Oxygen availability in the aggregation of offspring has been shown to be a critical factor affecting the survival and development of embryos in aquatic systems. It is not yet known, however, to what extent the capacity to provide O 2 to embryo aggregations may act on the time of parental protection (here encapsulation), ultimately determining indirect and direct embryonic development. We assessed O 2 conditions during encapsulation, the factors determining those conditions, and the consequences on embryo survival in 2 gastropod species with contrasting developmental modes: Crepidula fornicata, an indirect developer, and Crepidula coquimbensis, a direct developer showing adelphophagy. Results showed that intracapsular O 2 conditions decreased to almost hypoxic conditions throughout development in C. fornicata, in contrast to the oxygenated conditions observed in C. coquimbensis during the entire encapsulation period. These contrasting patterns between species were explained by: (1) differences in metabolic rate of the embryos, (2) differences in total metabolizing material per capsule throughout development, and (3) differences in wall thickness and rates of decay throughout development, which may affect O 2 diffusion. Moreover, when the low O 2 conditions observed at the end of the encapsulation period were maintained after hatching by artificially extending encapsulation for 3 d, a dramatic negative effect on embryonic survival was observed in the indirect developer. In contrast, no effect on juvenile survival was observed in the direct developer. We suggest that hatching at intermediate stages of embryonic development in C. fornicata may be a response to increased O 2 constraints during the encapsulated period.
KEY WORDS: Parental protection · Oxygen limitation · Developmental mode · Larvae · Crepidula · EncapsulationResale or republication not permitted without written consent of the publisher Mar Ecol Prog Ser 368: 197-207, 2008 aggregation in jelly masses or capsules to active guarding of the embryos by the parents (Pechenik 1979, Rumrill 1990. A wide range of physical factors can also influence offspring development and survival, determining the need for parental protection, affecting its extent, and ultimately driving its evolution (Williams 1966, Trivers 1972, Clutton-Brock 1991, Roff 2002. In aquatic systems, the low solubility and diffusion rate of O 2 in water seem to be major constraints of parental protection through offspring aggregation (in amphibians: Crump 1996; in marine invertebrates: Booth 1995, Cohen & Strathmann 1996, Dick et al. 1998, Lee & Strathmann 1998, Fernández et al. 2000. Embryo aggregations of marine invertebrate species exhibit low O 2 levels which negatively affect embryonic development and survival (Strathmann & Strathmann 1982, Chaffee & Strathmann 1984, Strathmann & Chaffee 1984, Cancino et al. 2000. However, the consequences of reduced development and mortality risks during the protected phase on the optimal time of parental protection in aquatic systems are sti...