First Report of Lecanicephalidean Tapeworms (Eucestoda) from Freshwater, Including Description of Three New Species of Tetragonocephalum Shipley and Hornell, 1905.

The lecanicephalidean cestodes parasitizing the spiral intestine of the endangered giant freshwater whipray, Urogymnus polylepis (Bleeker), are investigated for the first time. Eight host specimens were collected between 2002 and 2008 at 2 collecting sites off the eastern coast of Borneo: 6 from the Kinabatangan River (Malaysia) and 2 from a fish market in Tarakan (Indonesia). Two of these individuals were found to be infected with a total of 3 new species of TetragonocephalumShipley and Hornell, 1905. Tetragonocephalum georgei n. sp. and Tetragonocephalum opimum n. sp. were recovered from a host specimen from the Kinabatangan River, and Tetragonocephalum levicorpum n. sp. was found parasitizing a host specimen purchased at a fish market in Tarakan. Specimens of each of the new species were prepared for light microscopy; specimens of 2 of the new species were prepared for scanning electron microscopy, and histological sections were prepared for 1 of the new species. The 3 new species are distinct from the 9 valid species of Tetragonocephalum and the 1 species inquirendum based on, for example, total length, number of proglottids and testes, and size of the scolex and acetabula. Tetragonocephalum georgei n. sp. and T. levicorpum n. sp. are unusual among their congeners in that they are euapolytic (i.e., gravid proglottids were not observed) rather than apolytic. They differ from one another in scolex and acetabula size. Tetragonocephalum opimum n. sp. is unusual among its congeners in its possession of vitelline follicles arranged in 2, rather than 3, regions in the proglottid. These new species increase the total number of valid species of Tetragonocephalum to 12 and the total number of known cestodes from U. polylepis to 13 species across 6 genera in 4 orders. This is the first account of lecanicephalideans reported from freshwater. The taxonomic status of each of the 32 nominal taxa historically associated with Tetragonocephalum is re-assessed. Type host identities of all valid species are revised and discussed in light of recent taxonomic efforts in the Dasyatidae Jordan and Gilbert.