The ecology, functional morphology and taxonomy of Echeneibothrium Beneden, 1849 (Cestoda: Tetraphyllidea), a revision of the genus and comments on Discobothrium Beneden, 1870, Pseudantho-bothrium Baer, 1956, and Phormobothrium Alexander, 1963

Two new, nine little known species and nine unnamed ‘species’ of Echeneibothrium Beneden, 1850, have been investigated and observations have been made on the ‘related’ genera, Discobothrium Beneden, 1870, and Pseudanthobothrium Baer, 1956. Brief descriptions of the species are given, but a discussion of the history of each differs greatly from most previous accounts in that few or no synonyms have been listed for the species and in that nine forms are left unnamed; this approach differs greatly from that in a large number of taxonomic papers in helminthology. A detailed discussion of the reasons for adopting this procedure is, therefore, given. A provisional key to nineteen ‘species’ of Echeneibothrium is included. E. minutum sp.nov. closely resembles E. variabile var. exiguum of Euzet (1959), described in this paper as ‘Echeneibothrium sp. of R. clavata’, but occurs only in R. batis and differs in its very small size, in having only about ten proglottids, long peduncles to the bothridia and a myzorhynchus which remains evaginated in fixed specimens. It differs from E. demeusiae Euzet, 1959, mainly in the number of proglottids, testes and loculi. E. elongatum sp.nov. resembles E. dubium from which it differs in the length and form of the strobila, in having only six clearly visible loculi in each bothridium, about twenty testes and in that it occurs in R. circularis. It has, however, more characters in common with a species referred to in this paper as ‘Echeneibothrium sp. from Raja naevus’ and which possesses eight loculi. A list is given of twenty-seven species wrongly allocated by various authors to the genus Echeneibothrium. The comparative morphology of some species of the genus, their ecology and speculations on their life-histories, are described. It is shown for the first time that Echeneibothrium from species of Raja caught off the British Isles, and presumably from rays of other localities, can be separated into two distinct biological groups. One group includes species which possess shallow open bothridia covering comparatively large areas of the host's mucosa, a variable myzorhynchus which may be either rudimentary or very large and spherical eggs laid singly or in capsules; these occur mainly in rays from shallow waters and in which the mucosa of the intestine lacks prominent villi. The other group includes species in which each bothridium possesses a small opening adapted for attachment to a single villus, the myzorhynchus is consistently cylindrical, the eggs have long polar filaments and the species occur in deep water rays with well-defined villi on the intestinal mucosa. Studies on variation in the form of the eggs in different species of Echeneibothrium in relation to the behaviour and feeding habits of the various host-species suggest that the eggs are eaten by arthropods which are most likely to form part of the diet of the final host. A general trend towards an increase in the complexity of the bothridium and a decrease in the size of the myzorhynchus in Echeneibothrium is thought to be of considerable advantage to the species and an indication that Rhinebothrium, which is restricted to the Myliobatoidea, has evolved from a form like Echeneibothrium which is restricted to the Rajoidea. This supports the view that the Myliobatoidea have evolved from the Rajoidea. The results from this study of Echeneibothrium illustrate the following general rule which can be applied to the Tetraphyllidea, namely, those species which are abundant in any given host are very well adapted for attachment to the host's gut and cause little, if any, damage while rare species are often highly pathogenic. This work would not have been possible without generous financial aid from the Department of Scientific and Industrial Research and various kinds of other aid and encouragement from a number of persons, in particular Dr Gwendolen Rees. It was begun in 1958 at the University College of Wales, Aberystwyth, the British Museum (Natural History), University of Montpellier, France, University of Neuchatel, Switzerland, the Plymouth Marine Laboratory and the Aberdeen Marine Laboratory during my tenure of a D.S. I.R. Fellowship. I wish to record my gratitude to Dr Rees who has also suggested improvements to the manuscript and to the following persons: the late Professor T. A. Stephenson, F.R. S. , Dr E. E. Watkin, Mr S. Prudhoe, Dr L. Euzet, Professor Jean G. Baer, The Director, Dr F. S. Russell and Staff, in particular Mr J. Green and Mr A. Mattacola, of the Plymouth Marine Laboratory, and the Director and staff, in particular Dr B. B. Rae and Dr Z. Kabata, of the Aberdeen Marine Laboratory. The work was later continued at the University College, Cardiff, and I am most grateful to Professor James Brough for providing excellent research facilities and to Mr W. O'Grady for technical assistance. At the Bureau of Helminthology, St Albans, much encouragement from Professor R. T. Leiper, C.M. G. , F.R. S. , enabled me to put the work together while having free access to his invaluable collection of books and reprints on helminths; it became unnecessary to search elsewhere for the literature. Mr G. Dimmock gave technical assistance for which I am most grateful.