Towards a phylogenetic classification of dendrocoelid freshwater planarians (Platyhelminthes): a morphological and eclectic approach

We explore and review the taxonomic distribution of morphological features that may be used as supporting apomorphies for the monophyletic status of various taxa in future, more comprehensive phylogenetic analyses of the dendrocoelid freshwater planarians and their close relatives. Characters examined are: (i) musculature of the pharynx; (ii) anatomy of the eye; (iii) distribution of testis follicles along the longitudinal axis of the body; (iv) epidermis of the body wall; (v) musculature of the ventral body wall; (vi) presence or absence of a penis papilla; (vii) position of the mouth opening; and (viii) anterior adhesive organ. A phylogenetic hypothesis, in the form of a character state tree, is developed. This hypothesis suggests that the Kenkiidae should be resurrected as a separate family, sharing a sistergroup relationship with the Dendrocoelidae. This sistergroup relationship is supported by the shared presence of an apomorphic anterior adhesive organ. Although the shape of the anterior adhesive organ, both within and between kenkiids and dendrocoelids, may differ considerably between species, they are all constructed according to the same principles. The monophyly of the Kenkiidae is supported by the presence of an apomorphic thickening of the marginal zone of the epidermis and presumably also by the presence of prepharyngeal testes. Within the Kenkiidae, the genus Macrocotyla should be considered as a junior synonym of Kenkia, which is characterized by the presence of an apomorphic Ôplanariid pharynx-type 2Õ. The monophyly of the Dendrocoelidae is suggested by the presence of a dendrocoelid type of pharynx. It is shown that multicellular eye cups with numerous photoreceptor cells are not restricted to the Dugesiidae and the Terricola but are also characteristic for the Dendrocoelidae; the phylogenetic implications of this character state distribution are discussed. It is pointed out that in planarians there may be a relationship between: (i) body size, the size of an eye cup and the number of its constituent pigment cells; and (ii) the size of an eye cup and the number of photoreceptor cells that it contains. Some tentative phylogenetic groups are suggested on the basis of the following presumed apomorphic characters: presence of transverse fibres in the ventral subepidermal musculature, presence of an extra longitudinal muscle layer in the outer pharynx musculature, a mouth opening that has shifted anteriad. It is suggested that the presumed monophyly of the genus Bdellocephala is supported by the apomorphic absence of a penis papilla, combined with a penial lumen provided with many plicae.