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.