Podotreme affinities of Raninoides sp. and Lyreidus brevifrons: evidence from spermatozoal ultrastructure (Crustacea: Brachyura: Raninoidea)

Spermatozoal synapomorphies which singly or collectively distinguish Raninoidea are: (1) the presence of single (Ranina, Raninoides) or multiple (Lyreidus) keel-like projections of the acrosomal capsule; (2) a very large, weakly electron-opaque peripheral acrosomal zone (Ranina, Raninoides) and an homologous large outer zone in Lyreidus; (3) poor differentiation of the operculum from the capsule (autapomorphy); (4) a very well developed, perforate subopercular zone, of variable form; (5) presence of unique inward longitudinal projections (septa or corrugations) in the wall of the subacrosomal chamber (autapomorphy). Shared, presumably synapomorphic characters of Ranina and Raninoides but not of Lyreidus within the Raninidae, are: (1) branching of some of the subacrosomal septa (unbranched in Lyreidus); (2) the subspheroidal form of the acrosome in Ranina with a length: width ratio (L:W) of 0.76, and, although slightly more depressed, in Raninoides (L:W 0.73), considered apomorphic relative to the more depressed form in Lyreidus (L:W 0.52); (3) single or multiple coiled perforatorial filaments (Ranina, Raninoides) contrasting with a capitate perforatorium with “amoeboid” head in Lyreidus; (4) division of the capsule wall to give one posterior (Ranina) or multiple enclaves, plesiomorphically (?) absent in Lyreidus. Similarities of Lyreidus with other podotremes include the capitate perforatorium, questionably related to the radiate spiked-wheel structure of homolids in which acrosome proportions are similar or less closely to the bilateral capitate perforatorium of dromiids and dynomenids, and basal capsular projections as in the dromiid Stimdromia (=Petamolera) lateralis and in cyclodorippoids. No spermatozoal synapomorphies support a sister-group relationship of raninoids and heterotreme-thoracotreme crabs.