Spermatozoa of an Old World Ricinulei (Ricinoides karschii, Ricinoidae) with notes about the relationships of Ricinulei within the Arachnida

The ultrastructure of spermatozoa is a valuable tool for phylogenetic and systematic studies. Ricinulei are enigmatic and poorly studied arachnids. So far, spermatozoa are only known from New World ricinuleids. The goals were to study, by means of light and transmission electron microcopy, the spermatozoa of an Old World species with regard to their phylogenetic implications, e.g., does the sperm structure contribute to the debated sister-group relationship of Acari and Ricinulei. The spermatozoa are coiled-flagellate and characterized by a cap-like acrosomal vacuole covered by electron-dense material, an elongated nucleus covered by a manchette of microtubules during spermiogenesis, an axoneme with a 9+2 microtubular pattern, a nuclear tube and axonemal basis which both originate underneath the acrosomal vacuole and cleistospermia as transfer form equipped with three intracellular plates. The data of the present study did not support a close relationship of Ricinulei and Acari which have aflagellate sperm with various synapomorphies as e.g., lacking nuclear envelopes/membranes in Actinotrichida (very similar to Solifugae) or vacuolated spermatozoa in Anactinotrichida. Affinities of Ricinulei are discussed in the light of the ultrastructure of arachnid spermatozoa.