Ultrastructure of the female pedal gonad in Phoxichilidium femoratum (Chelicerata, Pycnogonida)

Phoxichilidium femoratum is a common species of sea spiders – a small and unique group of chelicerates with unusual adult anatomy. In particular, substantial parts of the reproductive system in pycnogonids (unlike euchelicerates) are located in the appendages. Existing studies of pycnogonid gonads are often limited to light-microscopic level, cover a small range of species, and focus on the contents of the gonad diverticula. Ultrastructural data are rare and contradictory, and the organisation of the gonad wall and the gonoducts is unknown. Here we present a detailed light and transmission electron microscopy-based examination of the pedal portion of the adult female reproductive system in Phoxichilidium femoratum Rathke, 1799. We describe its gross anatomy and the ultrastructure of the gonad diverticulum, oviduct and gonopore, as well as development of the oocytes. Each gonad diverticulum is enclosed in the extracellular matrix of the horizontal septum and bears some internal cellular lining. However, neither the gonad lining, nor the septum sheath cells, ever form a continuous epithelial layer. Oocytes, which undergo maturation in the diverticulum, remain, until very late in the process, attached to the gonad wall though specialised stalk cells. Interestingly, stalk cells do not participate in egg envelope or yolk formation: both are synthesized endogenously in the oocytes. The oviduct is supplied with musculature, which assists in egg transport to the gonopore, whereas the gonopore itself is surrounded by specialised glands.