Ultrastructure of sperm and spermathecae in Micromaldane spp. (Polychaeta: Capitellida: Maldanidae)

Assessing the possibility that external fertilization has re-evolved requires the study of monophyletic groups that exhibit various reproductive methods. Maldanid polychaetes show a range of reproductive mechanisms, though previous studies of reproduction have hitherto been restricted to larger species with external fertilization. Micromaldane pamelae Rouse and M. nutricula Rouse are small, gonochoristic maldanids that brood directly developing larvae. Both species have sperm with elongate nuclei and an acrosome extending down each side of the anterior end of the nucleus. A true midpiece is absent; two mitochondria extend along the posterior region of the nucleus. Spermatids develop synchronously in large clusters connected by a cytophore. In M. pamelae sperm are released into the water as spermatozeugmata. These are comprised of clusters of sperm with their tails oriented to the centre and the sperm heads facing outwards. Females of M. pamelae and M. nutricula bear pairs of spermathecae ventrally (M. pamelae three pairs, between setigers 10 and 11, 11 and 12 and 12 and 13 and M. nutricula two pairs, between setigers 10 and 11 and setigers 11 and 12). The blind sacs are epidermal invaginations bound closely together. The entrance to each spermatheca may only be 1 to 2 m across with each spermatheca holding several hundred sperm. This represents the first detailed study of spermathecae in the Capitellida. The occurrence and structure of spermathecae and spermatozeugmata in other groups are discussed and compared with Micromaldane spp. Comparisons are made with non-polychaetes with the purpose of discussing functional aspects of reproductive mechanisms in marine metazoans in general. Elongate sperm nuclei are associated with sperm storage and/or large egg size. The lack of an elongate sperm midpiece may be an indicator of having to swim in water but does not contraindicate sperm storage. Spermatozeugmata may serve as an indication of sperm storage and brooding of larvae. Speculations on the phylogenetic significance of these reproductive features are limited by the fact that supposedly modified (i.e., derived) states may reflect functional/structura, constraints of small body size.