The status of Rhoptrobothrium Shipley et Hornell, 1906 (Cestoda: Tetraphyllidea), with redescription of the type species, R. myliobatidis, and description of three new species from two species of Aetomylaeus (Myliobatiformes: Myliobatidae) from Malaysian Borneo

As part of a metazoan parasite survey of elasmobranchs from Malaysian Borneo, specimens of Rhoptrobothrium Shipley et Hornell, 1906 were collected from the eagle rays Aetomylaeus maculatus (Gray) and Aetomylaeus niehofii (Bloch et Schneider). The type species is redescribed from its type host, and a neotype specimen is designated. In addition, three new species of Rhoptrobothrium are described: R. chongi sp. n., R. gambangi sp. n. and R. limae sp. n. Rhoptrobothrium myliobatidis conspicuously differs from the three new species in its lack of a secondary areola; R. limae is distinguished from R. chongi and R. gambangi based on its greater total length; R. chongi possesses conspicuously stalked remi, while R. gambangi possesses short remi, often folded anteriorly. Rhoptrobothrium is somewhat unusual among tetraphyllideans in its possession of a "metascolex," a character it shares with other taxa in the Thysanocephalinae (i.e., Myzocephalus Shipley et Hornell, 1906, Myzophyllobothrium Shipley et Hornell, 1906 and Thysanocephalum Linton, 1889). The morphology of the "metascolex" of Rhoptrobothrium is investigated and new terminology is suggested to standardise the names given to structures constituting a metascolex. As a result, Rhoptrobothrium is considered to possess cephalic peduncle extensions, termed remi. In Rhoptrobothrium, each remus bears, at its distal end, a primary areola, and, in the case of the three new species, also a secondary areola proximal to the primary areola. Myzocephalus and Myzophyllobothrium are tentatively considered to possess remi; the configuration of the "metascolex" of Thysanocephalum, however, is not considered homologous to the condition in the other three genera currently placed in the Thysanocephalinae.