Paradorippe granulata - A crab with external fertilization and a novel type of sperm storage organ challenges prevalent ideas on the evolution of reproduction in Eubrachyura (Crustacea: Brachyura: Dorippidae)

Abstract

Two fundamentally different sperm storage organs occur in Brachyura. The probably paraphyletic podotremes show intersegmental spermathecae, which are distant from oviducts and coxal gonopores. Hence, fertilization is external. In contrast to this, the seminal receptacles of Eubrachyura are directly connected with the ovaries. Thus, at least initial fertilization is internal. This pattern has been interpreted as an apomorphy of Eubrachyura. To test this hypothesis, we studied the morphology of the reproductive organs of Paradorippe granulata, a representative of the putatively early diverging eubrachyuran lineage Dorippoidea. Applying histology, 3D-reconstructions and micro-computed-tomography we revealed a novel type of sperm storage organ. Female P.granulata lack the characteristic eubrachyuran seminal receptacle. Instead sperm is stored in four cuticle-lined bursae, two on each side of the paired oviducts. The elaborate bulbous male gonopod with several terminal processes is adapted to transferring sperm into the female twin bursae. Since oviducts and twin bursae are not directly connected, spermatozoa and oocytes mix when gametes pass through the sternal vulva. Thus, fertilization in P.granulata is external. Our finding of a eubrachyuran crab that lacks seminal receptacles and exhibits external fertilization calls prevailing concepts on the evolution of sperm storage in Eubrachyura into question.

Keywords: Bursa; Copulatory organs; Gonopod; Histology; Reproductive system; Seminal receptacle.