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The ability to withstand high summer temperatures is an extremely important determinant of species distributions in the intertidal zone; most marine organisms cannot live intertidally because of the harsh conditions experienced during aerial emersion. The members of some species, though, live both intertidally and subtidally, and their physiological tolerance to stressors may differ depending on their genetic connectivity, acclimatization, and differential post settlement mortality. Furthermore, tolerances of organisms at different life-history stages within both habitats may differ and will play an important role in determining adult distributions. To determine the effect of habitat and life-history stage on physiological tolerance and the likely contribution of upper thermal tolerance limits to the native distribution of an important invasive species, the slippershell snail Crepidula fornicata, we compared the thermal tolerance of intertidal and subtidal embryos and adults in relation to conditions that they experience in the field. We found C. fornicata living both intertidally and subtidally in Bissel Cove, Rhode Island, where they experienced drastically different thermal conditions. Intertidal individuals achieved body temperatures as high as 42°C, more than 15°C higher than those recorded in subtidal conspecifics. However, subtidal individuals had remarkably high thermal tolerances that were nearly identical to those of intertidal conspecifics (lethal range = 33−37°C following a 3 h exposure). Furthermore, embryos were even more tolerant to high thermal stress than adults. These results are surprising: the few previous studies that have compared thermal tolerances among individuals at different life-history stages have found differences among them, and early stages are generally more sensitive than adults. Interestingly, the markedly higher temperatures that intertidal animals experienced in this study had little effect on their thermal tolerance. Intertidal (but not subtidal) slippershell snails are now living dangerously close to their upper thermal limits and will almost certainly be relegated to the subtidal as global temperatures rise.

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Brooding in invertebrates serves to protect embryos from stressful external conditions by retaining progeny inside the female body, effectively reducing the risk of pelagic stages being exposed to predation or other environmental stressors, but with accompanying changes in pallial fluid characteristics, including reduced oxygen availability. Brooded embryos are usually immobile and often encapsulated, but in some Ostrea species the embryos move freely inside the female pallial cavity in close association with the mother’s gills for as long as eight weeks. We used endoscopic techniques to characterize the circulation pattern of embryos brooded by females of the oyster, Ostrea chilensis. Progeny at embryonic and veliger stages typically circulated in established patterns that included the use of dorsal and ventral food grooves (DFG, VFG) to move anteriorly on the gills. Both embryos and veligers accumulated around the mother’s palps, and remained there until an active maternal countercurrent moved them to the gill inhalant area. Both food grooves were able to move embryos, veligers, and food-particle aggregates anteriorly, but the DFG was more important in progeny transport; early embryos were moved more rapidly than veligers in the DFG. A microcirculation pattern of embryos was apparent when they were moved by gill lamellae: when they were close to the VFG, most embryos lost gill contact and ´´fell´´ down to the DFG. Those that actually reached the DFG moved anteriorly, but others came into contact with the base of the lamellae and again moved towards the VFG. The circulation pattern of the progeny appears well-suited for both cleaning them and directing them posteriorly to an area where there is more oxygen and food than in the palp region. This process for actively circulating progeny involves the feeding structures (gill and palps) and appears to be energetically costly for the female. It also interferes with feeding, which could explain the poor energy balance previously documented for brooding females of this species.

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Organisms that encounter stressful situations in nature often cope using behavioral (e.g., avoidance) or physiological tactics. In sessile mollusks, the only available behavioral option in dealing with salinity stress is to "clam up", isolating their tissues from the environment. Though effective in the short term, prolonged isolation can have detrimental physiological consequences, particularly for females brooding embryos in a mantle cavity that is isolated from the external environment. In the Quempillén estuary, the Chilean oyster, Ostrea chilensis, spent nearly one-third of its brooding season at salinities low enough to cause female isolation. When females thus isolated themselves, the dissolved oxygen in their mantle cavity fluid dropped to hypoxic levels within 10 min. In females that were brooding embryos, this depletion of oxygen was not uniform: oxygen was depleted more quickly in the palp region (where embryos accumulate) than in the inhalant region. Additionally, oxygen was reduced even more quickly in the palp region when females were brooding late-stage embryos, which consumed oxygen significantly more quickly than embryos in earlier developmental stages. Finally, O. chilensis used anaerobic metabolism to cope with the hypoxia induced by isolation, as lactate accumulated in the tissues of both females (brooding > non-brooding) and embryos (late stage > early stage). Our findings demonstrate the trade-off between an adaptive avoidance behavior (clamming up) and the potentially detrimental consequences brought on by such a behavior (hypoxia). Cycling of embryos throughout the mantle cavity by deliberate female pumping keeps them from accumulating in the area between the palps, forestalling the creation of hypoxic conditions there. In addition, the capacity for anaerobic metabolism by both females and their embryos should help them tolerate the low oxygen levels that do eventually arise when the pallial cavity is isolated from the surrounding environment during long periods of reduced ambient salinity.