of the hooks, size of the cirrus sac and testes in mature proglottids, and the total number of proglottids. The most prominent characteristic distinguishing A.pulidofloresaesp. nov. from other species of the Atlantic Ocean/Caribbean Sea/Gulf of Mexico is the form of the scolex, which has the "clover leaf" configuration. In addition, it can be distinguished by the total length of the worm, total number of proglottids, small accessory suckers, the form of the bothridia, length of the axial and abaxial prongs and total abaxial prong length of hooks, and the number of testes. According to the current category scheme, A.garciaprietoisp. nov. and A.pulidofloresaesp. nov., belong to categories 1 and 5, respectively.

Two New Cestode Species of Phoreiobothrium (Onchoproteocephalidea) from Two Carcharhinid Sharks of the Persian Gulf, with the First Report of Variation in Distribution of Vitelline Follicles of the Genus.

The aim of this work was the morphological study of the onchoproteocephalidean cestodes from two shark species of the Persian Gulf. Several specimens of Rhizoprionodon oligolinx, as well as of Carcharhinus dussumieri, were collected from the Persian Gulf, and the isolated cestodes from them were studied morphologically by light and scanning electron microscopy. Phoreiobothrium sarahae n. sp. and Phoreiobothrium iraniense n. sp. were isolated, respectively, from Rhizoprionodon oligolinx Springer and Carcharhinus dussumieri (Müller & Henle). Both species are quite distinct from their congeners, except for P. sorrahcola Masoumi Ganjgah & Haseli, 2020, in the distribution of the vitelline follicles (i.e., having more than two columns of vitelline follicles in each lateral band in the proglottids) and are distinguished from each other based on measurement, e.g., by the length of the cephalic peduncle, the width of the scolex at the level of hooks, and some measures related to the hooks. Phoreiobothrium iraniense n. sp. most closely resembles P. sorrahcola, earlier described from C. sorrah (Müller & Henle) from the Persian Gulf, with which it has an extensive overlap in the ranges of measurements and morphology. Nonetheless, it differs from P. sorrahcola in the length of the cephalic peduncle and possessing, rather than lacking, the gladiate spinitriches on the proximal bothridial surfaces. Along with the description of the two new species increasing the number of members of the genus in the Iranian coastal waters from three to five, for the first time a variation in distribution of the vitelline follicles is reported for Phoreiobothrium in this paper.

A revision of the Rhoptrobothriidae (Cestoda: Tetraphyllidea).

The Rhoptrobothriidae are one of the more enigmatic families of cestodes of elasmobranchs. Opinions on the taxonomic status of the familys three original genera (i.e., Myzophyllobothrium, Rhoptrobothrium, and Myzocephalus) have varied over the 115 years since they were erected. Some authors have considered all three valid, others have considered Rhoptrobothrium to be a synonym of Myzopyllobothrium or a genus inquirendum, yet others have considered Myzocephalus to be a synonym of the phyllobothriid genus Thysanocephalum. All three genera were established for specimens collected from eagle rays off Sri Lanka. The erection of Mixophyllobothrium for two specimens from a cowtail stingray off India three decades ago added additional confusion to the situation, with some authors considering it valid and others a synonym of Myzocephalus. These disagreements stem largely from differences in interpretation of the complex morphology of the scolex of members of these genera. Furthermore, with the exception of Rhoptrobothrium comprising four species, each genus is monotypic. All but Rhoptrobothrium has not been considered in detail for nearly a century, largely because of a lack of available material. The taxonomic status of these genera is assessed here based on light and scanning electron microscopy, and molecular data generated from new material collected from eagle rays off Indonesian and Malaysian Borneo, Japan, Sri Lanka, and Viet Nam. Morphological work indicates that the genera differ largely only in the degree of folding of the four remi that extend from the cephalic peduncle. A molecular phylogeny based on sequence data for the D1D3 region of the 28S rRNA gene, which include new data for eight specimens of four species, indicates that Myzophyllobothrium, Myzocephalus, and Rhoptrobothrium are not mutually monophyletic. The latter two genera and Mixophyllobothrium are considered synonyms of Myzophyllobothrium and five species are transferred to that genus. Myzophyllobothrium okamuri n. comb. is considered a species inquirendum. Myzophyllobothrium nagasawai n. sp. is described from Aetobatus narutobiei off Japan. Myzophyllobothrium narinari n. comb. is re-described based on newly collected cestodes from the type host and locality (i.e., Aetobatus ocellatus off Sri Lanka). Despite consisting of only a single genus, the family status of the group is retained in recognition of the unusual configuration of the scolex, which bears four biloculate bothridia and four remi extending from the cephalic peduncle. The ordinal placement of the family remains uncertain, but affinities with the Phyllobothriidea, rather than Tetraphyllidea are considered.

Three new species of 'tetraphyllidean' cestodes from an undescribed bamboo shark (Orectolobiformes: Hemiscylliidae) in Sri Lanka.

Examination of seven specimens of an undescribed species of bamboo shark (Orectolobiformes: Hemiscylliidae), currently referred to as Chiloscyllium sp. 1, from Sri Lanka yielded three new species of 'tetraphyllidean' cestodes, one each in the genera Carpobothrium Shipley et Hornell, 1906, Spiniloculus Southwell, 1925, and Yorkeria Southwell, 1927. Carpobothrium marjorieae Caira, Otto et Jensen sp. n. differs from its three valid congeners in total length, number of testes, and size of the apical sucker of its anterior bothridial flap. Like Carpobothrium eleanorae Koontz et Caira, 2016 it possesses spherical eggs with highly elongate bipolar filaments, but its eggs are conspicuously larger than those of the latter species. Spiniloculus akshayi Caira, Otto et Jensen sp. n. can be distinguished from its five congeners in total length and number of proglottids. Yorkeria sachiniae Caira, Otto et Jensen sp. n. differs from its 16 congeners in number of proglottids and testes, total length, terminal proglottid, cirrus sac, and cephalic peduncle length, hook size, genital pore position, and bothridial shape. The eggs of Y. sachiniae Caira, Otto et Jensen sp. n. are spindle-shaped and bear a single long, polar filament. All three species were found to exhibit microthrix patterns on their scolices like those of the majority of their congeners that have been examined to date with scanning electron microscopy. Microtriches on the distal surfaces of the bothridial pouches of a species of Carpobothrium were characterised for the first time; C. marjorieae Caira, Otto et Jensen sp. n. was found to possess a modified form of gladiate spinithrix on this surface. Among the seven members of the genus, this is the first species of Chiloscyllium Müller et Henle that has been found to host representatives of all three of these 'tetraphyllidean' genera. However, unlike most of its congeners, a representative of the non-hooked genus Scyphophyllidium Woodland, 1927 was not found infecting Chiloscyllium sp. 1.

Two new species of Caulobothrium (Cestoda: "Tetraphyllidea") from the duckbill eagle ray, Aetomylaeus bovinus (Myliobatiformes: Myliobatidae), off Senegal with new insights on morphological features of the genus.

Two new species of the cestode genus Caulobothrium, collected from the duckbill eagle ray, Aetomylaeus bovinus, off Senegal, are described. Although postulated as sister taxa in an earlier molecular phylogenetic analysis, Caulobothrium multispelaeum n. sp. and Caulobothrium katzi n. sp., respectively, are among the smallest and largest members of the genus. The smaller species is unique among its congeners in possessing unusual medial longitudinal grooves along the dorsal and ventral surfaces of its strobila that develop into a tandem series of elliptical apertures on the posterior proglottids. The inner surfaces of these apertures stained positively with McManus' periodic acid Schiff in a manner similar to that seen in members of the distantly related lecanicephalidean genus Elicilacunosus. The larger species differs from its congeners in size, number of proglottids, and arrangement of bothridial loculi. Both new species were found to possess a small apical sucker on the anterior margin of each of their bothridia. Scanning electron microscopy (SEM) and frontal sections of a bothridium of Caulobothrium tetrascaphium suggests that this species also bears an apical sucker. Examination of the hologenophore of the species provisionally referred to as Caulobothrium n. sp. 5 in the earlier molecular analysis indicates it is conspecific with the recently described Caulobothrium pedunculatum, which was also determined to possess bothridial apical suckers. This leads us to suspect that this feature may be found to occur in all members of the genus. SEM of specimens of Caulobothrium for the first time indicates their bothridial surfaces are covered with filitriches of various sizes but lack spinitriches; spinitriches were seen only on the cephalic peduncle of C. katzi n. sp. The geographic distribution and host associations of Caulobothrium are expanded to include data now available for all species. The diagnosis of the genus is revised to include all of this information.

A NEW SPECIES OF ACANTHOBOTHRIUM (CESTODA: ONCHOPROTEOCEPHALIDEA) FROM THE SMALLEYE PYGMY SHARK, SQUALIOLUS ALIAE (CHONDRICHTHYES: SQUALIFORMES: DALATIIDAE), FROM TAIWAN.

The cestode fauna of Squaliolus aliae was examined for the first time following the collection of elasmobranch specimens from Taiwan in 2005, 2013, and 2017. This small shark was found to host 2 tapeworm species. These consist of Acanthobothrium katherineae n. sp., which is new to science and is described herein, and a second species, in the genus Scyphophyllidium, which also appears to be new, but which is represented by insufficient material for formal description. Acanthobothrium katherineae is a category 5 species. It can be distinguished from 5 of the 19 other category 5 species in that it is apolytic, retaining proglottids on its strobila until they are gravid. This new species differs from the remaining 14 category 5 species in its combination of the following features: It is a smaller worm, has fewer than 100 proglottids, has a relatively short cephalic peduncle, and differs in bothridial size and loculus ratio. Sequence data for the D1-D3 region of the 28S rDNA gene were generated for one specimen of A. katherineae. This sequence, along with comparable sequence data for adults of 14 described and 2 undescribed species as well as specimens of 6 undescribed larval members of the genus, was included in a maximum likelihood phylogenetic analysis. The resulting tree places the shark-hosted A. katherineae within a clade of stingray-hosted species, with Acanthobothrium romanowi as its sister taxon. Acanthobothrium katherineae is 1 of only 19 Acanthobothrium species known to parasitize sharks. The tree resulting from this study, which is preliminary given the relatively poor taxon sampling of the diversity in the genus, included 3 of the shark-parasitizing Acanthobothrium species and suggests that all 3 represent host-switching events. This is the first report of an Acanthobothrium species from the family Dalatiidae and the first report of a Scyphophyllidium species from the order Squaliformes. These findings suggest that other members of the Squaliformes, many of which have not previously been examined for parasites, may host additional novel cestode taxa.

Four new species of Acanthobothrium van Beneden, 1849 (Cestoda: Onchoproteocephalidea) from the spotted skate, Raja straeleni Poll, off the Western Cape, South Africa.

The examination of eight spotted skates, Raja straeleni Poll, resulted in the discovery of four new species of Acanthobothrium van Beneden, 1849, namely A. microhabentes sp. n., A. microtenuis sp. n., A. crassus sp. n., and A. dolichocollum sp. n., located off the Western Cape of South Africa. With a total of over 200 valid species of Acanthobothrium recognised worldwide, the use of an integrative approach becomes imperative in the interest of simplifying interspecific comparisons between congeners. In accordance with this, the four new species were incorporated into the category classification system established by Ghoshroy and Caira in 2001, where they were identified as category 2 species, which, at present, includes 47 recognised species of Acanthobothrium. Nevertheless, each of the four new species exhibits postovarian testes, a most intriguing and highly unusual feature among Acanthobothrium, instantly differentiating them from most congeners. This feature has been reported in 12 congeners, which have previously been considered to be restricted to waters of the Indo-Pacific Ocean. Not only do the four new congeners represent the first species of Acanthobothrium reported from southern Africa, but they also represent the first reported species with postovarian testes from the southern Atlantic Ocean. Regarding the legitimacy of the four new species, only two other category 2 species are reported to exhibit this feature, namely A. popi Fyler, Caira et Jensen, 2009, and A. bobconniorum Fyler et Caira, 2010, to which the four congeners were compared to. Acanthobothrium microhabentes sp. n. is the smallest of the congeners and differs from A. popi and A. bobconniorum by having fewer testes and postovarian testes, a shorter body, fewer proglottids, a shorter scolex, and longer cephalic peduncle. Acanthobothrium microtenuis sp. n. differs from A. popi and A. bobconniorum by having fewer testes and postovarian testes, a shorter scolex, longer cephalic peduncle, and the possession of columnar spinitriches on the anterior region of the terminal proglottid. Acanthobothrium crassus sp.n. differs from A. popi and A. bobconniorum by having fewer postovarian testes, a narrower cirrus-sac, larger vitelline follicles, and a longer cephalic peduncle. Acanthobothrium dolichocollum sp. n. is the longest of the four new species and differs from A. popi and A. bobconniorum by having fewer postovarian testes, more postporal testes, a larger body, more proglottids, larger testes and vitelline follicles, and an exceptionally long cephalic peduncle. Apart from differences in overall size, the four new species differ in a combination of measurements for the scolex, vitelline follicles, muscular pad and cephalic peduncle, and the number of proglottids and testes. The four species were recovered from a previously unexplored host and locality, expanding the host associations and geographical distribution of the genus.

Two new species of Acanthobothrium Blanchard, 1848 (Cestoda: Onchoproteocephalidea) from rajiform batoids off Argentina.

Two new species of Acanthobothrium Blanchard, 1848 were collected from rajiform batoids along the coast of Argentina. They are Acanthobothrium carolinae sp. n. from Bathyraja magellanica (Philippi) and Acanthobothrium domingae sp. n. from Dipturus brevicaudatus (Marini). Both belong to category 1 and 2 because their total length is lower than 15 mm, they have fewer than 50 proglottids, fewer than 80 testes and a symmetric or asymmetric ovary. Acanthobothrium carolinae sp. n. differs from all congeners by the following combination of features: proglottid apolysis, hook morphology, microthrix pattern on the cephalic peduncle and distribution of the vitelline follicles. Acanthobothrium domingae sp. n. is unique in the combination of proglottid apolysis, total length, number of proglottids, hook and septal morphology, microthrix pattern on the cephalic peduncle, number and distribution of testes, cirrus sac length, distribution of the vitelline follicles and genital pore position. The discovery of these species increases the number of species of Acanthobothrium parasitising rajiform batoids off Argentina from one to three. All three species have a unique pattern of distribution of the gladiate spinitriches along the length of the cephalic peduncle, i.e., gladiate spinitriches being interrupted in a medial line both dorsally and ventrally.

A New Onchoproteocephalidean Cestode Species from the Spot-Tail Shark, Carcharhinus sorrah (Müller et Henle), in the Persian Gulf.

Members of the onchoproteocephalidean genus Phoreiobothrium Linton, 1889 infect carcharhinid and sphyrnid sharks. Although the Persian Gulf harbors a rich fauna of sharks, the taxonomic diversity of Phoreiobothrium in this region has been explored little. A new species of Phoreiobothrium is herein described from Carcharhinus sorrah (Müller et Henle) off southern Iran. Cestodes were isolated from the spiral intestine of Carcharhinus sorrah, fixed in 10% seawater-buffered formalin, and stored in 70% ethanol. For morphological examinations, worms were stained using acetic carmine; a subset was prepared for SEM. Phoreiobothrium sorrahcola n. sp. is the first record of the genus from C. sorrah and most closely resembles Phoreiobothrium rozatii Alijanpour Darvishi et Haseli, 2019 recently described from the Gulf of Oman. However, it differs from P. rozatii in its possession of the rectangular, rather than triangular, bothridia, a longer cephalic peduncle (878-1300 vs 466-841), the longer axial prongs of lateral and medial hooks (B 56-80 vs 43-52; B' 66-98 vs 46-62), and a greater maximum width of worm at level of scolex (302-415 vs 215-298). With the description of this new species, three species of Phoreiobothrium have been reported so far from the southern waters of Iran. Phoreiobothrium now includes 17 valid species infecting 16 shark species.

Insights from new cestodes of the crocodile shark,Pseudocarcharias kamoharai(Lamniformes: Pseudocarchariidae), prompt expansion ofScyphyophyllidumand formal synonymization of seven phyllobothriidean genera – at last!

Three new cestode species are described from the crocodile shark (Pseudocarcharias kamoharai) in Ecuador. All three were examined with light and scanning electron microscopy. The unique combination of morphological features in one of the new species prompted formal investigation of the non-monophyly of Paraorygmatobothrium relative to the morphologically similar genera Doliobothrium, Guidus, Marsupiobothrium, Nandocestus, Orectolobicestus, Ruhnkecestus and Scyphophyllidium. Sequence data generated for part of the 28S rDNA gene were subjected to maximum likelihood (ML) analysis. The resulting tree led to the synonymization of six of these seven genera with Scyphophyllidium, and transfer of their species to the latter genus. With the new species, the number of described members of Scyphophyllidium is now 45. The diagnosis of Scyphophyllidium is revised to accommodate these species. In addition, to expedite future descriptions, eight categories of Scyphophyllidium species are circumscribed, based largely on bothridial features. Scyphophyllidium timvickiorum n. sp. is a category 1 species. Beyond being the smallest category 1 species, it bears, rather than lacks, apical suckers and lacks, rather than bears, strobilar scutes. The two other new species are members of Clistobothrium. Clistobothrium amyae n. sp. differs from its congeners in bothridial shape, elongate cephalic peduncle and tiny size. Clistobothrium gabywalterorum n. sp. differs from the two of its congeners that also possess foliose bothridia in overall size and testis number. Despite their substantial morphological differences, the ML tree indicates they are sister taxa. Both are unique among their congeners in possessing cephalic peduncle spinitriches. The diagnosis of Clistobothrium is revised accordingly.

Five new species of Acanthobothrium (Cestoda: Onchoproteocephalidea) from the long-tailed butterfly ray, Gymnura cf. poecilura (Elasmobranchii: Gymnuridae), from the Persian Gulf and Gulf of Oman.

Five new species of Acanthobothrium Blanchard, 1848 are described from Gymnura cf. poecilura from the Gulf of Oman and Persian Gulf. They all belong to the Category 1 and can be differentiated from all congeners by a combination of characters, including marginal lappets on the bothridial rim, the lack of spinitrich microtriches on the proximal bothridial surfaces, the position of the genital pore in the posterior one fifth of the proglottid, the direction of the cirrus sac parallel and clinging to the ovarian lobe, the lack of post-vaginal testes, and the interruption of the vitelline follicles by the ovary. The five new species are morphologically similar to each other but differ among each other in their cephalic peduncle length, proglottid and testes number, and the apolysis status. The most similar species to this new group is Acanthobothrium fogeli Gloldstein, 1964 from the Gulf of Mexico. The new species differ from A. fogeli by the muscular pad size, cephalic peduncle length and having marginal lappets on the bothridial rim. the species of Acanthobothrium occurs in three families of elasmobranchs in the Gulf of Oman and Persian Gulf (Dasyatidae, Rhynchobatidae and Gymnuridae). The true identity of many hosts in the region is ambiguous. Therefore, we designated the sampled elasmobranch as G cf. poecilura in accordance to the previously molecular study on a few individuals from the region.

Orygmatobothrium persiense n. sp. (Cestoda: Phyllobothriidea) from the Arabian Smooth-Hound Shark Mustelus mosis (Triakidae) in the Persian Gulf.

A new species of Orygmatobothrium Diesing, 1863 was discovered from Mustelus mosis Hemprich et Ehrenberg in the north-eastern Persian Gulf. Orygmatobothrium persiense n. sp. is the fourth valid species of its genus and differs from its congeners by the number of the proglottids. In addition, the testes distributed in 3-4 rows deep in cross sections, the length of the cephalic peduncle, and a conspicuous bothridial cleft above the apical sucker distinguish the new species, respectively, from O. juani, O. musteli, and O. schmittii. This is the first time that a species of Orygmatobothrium is described and reported outside the Atlantic Ocean.

Ultrastructure of cell types within the scolex and cephalic peduncle of the litobothriidean tapeworm, Litobothrium aenigmaticum

AbstractLitobothrium aenigmaticum Caira, Jensen, Waeschenbach, & Littlewood 2014 is a cestode species that parasitizes pelagic thresher sharks in Taiwan and the Gulf of California. A previous study using light microscopy suggested the scolex contains four types of unusual tissues, one of which was considered to aid in adhesion of the worm to the mucosa of the spiral intestine. The function of the other tissues was unknown. Also unknown was the function of two laterally paired ducts found extending throughout the length of the worm. The goal of the present study was to use transmission electron microscopy (TEM) to examine the internal anatomy of the scolex and cephalic peduncle of this tapeworm in more detail. Three specimens from Taiwan were examined with TEM and two with light microscopy. Transmission electron microscopy revealed that the laterally paired ducts were excretory, since they were closely associated with numerous protonephridia. The number of cell types, previously referred to as tissues, in the scolex was expanded to 11 based on a combination of features including cell size, nucleus size, ratio of the maximum diameter of the nucleus to maximum diameter of the whole cell, and types of observed organelles, as well as the presence or absence of cytoplasmic electron‐dense and electron‐lucent vesicles. The cytoplasm of all 11 cell types was found to include electron‐dense vesicles, and all types were periodic acid‐Schiff (PAS) positive. This suggests that the electron‐dense vesicles may contain glycoproteins and/or mucoproteins. As all 11 cell types occurred within the region of the scolex that is surrounded by a papilla of host mucosa at the site of tapeworm attachment, we hypothesize that one or more of these cell types may produce a substance that induces this inflammatory reaction. The specific products in the secretory vesicles and how those products are released are still unclear, as are any relationships between or among the cell types in terms of products produced. Further research will be necessary to fully understand the complexity of this organism.

Diphyllidean cestodes from the bigeye houndshark Iago omanensis (Norman) (Carcharhiniformes: Triakidae) in the Gulf of Oman, with the description of Coronocestus ehsanentezarii sp. nov. (Echinobothriidae).

A new species of Coronocestus Caira, Marques, Jensen, Kuchta and Ivanov, 2013 is described from Iago omanensis (Norman) from the Gulf of Oman. Coronocestus ehsanentezarii sp. nov. differs from C. musteli (Pintner, 1889) Caira, Marques, Jensen, Kuchta and Ivanov, 2013 in that its ovary is H- rather than U-shape. The new species is easily distinguished from C. diamanti (Ivanov and Lipshitz, 2006) Caira, Marques, Jensen, Kuchta and Ivanov, 2013 by the number of spines per column on the cephalic peduncle (24-36 vs 95-118). It differs from C. hormozganiense (Haseli, Malek, Palm and Ivanov, 2012) Caira, Marques, Jensen, Kuchta and Ivanov, 2013 based on a greater number of spines per column on the cephalic peduncle (24-36 vs 18-21). The new species differs from C. notoguidoi (Ivanov 1997) Caira, Marques, Jensen, Kuchta and Ivanov, 2013 in that of its scolex is craspedote rather than acraspedote. Unlike C. coronatum (Robinson 1959) Caira, Marques, Jensen, Kuchta and Ivanov, 2013, C. ehsanentezarii sp. nov. possesses 29-35 rather than 20 apical hooks in each dorso-ventral group. The new species possesses lateral hooklets with two rows (a and b designations) in each cluster. This character had been presented earlier only for Andocadoncum Abbott and Caira, 2014. Furthermore, a new locality record is presented for C. diamanti from Iago omanensis in the Gulf of Oman. Thus, I. omanensis certainly hosts two diphyllidean species simultaneously in the Gulf of Oman. The generic diagnosis of Coronocestus is also revised to include new data.

Halysioncum kishiense sp. n. and Echinobothrium parsadrayaiense sp. n. (Cestoda: Diphyllidea) from the banded eagle ray, Aetomylaeus cf. nichofii off the Iranian coast of the Persian Gulf

Two new species of two genera of the order Diphyllidea van Beneden in Carus, 1863, Halysioncum Caira, Marques, Jensen, Kuchta et Ivanov, 2013 and Echinobothrium van Beneden, 1849 sensu stricto are described from Aetomylaeus cf. nichoffi (Bloch et Schneider) off the Iranian coast of the Persian Gulf. Halysioncum kishiense sp. n. differs from all other congeners in the number of apical hooks with the exception of H. hoffmanorum (Tyler, 2001) and H. pigmentatum (Ostrowski de Núñez, 1971). Halysioncum kishiense sp. n. can be easily differentiated from H. hoffmanorum and H. pigmentatum by the number of hooklets and testis numbers. Echinobothrium parsadrayaiense sp. n. is differentiated from all its congeners except for E. acanthinophyllum Rees, 1961 by its hook formula. The number of spines per column on the cephalic peduncle, the number of testes and possession of a thick-walled rather than thin-walled vagina distinguish E. parsadrayaiense sp. n. from E. acanthinophyllum. To date, with these two new species, five species of Diphyllidea have been reported from the Persian Gulf.

A new combination and a new species of onchobothriid tapeworm (Cestoda: Tetraphyllidea: Onchobothriidae) from triakid sharks

A new combination and a new species of onchobothriid tapeworm are described from triakid sharks. We found Platybothrium xiamenensis Wang & Yang, 2001 and Erudituncus musteli (Yamaguti, 1952) from Hemitriakis japanica (Müller & Henle). Based on the morphology of the hooks P. xiamenensis is transferred to the genus Erudituncus Healy, Scholz & Caira, 2001. The specimens studied by us differ from the original description in the number of proglottids and testes and in the size of the cirrus-sac. However, we consider them conspecific with E. xiamenensis due to the consistent hook morphology and laciniations in both descriptions and believe the differences reflect intraspecific variation. The type-host of E. xiamenensis was reported as Mustelus griseus Pietschmann. However, in the present study, this parasite was found only in H. japanica and never in M. griseus although many specimens of the latter host were examined. This suggests that the type-host in the original description has probably been misidentified. We found another undescribed species in M. griseus, Calliobothrium shirozame n. sp., which is distinguished from the congeners by having a unique combination of the number of laciniations: four in the cephalic peduncle, six in the immature proglottids and four in the mature proglottids.

Morphology Meets Molecules: A New Genus and Two New Species of Diphyllidean Cestodes From the Yellowspotted Skate,Leucoraja wallacei, from South Africa

Two morphologically disparate undescribed species of diphyllidean cestodes from the Yellowspotted skate, Leucoraja wallacei , from South Africa were included in a recent molecular phylogenetic study aimed at revising diphyllidean classification. From a molecular standpoint, these species were determined to be only distantly related to one another. One (originally referred to as Echinobothrium n. sp. 2) showed affinities with species of the batoid-parasitizing Echinobothrium sensu stricto and is described here as Echinobothrium marquesi n. sp. This species most closely resembles Echinobothrium joshuai, an affinity supported by the previous molecular study, but differs in the form of its "B" hooks and degree of overlap between bothria and cephalic peduncle. The other species (originally referred to as New genus n. sp. 1), although exhibiting the full complement of scolex armature, grouped among primarily shark-hosted genera, most of which lack or exhibit reduced scolex armatures. That unexpected result suggested that erection of a novel genus might be warranted, but morphological grounds supporting the action were not apparent at that time. The present study aimed to explore the morphology of this taxon in more detail. Light and scanning electron microscopy revealed this taxon to be exceptional in that its lateral hooklets, which are arranged in 2 clusters like those of Echinobothrium and Coronocestus, are unique in being arranged in anterior and posterior rows, rather than in a single row. Andocadoncum n. gen. is erected, with Andocadoncum meganae n. sp. as its type, to accommodate this taxon. A minor adjustment to the existing hook formula by presenting counts for the anterior hooklets (a) separately from the posterior hooklets (b) readily accommodates this new form. Leucoraja wallacei is distinctive among skates in hosting 2 different genera of diphyllideans. In the context of the previous molecular phylogenetic analyses, it appears that the presence of Andocadoncum n. gen. in this skate species may be the result of a host-switching event involving a weakly or unarmed shark-hosted taxon moving to a batoid-a shift that was likely accompanied by a return to the fully armed condition seen, for example, in other batoid-parasitizing members of the order.

An enigmatic new tapeworm, Litobothrium aenigmaticum, sp. nov. (Platyhelminthes : Cestoda : Litobothriidea), from the pelagic thresher shark with comments on development of known Litobothrium species

An enigmatic new tapeworm is described from pelagic thresher sharks in México and Taiwan. While lsrRNA (D1-D3) data robustly place it in the Litobothriidea, it bears essentially no morphological resemblance to other members of the order. Instead it superficially resembles the freshwater fish-inhabiting Caryophyllidea. Its scolex consists of a simple dome-shaped scolex proper and an extensive cephalic peduncle housing four distinct tissue types. It is hyperapolytic, thus reproductive anatomy is unknown. Developmental data show typical litobothriideans bear basic elements of their adult scolex upon entering the definitive host, undermining the notion that the new cestode represents a distinct litobothriidean life cycle stage. Scanning electron microscopy revealed the new species shares bands of distinctive microtriches with its congeners. In combination these data justify establishment of Litobothrium aenigmaticum, sp. nov.; the generic, familial and ordinal diagnoses are emended accordingly. Unlike typical litobothriideans, each worm is associated with a mucosal expansion at its attachment site, like those seen in some caryophyllideans. This pathological change may represent a worm-induced host response serving to reinforce attachment of the simple scolex to the mucosa. If so, the convergence of this litobothriidean on a morphology like that seen in the distantly related Caryophyllidea is a result of similarity in mode of attachment.