Phylogenetic Implications of Sperm Storage in Podotremata: Histology and 3D-Reconstructions of Spermathecae and Gonopores in Female Carrier Crabs (Decapoda: Brachyura: Homoloidea)

Female reproductive systems are important characters for understanding the evolution of Brachyura and resolving its phylogenetic relationships. We herein investigate a podotreme brachyuran reproductive system comprehensively for the first time studying spermathecae and gonopores of Homoloidea with histological methods, micro-computer tomography and scanning electron microscopy. Our results show that spermathecal apertures are species-specific and their shape corresponds closely to that of male copulatory organs. Apertures were either enclosed by membranous cuticle areas or otherwise occluded preventing direct access into spermathecae. 3D-reconstructions reveal that spermathecae differ between the species Paromola cuvieri and Homola barbata with regard to the involvement of sternite 7 and 8, respectively, in forming the sperm storage chamber. The cuticle epithelium that lines the spermathecal chamber is irregular and distinct from the remaining cylindrical cuticle epithelium. A first uniramous pleopod was present in all homoloids studied and always held in a position to cover spermathecal apertures. Specific pulvinated cuticle structures present on both sides of the first pleopod are herein interpreted as adhesive structures functioning in reproductive processes. The coxal gonopores were enclosed by a laterally arising muscular mobile operculum that resembles opercula described in eubrachyuran vaginae, which raises the question whether these two structures are homologous. Our results are compared with data available for other brachyuran groups and discussed in terms of phylogenetic relationships within the Brachyura and possible functions in insemination and fertilization in Podotremata. (C) 2016 Wiley Periodicals, Inc.