Insect-like olfactory adaptations in the terrestrial giant robber crab

The robber crab (Birgus latro), also known as the coconut crab, is the world's largest land-living arthropod, with a weight reaching 4 kg and a length of over half a meter [1]. Apart from the marine larval stage 2, 3, this crab is fully terrestrial, and will actually drown if submerged in water [4]. A transition from sea to land raises dramatically new demands on the sensory equipment of an animal. In olfaction, the stimulus changes from hydrophilic molecules in aqueous solution to mainly hydrophobic in the gaseous phase [5]. The olfactory system of land crabs thus represents an excellent opportunity for investigating the effects of the transition from sea to land. Have land crabs come to the same solutions as other terrestrial animals, or is their olfactory sense characterized by unique innovations? Here, we show that the robber crab has evolved an olfactory sense with a high degree of resemblance to the insect system. The similarities extend to physiological, behavioral, and morphological characters. The insect nose of the robber crab is a striking example of convergent evolution and nicely illustrates how similar selection pressures result in similar adaptation     

Land hermit crabs use odors of dead conspecifics to locate shells

A series of experiments at two tropical locations tested the ability of land hermit crabs Coenobita perlatus (H. Milne Edwards) and Coenobita compressas (H. Milne Edwards) to detect and respond to odors of dead conspecifics. An attraction array compared numbers of crabs attending hidden food odors and dead conspecific odors. Pit experiments tested crab shell-acquisition behaviors at different hidden odors. Bucket experiments confined crabs collected from various categories (feeding crabs, wandering crabs and crabs aggregated at dead conspecific odors) and tested behavioral responses to odors and an empty shell. Land hermit crab behavior at both sites was similar. Crabs were attracted to dead conspecific odors up to 10 times more than to food odors. Crabs attracted to dead conspecifics displayed significantly more shell-acquisition behaviors: touching other crab's shells in an exploratory manner and switching shells if an empty shell was available. In buckets, crabs from each category switched into shells. Results are compared to previous reports of similar shell-seeking behaviors by marine hermit crabs in response to dead conspecific odors. It is suggested that responding to dead conspecific odors for shell source location is an evolutionarily conserved behavior developed before hermit crabs became terrestrial.     

A Spatially Explicit Model of Synchronization in Fiddler Crab Waving Displays

Fiddler crabs (Uca spp., Decapoda: Ocypodidae) are commonly found forming large aggregations in intertidal zones, where they perform rhythmic waving displays with their greatly enlarged claws. While performing these displays, fiddler crabs often synchronize their behavior with neighboring males, forming the only known synchronized visual courtship displays involving reflected light and moving body parts. Despite being one of the most conspicuous aspects of fiddler crab behavior, little is known about the mechanisms underlying synchronization of male displays. In this study we develop a spatially explicit model of fiddler crab waving displays using coupled logistic map equations. We explored two alternative models in which males either direct their attention at random angles or preferentially toward neighbors. Our results indicate that synchronization is possible over a fairly large region of parameter space. Moreover, our model was capable of generating local synchronization neighborhoods, as commonly observed in fiddler crabs under natural conditions.     

Sexual selection and the physiological consequences of habitat choice by a fiddler crab

In mid-Atlantic salt marshes, reproductively active male sand fiddler crabs, Uca pugilator, use a single greatly enlarged major claw as both a weapon to defend specialized breeding burrows from other males and an ornament to attract females for mating. During the summer breeding season, females strongly prefer to mate with males controlling burrows in open areas high on the shore. Food availability decreases while temperature and desiccation stress increase with increasing shore height, suggesting that the timing and location of fiddler crab mating activity may result in a potential trade-off between reproductive success and physiological condition for male crabs. We compared thermal preferences in laboratory choice experiments to body temperatures of models and living crabs in the field and found that from the perspective of a fiddler crab, the thermal environment of the mating area is quite harsh relative to other marsh microhabitats. High temperatures significantly constrained fiddler crab activity on the marsh surface, a disadvantage heightened by strongly reduced food availability in the breeding area. Nevertheless, when the chance of successfully acquiring a mate was high, males accepted a higher body temperature (and concomitantly higher metabolic and water loss rates) than when the chances of mating were low. Likewise, experimentally lowering costs by adding food and reducing thermal stress in situ increased fiddler crab waving display levels significantly. Our data suggest that fiddler crabs can mitigate potential life history trade-offs by tuning their behavior in response to the magnitude of both energetic and non-energetic costs and benefits.     

Predator-prey dynamics between recently established stone crabs (Menippe spp.) and oyster prey (Crassostrea virginica)

Range expansion and population establishment of individual species can have significant impacts on previously established food webs and predator-prey dynamics. The stone crab (Menippe spp.) is found throughout southwestern North Atlantic waters, from North Carolina through the Gulf of Mexico and the Central American Caribbean, including the Greater Antilles. Recent observations suggest that stone crabs have become better established on certain oyster reefs in North Carolina than in the early 1900s when they we first observed in NC. To assess the predatory impact of stone crabs on oysters, we (1) quantified stone crab densities on subtidal oyster reefs in Pamlico Sound, NC using scuba surveys, and (2) conducted laboratory predation experiments to assess the functional response of stone crabs to varying densities of oysters. We then (3) analyzed previously unpublished functional response data on another important oyster predator, the mud crab Panopeus herbstii. Finally, we (4) compared and contrasted potential predatory impacts of stone, mud and blue crabs (Callinectes sapidus). The functional response data and analyses for both stone crabs and mud crabs were consistent with a type II functional response. Mud crabs, on a m² basis, inflicted the highest proportional mortality on oysters over a 24 hour period, followed by stone and then blue crabs. Proportional mortality did not vary significantly with oyster size; however, relatively small and large oysters were consumed disproportionately less than medium-sized oysters, likely due to the mechanical inability of stone crabs to handle small oysters, and the inability to crush large oysters. Although stone crabs appear to be established in Pamlico Sound at densities equivalent to densities in other systems such as the U.S. Florida Panhandle, their predatory activities on oysters are not expected to have as significant a negative impact on oyster populations compared to other resident predators such as mud crabs.     

The effect of audience on intrasexual interaction in the male fiddler crab, <Emphasis Type="Italic">Uca maracoani</Emphasis> (Latreille 1802–1803, Decapoda,Ocypodidae)

Animal communication occurring in wide networks can involve signals sent to several receivers. The animals composing the audience may affect how individuals display during an interaction and may change the message that is sent. In this study, we investigated whether the presence of a conspecific affected the intensity of agonistic interaction between male fiddler crabs, Uca maracoani. Pairs of males of the same size were observed when in the presence of a male, a female or no crab as audience. We found that if there is a female audience, males became more aggressive and interacted the most. Also, the female audience leads to an increase in incidence of male foaming, possibly indicating predisposition for mating. If the audience was a male or if there was no audience, there was no significant difference in interaction. These results indicate that the presence of an audience affects the way male fiddler crabs interact and the type of displays they show. Therefore, the context seems to guide the fiddler crab behavior in terms of how they perform in order to send information about themselves to conspecifics.     

A comparison of reproductive patterns and adult dispersal in sympatric introduced and native marine crabs: implications for species characteristics of invaders

The introduction and translocation of nonindigenous marine species is widespread and can pose severe threats to biodiversity and ecosystem functioning. Predicting which species are potential invaders is of particular interest to ecologists. One approach is to identify characteristics that predispose a species to becoming a successful invader. Since its introduction in the 1980’s, the invasive Asian shore crab, Hemigrapsus sanguineus, has shown a remarkable ability to colonize rocky intertidal habitats along the east coast of the United States. In Long Island Sound H. sanguineus occurs sympatrically with the functionally equivalent, but non-invasive, native Atlantic mud crab, Panopeus herbstii. The presence of both species at the same site allowed us to make a detailed, simultaneous assessment of life history traits and adult dispersibility of co-occurring invading and native crab species. We investigated fecundity and maturation rates, length of breeding season and brood production for both species, and conducted field experiments using mark–recapture techniques to determine mobility patterns. Our results show that the nonindigenous Asian crab has a greater reproductive potential than the native mud crab as evidenced by a longer breeding season, multiple brood production and higher fecundity rates. Field experiments confirmed previous studies indicating H. sanguineus is a highly mobile crab, and further demonstrated that adult Asian crabs are more likely than mud crabs to disperse from their shelter/refuge sites. Recovery rates for native mud crabs were significantly higher than those for Asian crabs in three experimental trials, across sites and years. This work provides new information about life history characteristics of both species and supports the hypothesis that high reproductive potential combined with high adult dispersal ability may be important factors associated with the invasion/establishment success of the Asian shore crab. More study is needed, however, to determine the applicability of these findings to other highly successful marine invaders.     

Density affects mating mode and large male mating advantage in a fiddler crab

Fiddler crabs show two different mating modes: either females search and crabs mate underground in male burrows, or males search and crabs mate on the surface near female burrows. We explored the relationship between crab density, body size, the searching behavior of both sexes, and the occurrence of both mating modes in the fiddler crab Uca uruguayensis. We found that crabs change their mating mode depending on their size and crab density. Crabs mated mostly on the surface at low densities, and underground at high densities. The proportion of wandering receptive females but not courting males accounted for the variation in mating modes. This suggests that whether crabs mate underground (or on the surface) is determined by the presence (or absence) of searching females. We found that the change in the mating mode affected the level of assortative mating; males mating underground were bigger than those mating on the surface, suggesting active female choice. Given that fiddler crabs experience multiple reproductive cycles, they are prone to showing behavioral plasticity in their mating strategy whenever the payoffs of using different mating modes differ between reproductive events. Our results suggest that the incorporation of different levels of environmental variability may be important in theoretical models aimed at improving our understanding of the evolution of alternative mating tactics and strategies.     

Movement,activity pattern,and rôle of a hermit crab population in a sublittoral epifaunal community

The movements and activity of hermit crabs in a sublittoral soft-bottom community in the North Adriatic Sea (Gulf of Trieste) were monitored by means of observations by a SCUBA diver and time-lapse photography. Tagging experiments showed that the hermit crabs exhibit a pattern of movement which causes them to remain in a defined area for extended periods. During a two-year period, the radius of activity of a crab is estimated to be under 15 m. This leads to a high recovery rate of tagged shells. The average speed of a hermit crab was 2.1 m/h and the average distance travelled per day was 21.6 m as calculated from the analysis of the time-lapse films. This distance is an order of magnitude larger than that estimated from the tagging experiments (1.1 m/day). The activity of the crabs was restricted to the sediment surface between multi-species clumps. The approximate area of the sediment surface traversed by a crab is 0.4 m²/day. Many inter- and intraspecific encounters were observed, particularly among hermit crabs and the brittle stars Ophiothrix quinquemaculata (D.Ch.) and Ophiura lacertosa (Pennant). The daily activity pattern of the crabs is related to the light and dark phases on the sediment surface. The high density and conspicuous activity of the hermit crabs point to the important rôle of the pagurids in the investigated community.     

Experimental studies on the foraging behavior of the sand fiddler crab Uca pugilator (Bosc, 1802)

The purpose of this study was to test the hypothesis that foraging sand fiddler crabs. Uca pugilator (Bosc), move through the habitat in response to low substratum food levels even though these movements may take the crabs considerable distances from the safety of the burrow area. Chl a and ATP concentrations were used as measures of food density in foraged and unforaged substratum. Field and laboratory feeding experiments showed that crab foraging intensity in a habitat patch was directly correlated with food density in the patch either in the presence or absence of alternative food patches. Other experiments showed that sand fiddlers can respond to differences in food level on a scale of millimeters and do this by probing the substratum with minor chelae. Food levels in aggregations of non-ingested particles harvested by sand fiddlers, feeding pellets, correspond to low foraging intensities predicted from foraging experiments and crabs exhibit low foraging intensities on substratum patches derived from feeding pellets. Substratum food levels in two distinct areas corresponded to high predicted foraging intensities and there was no consistent trend in the level of food in the burrow vs. the nonburrow microhabitats. These results suggest that the movements of foraging sand fiddlers are to some extent controlled by the reduction in substratum food levels due to feeding during a single foraging episode. Sand fiddlers can extract over 70% of the food from harvested substratum over a broad range of substratum food densities but harvest only 42% of the available substratum.     

On the growth of the blue king crab (<Emphasis Type="Italic">Paralithodes platypus</Emphasis>) population in the Peter the Great Bay of the sea of Japan

The causes of the appearance of large blue king crabs (Paralithodes platypus) in Peter the Great Bay for the last decade are discussed. This species is an important commercial resource in the waters of Russian Far Eastern seas, and its general concentrations are related mainly to the sublittoral and upper bathyal zones of the northwestern Bering Sea and the northern Sea of Okhotsk. Until recently, this species has been observed in areas along the continental coast of the northwestern Sea of Japan up to the Peter the Great Bay, where it incidentally showed up in red king crab (P. camtschaticus) and snow crab (Chionoecetes opilio) catches but was also commercially used. This area was considered as the southern periphery of the species range. Since the late 1990s, both male and female blue king crabs have been recorded in trawl and trap catches during research works conducted within the Peter the Great Bay. Since 2002, any commercial catches of shelf crab species are prohibited in the waters south of 47°20′ N because of a dramatic decline in their populations. Since then all the illegally caught crabs, including blue king crabs that are seized live from poachers, are released back into the water in certain places of the bay. In total, at least 29 503 blue king crabs, including egg-bearing females, were released within the period from 2002 to November 2009. At present, the overall blue king crab abundance in Peter the Great Bay, estimated based on the trap catches over an area of 7048 km², is 50500, the abundance of commercial-size males (with a carapace width over 130 mm) is 7500, and the male to female ratio is 1.00: 1.35. The increase in the blue king crab population observed in the bay is the result of the immigration of mature and viable individuals from other areas of its range. After this “uncontrolled introduction” blue king crabs adapted to new conditions, and then began breeding and spreading over the entire area of the bay.     

Reproductive biology and genetic diversity of the Sea Anemone Aulactinia stella (Verrill, 1864)

Artificial reef areas can be used for management and conservation of commercially exploited crustacean decapods but their behaviour in these environments is poorly characterised. Acoustic tags were used to study the behaviour of 3 adult spiny lobsters (Palinurus mauritanicus) and 3 adult spider crabs (Maja squinado) over a period of 64 days in summer, evaluating the use of artificial reef areas as suitable sites for re-stocking of overfished decapods. For this purpose, animals were released in a western Mediterranean artificial reef located at 20 m depth, close to a cabled seafloor observatory, which simultaneously recorded temperature, salinity, current direction, current speed and light intensity over the study period. Spiny lobsters lingered in the reef area, whereas the spider crabs left the area 21–45 h after release. These behavioural differences suggested that artificial reefs might be a good area to deploy adult lobsters for re-stocking purposes. The movements displayed by the lobsters during this experiment were not influenced by any of the measured environmental factors, whereas spider crab movements occurred against major current direction, when current speed was intense.     

Ecology and behavior of Gecarcoidea natalis, the Christmas Island red crab, during the annual breeding migration.

The terrestrial crab Gecarcoidea natalis is endemic to the forests of Christmas Island but must migrate each year to the coast to breed. During 1993 and 1995, radio-tracking, mark and recapture, and counting methods were used to establish the routes, walking speeds, direction of travel, and destinations of migrating crabs, as well as crab numbers and distribution. The density of crabs ranged from 0.09 to 0.57 crabs per square meter, which gave a population estimate of 43.7 million adult crabs on the island. During the dry season the crabs were relatively inactive but on arrival of the wet season immediately began their migration. The crabs generally walked in straight lines, and most crabs from around the Island traveled toward the northwest shore instead of simply walking toward the nearest shore. The maximum recorded distance walked by a red crab in one day was 1460 m, but the mean was 680 m per day in 1993 and 330 m in 1995. Comparing the 1993 and 1995 study seasons, there was a 3-week difference in the timing of the start of the migration, but the spawning date was fixed by the lunar phase and took place 17 to 18 days after mating. In 1993, late rain prompted a "rushed" migration and crabs walked directly to their shore destinations; in contrast, in 1995 most crabs made stops of 1 to 7 days during the downward migration. By giving the crabs a chance to feed along the way and minimizing the time that the population was concentrated near the shore, these stops may be important in ensuring that the animals have enough food after the long dry season. Furthermore, this behavior implies that the crabs are able to judge how far away they are from the shore during the migration.     

Behavioral and electrophysiological experiments suggest that the antennular outer flagellum is the site of pheromone reception in the male helmet crab Telmessus cheiragonus

Sexually competent females of Telmessus cheiragonus (helmet crab) release two pheromones that elicit grasping and copulation behaviors in males (Kamio et al., 2000, 2002, 2003). Our study aimed to use behavioral and electrophysiological techniques to identify the site of reception of these sex pheromones. In behavioral experiments, either the inner or the outer flagella of the antennules were ablated bilaterally from male crabs, and responses of male crabs to female odor were examined. When the inner flagella were surgically ablated, the sexual response (i.e., grasping and copulation behavior) of male crabs was not significantly changed relative to control animals that had their second antennae ablated. In contrast, the sexual response was significantly reduced when the outer flagella of the antennules were ablated, suggesting that the outer flagellum is the receptor organ that detects the sex pheromones. In electrophysiological experiments, urine, which in females contains the pheromone that elicits grasping behavior by males but does not contain the pheromone eliciting copulation, whose release site is not known, was tested. Female and male urine as well as shrimp extract evoked phasic responses of chemosensory afferents innervating aesthetasc sensilla on the outer flagellum of male crabs. The response of the afferents had significantly higher magnitude and lower threshold when female urine was applied. Thus, behavioral and electrophysiological observations suggest that in male helmet crabs, the outer flagellum of the antennule is the chemosensory organ that detects female sex pheromone.     

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

Molecular keys unlock the mysteries of variable survival responses of blue crabs to hypoxia

Hypoxia is a major stressor in coastal ecosystems, yet generalizing its impacts on fish and shellfish populations across hypoxic events is difficult due to variability among individuals in their history of exposure to hypoxia and related abiotic variables, and subsequent behavioral and survival responses. Although aquatic animals have diverse physiological responses to cope with hypoxia, we know little about how inter-individual variation in physiological state affects survival and behavioral decisions under hypoxic conditions. Laboratory experiments coupled with molecular techniques determined how extrinsic factors (e.g., water body and temperature) and respiratory physiology (hemocyanin concentration and structure) affected survival and behavior of adult blue crabs (Callinectes sapidus) exposed to different levels of hypoxia over a 30-h time period. Nearly 100% of crabs survived the 1.3 mg dissolved oxygen (DO) l^?1 treatment (18.4% air saturation), suggesting that adult blue crabs are tolerant of severe hypoxia. Probability of survival decreased with increasing hypoxic exposure time, lower DO, and increasing temperature. Individual-level differences in survival correlated with water body and crab size. Crabs collected from the oligo/mesohaline and hypoxic Neuse River Estuary (NRE), North Carolina, USA survived hypoxic exposures longer than crabs from the euhaline and normoxic Bogue and Back Sounds, North Carolina. Furthermore, small NRE crabs survived longer than large NRE crabs. Hemocyanin (Hcy) concentration did not explain these individual-level differences, however, hypoxia-tolerant crabs had Hcy structures indicative of a high-O₂-affinity form of Hcy, suggesting Hcy “quality” (i.e., structure) may be more important for hypoxia survival than Hcy “quantity” (i.e., concentration). The geographic differences in survival we observed also highlight the importance of carefully selecting experimental animals when planning to extrapolate results to the population level.     

The ectosymbiont crab Dissodactylus mellitae-sand dollar Mellita isometra relationship

The presence of the ectosymbiont Dissodactylus mellitae on Mellita isometra was studied at Tybee Island in Georgia. The number of crabs found on sand dollars, stage of maturity, sex, numbers of eggs, and size of eggs produced by crabs were noted. The test diameter of sand dollars, and the number and diameter of eggs produced were also noted. Infestation rates of crabs on sand dollars varied over time. The maximum number of crabs found on a sand dollar was 10. The three types of population dispersion, random, uniform, and clumped, were observed for crabs on sand dollars throughout the sampling period. Clumped or gregarious settling was observed when juvenile crabs were abundant, uniform and random distribution when mature crabs were abundant. Female crabs were significantly larger than male crabs, with carapace width of the largest mature female crab being 4.6 mm and the largest male 3.1 mm. Crabs produced between 80 and 300 eggs from 0.188 to 0.291 mm in diameter. Bigger crabs produced significantly more eggs than smaller crabs. Sand dollar sizes were 50-110 mm, with a mode between 60 and 70 mm. The effect of crab burden on egg production in the sand dollar was time-dependent. The presence of crabs on sand dollars correlated with total egg production of sand dollars in May, the peak of the spawning season, with sand dollars carrying one or two crabs having a lower egg production than those without crabs. Overall, variation in egg size was attributed to variation among females followed by variation between seasons and crab burden.     

A comparison of the predatory impacts of an invasive and native crab species using a functional response approach

The European green crab (Carcinus maenas) is invasive on the West coast of North America, but the ecological consequences of this invasion remain poorly understood. Comparative functional response analysis has arisen as a method of elucidating ecological consequences of invasive species by comparing the impact of these species to native analogues. Through comparative functional response experiments of green crabs and native red rock crabs (Cancer productus) we found that green crab predation increased asymptotically (Type II functional response) when fed increasing densities of Pacific oysters (Magallana gigas), while red rock crab predation displayed a sigmoidal (Type III) response. At high oyster densities red rock crabs consume more Pacific oysters than green crabs do, due to their reduced handling time, though green crabs consume more Pacific oysters relative to their size than red rock crabs. However, compared to red rock crabs, green crabs consume more oysters at low prey densities, which implies that they have a larger, potentially destabilizing impact on low densities of Pacific oysters. As green crabs continue to spread across the West coast of North America, Pacific oysters will face increased predation pressure. Our results show the advantage of using functional response analysis to compare density dependent predation between an invasive species and a native species to predict the ecological consequences of invasions.

     

Habitat partitioning between prey soldier crab Mictyris brevidactylus and predator fiddler crab Uca perplexa

Interaction and habitat partition between the soldier crab Mictyris brevidactylus (prey) and the fiddler crab Uca perplexa (predator) were examined at a sandy tidal flat on Okinawa Island, Japan, where they co-occur. Both live in dense colonies. When the soldier crabs were released in the densely populated habitat of the fiddler crab, male fiddler crabs, which maintain permanent burrows in hard sediment, preyed on small soldier crabs and repelled large ones. Thus, the fiddler crabs prevented the soldier crabs from trespassing. It was also observed whether soldier crabs burrowed successfully when they were released 1) where soldier crab burrows just under the sand were abundant, 2) in a transition area between the two species, 3) an area without either species, and 4) where artificial tunnels simulated soldier crabs' feeding tunnels were made by piling up sand in the area lacking either species. In contrast to the non-habitat area, many soldier crabs burrowed in the sediment near the release point in the tunnel, transition and artificial tunnel areas. This indicates that the feeding tunnels on the surface attracted other crabs after emergence. When the large male fiddler crabs were transplanted into the artificial burrows made in soft sediment of the soldier crab habitat, all left their artificial burrows by 2 days. In the fiddler crab habitat, however, about one-third of the transplanted male fiddler crabs remained in the artificial burrows after 3 days. The soldier crabs regularly disturb the sediment by the up and down movement of their burrow (small air chamber) between tides. This disturbance probably prevents the fiddler crab from making and occupying permanent burrows. Thus, it appears that these crabs divide the sandy intertidal zone by sediment hardness and exclude each other by different means.     

Ontogenetic niche specialization of the spider crab Libinia ferreirae associated with the medusa Lychnorhiza lucerna

Symbiotic relationships in marine environments are not fixed and can change throughout the animal’s life. This study investigated the ontogeny of symbiosis of the spider crab Libinia ferreirae with the host medusa Lychnorhiza lucerna. We described the type of relationship, the temporal correlation among species, and food habits. More than 50% of the sampled crabs were symbionts, most in early life stages. The highest number of crabs found in a single medusa was 11. Symbiosis was observed throughout most of the year but was more evident in warm periods. The crab has many benefits in this relationship with a medusa. One is the use of food resources captured by the medusa, primarily copepods. Because the crab steals the medusa’s food, it is a kleptoparasitic relationship. There is a niche partition between symbiont and the free-living crabs as they occupy different habitats and use nonoverlapping food resources. Previous research reported that symbiosis first developed during the crab’s last larval phase (megalopa) when crab and medusa are in the same habitat. Observation of the crab’s behavior shows that symbiosis occurs when the crab can grab to the medusa when the host touches the sea bottom. The crab also took advantage of water currents, releasing itself from the substrate and then drifting toward the medusa. The symbiotic relationship that crabs have with the medusa provides then with a nursery, food resources, shelter, dispersion, and decreased competition with free-living adult crabs, all essential for the crab’s survival.