For many decades it has been accepted that marine turtle hatchlings from the same nest generally emerge from the sand together. However, for loggerhead turtles (Caretta caretta) nesting on the Greek Island of Kefalonia, a more asynchronous pattern of emergence has been documented. By placing temperature loggers at the top and bottom of nests laid on Kefalonia during 1998, we examined whether this asynchronous emergence was related to the thermal conditions within nests. Pronounced thermal variation existed not only between, but also within, individual nests. These within-nest temperature differences were related to the patterns of hatchling emergence, with hatchlings from nests displaying large thermal ranges emerging over a longer time-scale than those characterised by more uniform temperatures. In many egg-laying animals, parental care of the offspring may continue while the eggs are incubating and also after they have hatched. Consequently, the importance of the nest site for determining incubation conditions may be reduced since the parents themselves may alter the local environment. By contrast, in marine turtles, parental care ceases once the eggs have been laid and the nest site covered. The positioning of the nest site, in both space and time, may therefore have profound effects for marine turtles by affecting, for example, the survival of the eggs and hatchlings as well as their sex (Janzen and Paukstis 1991). During incubation, sea turtle embryos grow from a few cells at oviposition to a self-sufficient organism at hatching some 50-80 days later (Ackerman 1997). After hatching, the young turtles dig up through the sand and emerge typically en masse at the surface 1-7 nights later, with a number of stragglers following over the next few nights (Christens 1990). This contrasts with the frequently observed pattern of hatching asynchrony in birds. It has been suggested that the cause of mass emergence in turtles is that eggs within a clutch are fertilised within a short period of time and then, when thermal conditions within the nest are uniform, develop at very similar rates and hence hatch and emerge together (Porter 1972). As a corollary of this idea, it would be predicted that when there are pronounced within-nest thermal gradients, development rates of siblings will be different and hence asynchronous hatching and emergence might occur. While it may be energetically beneficial for hatchlings to emerge in a group (Carr and Hirth 1961), if the extent of hatching asynchrony is marked then there may be severe costs for individuals if they wait for all their siblings to hatch before attempting to dig out of the sand (Hays and Speakman 1992). Under such conditions, the protracted emergence of small groups of hatchlings over several nights may be favoured. Examination of the literature suggests that emergence asynchrony may be more widespread than generally considered. For example, Witherington et al. (1990) described loggerhead turtle hatchlings (Caretta caretta) emerging over 4 days in Florida; for green turtles (Chelonia mydas), Hendrickson (1958) documented that nests in Malaysia and Sarawak produced hatchlings for up to 8 days; whilst Diamond (1976) found that hawksbill (Eretmochelys imbricata) nests on Cousin Island, Seychelles, were active for up to 4 days. Similarly, on the Greek Island of Kefalonia, we have shown that emergence from individual loggerhead turtle nests may occur on up to 11 nights (Hays and Speakman 1992). It is logical to suppose that asynchronous emergence relates to thermal gradients within nests, since the incubation duration of sea turtle eggs is related to temperature, with eggs hatching quicker when the temperature is higher. Here we test this hypothesis by measuring thermal variations within loggerhead turtle nests and comparing these variations to the patterns of hatchling emergence.
Climate change is a clear and present threat to species survival. For species with temperature-dependent sex determination, including all sea turtles, it has been hypothesised that climate change may drive the creation of sex-ratio biases leading to population extinctions1. Through a global analysis across multiple species, we present the first direct empirical evidence for a demographic consequence of male scarcity in sea turtle populations, with a lower incidence of multiple paternity being found in populations with more extreme female-biased hatchling sex-ratio skews. For green turtles, when the female bias in hatchling sex ratio was >90%, the incidence of multiple paternity was low compared to other nesting sites, being 24.5% in the eastern Mediterranean (Cyprus), 36.4% on Redang Island (Malaysia) and 15.4% on the southern Great Barrier Reef (Heron Island, Australia) compared to higher values (range 61.1-91.7%) at other sites globally. These results suggest that a low incidence of multiple paternity may serve as a harbinger of future problems with egg fertility if males become even scarcer. Assessments of the incidence of multiple paternity at sites where adult males are expected to become scarce, such as Raine Island on the northern Great Barrier Reef in Australia, may help to identify when a lack of males raises the threat of local extinctions. In such cases, intervention to increase the production of male hatchlings may be needed.