Coenobita Rugosus Research Articles

Seasonal variation in the land hermit crab, Coenobita rugosus H. Milne Edwards, 1837 (Decapoda, Anomura) and a preliminary study of land hermit crabs as a bioindicator of turtle nesting at Penghu County Wang-An Island green turtle nesting refuge, Taiwan

Abstract The present study surveyed the land hermit crab species, body size, shell type, and shell suitability, at five sites over four seasons in the Penghu County Wang-an Island Green Turtle Nesting Refuge. Coenobita rugosus H. Milne Edwards, 1837 was the only species recorded and found at three sites (Tiantai Hill, Xi’an Reservoir, and Tudigong). The density index of hermit crabs exhibited notable variations across the sites, showing no significant variation with the changing seasons. The bush habitat displayed a higher density index compared to the sand habitat. Additionally, the sites differed significantly in terms of the shell type and body size of the hermit crabs. Further, no significant relationship was established between abundance and weather factors or between crab abundance and sea turtle nesting. This investigation is a preliminary examination of the possibility of land hermit crabs being used as bioindicators for breeding of sea turtles.

Polarization vision in terrestrial hermit crabs.

Polarization vision is used by a wide range of animals for navigating, orienting, and detecting objects or areas of interest. Shallow marine and semi-terrestrial crustaceans are particularly well known for their abilities to detect predator-like or conspecific-like objects based on their polarization properties. On land, some terrestrial invertebrates use polarization vision for detecting suitable habitats, oviposition sites or conspecifics, but examples of threat detection in the polarization domain are less well known. To test whether this also applies to crustaceans that have evolved to occupy terrestrial habitats, we determined the sensitivity of two species of land and one species of marine hermit crab to predator-like visual stimuli varying in the degree of polarization. All three species showed an ability to detect these cues based on polarization contrasts alone. One terrestrial species, Coenobita rugosus, showed an increased sensitivity to objects with a higher degree of polarization than the background. This is the inverse of most animals studied to date, suggesting that the ecological drivers for polarization vision may be different in the terrestrial environment.

Taxonomic status and phylogenetic relationship of Anomura (Crustacea: Decapoda) based on mitochondrial sequences and gene order rearrangements

The complete mitochondrial genome (mitogenome) contributed crucial information, which could improve the better understanding of molecular phylogenetic analysis, evolution, and gene rearrangements. However, only a few Coenobitidae and Albuneidae mitogenomes have been described, and controversies about the Anomuran phylogeny remain. Here, we determined three Coenobitidae species (Coenobita perlatus, Coenobita rugosus, Coenobita clypeatus) and one Albuneidae species (Emerita talpoida). As the representative species of genera or families, and even the first species of the genus, supplementing these four species is essential to study the phylogenetic relationships within Anomura. Most of the four novel mitogenomes have the typical 37 genes, except for E. talpoida, which lacked trnF. Through combining these new data with 31 Anomuran mitogenomes from Genbank, different gene rearrangement patterns and internal phylogenetic relationships of Anomura were investigated. In our phylogeny analysis, all the Anomuran species were clustered into one group, and the polyphyly of Paguroidea was well supported. Moreover, various peculiar mitochondrial gene orders (MGOs) were summarized among Anomura and preliminary determined their rearrangement mechanisms through CREx. Twenty different MGOs were suggested in our research, and we were focused seven MGO patterns (Pattern A-J) in the process of discussing the gene rearrangement mechanism. Currently making full use of the large taxon sampling, our results provide valuable information on the evolutionary status of Anomuran species and are available for systematic rearrangement and phylogenetic analyses.

Shell resource partitioning as a mechanism of coexistence in two co-occurring terrestrial hermit crab species

BackgroundCoexistence is enabled by ecological differentiation of the co-occurring species. One possible mechanism thereby is resource partitioning, where each species utilizes a distinct subset of the most limited resource. This resource partitioning is difficult to investigate using empirical research in nature, as only few species are primarily limited by solely one resource, rather than a combination of multiple factors. One exception are the shell-dwelling hermit crabs, which are known to be limited under natural conditions and in suitable habitats primarily by the availability of gastropod shells. In the present study, we used two co-occurring terrestrial hermit crab species, Coenobita rugosus and C. perlatus, to investigate how resource partitioning is realized in nature and whether it could be a driver of coexistence.ResultsField sampling of eleven separated hermit crab populations showed that the two co-occurring hermit crab species inhabit the same beach habitat but utilize a distinct subset of the shell resource. Preference experiments and principal component analysis of the shell morphometric data thereby revealed that the observed utilization patterns arise out of different intrinsic preferences towards two distinct shell shapes. While C. rugosus displayed a preference towards a short and globose shell morphology, C. perlatus showed preferences towards an elongated shell morphology with narrow aperture.ConclusionThe two terrestrial hermit crab species occur in the same habitat but have evolved different preferences towards distinct subsets of the limiting shell resource. Resource partitioning might therefore be the main driver of their ecological differentiation, which ultimately allowed these co-occurring species to coexist in their environment. As the preferred shell morphology of C. rugosus maximizes reproductive output at the expense of protection, while the preferred shell morphology of C. perlatus maximizes protection against predation at the expense of reproductive output, shell resource partitioning might reflect different strategies to respond to the same set of selective pressures occurring in beach habitats. This work offers empirical support for the competitive exclusion principle-hypothesis and demonstrates that hermit crabs are an ideal model organism to investigate resource partitioning in natural populations.

Annual and lunar breeding rhythmicity in females of the terrestrial hermit crab Coenobita rugosus H. Milne Edwards, 1837 (Decapoda: Anomura: Coenobitidae) in Okinawa, Japan

Abstract We investigated the reproductive characteristics of a terrestrial hermit crab (Coenobita rugosus H. Milne Edwards, 1837) inhabiting Okinawa-Jima. Monthly changes in the gonadosomatic index and ovarian histology revealed that this species had fully developed oocytes from May to October. Data showing that females with (ovigerous females) and without (non-ovigerous females) embryos on their pleopods appeared simultaneously on the shore from June to September 2014 demonstrates that this species undergoes larval release several times for four months. When non-ovigerous females were reared under a combination of two day lengths and two temperatures, oocyte development was induced in the high-temperature group, suggesting that temperature is a proximate factor in ovarian development. The weekly collection of females during the spawning season revealed that the proportion of fully developed oocytes in an ovary increased toward the time of the new moon and decreased around the first-quarter moon, suggesting that this species is a lunar-spawner with a new-moon preference. Some females releasing larvae around the new or full moon were recaptured at approximately 30 d intervals, suggesting at least two major groups with lunar-synchronized larval release. The ovarian development and larval release of C. rugosus are likely entrained to external factors, which change periodically in their habitats.

Improving the survival rate of land hermit crabs ( Coenobita rugosus) through artificial habitat design

Coenobita rugosus is one of the most popular hermit crab species traded as pets due to its attractive colors, easy maintenance, and especially by its uniqueness in using others’ shells as their home. The aims of this experiment were to improve survival rate of hermit crabs by providing their ecological needs in artificial habitat and to observe their behaviour in the designed artificial habitat. The experiment was conducted by designing five types of terrarium i.e. empty terrarium as negative control (TC); terrarium with separating system (T1); terrarium with separating system, hiding, and climbing space (T2); terrarium with separating system and burrowing space (T3); and complete terrarium with separating system and all spaces mentioned (T4). The best survival and growth rate were shown by T4 followed by T2 terrarium. T2 and T4 survival rate were significantly greater (p < 0.05) from the three other treatments. Despite the low survival rate, terrariums with separating systems could maintain the survival rate of land hermit crab in the first 6 weeks of the experiment compared to control terrariums. This proved that a separating system holds an important role in keeping terrarium’s cleanliness thus decrease the mortality.

What are the sympatric mechanisms for three species of terrestrial hermit crab (Coenobita rugosus, C. brevimanus, and C. cavipes) in coastal forests?

Terrestrial hermit crabs play a significant role in coastal ecology. For example, as seed dispersers and debris scavengers in coastal forests, they accelerate the decomposition of organic substances. In the Indo-Pacific Ocean, Coenobita rugosus, C. brevimanus, and C. cavipes are the three most common species of terrestrial hermit crab. Because the mechanisms that contribute to the sympatry of these three species of crab have not been identified, this study investigated the three most likely explanations: niche differences, competition, and predation. The results showed that the three species displayed niche differences in terms of seasonal activity, habitat, utilization of shells, and food preference, suggesting that competition for resources is avoided. The habitat of terrestrial hermit crabs in Taiwan is closely associated with that of humans. Our study helps improve our understanding of the ecology of terrestrial hermit crabs and their conservation.

Recruitment to adult habitats in terrestrial hermit crabs on the coast of Ishigakijima Island, Ryukyu Archipelago, Japan

AbstractTerrestrial hermit crabs in the family Coenobitidae (genera Coenobita and Birgus) are distributed in tropical and subtropical regions. They occupy various habitats ranging from shore to inland forests, and the two shore‐dwelling species, Coenobita rugosus and C. violascens, possess different distributional characteristics on Ishigakijima Island, Ryukyu Archipelago, Japan. Coenobita rugosus is distributed throughout the coast of the island and is abundant in beach areas, whereas C. violascens has mainly been found in river mouth areas. However, very little is known about the habitats used by the early life stages of coenobitid crabs because identifying the species of recently landed early juveniles is difficult. We tested whether the species compositions of early juveniles of coenobitids differed between beach and river mouth sites on Ishigakijima Island. We collected and identified the early stage coenobitids using PCR–RFLP techniques. A total of 576 early juveniles of five Coenobita species were collected, of which 0.7% were C. brevimanus, 7.3% were C. cavipes, 0.2% were C. purpureus, 70.1% were C. rugosus, and 21.7% were C. violascens. The early juveniles of Birgus latro were not found. The early juveniles of C. rugosus occurred at both beach and river mouth sites, and they were abundant at beach sites. The early juveniles of C. violascens were only found at river mouth sites. These findings indicate that C. rugosus and C. violascens complete their life cycles on land near the localities where they land. The early juveniles of the inland‐dwelling species, C. cavipes, were also mainly collected from river mouth sites, which suggested that juveniles of C. cavipes selected landing sites near river mouth areas and then migrated into the inland forests, passing through riverside areas. Our results highlighted the importance of river mouth areas for recruitment to adult habitats by some coenobitid species.

Mechanisms causing size differences of the land hermit crab Coenobita rugosus among eco-islands in Southern Taiwan.

Numerous environmental factors can influence body size. Comparing populations in different ecological contexts is one potential approach to elucidating the most critical of such factors. In the current study, we found that the body size of the land hermit crab Coenobita rugosus was significantly larger on Dongsha Island in the South China Sea than on other eco-islands around Southern Taiwan. We hypothesized that this could be due to differences in (1) shell resources, (2) parasite impact, (3) competition, (4) predation, and (5) food. We found no supporting evidence for the first three hypotheses; the shells used by the hermit crabs on Dongsha were in poorer condition than were those used elsewhere, extremely few individuals in the region had ectoparasites, and the density of hermit crabs varied considerably among localities within each island. However, significantly higher percentages of C. rugosus reached age 3 years on Dongsha than at Siziwan bay in Taiwan. Two growth rate indices inferred from size structures suggested faster growth on Dongsha than at Siziwan. The condition index (i.e., the body mass/shield length ratio of C. rugosus) was also greater on Dongsha than at Siziwan. Therefore, Dongsha hermit crabs seem to have superior diet and growth performance. Seagrass debris accumulation at the shore of Dongsha was considerable, whereas none was observed at Siziwan or on the other islands, where dicot leaves were the dominant food item for the vegetarian hermit crabs. We then experimentally evaluated the possible role of seagrass as food for C. rugosus. The crabs on Dongsha preferred seagrass to dicot leaves, and their growth increment was faster when they fed on seagrass than when they fed on dicot leaves; no such differences were found in the Siziwan hermit crabs. The aforementioned results are compatible with the food hypothesis explaining the size differences among the islands. The predator hypothesis could explain the greater life span but not the other findings. Populations of C. rugosus on islands with seagrass debris piles probably contribute more to the gene pool of the species because higher proportions of these populations could achieve high fecundity. The fate of these terrestrial hermit crabs may rely on the health of underwater seagrass ecosystems that are under threat from global change.

Phylogeny and phylogeography of the land hermit crab Coenobita purpureus (Decapoda: Anomura: Coenobitidae) in the Northwestern Pacific Region

AbstractHermit crabs of the family Coenobitidae comprise the land hermit crabs, with 16 Coenobita species, and the coconut crab Birgus latro. They are terrestrial but spend their marine life as planktonic larvae. Some coenobitid crabs are widely distributed in the Indo‐West Pacific region, and some species occupy narrower ranges. To improve our knowledge of coenobitid crab speciation and geographical distribution patterns, we examined the phylogenetic relationship between Coenobita purpureus, which has a narrow distribution in the Northwestern Pacific, and its more widely distributed coenobitids including B. latro, Coenobita brevimanus, Coenobita cavipes, Coenobita perlatus, Coenobita rugosus and Coenobita violascens based on the mtDNA cytochrome c oxidase subunit I (COI) and 16S rDNA genes. We also assessed the phylogeography of Co. purpureus populations based on the COI gene. Our phylogenetic and phylogeographical analyses revealed that (i) Co. purpureus clustered with Co. rugosus and (ii) the Co. purpureus population in the Ogasawara Islands was genetically distinct from those in other regions, which showed genetic panmixia. It has been hypothesized that Co. purpureus evolved in the isolated landmasses of the Ryukyu region during the Pliocene, and that its population expanded and colonized the Ogasawara Islands in the late Pleistocene. Further phylogeographical studies on Coenobita species with relatively narrow distributions coupled with characterization of their phylogenetic relationships with widely distributed congeners will advance our knowledge of the speciation and geographical distribution history of coenobitid crabs.

A tale of two antennules: the performance of crab odour-capture organs in air and water.

Odour capture is an important part of olfaction, where dissolved chemical cues (odours) are brought into contact with chemosensory structures. Antennule flicking by marine crabs is an example of discrete odour capture (sniffing) where an array of chemosensory hairs is waved through the water to create a flow-no flow pattern based on a narrow range of speeds, diameters of and spacings between hairs. Changing the speed of movement and spacing of hairs at this scale to manipulate flow represents a complicated fluid dynamics problem. In this study, we use numerical simulation of the advection and diffusion of a chemical gradient to reveal how morphological differences of the hair arrays affect odour capture. Specifically, we simulate odour capture by a marine crab (Callinectes sapidus) and a terrestrial crab (Coenobita rugosus) in both air and water to compare performance. We find that the antennule morphologies of each species are adaptions to capturing odours in their native habitats. Sniffing is an important part of odour capture for marine crabs in water where the diffusivity of odorant molecules is low and flow through the array is necessary. On the other hand, flow within the hair array diminishes odour-capture performance in air where diffusivities are high. This study highlights some of the adaptations necessary to transition from water to air.

Do terrestrial hermit crabs sniff? Air flow and odorant capture by flicking antennules.

Capture of odorant molecules by olfactory organs from the surrounding fluid is the first step of smelling. Sniffing intermittently moves fluid across sensory surfaces, increasing delivery rates of molecules to chemosensory receptors and providing discrete odour samples. Aquatic malacostracan crustaceans sniff by flicking olfactory antennules bearing arrays of chemosensory hairs (aesthetascs), capturing water in the arrays during downstroke and holding the sample during return stroke. Terrestrial malacostracans also flick antennules, but how their flicking affects odour capture from air is not understood. The terrestrial hermit crab, Coenobita rugosus, uses antennules bearing shingle-shaped aesthetascs to capture odours. We used particle image velocimetry to measure fine-scale fluid flow relative to a dynamically scaled physical model of a flicking antennule, and computational simulations to calculate diffusion to aesthetascs by odorant molecules carried in that flow. Air does not flow into the aesthetasc array during flick downstrokes or recovery strokes. Odorants are captured from air flowing around the outside of the array during flick downstrokes, when aesthetascs face upstream and molecule capture rates are 21% higher than for stationary antennules. Bursts of flicking followed by pauses deliver discrete odour samples to olfactory sensors, causing intermittency in odour capture by a different mechanism than aquatic crustaceans use.

A glycosyl hydrolase family 16 gene is responsible for the endogenous production of β-1,3-glucanases within decapod crustaceans

To identify the gene responsible for the production of a β-1,3-glucanase (laminarinase) within crustacea, a glycosyl hydrolase family 16 (GHF16) gene was sequenced from the midgut glands of the gecarcinid land crab, Gecarcoidea natalis and the freshwater crayfish, Cherax destructor. An open reading frame of 1098bp for G. natalis and 1095bp for C. destructor was sequenced from cDNA. For G. natalis and C. destructor respectively, this encoded putative proteins of 365 and 364 amino acids with molecular masses of 41.4 and 41.5kDa. mRNA for an identical GHF16 protein was also expressed in the haemolymph of C. destructor. These putative proteins contained binding and catalytic domains that are characteristic of a β-1,3-glucanase from glycosyl hydrolase family 16. The amino acid sequences of two short 8–9 amino acid residue peptides from a previously purified β-1,3-glucanase from G. natalis matched exactly that of the putative protein sequence. This plus the molecular masses of the putative proteins matching that of the purified proteins strongly suggests that the sequences obtained encode for a catalytically active β-1,3-glucanase. A glycosyl hydrolase family 16 cDNA was also partially sequenced from the midgut glands of other amphibious (Mictyris platycheles and Paragrapsus laevis) and terrestrial decapod species (Coenobita rugosus, Coenobita perlatus, Coenobita brevimanus and Birgus latro) to confirm that the gene is widely expressed within this group. There are three possible hypothesised functions and thus evolutionary routes for the β-1,3-glucanase: 1) a digestive enzyme which hydrolyses β-1,3-glucans, 2) an enzyme which cleaves β-1,3-glycosidic bonds within cell walls to release cell contents or 3) an immune protein which can hydrolyse the cell walls of potentially pathogenic micro-organisms.

Scaling of olfactory antennae of the terrestrial hermit crabs Coenobita rugosus and Coenobita perlatus during ontogeny.

Although many lineages of terrestrial crustaceans have poor olfactory capabilities, crabs in the family Coenobitidae, including the terrestrial hermit crabs in the genus Coenobita, are able to locate food and water using olfactory antennae (antennules) to capture odors from the surrounding air. Terrestrial hermit crabs begin their lives as small marine larvae and must find a suitable place to undergo metamorphosis into a juvenile form, which initiates their transition to land. Juveniles increase in size by more than an order of magnitude to reach adult size. Since odor capture is a process heavily dependent on the size and speed of the antennules and physical properties of the fluid, both the transition from water to air and the large increase in size during ontogeny could impact odor capture. In this study, we examine two species of terrestrial hermit crabs, Coenobita perlatus H. Milne-Edwards and Coenobita rugosus H. Milne-Edwards, to determine how the antennule morphometrics and kinematics of flicking change in comparison to body size during ontogeny, and how this scaling relationship could impact odor capture by using a simple model of mass transport in flow. Many features of the antennules, including the chemosensory sensilla, scaled allometrically with carapace width and increased slower than expected by isometry, resulting in relatively larger antennules on juvenile animals. Flicking speed scaled as expected with isometry. Our mass-transport model showed that allometric scaling of antennule morphometrics and kinematics leads to thinner boundary layers of attached fluid around the antennule during flicking and higher odorant capture rates as compared to antennules which scaled isometrically. There were no significant differences in morphometric or kinematic measurements between the two species.

Isolation increases time to emerge from shells in two Coenobita hermit crab species

Isolation increases activity in individuals of some species, whereas it decreases activity in individuals of other species. Furthermore, as the length of isolation increases, behavioral changes can become more pronounced. Here, we examined effects of short-term isolation on latencies for individuals to emerge from their shells in two species of terrestrial hermit crab (Coenobita rugosus and Coenobita brevimanus) over a 4-week period. Within each species, crabs were housed individually (isolate context) or socially in groups of four (social context). In each week, crabs were exposed to an emergence test, which measured the latency to emerge from the shell following a potentially threatening stimulus. We obtained a significant context × week interaction. Although socially housed crabs showed no change in emergence over the 4 weeks of the study, isolate crabs took longer to emerge from their shells as the period of isolation increased. Our study corroborates findings in other invertebrate species, as well as in vertebrate species, that isolation generally decreases activity levels and may increase fear levels, in individuals.

Hermit crabs perceive the extent of their virtual bodies

A flexible body image is required by animals if they are to adapt to body changes and move effectively within a structurally complex environment. Here, we show that terrestrial hermit crabs, Coenobita rugosus, which frequently change shells, can modify walking behaviour, dependent on the shape of the shell. Hermit crabs walked along a corridor that had alternating left and right corners; if it was narrow at the corner, crabs rotated their bodies to avoid the wall, indicating an awareness of environmental obstacles. This rotation increased when a plastic plate was attached to the shell. We suggest that the shell, when extended by the plate, becomes assimilated to the hermit crab's own body. While there are cases of a tool being assimilated with the body, our result is the first example of the habitat where an animal lives and/or carries being part of a virtual body.

Can land crabs be used as a rapid ecosystem evaluation tool? A test using distribution and abundance of several genera from the Seychelles

Habitat destruction and the introduction of exotic species are the primary causes of biodiversity loss in tropical island ecosystems. Conservation efforts on oceanic islands are often biased towards charismatic vertebrate faunas, neglecting invertebrate assemblages. We sampled the land crab community on five islands in the central Seychelles; in the intertidal zone, in the supralittoral zone and in nine different inland habitat types to explore the impacts of exotic vegetation and environmental variables on land crab abundance and community composition, and investigate whether land crabs can be used as a tool for the rapid assessment of habitat quality on tropical oceanic islands. We found that species richness and the abundance of the dominant ghost crab Ocypode cordimana was higher in native habitat types than habitats dominated by exotic vegetation. Available ground substrate suitable for burrowing may be a limiting factor for O. cordimana in exotic habitat types. Coenobita rugosus, the dominant crab in the supralittoral zone is largely absent where there is no supralittoral vegetation. These results suggest that land crabs could be reliable indicators of habitat quality on oceanic islands. The abundance of land crabs could be used in the rapid assessment of ecosystem perturbation and identification of sites requiring restoration or management.

Observation of intersexuality in land hermit crabs (Anomura: Coenobitidae)

We report here the first observation of intersexuality in a land hermit crab (Coenobitidae). Three species of land hermit crabs from Okinawa were investigated. In the population of the terrestrial hermit crab Coenobita rugosus two per cent of males were found to be intersex. Besides the true male gonopores on the coxae of the 5th pair of pereopods, they had additional openings on the coxae of the 3rd pair of pereopods. Both specimens examined had a normally developed male reproductive system. In the populations of two other species studied, C. brevimanus and C. purpureus, however, no intersex individuals have been found. Examples of intersexuality in decapod crustaceans are discussed and putative explanations of this phenomenon in land hermit crabs are proposed.

Resource availability: Ancient homes for hard-up hermit crabs.

Mollusc shells are a vital but sometimes scarce resource for hermit crabs, protecting them from mechanical damage and desiccation, but they require continual replacement as the crab grows. I have discovered that Coenobita rugosus, a large, tropical, semi-terrestrial hermit crab, will resort to using fossil shells when no other suitable casing is available. These unlikely mobile homes fall out of coastal limestone as it is eroded by the sea in southwestern Madagascar, placing the occupants alongside Homo sapiens as resourceful exploiters of prehistoric animal remains.

Reproductive Biology of Three Land Hermit Crabs (Decapoda: Anomura: Coenobitidae) in Okinawa, Japan

Reproductive ecological research on three land hermit crabs, Coenobita rugosus, C. purpureus, and C. cavipes, was conducted in the southern part of Okinawa-jima island in 1985, 1986, 1987, and for a short period in 1999. Size (carapace length) of the smallest ovigerous female was 3.93 mm for C. rugosus, 3.83 mm for C. purpureus, and 9.49 mm for C. cavipes. Breeding season is late May to November for C. rugosus, late May to mid-September for C. purpureus, and mid-May to late August for C. cavipes. Some females of all three species probably produced at least two broods during the breeding season. The smallest males in which spermatophores were present in dissected vas deferens were 4.24 mm for C. rugosus and 4.94 mm for C. purpureus. Coenobita cavipes females produced more, smaller eggs in comparison with C. purpureus. My observations suggest that coenobitid crabs living in areas with a low supply of shells or with poor shells reproduce at smaller sizes, as is the case in marine hermit crabs. Time of onset of larval release by C. rugosus, with its protracted breeding season, varied according to the seasonal shift in time of sunset. The period during which females of C. rugosus released larvae was about 2 hr in spring tides but was much longer (3 to 5 hr) during neap tides. Larger females of C. purpureus occupied shells derived from the land snail Achatina fulica; smaller ones used shells from the marine snail Lunella granulata. Use of mutually exclusive larval release sites by the larger and smaller females of C. purpureus remained unchanged over 13 yr, from 1986 to 1999. This behavioral difference may be related to the differences in their habitats (i.e., inland versus shore) and to the route traveled by the larger crabs in reaching the sea from inland sites.