Ultrastructure of spermatophoral sperm in the freshwater gastropod Thiara amarula (Linnaeus, 1758) (Cerithioidea: Thiaridae): potential taxonomic features including eusperm nuclear content differentiation

The ultrastructure of spermatophoral sperm is investigated for the first time in a cerithioidean gastropod. Thiara amarula (Linnaeus, 1758), a widespread Indo-West Pacific species of the Thiaridae, produces uniaxonemal euspermatozoa and multiaxonemal paraspermatozoa. Euspermatozoa possess a laterally-flattened acrosomal vesicle (with eccentrically positioned invagination and subacrosomal material), a tubular, laterally-compressed nucleus, an elongate midpiece (with four, straight, equal-sized mitochondria each with parallel cristae), an elongate glycogen piece and an end piece. The eusperm nucleus exhibits or is associated with a number of unusual features including: (1) nuclear contents differentiated into two components (fibrous, highly electron-dense material, enclosing a pair of less-dense, finely granular tracts); (2) nuclear invagination with two elongate grooves aligned at 90 degrees to the axonemal central microtubules and parallel with the 1 and 5 axonemal doublets; (3) a periodically-banded rootiet associated with the centriolar complex near the nuclear apex; (4) an electron-dense structure attaching the nucleus to the axoneme at the nucleus-midpiece junction. Feature (1) has been reported elsewhere in Melanoides tuberculata (Thiaridae), but in no other examined gastropods (or other molluscs) and its purpose is unknown. Paraspermatozoa of T. amarula possess an elongate head region (up to 14 axonemes and large, irregular mitochondria enclosed by sheath of spherical, dense vesicles), a short 'glycogen' region and a posterior tuft of glycogen tails. The two sperm types do not form spermatozeugmata and observations suggest that neither type undergoes any noticeable structural change while inside the spermatophore. Sperm features suggest a close relationship between T. amurula and M. tuberculata and more broadly with other 'Group T cerithioidean families exhibiting four equal-sized eusperm mitochondria, including Potamididae, Modulidae, Scaliolidae, Melanopsidae, Semisulcospiridae and Pleuroceridae.