Brood care in freshwater crayfish and relationship with the offspring's sensory deficiencies.

Prolonged brood care is one of the evolutionary clues for the successful colonization of freshwater habitats by freshwater crayfish (Astacida). By means of macrophotography, light microscopy, and scanning electron microscopy we investigated all phases of brood care in freshwater crayfish, with particular emphasis on the morphological structures involved. We selected the recently discovered parthenogenetic marbled crayfish (species identity not yet known) as a model organism due to its fast reproduction and high resistance to handling stress. In order to examine if there is a causal relationship between brood care and the developmental status of the offspring's sensory apparatus, we additionally investigated major sense organs of juvenile Stages 1-5 in comparison with those of the adults. Brood care in the marbled crayfish is characterized by initial and final "active" phases dominated by specific maternal or juvenile behavior and a medial "passive" phase based more on the action of temporarily developed structures rather than on behavior. The most remarkable feature of this period, which includes permanent carrying of the eggs and the first two juvenile stages under the mother's abdomen, is safeguarding of hatching by a telson thread that keeps the helpless newborn hatchlings linked to the egg cases on the maternal pleopods and thus prevents them from being lost. Further important transient structures are the recurved hooks on the first pereiopods of Stage 1 and 2 juveniles that are used to firmly attach these nonfeeding stages to the mother's abdomen. In hatchlings all sense organs necessary for an independent life, such as eyes, olfactory aesthetascs, gustatory fringed setae, hydrodynamic receptor hairs, and statocysts are not developed or are underdeveloped, making brood care indispensable. Most of these sense organs appear in Stage 2 juveniles, but only from Stage 3, the first freelancing and feeding stage, are all sense organs well developed and operating, thus reducing brood care in this final period to temporary provisioning of shelter. Brooding of the eggs and postembryonic brood care are to some extent also found in other freshwater Decapoda like freshwater crabs and aeglid anomurans, but safeguarding of hatching is confined to the Astacida only. This sophisticated mode of passive brood care is unique in the animal kingdom and is apparently related to the sensory deficiencies of the first juvenile stage.

Life stages and reproductive components of the Marmorkrebs (marbled crayfish), the first parthenogenetic decapod crustacean.

Recently, we briefly reported on the first case of parthenogenesis in the decapod Crustacea which was found in the Marmorkrebs or marbled crayfish, a cambarid species of unknown geographic origin and species identity. Curiously, this animal is known only from aquarium populations, where it explosively propagates. By means of light and electron microscopic techniques we have now investigated the reproductive components of this crayfish, using more than 100 specimens ranging from hatchling to repeatedly spawned adult. Additionally, we documented its principal life stages. Our results revealed that the external sexual characters and also the gonads of the marbled crayfish are purely female, making this fast-reproducing species a good model for investigating female reproductive features in crayfish. Testicular tissues, ovotestes, or male gonoducts, gonopores, or gonopods were never found, either in small juveniles or large adult specimens, confirming the parthenogenetic nature of this crayfish. Parthenogenesis may have arisen spontaneously or by interspecific hybridization since Wolbachia-like feminizing microorganisms were not found in the ovaries. The external sexual characters of the marbled crayfish are first recognized in Stage 4 juveniles and are structurally complete approximately 2 months after hatching in specimens of approximately 2 cm total length. In the same life stage the ovary is fully differentiated as well, although the oocytes are in previtellogenic and primary vitellogenic stages only. The architecture of the mature ovary and also the synchronous maturation of cohorts of primary vitellogenic oocytes by secondary vitellogenesis are in general agreement with data published on ovaries of bisexual crayfish. New results were obtained with respect to the muscular nature of the ovarian envelope and its extensive proliferation after the first spawning, the distribution of hemal sinuses in the ovarian envelope and in the interstitium around the oogenetic pouches, the high transport activity of the follicle cells, and the colonization of oogenetic pouches by previtellogenic oocytes that originate in the germaria. Investigation of the nuclei of oocytes in the germaria and oogenetic pouches revealed no signs of meiosis, as usually found in females of bisexual decapods, suggesting that parthenogenesis in the marbled crayfish might be an apomictic thelytoky. The detection of new rickettsial and coccidian infections in the ovary and further organs raises fears that the marbled crayfish might endanger native European species by transmission of pathogens once escaped into the wild.