Thermal threshold and interspecific competition help explain intertidal hermit crab assemblages.

Habitat heterogeneity promotes species diversity by providing a variety of abiotic and biotic conditions, whose impact on performance varies among species. Then, mobile species would be expected to move to areas whose conditions maximize their fitness. However, biotic pressures such as interspecific competition can push subordinate species into suboptimal areas, impeding this matching. The tropical hermit crab Clibanarius albidigitus occupies mostly upper intertidal sites where they can experience extreme environmental conditions. Meanwhile, its stronger agonistic competitor, Calcinus californiensis, mostly inhabits more moderate conditions at the mid intertidal. We estimated the avoidance threshold of the two hermit crab species to increasing water temperatures to help explain their intertidal distribution. We also compared the avoidance threshold of Cli. albidigitus to rising temperatures when presented alone and in the presence of chemical cues of its competitor to assess potential competitive niche exclusion. The avoidance threshold was measured in experimental tanks with a ramp that led from the water to an air-exposed platform; the threshold was defined as the temperature at which individuals emerged and remained air-exposed. Clibanarius albidigitus emerged at a higher temperature than its competitor, showing a higher thermal tolerance and potentially explaining its distribution in the upper intertidal. In the presence of Cal. californiensis, Cli. albidigitus emerged at lower temperature than when alone, likely as a strategy to reduce competition with stronger agonistic competitors, even at the cost of coping with harsh conditions. Our results support the hypothesis that competitive habitat exclusion contributes to explaining hermit crab assemblages.

Metabolic and behavioural effects of hermit crab shell removal techniques: Is heating less invasive than cracking?

Hermit crabs (Paguroidea; Latreille 1802) offer great opportunities to study animal behaviour and physiology. However, the animals' size and sex cannot be determined when they are inside their shell; information crucial to many experimental designs. Here, we tested the effects of the two most common procedures used to make crabs leave their shells: heating the shell apex and cracking the shell with a bench press. We compared the effects of each of the two procedures on the metabolic rate, hiding time, and duration of the recovery time relative to unmanipulated hermit crabs. The hermit crabs forced to abandon their shell through heating increased their respiratory rate shortly after the manipulation (1 h) and recovered their metabolic rate in less than 24 h, as occurs in individuals suddenly exposed to high temperatures in the upper-intertidal zone. Hermit crabs removed from their shells via cracking spent more time hiding in their new shells; this effect was evident immediately after the manipulation and lasted more than 24 h, similar to responses exhibited after a life-threatening predator attack. Both methods are expected to be stressful, harmful, or fear-inducing; however, the temperature required to force the crabs to abandon the shell is below the critical thermal maxima of most inhabitants of tropical tide pools. The wide thermal windows of intertidal crustaceans and the shorter duration of consequences of shell heating compared to cracking suggest heating to be a less harmful procedure for removing tropical hermit crabs from their shells.

The energetic costs of living in the surf and impacts on zonation of shells occupied by hermit crabs.

Crashing waves create a hydrodynamic gradient in which the most challenging effects occur at the wave breaking zone and decrease towards the upper protected tide pools. Hydrodynamic forces depend on the shape of the submerged body; streamlined shapes decrease drag forces compared with bluff or globose bodies. Unlike other animals, hermit crabs can choose their shell shape to cope with the effects of water flow. Hermit crabs occupy larger and heavier shells (conical shape) in wave-exposed sites than those used in protected areas (globose shape). First, we investigated whether a behavioral choice could explain the shells used in sites with different wave action. Then, we experimentally tested whether the shells most frequently used in sites with different wave action reduce the energetic cost of coping with water flow. Metabolic rate was measured using a respirometric system fitted with propellers in opposite walls to generate bidirectional water flow. The choice of shell size when a large array of sizes are available was consistent with the shell size used in different intertidal sites; hermit crabs chose heavier conical shells in water flow conditions than in still water, and the use of heavy conical shells reduced the energetic cost of coping with water motion. In contrast to conical shells, small globose shells imposed lower energy costs of withstanding water flow than large globose shells. The size and type of shells used in different zones of the rocky shore were consistent with an adaptive response to reduce the energetic costs of withstanding wave action.