Coenobita Research Articles

Fine structural organization of the hemiellipsoid body of the land hermit crab, Coenobita clypeatus

Electron microscopical observations of the hemiellipsoid bodies of the land hermit crab Coenobita clypeatus resolve microglomerular synaptic complexes that are comparable to those observed in the calyces of insect mushroom bodies and which characterize olfactory inputs onto intrinsic neurons. In an adult hermit crab, intrinsic neurons and one class of efferent neurons originate from neuronal somata of globuli cells covering the hemiellipsoid bodies. Counts of their nucleoli show that about 120,000 globuli cells supply each hemiellipsoid body in an adult hermit crab. This number is comparable to the number of globuli cells supplying mushroom bodies of certain insects, such as honey bees and cockroaches. Counts of axons in tracts leading from the olfactory lobes to the hemiellipsoid bodies resolve 20,000 afferent axons, however, an order of magnitude greater than known for any insect. These afferent axons provide numerous swollen varicosities, each presynaptic to many small profiles, and thus comparable to the microglomeruli that characterize insect mushroom body calyces. Also, common to mushroom bodies and hemiellipsoid bodies are arrangements of intrinsic neurons, afferent neurons containing dense core vesicles, and systems of serial synaptic complexes that relate to postsynaptic profiles of efferent neurons. Together, the ultrastructural organization of the hemiellipsoid bodies of C. clypeatus supports the proposition that this center may share a common origin with the insect mushroom body despite obvious divergent evolution of overall shape.

Neuronal organization of the hemiellipsoid body of the land hermit crab, Coenobita clypeatus: Correspondence with the mushroom body ground pattern

Malacostracan crustaceans and dicondylic insects possess large second-order olfactory neuropils called, respectively, hemiellipsoid bodies and mushroom bodies. Because these centers look very different in the two groups of arthropods, it has been debated whether these second-order sensory neuropils are homologous or whether they have evolved independently. Here we describe the results of neuroanatomical observations and experiments that resolve the neuronal organization of the hemiellipsoid body in the terrestrial Caribbean hermit crab, Coenobita clypeatus, and compare this organization with the mushroom body of an insect, the cockroach Periplaneta americana. Comparisons of the morphology, ultrastructure, and immunoreactivity of the hemiellipsoid body of C. clypeatus and the mushroom body of the cockroach P. americana reveal in both a layered motif provided by rectilinear arrangements of extrinsic and intrinsic neurons as well as a microglomerular organization. Furthermore, antibodies raised against DC0, the major catalytic subunit of protein kinase A, specifically label both the crustacean hemiellipsoid bodies and insect mushroom bodies. In crustaceans lacking eyestalks, where the entire brain is contained within the head, this antibody selectively labels hemiellipsoid bodies, the superior part of which approximates a mushroom body's calyx in having large numbers of microglomeruli. We propose that these multiple correspondences indicate homology of the crustacean hemiellipsoid body and insect mushroom body and discuss the implications of this with respect to the phylogenetic history of arthropods. We conclude that crustaceans, insects, and other groups of arthropods share an ancestral neuronal ground pattern that is specific to their second-order olfactory centers.

Quantifying personality in the terrestrial hermit crab: Different measures, different inferences

There is much interest in studying animal personalities but considerable debate as to how to define and evaluate them. We assessed the utility of one proposed framework while studying personality in terrestrial hermit crabs (Coenobita clypeatus). We recorded the latency of individuals to emerge from their shells over multiple trials in four unique manipulations. We used the specific testing situations within these manipulations to define two temperament categories (shyness-boldness and exploration-avoidance). Our results identified individual behavioral consistency (i.e., personality) across repeated trials of the same situations, within both categories. Additionally, we found correlations between behaviors across contexts (traits) that suggested that the crabs had behavioral syndromes. While we found some correlations between behaviors that are supposed to measure the same temperament trait, these correlations were not inevitable. Furthermore, a principal component analysis (PCA) of our data revealed new relationships between behaviors and provided the foundation for an alternate interpretation: measured behaviors may be situation-specific, and may not reflect general personality traits at all. These results suggest that more attention must be placed on how we infer personalities from standardized methods, and that we must be careful to not force our data to fit our frameworks.

Transition from sea to land: olfactory function and constraints in the terrestrial hermit crabCoenobita clypeatus

The ability to identify chemical cues in the environment is essential to most animals. Apart from marine larval stages, anomuran land hermit crabs (Coenobita) have evolved different degrees of terrestriality, and thus represent an excellent opportunity to investigate adaptations of the olfactory system needed for a successful transition from aquatic to terrestrial life. Although superb processing capacities of the central olfactory system have been indicated in Coenobita and their olfactory system evidently is functional on land, virtually nothing was known about what type of odourants are detected. Here, we used electroantennogram (EAG) recordings in Coenobita clypeatus and established the olfactory response spectrum. Interestingly, different chemical groups elicited EAG responses of opposite polarity, which also appeared for Coenobita compressus and the closely related marine hermit crab Pagurus bernhardus. Furthermore, in a two-choice bioassay with C. clypeatus, we found that water vapour was critical for natural and synthetic odourants to induce attraction or repulsion. Strikingly, also the physiological response was found much greater at higher humidity in C. clypeatus, whereas no such effect appeared in the terrestrial vinegar fly Drosophila melanogaster. In conclusion, our results reveal that the Coenobita olfactory system is restricted to a limited number of water-soluble odourants, and that high humidity is most critical for its function.

Homes for hermits: temporal, spatial and structural dynamics as transportable homes are incorporated into a population

AbstractUnderstanding the process by which limiting resources are incorporated into populations is a major goal of ecology. While many studies have examined this dynamic process using essential resources like homes, few of these studies have involved homes that can be transported by their occupants. This study introduced over a thousand transportable homes into a population of terrestrial hermit crabs Coenobita compressus, animals that carry their homes with them wherever they travel. These new homes were tracked between years to test key predictions about the temporal dynamics the homes would generate, and the spatial and structural changes the homes would undergo as they were used by the population. When moving into new homes, crabs dropped off their old homes directly at the exchange site, and the number of such traded‐in homes peaked rapidly in time. Traded‐in homes were under half the diameter of new homes, a difference apparently magnified by social formations involving vacancy chains. After crabs moved into new homes, they carried the homes away from the exchange site. The following year, these homes were displaced a distance four orders of magnitude times their diameter, thus penetrating extensively through the population. Between years, crabs also remodeled the internal architecture of the homes, creating homes that were more spacious and less of a burden to carry. These results suggest that transportable homes generate novel ecological dynamics along temporal, spatial and structural dimensions, which are a direct consequence of their transportability.

Egg production and shell relationship of the land hermit crab Coenobita scaevola (Anomura: Coenobitidae) from Wadi El-Gemal, Red Sea, Egypt

The aim of the present study is to characterize the fecundity of the land hermit crab Coenobita scaevola as well as the influence of shell type on fecundity using morphometric relationships. Hermit crabs were collected monthly from January to December 2007 from the protected area of Wadi El-Gemal, at Marsa Alam on the Red Sea, and ovigerous females were selected. Hermit crab wet weight and the gastropod shell weight were recorded. The number of eggs carried by females of several sizes (CL, carapace length), stages of development and egg size were determined. Shells of eight gastropod species were occupied by ovigerous females of C. scaevola. Shells of Nerita undata was the most occupied (65.7%), particularly by individuals falling within the size range 5.0–7.0mm CL. Only 35 berried females were recorded during May, July and September and the mean fecundity was 679.8±140 eggs. Fecundity was found positively correlated with crab size and shell dimensions. The relationship between fecundity and the internal volume of the occupied shell was ranked as the most correlated. The impact of shell utilization on hermit crab fecundity is discussed.

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.

Terrestrial hermit crabs (Anomura: Coenobitidae) as taphonomic agents in circum-tropical coastal sites

Hermit crabs are ever alert for more suitable shells to inhabit, but what this may mean for coastal shell middens has rarely been considered. Here, the impact of the most landward-based of hermit crab families, the tropical Coenobitidae, upon archaeological shell-bearing deposits is assessed using a case study: the Neolithic Ugaga site from Fiji. At Ugaga, hermit crabs were found to have removed the majority of shells from the midden and had deposited their old, worn shells in return. The behavioural ecology of genus Coenobita suggests a mutualistic interaction whereby humans make available shell and food resources to hermit crabs, which in turn provide a site cleaning service by consuming human and domestic waste. Diagnostic indicators of terrestrial hermit crab wear patterns on gastropod shells are outlined and the conditions under which extensive ‘hermitting’ of shell midden deposits may occur are investigated. The ability to recognise hermit crab modification of shells is considered not only important for archaeomalacologists analysing tropical shell deposits, but also for field archaeologists wishing to select suitable shell samples for radiocarbon dating.

Relative Growth of the Land Hermit Crab, Coenobita Clypeatus (Anomura, Coenobitidae) from a Coral Reef Island, Southern Gulf of Mexico

Information on growth in hermit crabs is very scarce, possibly due to the inherent difficulties of extracting them from their shells. In this work we document the relative growth, sexual size dimorphism, and level of chelae asymmetry of the land hermit crab Coenobita clypeatus (Fabricius, 1787) collected on a coral reef island in the National Park Arrecife Alacranes, southern Gulf of Mexico, during December 2009 and September 2010. Specimens examined were anaesthetized in a clove oil-water solution to avoid unnecessary crab mortality. A total of 387 individuals was measured on six external variables: cephalothorax length and width, and both chelae in length and width. Taking the cephalothorax length as the reference dimension, the remaining measurements were fitted to a power function to examine the degree of allometry in growth and make comparisons between sexes. For each sex, an asymmetry index (AI) was calculated to determine the degree of heterochely, and the significance of its value was evaluated by means of null models. Numerically, males dominated the crab population in the largest size classes (>12.6 mm CL) in both seasons. As in most anomurans, C. clypeatus males attained larger body sizes than females, but their relative growth was most similar. For both sexes, the growth of chela width was positively allometric (t-test, p < 0.05). Also, AI values showed a heterocheleous condition, with the left chela significantly higher (p < 0.001) than the right. Regarding our own results and previous reports, we propose that heterochely could be triggered during the glaucothoal and juvenile stages. Numerical dominance of males in the larger size classes is probably due to the segregation of ovigerous females from the main crab group, searching suitable areas for egg deposition, but also to a better condition of males to carry heavier shells than females, which in their turn have to invest a considerable amount of energy in reproduction.

Culture and Maintenance of Selected Invertebrates in the Laboratory and Classroom

Invertebrate species have been used for many years in the laboratory and teaching environment. We discuss some of the most commonly maintained invertebrates--the nematode (Caenorhabditis elegans), the California sea hare (Aplysia californica), the fruit fly (Drosophila melanogaster), terrestrial hermit crabs, the horseshoe crab (Limulus polyphemus), and cephalopods--and briefly describe general techniques for culturing them in captivity. The aim of this article is to give potential users an idea of the materials, methods, and effort required to maintain each type of organism in a laboratory or classroom setting.

Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers

Hermit crabs are critically dependent upon gastropod shells for their survival and reproductive fitness. While anecdotal reports have suggested that hermit crabs may be capable of removing live gastropods from their shells to access the essential shell resource, no systematic experiments have been conducted to investigate this possibility. This paper reports experiments on both marine ( Pagurus bernhardus) and terrestrial ( Coenobita compressus) hermit crabs in which crabs were paired in the laboratory with the gastropods whose shells they inhabit in the field. Pairings included both shelled and naked crabs and spanned the full range of the gastropod life cycle. Neither marine nor terrestrial hermit crabs were successful at removing live gastropods from their shells. Furthermore, only a small fraction of the crabs (5.7%) were capable of accessing shells in which the gastropod had been killed in advance, with its body left intact inside the shell. Finally, although hermit crabs readily entered empty shells positioned on the surface, few crabs (14.3%) were able to access empty shells that were buried just centimeters beneath them. These results suggest that hermit crabs are constrained consumers, with the shells they seek only being accessible during a narrow time window, which begins following natural gastropod death and bodily decomposition and which typically ends when the gastropod's remnant shell has been buried by tidal forces. Further experiments are needed on more species of hermit crabs as well as fine-grained measurements of (i) the mechanical force required to pull a gastropod body from its shell and (ii) the maximum corresponding force that can be generated by different hermit crab species' chelipeds.

Increased amplitude and duration of acoustic stimuli enhance distraction

Extraneous sounds have a variety of effects on animals; they may interfere with communication, cause physical harm, increase wariness, influence settlement decisions, or they may cause distractions in ways that increase vulnerability to predation. We designed a study to investigate the effects of changing both the amplitude and duration of an acoustic stimulus on distraction in a terrestrial hermit crab (Coenobita clypeatus). In experiment 1, we replicated the key findings from a field result: crabs hid more slowly in response to a silent visual stimulus when we simultaneously broadcast a white noise than they did when in a silent condition. In experiment 2, we altered the noise duration and found that a long noise generated greater latencies to hide than a short noise. In experiment 3, we increased the noise amplitude and found that hide latency increased with higher-intensity auditory stimuli. These experiments demonstrate a variety of stimulus factors that influence distraction. Our results suggest that prey animals could be in greater danger from predators when in an environment with auditory distractions.

Brain architecture of the largest living land arthropod, the Giant Robber Crab Birgus latro (Crustacea, Anomura, Coenobitidae): evidence for a prominent central olfactory pathway?

BackgroundSeveral lineages within the Crustacea conquered land independently during evolution, thereby requiring physiological adaptations for a semi-terrestrial or even a fully terrestrial lifestyle. Birgus latro Linnaeus, 1767, the giant robber crab or coconut crab (Anomura, Coenobitidae), is the largest land-living arthropod and inhabits Indo-Pacific islands such as Christmas Island. B. latro has served as a model in numerous studies of physiological aspects related to the conquest of land by crustaceans. From an olfactory point of view, a transition from sea to land means that molecules need to be detected in gas phase instead of in water solution. Previous studies have provided physiological evidence that terrestrial hermit crabs (Coenobitidae) such as B. latro have a sensitive and well differentiated sense of smell. Here we analyze the brain, in particular the olfactory processing areas of B. latro, by morphological analysis followed by 3 D reconstruction and immunocytochemical studies of synaptic proteins and a neuropeptide.ResultsThe primary and secondary olfactory centers dominate the brain of B. latro and together account for ca. 40% of the neuropil volume in its brain. The paired olfactory neuropils are tripartite and composed of more than 1,000 columnar olfactory glomeruli, which are radially arranged around the periphery of the olfactory neuropils. The glomeruli are innervated ca. 90,000 local interneurons and ca. 160,000 projection neurons per side. The secondary olfactory centers, the paired hemiellipsoid neuropils, are targeted by the axons of these olfactory projection neurons. The projection neuron axonal branches make contact to ca. 250.000 interneurons (per side) associated with the hemiellipsoid neuropils. The hemiellipsoid body neuropil is organized into parallel neuropil lamellae, a design that is quite unusual for decapod crustaceans. The architecture of the optic neuropils and areas associated with antenna two suggest that B. latro has visual and mechanosensory skills that are comparable to those of marine Crustacea.ConclusionsIn parallel to previous behavioral findings that B. latro has aerial olfaction, our results indicate that their central olfactory pathway is indeed most prominent. Similar findings from the closely related terrestrial hermit crab Coenobita clypeatus suggest that in Coenobitidae, olfaction is a major sensory modality processed by the brain, and that for these animals, exploring the olfactory landscape is vital for survival in their terrestrial habitat. Future studies on terrestrial members of other crustacean taxa such as Isopoda, Amphipoda, Astacida, and Brachyura will shed light on how frequently the establishment of an aerial sense of olfaction evolved in Crustacea during the transition from sea to land. Amounting to ca. 1,000,000, the numbers of interneurons that analyse the olfactory input in B. latro brains surpasses that in other terrestrial arthropods, as e.g. the honeybee Apis mellifera or the moth Manduca sexta, by two orders of magnitude suggesting that B. latro in fact is a land-living arthropod that has devoted a substantial amount of nervous tissue to the sense of smell.

Social context of shell acquisition in Coenobita clypeatus hermit crabs

Vacancy chains involve unique patterns of resource acquisition behaviors that determine how reusable resources are distributed through animal populations. Shell vacancy chains have been described for several hermit crab species, both terrestrial and marine, but little is known about the ecological and behavioral dynamics of shell choice in social versus solitary contexts. Here, we present a novel conceptual framework that differentiates 2 types of shell vacancy chain in hermit crabs and discuss fundamentally distinct predictions concerning the behavioral and ecological costs and benefits associated with synchronous versus asynchronous vacancy chains. In laboratory studies of the terrestrial hermit crab Coenobita clypeatus, we found support for the prediction that social context alters shell acquisition behaviors. Field observations demonstrated that both synchronous and asynchronous vacancy chains are common and revealed previously undescribed waiting and piggybacking behaviors that appear to facilitate synchronous vacancy chains. Additionally, simulation results from an agent-based model showed that population density and waiting behaviors can both influence the likelihood of synchronous vacancy chains. Together, these results indicate that better understanding of hermit crab resource acquisition requires studying social behaviors, including vacancy chain formation.

How rugged individualists enable one another to find food and shelter: field experiments with tropical hermit crabs

Animals from invertebrates to humans benefit from information conspecifics make available, including information produced inadvertently. While inadvertent social information may frequently be exploited in nature, experiments have rarely been conducted in the wild to examine how such information helps animals in their natural ecology. Here I report a series of field experiments on free-living terrestrial hermit crabs (Coenobita compressus), showing how these asocial invertebrates learn the locations of their most essential resources, food and shelter, using inadvertent cues from conspecific competitors. Crabs have limited abilities to locate resources individually, but as they coalesce on a resource, their aggregation can be noticed by passing foragers, tipping them off about the discovery. Foragers were strongly attracted to experimentally simulated aggregations in which crabs were tethered to the same spot and in which the resources normally found beneath aggregations were excluded. Simulated aggregations of crabs whose shells were removed were likewise attractive, more than even these sought-after-shelters themselves. Experiments that simulated the chemical and visual cues of aggregations independently revealed that foragers oriented to aggregations primarily by sight, cueing in on the jostling competitive activity of the aggregation. Although crabs have not been selected to recruit others to newly discovered resources, their natural ecology has provided a setting where competitors regularly help one another by means of inadvertent social information.

3-Decanol in the haemolymph of the hermit crab Clibanarius vittatus signals shell availability to conspecifics

Hermit crabs with poor fitting shells are chemically attracted to dying gastropods and conspecifics where a shell may become available. For land hermit crabs, the shell cue is a volatile compound found in the haemolymph. Based on this knowledge, we tested the hypothesis that shell investigation behavior in aquatic hermit crabs, the ancestral predecessors of terrestrial hermit crabs, is also triggered by volatile cues. Volatile compounds from haemolymph of Clibanarius vittatus and Pagurus pollicaris and brachyuran decapod crustaceans were purged from a water–haemolymph solution, trapped in seawater and tested for induction of shell investigation behavior with juvenile C. vittatus. Only volatiles from C. vittatus haemolymph stimulated shell investigation. Volatile compounds were isolated from haemolymph by headspace solid-phase microextraction (SPME) and analyzed by coupled gas chromatography–mass spectrometry (GC–MS). Two prominent compounds were identified, 3-decanol, which was unique to C. vittatus haemolymph, and 2-ethyl-1-hexanol, which was present in the haemolymph of all 4 crustacean species. In shell investigation bioassays, 3-decanol from C. vittatus haemolymph stimulated shell investigation behavior, while 2-ethyl-1-hexanol did not. In bioassays with synthetic 1-, 2-, 4-, and 5-decanol, shell investigation behavior was evoked by 1-decanol, 5-decanol and 3-undecanol. There was no response to 2- and 4-decanol. The response of C. vittatus to volatile shell cues supports the hypothesis that volatile cue detection evolved prior to the occupation of terrestrial niches by crustaceans.

Vacancy Chains Provide Aggregate Benefits To Coenobita clypeatus Hermit Crabs

AbstractVacancy chain theory describes a unique mechanism for the sequential distribution of animal resources across multiple individuals. This theory applies to any resources, such as shelters or nest sites, that are discrete, reusable, and limited in use to single individuals or groups at one time. Hermit crabs rely on gastropod shells for shelter, and a single vacant shell can initiate a chain of sequential shell switches that distributes new resources across many individuals. Using the terrestrial hermit crab Coenobita clypeatus, we examined the previously untested theoretical prediction that this process will yield trickle‐down resource benefits to vacancy chain participants (aggregate benefits). In laboratory experiments, we measured improvements in shell quality when a single vacant shell was provided to groups of eight crabs. We found that crabs participating in vacancy chains (averaging 3.2 individuals) gained significant reductions in their shell crowding. In addition, vacancy chains terminated early when experimental groups included a single crab occupying a damaged shell, because damaged vacancies always remained unoccupied. Hermit crabs in damaged shells were more likely to win resource contests for high quality shells against size‐matched hermit crabs in crowded shells. Finally, field additions of many new shells to an island population of C. clypeatus hermit crabs reduced average shell crowding for crabs of all sizes, possibly from propagation of benefits through vacancy chains. These results provide empirical support for the theoretical prediction that vacancy chains should provide benefits distributed across many vacancy chain participants. Since shelter‐based vacancy chains likely occur in other animals, additional studies of vacancy chain processes should provide new insights into resource acquisition behaviors in diverse animal groups.

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to delta(15) N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in delta(15) N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation.

Weight difference threshold during shell selection relates to growth rate in the semi-terrestrial hermit crab Coenobita compressus

[To minimize the energetic expenditure imposed by carrying a protective gastropod shell, terrestrial hermit crabs should discriminate the weight difference between available shells; if the difference exceeds a certain amount — the threshold — the lighter shell should be accepted because of the cost of growing in a shell that is too heavy, while swapping between similar shells (less different than the threshold) may be inconsequential. We assigned shells of different weight adjusted to the crab's size to individually-housed crabs to (1) establish the weight difference at which crabs systematically reject a heavy shell and (2) compare growth rates of crabs confined to shells that were lighter- or heavier than the weight differential threshold. Crabs were significantly more likely to reject a heavy shell over a lighter one if the difference between the two was >50%, and when the shell weight exceeded 60% of the crab's weight. Crabs living in shells 50% heavier. This suggests that hermit crabs reject a heavier shell if a meaningful threshold is exceeded: this finding should prompt research into the neurophysiologic mechanism involved in shell assessment., To minimize the energetic expenditure imposed by carrying a protective gastropod shell, terrestrial hermit crabs should discriminate the weight difference between available shells; if the difference exceeds a certain amount — the threshold — the lighter shell should be accepted because of the cost of growing in a shell that is too heavy, while swapping between similar shells (less different than the threshold) may be inconsequential. We assigned shells of different weight adjusted to the crab's size to individually-housed crabs to (1) establish the weight difference at which crabs systematically reject a heavy shell and (2) compare growth rates of crabs confined to shells that were lighter- or heavier than the weight differential threshold. Crabs were significantly more likely to reject a heavy shell over a lighter one if the difference between the two was >50%, and when the shell weight exceeded 60% of the crab's weight. Crabs living in shells 50% heavier. This suggests that hermit crabs reject a heavier shell if a meaningful threshold is exceeded: this finding should prompt research into the neurophysiologic mechanism involved in shell assessment.]

Brain architecture in the terrestrial hermit crab Coenobita clypeatus(Anomura, Coenobitidae), a crustacean with a good aerial sense of smell

BackgroundDuring the evolutionary radiation of Crustacea, several lineages in this taxon convergently succeeded in meeting the physiological challenges connected to establishing a fully terrestrial life style. These physiological adaptations include the need for sensory organs of terrestrial species to function in air rather than in water. Previous behavioral and neuroethological studies have provided solid evidence that the land hermit crabs (Coenobitidae, Anomura) are a group of crustaceans that have evolved a good sense of aerial olfaction during the conquest of land. We wanted to study the central olfactory processing areas in the brains of these organisms and to that end analyzed the brain of Coenobita clypeatus (Herbst, 1791; Anomura, Coenobitidae), a fully terrestrial tropical hermit crab, by immunohistochemistry against synaptic proteins, serotonin, FMRFamide-related peptides, and glutamine synthetase.ResultsThe primary olfactory centers in this species dominate the brain and are composed of many elongate olfactory glomeruli. The secondary olfactory centers that receive an input from olfactory projection neurons are almost equally large as the olfactory lobes and are organized into parallel neuropil lamellae. The architecture of the optic neuropils and those areas associated with antenna two suggest that C. clypeatus has visual and mechanosensory skills that are comparable to those of marine Crustacea.ConclusionIn parallel to previous behavioral findings of a good sense of aerial olfaction in C. clypeatus, our results indicate that in fact their central olfactory pathway is most prominent, indicating that olfaction is a major sensory modality that these brains process. Interestingly, the secondary olfactory neuropils of insects, the mushroom bodies, also display a layered structure (vertical and medial lobes), superficially similar to the lamellae in the secondary olfactory centers of C. clypeatus. More detailed analyses with additional markers will be necessary to explore the question if these similarities have evolved convergently with the establishment of superb aerial olfactory abilities or if this design goes back to a shared principle in the common ancestor of Crustacea and Hexapoda.