This review brings together decades of work on elasmobranch tapeworms. The field has advanced significantly over the past 20 yr, with an emphasis on the discovery and description of novel taxa, and the establishment of phylogenetic frameworks for individual orders and their interrelationships. Tapeworms parasitizing elasmobranchs represent 9 orders and include 977 species and 201 genera-over 250 species and 50 genera are new within the last 2 decades. The 9 orders are treated individually, highlighting recent assessments of phylogenetic relationships informed by molecular sequence data. All but the "Tetraphyllidea" are monophyletic. Although much remains to be learned about their interrelationships, existing phylogenetic hypotheses suggest that elasmobranch tapeworms have played a key role in the evolution of the cestodes of essentially all other vertebrate groups. The apical organ is a defining feature (i.e., a synapomorphy) of a clade consisting of acetabulate taxa and Litobothriidea. Novel hook amino acid composition data support the independent origin of hooks in the various groups of hooked tapeworms. Cestode records exist for representatives of most of the major groups of elasmobranchs, however skates (Rajiformes) and catsharks ("Scyliorhinidae") are particularly neglected in terms of species sampled. The majority of tapeworm species are extremely host-specific exhibiting species-specific (i.e., oioxenous) associations with their hosts. Rapid advancements in elasmobranch taxonomy, with over 300 of the 1,200 species appearing new in the past 20 yr, signal the need for careful attention to be paid to host identifications; such identifications are best documented using a combination of specimen, photographic, and molecular data. Above the species level, many cestode taxa are restricted to host orders, families, or even genera. Documentation of these affiliations allows robust predictions to be made regarding the cestode faunas of unexplored elasmobranchs. Trypanorhynchs are the notable exceptions. Life cycles remain poorly known. Recent applications of molecular methods to larval identifications have reinvigorated this area of research. Tapeworms are more diverse in elasmobranchs of tropical and subtropical waters, but they occur globally not only at the poles and in deep waters, but also in freshwaters of South America and Southeast Asia. The cestode faunas of batoids are much more speciose and complex than those of sharks. The faunas of deeper water sharks are particularly depauperate. The tapeworms of elasmobranchs and their hosts are now among the most well documented host-parasite systems in existence. This system has not yet reached its potential as a resource for investigations of basic ecological and evolutionary principles.

https://bioone.org/journals/journal-of-parasitology/volume-100/issue-4/14-516.1/A-Digest-of-Elasmobranch-Tapeworms/10.1645/14-516.1.pdf

A digest of elasmobranch tapeworms.

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.

https://bioone.org/journals/journal-of-parasitology/volume-90/issue-2/GE-3128/FIVE-NEW-SPECIES-OF-PEDIBOTHRIUM-TETRAPHYLLIDEA--ONCHOBOTHRIIDAE-FROM-THE/10.1645/GE-3128.pdf

Five new species of pedibothrium (tetraphyllidea: onchobothriidae) from the tawny nurse shark, nebrius ferrugineus, in the pacific ocean

The first records of the tapeworm genus Pedibothrium from nurse sharks ( Ginglymostoma cirratum ) in the eastern Pacific and eastern Atlantic Oceans provide new insights on the age of the association between these tapeworms and this host. Four individuals of G. cirratum examined from the southern tip of the Baja Peninsula, Mexico, were found to host P. manteri and P. brevispine . Tapeworm specimens discovered in the collection of J. Cadenet, taken from G. cirratum off Goree near Dakar, Senegal in 1947, were of P. globicephalum and P. manteri . These tapeworm faunas are consistent with those found previously in small and large nurse sharks, respectively, in the western Atlantic Ocean. The distributional data, combined with the non-vagile habits of the nurse shark, indicate this host–parasite association dates from the most recent completion of the Panamanian Isthmus, approximately 2 Mya. This suggests that P. manteri and P. brevispine have remained unchanged for at least 2 Myr. The data from the eastern Atlantic Ocean are less informative. The amphi-Atlantic distribution of P. manteri and P. globicephalum in G. cirratum may have resulted from vicariant events associated with the formation of the Atlantic Ocean. However, fossils of at least two extinct species of Ginglymostoma are also known from both sides of the Atlantic Ocean. Short of parallel evolution of three different species of Ginglymostoma on either side of the Atlantic Ocean, these data seem more consistent with dispersal as an explanation for this disjunction and are thus of limited utility for extending the age of this host–parasite association at this time. Analysis of the degree of genetic divergence among individuals of G. cirratum and among individuals of the broadly distributed species of Pedibothrium from all three localities would be extremely interesting.

Age of association between the nurse shark, Ginglymostoma cirratum, and tapeworms of the genus Pedibothrium (Tetraphyllidea: Onchobothriidae): implications from geography

In an effort to expand knowledge of Clade 3-one of the ten clades that compose the non-monophyletic order 'Tetraphyllidea' all current members of which parasitise orectolobiform sharks-we targeted species of orectolobiform sharks that had not previously been examined for 'tetraphyllidean' cestodes. That work led to the discovery of three new species off Australia and Taiwan. Ambitalveolus gen. n. was erected to accommodate these species. Ambitalveolus costelloae gen. n. et sp. n., Ambitalveolus kempi sp. n., and Ambitalveolus penghuensis sp. n. differ from one another in scolex size, genital pore position, and number of marginal loculi, proglottids, and testes. Among 'tetraphyllideans', the new genus most closely resembles the two other genera in Clade 3. It differs from Carpobothrium Shipley et Hornell, 1906 in lacking anterior and posterior flap-like extensions of its bothridia; instead, its bothridia are essentially circular. It differs from Caulopatera Cutmore, Bennett et Cribb, 2010 in that its vitelline follicles are in two lateral bands, rather than circum-medullary, and in that its bothridia bear, rather than lack, conspicuous marginal loculi. A key to the three genera in Clade 3 is provided. A phylogenetic analysis including new sequence data for one of the three new species of Ambitalveolus gen. n., the only species of Caulopatera, and all four described species and one undescribed species of Carpobothrium supports previously hypothesised close affinities between Caulopatera and Carpobothrium, with the new genus as their sister group. This is the first report of 'tetraphyllidean' cestodes from the orectolobiform shark family Brachaeluridae Applegate. The association of the new species with orectolobiform sharks is consistent with those of the other members of Clade 3. However, whereas species of Carpobothrium and Caulopatera parasitise members of the hemiscylliid genus Chiloscyllium Müller et Henle, species of Amitalveolus gen. n. parasitise members of the Brachaeluridae and Orectolobidae Gill.

Diversity and phylogenetic relationships of 'tetraphyllidean' Clade 3 (Cestoda) based on new material from orectolobiform sharks in Australia and Taiwan.

A new species of Pedibothrium (Cestoidea: Tetraphyllidea) from the short-tail nurse shark, Pseudoginglymostoma brevicaudatum (Elasmobranchii: Orectilobiformes), from southwest Madagascar

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A new species of Pedibothrium (Cestoidea: Tetraphyllidea) from the short-tail nurse shark, Pseudoginglymostoma brevicaudatum (Elasmobranchii: Orectilobiformes), from southwest Madagascar