Intraspecific competition and aggression for shells in the hermit crab Pagurus samuel1s

Many studies have investigated shell‐related behaviour in hermit crabs. Few studies, however, have focused specifically on the intraspecies aggression associated with shell competition. We examined intraspecies aggression in hermit crab (Pagurus samuelis) pairs as it relates to competition for a limiting resource, gastropod shells. Pairs of hermit crabs were observed in the laboratory in four different treatments that varied the presence or absence of shells for one or both of the crabs. Measurements of the latency to respond, the number of bouts, and the fight durations were recorded. There was a significant difference among treatments for all three measurements, and naked hermit crabs were much more aggressive than housed hermit crabs. There was no significant difference in aggression between males and females in any of the three treatments. The heightened aggression observed in naked P. samuelis is likely in service of acquiring a protective shell.     

Effects of chemical context on shell investigation behavior in hermit crabs

Specific chemicals in the environment evoke significant changes in the behavior of many aquatic organisms. We studied in the laboratory whether satiated individuals of the hermit crab, Pagurus longicarpus Say 1817, adjust their investigatory behavior towards an empty, optimal gastropod shell according to differences of chemical context. We also explored to what extent shell investigation by a crab in the same hunger state was affected by occupying an inadequately sized shell. Our results confirmed in part previous findings that crabs can discriminate the odor of freshly dead snails from the odor of freshly dead conspecifics. In the presence of the former odor, crabs inhabiting shells of inadequate size were more responsive and active than those in better-fitting shells. To the contrary, regardless of the quality of the inhabited shell, P. longicarpus remained practically motionless when presented with the odor of freshly dead conspecifics, possibly because the risks of incurring in predators would outweigh the benefits of acquiring a new shell. Unexpectedly, we found that crabs in both types of shell quality exhibited nearly the same behavior in control water, while crabs in adequate shells were more responsive in the presence of food odor. Individuals appeared insensitive to the odor of live snails; indeed, only one hermit crab species has been seen removing living snails from their shells. An intriguing result was that water conditioned by the odors of live conspecifics exerted a strong effect on all the individuals by inducing an intense shell investigation. Our study underlines the central role exerted by chemical detection in hermit crabs' behavior and demonstrates the existence of a complex interplay among chemical context, the physiological state of the animal, and the ecological pressures of the habitat.     

The effects of shell size and coil orientation on reproduction in female hermit crabs, Clibanarius vittatus

We investigated the effects of shell coil orientation and shell size on reproduction in field populations of the hermit crab, Clibanarius vittatus. Females were collected in the intertidal in Beaufort, NC. Shell parameters were measured and size (cephalothorax length) and reproductive status were determined for 70 females occupying Busycon shells. Crabs were categorized as berried (eggs on the pleopods), mature ovaries, or non-reproductive (no eggs). For berried females, the number of eggs was recorded. By offering a separate group of females access to empty shells, it was possible to calculate optimal shell size and the deficit in shell size for field-collected animals.Females that were berried were in shells closer to the optimal shell size than females with mature ovaries, both for shell weight and shell volume. And females with mature ovaries were in shells that were closer to the optimal size than females that were non-reproductive. For both categories of females without eggs on the pleopods, the majority of females were in shells that were too big (in weight and internal volume). While the percentage of berried females did not differ between dextral (Busycon carica) and sinistral (Busycon sinistrum) shells, the non-reproductive females had a much smaller deficit in volume in sinistral shells compared to dextral shells. For berried females, there was no relationship between the magnitude of their shell deficit and the number of eggs carried. Our results suggest that reproduction is inhibited when females occupy shells sufficiently greater than the optimal shell size.     

Shell shape affects movement patterns and microhabitat distribution in the hermit crabs Calcinus elegans, C. laevimanus and C. latens

This study examined the influence of shell shape on the distribution and movement patterns of three species of Hawaiian hermit crabs: Calcinus elegans, C. laevimanus, and C. latens. Field surveys showed strong differences in shell use depending on habitat. Individuals of C.elegans and C. latens were more frequently in unusual shapes of shells (the cowrie Cypraea caputserpentis and the variable worm shell Serpulorbis variabilis) when in tide pools and in more standard gastropod shells, such as the dog whelk Nassarius papillosus, when found in the subtidal. In addition, for both C.elegans and C. latens in tide pools, most crabs in unusual shaped shells were out on top of rocks, whereas most crabs in shells that were standard shapes were under rocks.In the laboratory, individuals of C.elegans and C. laevimanus in unusual shells initiated more shell exchanges and when given empty shells crabs readily occupied the standard shaped shells, but crabs did not move into the unusual shaped shells. Mark-recapture experiments in the field showed that C. elegans in standard shaped shells moved out of tide pools and stayed longer when placed on subtidal coral heads, whereas crabs in unusual shaped shells stayed in tide pools and did not stay on subtidal coral heads (in part due to predation). Laboratory tests showed that C. elegans in unusual shaped shells were more readily dislodged by surge than crabs in standard shaped shells. Thus, the difference in movement patterns in preferred vs. unpreferred shell shapes is an important factor influencing the microhabitat distribution of these hermit crabs.     

Auditory stimulation dishabituates anti-predator escape behavior in hermit crabs (Coenobita clypeatus)

Responses to innocuous stimuli often habituate with repeated stimulation, but the mechanisms involved in dishabituation are less well studied. Chan et al. (2010b) found that hermit crabs were quicker to perform an anti-predator withdrawal response in the presence of a short-duration white noise relative to a longer noise stimulus. In two experiments, we examined whether this effect could be explicable in terms of a non-associative learning process. We delivered repeated presentations of a simulated visual predator to hermit crabs, which initially caused the crabs to withdraw into their shells. After a number of trials, the visual stimulus lost the ability to elicit the withdrawal response. We then presented the crabs with an auditory stimulus prior to an additional presentation of the visual predator. In Experiment 1, the presentation of a 10-s, 89-dB SPL noise produced no significant dishabituation of the response. In Experiment 2 we increased the duration (50 s) and intensity (95 dB) of the noise, and found that the crabs recovered their withdrawal response to the visual predator. This finding illustrates dishabituation of an antipredator response and suggests two distinct processes—distraction and sensitization—are influenced by the same stimulus parameters, and interact to modulate the strength of the anti-predator response.     

Interference and exploitation components in interespecific competition between sympatric intertidal hermit crabs

This study was designed to evaluate the effect of interference and exploitation competition in shell partitioning between two hermit crab species (Pagurus criniticornis and Clibanarius antillensis). Field samples revealed that shells of the gastropod Cerithium atratum were the main resource used by both hermit crab species and that Pagurus used eroded or damaged shells in higher frequency than Clibanarius. The exploitative ability of each species was compared between species in the laboratory using dead gastropod (Cerithium) baits to simulate predation events and signalize newly available shells to hermit crabs. Pagurus reached the baits more rapidly than Clibanarius, but this higher exploitative ability did not explain shell utilization patterns in nature. Another experiment evaluated the dominance hierarchy between these two hermit crab species and revealed that Clibanarius was able to outcompete Pagurus for higher quality shells in agonistic encounters. This higher interference competitive ability of Clibanarius in relation to Pagurus may explain field observations. Nevertheless, Pagurus may be responsible to enhance shell availability to other hermit crab species that have lower ability to find and use newly available shells. Differently, the poorer condition of shells used by Pagurus, the higher ability of this species to attend gastropod predation events and its higher consumption rate by shell-breaking crabs (Menippe nodifrons) may increase its predation risks, thus revealing the disadvantages of such an exploitative competitive strategy for hermit crabs.     

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.     

The use of pleopods for shell water circulation and respiration by hermit crabs

The use of the pleopods was investigated in two species of Diogenid hermit crabs, Dardanus arrosor and D. calidus. A transparent glass shell was used to observe the movements of the pleopods in hermit crabs. The movements consisted of periodic, irregular beating, which generated an irregular flow of the water within the shell.

A strong, regular beating movement was elicited when there was detritus or faeces in the shell, as was occupation of a new shell. Injection of water with a low oxygen concentration into the shell failed to induce any variation in the pleopod beating frequency (PBF), while the injection of water with a high CO₂ concentration induced a sudden PBF increase. When pH was varied and CO₂ concentration held at normal atmospheric level, there was a change in PBF. However, CO₂ variation alone, at constant pH, did not trigger any visible reaction. This suggests that there is a receptor for pH, which acts independently of the CO₂ concentration.

The function of males hermit crabs pleopods has always been obscure. However, at least in Dardanus, they are actively involved in water circulation within the shell for shell-cleaning and probably for ventilation.     

Stimulus concordance and risk-assessment in hermit crabs (Coenobita clypeatus): implications for attention

Recent research has demonstrated that the topography of defensive reactions depends on factors that are extraneous to the stimulus that elicits the defensive response. For example, hermit crabs will withdraw more slowly to the approach of a simulated visual predator (i.e., the eliciting stimulus) when in the presence of a coincident acoustic stimulus. Multiple properties related to the magnitude (e.g., duration, amplitude) of the acoustic stimulus have been found to modulate the crabs' withdrawal response (Chan et al., 2010b). We demonstrate that the proximity in spatial location between a threatening visual stimulus and a potentially distracting extraneous auditory stimulus is an important determinant of anti-predator behavior in hermit crabs. We suggest that a distal relationship between the eliciting stimulus and an unrelated signal may produce greater distraction. This marks the first reported experimental evidence of this relationship in an invertebrate species.     

Desiccation tolerance of four sympatric tropical intertidal hermit crabs (Decapoda,Anomura)

This study describes and quantifies the air exposure tolerance of four sympatric hermit crabs (Pagurus criniticomis, Clibanarius antillensis, C. sclopetarius, and C. vittatus) in an intertidal area in southeastern Brazil. We report on survivorship, percentage and rate of weight and water loss until death, and overall body water content. The coexisting populations showed similar percentages of overall body water content and weight and water loss until death. Survivorship depended on crab size rather than species, and was positively correlated with crab size within each species (linear relationship) and with the size of all individuals of this hermit crab assemblage (exponential relationship). Each species had a characteristic rate of weight and water loss, indicating the existence of different physiological adaptations to resist desiccation. These differences were directly related to the species’ distribution patterns in the intertidal zone.     

Shell recruitment in the Mediterranean hermit crab Clibanarius erythropus

Gastropod shells are vital for the majority of hermit crab species, being essential for their survival, growth, protection, and reproduction. Given their importance, shells are acquired and transferred between crabs through several modalities. We conducted observations and experiments at the Asinara Island (Sardinia, Italy) to investigate the efficacy of the different behavioral tactics adopted by the hermit crab Clibanarius erythropus to acquire shells, such as: (1) locomotion and activity at different tidal phases; (2) attendance at shell-supplying sites (simulated predation sites with five different odors: live and dead gastropods, live and dead crabs, predator); and (3) interactions with conspecifics in aggregations on simulated gastropod predation sites. In each tidal phase, locomotion was slow (0.7 cm min^− 1) and, as a consequence, the probability of encountering empty shells and conspecifics was low. Simulated gastropod predation sites quickly attracted a larger number of hermit crabs than the other sites tested. Aggregations seemed to function as shell exchange markets, as previously suggested for other species: the first attendant took the experimental shell and a chain of shell exchanges among conspecifics followed. Our results show that, in C. erythropus, aggregation is the most efficient tactic for the acquisition of new shells, whereas in other species, such as Pagurus longicarpus, it is associated with exploitation ability due to the intense locomotion. The interspecific plasticity in hermit crabs' behavior is confirmed.     

Morphological traits determine the winner of “symmetric” fights in hermit crabs

This study aimed at investigating in the hermit crab Pagurus longicarpus whether some morphological traits and their slight variations might determine the winner of a contest in symmetric pairs, i.e. pairs composed of individuals matched by their overall size. In the pre-experimental phase, 400 crabs were individually kept in isolation for a week to eliminate the effects of their previous social experiences and were allowed to enter adequate shells to equalize their motivation to fight. Then, we formed 200 pairs matched for shell and body size and observed their agonistic behaviour for 15 min. Alphas (and betas) were deemed as those individuals that won more than half of the recorded fights. Finally, crabs were sacrificed and sexed, and 6 and 8 measures were taken of their shells and bodies, respectively. Raw data were converted to compositional data and subjected to an isometric log-ratio transformation prior to statistical analysis. Alphas were found to occupy heavier and wider shells than betas, both characteristics that, together with the correlated high level of encrustation, make the apparent dimension of the crab larger and thus favour that individual during agonistic encounters. As expected, hermit crabs were sexually dimorphic in their chelae, but sexes differed also in the length of the dactylus of their third walking leg and in their body weight. Alphas were heavier than betas and had longer propodi and wider dactyli on the third walking legs than betas. The latter character might be advantageous during shell fights when the walking legs of the attacker are used to hold the defender's shell in the spasmodic shaking and their wider dactyli might assure a firmer seizing of it. Indeed, a correct execution of this pattern allows for the eviction of the defender. Our results suggest that more attention towards variations in morphological traits other than body size might improve our understanding of the factors that affect individual Resource Holding Power and fight dynamics.     

Resource partitioning and competition for shells between intertidal hermit crabs on the outer coast of Washington

Summary Resource partitioning was quantified for 6 species of intertidal hermit crabs in the genus Pagurus, that occur on the outer coast of Washington. This, together with field evidence of shell shortage and with laboratory experiments to quantify the mechanism of interactions for shells, allowed estimation of the relative intensities of inter-and intraspecific competition between these species. The findings were that: (1) the magnitude of intraspecific competition was greater than any single interspecific competitive effect for all of the species; and (2) the relative proportion of intraspecific competition was greater for the middle and upper intertidal species than for the lower intertidal species. Studies at several outer coast sites supported these generalizations. Both of these findings are consistent with the hypothesis that competitive divergence has occurred in the past. The structure of the outer coast hermit crab assemblage is compared with that of the San Juan Archipelago hermit crab assemblage. Differences between the two do not seem to be the result of adaptive responses to the presence of more competing species in the former group.     

Shell selection behaviour and spatial distribution of three species of intertidal hermit crabs from Hormuz Island,Persian Gulf (Crustacea: Paguroidea)

Shell selection behaviour and spatial distribution of three hermit crab species, Diogenes avarus, D. karwarensis, and Areopaguristes perspicax, were studied at six sites along the intertidal zones of Hormuz Island in the Persian Gulf. 1025 specimens were collected occupying altogether 31 shell species (D. avarus 28 species, A. perspicax 22 species, and D. karwarensis 8 species). Diogenes avarus was found to be by far the most abundant of these three crab species, and Cerithidea cingulata the dominant shell occupied by these hermit crabs. The distribution of the hermit crabs significantly varied (p<0.05) among the sites. The number and the wide diversity of shells occupied in different sites show that the main factor in shell selection for these hermit crabs is the abundance and distribution of shell species in the field.     

Why hummingbirds have such large crops

Summary Male Anna's Hummingbirds (Calypte anna) defend territories that contain a predictable food source, floral nectar. For such a hummingbird, the meal size that maximizes long-term net energy intake is less than the maximal crop volume. Smaller meals must be consumed more frequently, but larger meals increase body mass and therefore flight cost. Individuals without territories or with inadequate territories do not have easy access to nectar and intrude on territories owned by otherC. anna, where they may be chased at nay time. It was predicted that these intruders should minimize the number of potentially risky intrusions necessary for maintenance by ingesting as much nectar as possible whenever they manage to feed without being chased (usually when owners are temporarily absent). Therefore, relative to uninterrupted feeding by owners, uninterrupted intruders should feed longer and take larger meals. Field observations supported these predictions. Intruders apparently filled their crops in all seasons, whereas owners ingested smaller amounts (0.21–0.22 ml) and fed for lengths of time consistent with the prediction of an optimization model (0.21 ml). Thus, owners may energetically optimize meal size whereas intruders fill their crops whenever they are not chased. Under most conditions, hummingbirds only fill their crops one-tenth to one-third full, leading to the question why hummingbirds have such large crops. This study demonstrates that a large crop volume may be of survival value when an individual lacks a territory or has inadequate access to resources and must poach on others' territories.     

Sex-specific differences and the role of predation in the interaction between the hermit crab, Pagurus longicarpus, and its epibiont, Hydractinia symbiolongicarpus

The interaction between the hermit crab, Pagurus longicarpus, and the shell epibiont, Hydractinia symbiolongicarpus, varies from mutualism to parasitism based on the environmental context. We tested the hypothesis that this interaction also varies as a function of hermit crab sex. Given that recent work showed a negative effect of Hydractinia on female reproduction, we predicted a greater frequency of males in Hydractinia shells in the field and a stronger preference by males than females for shells with Hydractinia. Field collections documented a significantly greater proportion of males than females occupying shells with Hydractinia, and a significantly greater proportion of ovigerous females than non-ovigerous females in shells with Hydractinia. In laboratory shell-switching experiments, a greater proportion of males than females chose to enter shells with Hydractinia, but there was no difference in the proportions of males and females that vacated shells with Hydractinia.We examined whether the presence of Hydractinia influenced predation rates. Blue crabs fed on more than twice as many hermit crabs in shells with Hydractinia as compared to bare shells, but there was no significant difference for stone crabs. Laboratory experiments showed that the force required to crush shells was significantly greater for shells without Hydractinia. Thus, the lower occupancy and preference exhibited by females than males for shells with Hydractinia appears to result both from the decreased reproduction shown in past studies and an increase in predation risk.     

埃塞俄比亚Alatish国家公园大型哺乳动物多样性、分布及其与栖息地的联系

于2004年8月-2005年3月间研究了Alatish国家公园在雨季和旱季期间大型哺乳动物的多样性和分布.基于地形图、卫片和预调查结果,在4种生境类型中识别出37个100 km2或者更小的斑块.在4个植被类型中随机选择了11个斑块,沿随机选择的样线进行了调查,样线10 km 长,0.2 km宽,覆盖了该区域面积的20%.使用Shannon-Wiener多样性指数、DISTANCE 5.0 软件以及 Simpson相似性指数分析了大型哺乳动物的多样性以及分布的相似性.记录了20种体型较大的哺乳动物;河边林地在雨季和旱季都有最高的多样性,林地的多样性最低;旱季中4种类型的栖息地具有相似的分布型;由于栖息地退化不能确定物种的分布格局.联合距离取样分析揭示出,旱季和雨季大型哺乳动物的密度较低,分别为4.067只/km2和6.263只/km2.所观察的哺乳动物可分为草食动物(15%)、游牧动物(30%)、杂食动物(25%)与食肉动物(30%).     

Population structure of large yellow croaker (Larimichthys crocea) revealed by single nucleotide polymorphisms

The large yellow croaker, Larimichthys crocea, is one of the most commercially important marine species native to China Seas. In this study, Sequenom MassARRAY was used to analyze 120 individuals from three geographic populations in the East and South China Seas and a cultured stock. Forty-four of 60 single nucleotide polymorphism (SNP) loci were ascertained. Eleven loci, scf117, scf708, scf247, scf595, scf100, scf166, scf305, scf664, scf117, scf187 and scf511 each showed population-specific genotypes. A total of 127 alleles were detected. Averagely, the effective number of alleles was between 1.328 and 1.341, expected heterozygosity and observed heterozygosity ranged from 0.273 to 0.320, and 0.190 to 0.235, respectively, Pic value ranged from 0.215 to 0.247, and F_IS ranged from 0.174 to 0.279. The neighbor-joining tree based on calculated genetic distances showed a concordant topology with the traditional morphological characterization. When the population size of wild large yellow croaker drastically has shrunk over the past 20 years, the genetic diversity in both wild and farmed stocks has declined as well.