Like night and day: Reversals of thermal gradients across ghost crab burrows and their implications for thermal ecology

Abstract Ghost crabs, Ocypode cordimanus , inhabit relatively hostile environments subject to thermal fluctuations, including both diurnal and seasonal cycles. For many ectotherms, including ghost crabs, a major challenge is to remain cool during hot daytime temperatures. This can be achieved by adopting a fossorial lifestyle, taking advantage of thermal refuge afforded by burrows of sufficient depth. Another consideration, often overlooked, is the potential advantage associated with ready access to a thermal energy source (a “charging station”) when surface temperatures are cooler. Being able to rapidly elevate body temperature during cool periods would enhance the crab's ability to maintain rate processes and carry out essential activities. We have measured ghost crab burrow temperature profiles at two times of the day with contrasting sun exposure (06:00 and 14:00), demonstrating how effective burrow depth (up to a maximum of 40 cm) provides thermal regulation below the surface of the sand (e.g., at dawn (06:00) and early afternoon (14:00) at a depth of 5 cm, temperatures (±SD) of 16.32 ± 0.96 °C and 25.04 ± 1.47 °C were recorded, respectively. Corresponding temperatures at a depth of 30 cm were 19.17 ± 0.59 °C and 19.78 ± 1.60 °C, respectively). This demonstrates that while temperature conditions at the surface vary dramatically from night to day, ghost crab burrows can maintain relatively constant temperatures at the burrow base throughout the diurnal cycle, at least during winter. As a consequence, the burrow heat signatures undergo a corresponding thermal gradient reversal between night and day, as revealed by infra-red photography. Complementing these field observations, we also determined heating and cooling times/constants for O. cordimanus in the laboratory (τ = 17.54 and 16.59 JK -1 , respectively), and analysed chemical composition of their carapace (external (with β Chitin evident) and internal (predominance of α Chitin)), which is the primary thermal interface with the environment. We find that ghost crabs both gain and lose heat relatively rapidly, which likely affects the range and duration of surface activities under different thermal conditions, and renders the thermal characteristics of their burrows vital for their persistence on beaches. Finally, we speculate that the distinctly contrasting thermal signatures of ghost crab burrows in comparison to the surrounding sand could in principle be used by crabs as spatial markers for navigation and to identify holes on return from nightly excursions, being identified either by direct thermal sensing or odours rising from the burrow base as a consequence of the thermal flux.