Characterization of the conspecific metamorphic cue for Hemigrapsus sanguineus (De Haan)

The Asian shore crab, Hemigrapsus sanguineus, is one of the most abundant invasive crabs along the east coast of the United States. Larval stages are generally planktonic, but the megalopa stage settles to the substratum near the time of metamorphosis. Reducing the time to metamorphosis may result in higher recruitment and survival. Previous work has shown that a water-soluble cue produced by adult H. sanguineus can induce metamorphosis of conspecific megalopae. Here we report the results of experiments in which megalopae were exposed to cues produced by different life stages of H. sanguineus. We also provide data from experiments that investigated the temporal stability, detection threshold, and chemical classification of the cue. Our results indicate that an active cue is produced by juveniles as well as adults. The cue is proteinaceous and begins to degrade within 2 days of production. The threshold for detection of the cue by megalopae lies between 0.1 and 0.01 µg of protein per ml.     

Adult fiddler crabs Uca pugnax (Smith) enhance sediment-associated cues for molting of conspecific megalopae

Adult-associated chemical cues can stimulate settlement and metamorphosis of invertebrate larvae into habitats with an enhanced likelihood of juvenile and adult survival. For example, sediments from adult fiddler crab habitat stimulate fiddler crab megalopae to metamorphose (molt) sooner than sediments without adult cues. A similar stimulation of molting occurs after exposure to waterborne chemical cues from adult habitats and to exudates and extracts of adult crabs. We tested whether sediments from habitats without adult Uca pugnax (Smith), which do not stimulate molting of their megalopae, could become stimulatory through brief exposure to adult crabs. Sediments were collected from tidal flats at several distances (∼ 1 m, ∼ 50 m, and ∼ 5.4 km) from adult habitats, and incubated for 24 h with or without adult crabs. Molting rates of laboratory-reared megalopae exposed for 48 h to adult-conditioned sediments were compared to those for untreated controls. Sediments collected in or within 1 m of adult habitat elicited the highest molting rates, and natural sediments from 50 m and 5.4 km had little or no effect on molting. However, incubating sediments collected away from adult habitat with adult crabs produced a higher molting response, and the magnitude of the enhancement increased with distance from adult habitat. Results suggest that the chemical cues that adult crabs release are retained by sediments and consequently stimulate molting of megalopae, regardless of the nature of the sediments themselves. Lack of chemical cues may retard colonization of newly created or heavily disturbed habitats that are otherwise suitable settlement and adult habitat.     

Metamorphosis of mangrove crab megalopae, Ucides cordatus (Ocypodidae): Effects of interspecific versus intraspecific settlement cues

It has recently been shown that metamorphosis of Ucides cordatus megalopae is triggered by substrata from the mangrove forest habitat, and, in particular, adult conspecific odours. Here we demonstrate that the gender of the odour-emitting crabs is insignificant for the metamorphic response in this species. We further investigate whether other estuarine crabs (Goniopsis cruentata, Uca spp., and Callinectes danae) also induce settlement and metamorphosis of U. cordatus megalopae. This is of special interest for population recovery in areas hit by lethargic crab disease (LCD), a fungus that selectively kills U. cordatus but not co-occurring species. Ucides megalopae were reared in four treatments with interspecific-conditioned seawater and tested against the effects of conspecific-conditioned seawater (positive control) and pure seawater (negative control). All megalopae in the positive control metamorphosed successfully, while only one (2%) moulted in the negative control, with a delay of 10 days compared with the latest metamorphosis in the former treatment. In seawater conditioned with U. maracoani and C. danae, which occur on sediment banks and in tidal creeks respectively, all larvae died before reaching the juvenile stage. In the treatments with odours of species that share the same mangrove forest microhabitat as U. cordatus, i.e. G. cruentata and a group of five fiddler crab species (mixed-odour treatment), 20 and 10% respectively of the megalopae moulted with a delay of up to 11 days. No specimens metamorphosed after day 39, but megalopae lived up to 93 days. Since only the conspecific- and coexisting-species treatments stimulated development, we hypothesize that Ucides megalopae are able to precisely identify species-habitat-specific settlement cues. This will be investigated in more detail in future studies, which will also test the effects of the odours of the five forest fiddler crab species separately. The impact of the interspecific odour treatments was much smaller than that of the conspecific odours, nevertheless elevated moulting rates of up to 18% relative to seawater may still significantly accelerate the repopulation of U. cordatus in areas lacking conspecifics, e.g. after massive crab mortalities or at first colonization.     

Getting a grip on the intertidal: flow microhabitat and substratum type determine the dislodgement of the crab Pachygrapsus crassipes (Randall) on rocky shores and in estuaries

Hydrodynamic forces can affect survival as well as limit the movement of motile benthic animals. An animal's danger of dislodgement depends on the hydrodynamic forces it experiences in its microhabitat relative to the force required to dislodge it (tenacity) from the substratum. We measured water flow and substratum characteristics in two different habitats of the shore crab Pachygrapsus crassipes: a wave-swept rocky shore and an intertidal mudflat. The maximum water velocities and accelerations in the microhabitats of the crabs at the wave-swept site were three times and two times greater, respectively, than at the mudflat site. In the laboratory, we measured the tenacity of crabs of various sizes on different substrata, and also measured their drag, lift and added-mass coefficients. Using these data, we calculated the flow conditions under which crabs would be overturned or sheared off the substratum in their two habitats. The net horizontal force (drag plus acceleration reaction) required to dislodge a crab on a rugose rock substratum was an order of magnitude greater than on smooth rock and two orders of magnitude greater than on mud. Our calculations indicate that, under non-storm conditions, crabs will not be dislodged from the substratum in either the mudflat or the wave-swept habitat when grasping the substratum with maximum tenacity. Moving crabs have lower tenacity and our calculations predict that hydrodynamic forces will restrict the mobility of large crabs more than that of small ones on smooth, but not on rugose rock.     

Home and Away: Comparisons of Resource Utilization by a Marine Species in Native and Invaded Habitats

The exotic Asian shore crab, Hemigrapsus sanguineus, was recently introduced to the northeastern coast of North America and during the 1990's breeding populations were established throughout southern New England. In 1997–1998, ecological studies of several co-occurring brachyuran crabs were conducted and in native (Tanabe Bay, Japan) and invaded (Long Island Sound, USA) habitats of H. sanguineus. Standardized comparisons of H. sanguineus were made between the 2 habitats using data on crab sizes, utilization of space, and food habits. Results revealed that (1) the resource use of H. sanguineus was quite different from that of other resident species in its invaded habitat, and (2) there were no substantial changes in resource utilization by H. sanguineus after it became established in the invaded habitat (relative to native Tanabe Bay). Differing patterns of resource use by H. sanguineus and other crabs in the invaded habitat, the lack of restriction in resource use by H. sanguineus following its introduction, and the climatological and physical similarities between native and invaded regions likely contributed to the successful invasion of H. sanguineus into rocky intertidal habitats in southern New England.     

Behavioural response thresholds in New Zealand crab megalopae to ambient underwater sound

A small number of studies have demonstrated that settlement stage decapod crustaceans are able to detect and exhibit swimming, settlement and metamorphosis responses to ambient underwater sound emanating from coastal reefs. However, the intensity of the acoustic cue required to initiate the settlement and metamorphosis response, and therefore the potential range over which this acoustic cue may operate, is not known. The current study determined the behavioural response thresholds of four species of New Zealand brachyuran crab megalopae by exposing them to different intensity levels of broadcast reef sound recorded from their preferred settlement habitat and from an unfavourable settlement habitat. Megalopae of the rocky-reef crab, Leptograpsus variegatus, exhibited the lowest behavioural response threshold (highest sensitivity), with a significant reduction in time to metamorphosis (TTM) when exposed to underwater reef sound with an intensity of 90 dB re 1 μPa and greater (100, 126 and 135 dB re 1 μPa). Megalopae of the mud crab, Austrohelice crassa, which settle in soft sediment habitats, exhibited no response to any of the underwater reef sound levels. All reef associated species exposed to sound levels from an unfavourable settlement habitat showed no significant change in TTM, even at intensities that were similar to their preferred reef sound for which reductions in TTM were observed. These results indicated that megalopae were able to discern and respond selectively to habitat-specific acoustic cues. The settlement and metamorphosis behavioural response thresholds to levels of underwater reef sound determined in the current study of four species of crabs, enables preliminary estimation of the spatial range at which an acoustic settlement cue may be operating, from 5 m to 40 km depending on the species. Overall, these results indicate that underwater sound is likely to play a major role in influencing the spatial patterns of settlement of coastal crab species.     

Metamorphosis in a semiterrestrial crab, Sesarma curacaoense: intra- and interspecific settlement cues from adult odors

The larvae of many marine invertebrate species are able to delay their settlement and metamorphosis in the absence of characteristic cues from the adult habitat. This phenomenon was experimentally studied in the megalopa stage of Sesarma curacaoense de Man, 1892, a semiterrestrial grapsid crab that lives in the shallow coastal mangrove habitats in the Caribbean region. Duration of the development and survival to metamorphosis to the first juvenile crab stage were compared between experimental treatments, where the water was conditioned with adult crabs (“adult-conditioned water,” ACW) and control groups reared in filtered seawater. In the experiments with larvae from two different females, development duration was significantly shorter and mortality lower in water conditioned with conspecific adults. In the two control groups, the effects of supply with an artificial substrate (nylon gauze) were tested. This comparison showed that the presence of substrate did not significantly influence the time to metamorphosis, but did reduce the mortality rate. In all later experiments, the megalopae were thus routinely provided with nylon gauze as a substrate. In each of the three subsequent replicate experiments conducted with larvae from different females, survival rate and development time to metamorphosis were compared between one control group and four treatments with ACW. The effectiveness of conspecific (S. curacaoense) adult odors as metamorphosis-stimulating cue was, in these experiments, compared with that of ACW from one congener (S. rectum) and two species belonging to different genera within the Grapsidae (Armases miersii, Chasmagnathus granulata). While the rate of survival showed inconsistent patterns among repeated experiments, the development was consistently fastest with conspecific ACW, followed by ACW from S. rectum, A. miersii and C. granulata. Only the conspecific and congeneric cues had statistically significant effects (i.e. shorter development than in the controls). These response patterns suggest that chemically similar factors (presumably pheromones) are produced by closely related species and, thus, their chemical structure may reflect phylogenetical relationships within a clade.     

Developmental shift in the selective tidal-stream transport behavior of larvae of the fiddler crab Uca minax (LeConte)

Following hatching, larvae of the fiddler crab Uca minax (La Conte) are exported from the adult habitat in estuaries to coastal and shelf waters where they undergo development prior to re-entering estuaries as postlarvae (megalopae). Studies of the spatial distribution of both newly hatched zoeae (Stage I) and megalopae indicate they undergo rhythmic vertical migrations associated with the tides for dispersal and unidirectional transport (selective tidal-stream transport) both within estuaries and between estuaries and the nearshore coastal ocean. We tested the hypothesis that U. minax zoeae possess a circatidal rhythm in vertical migration that facilitates offshore transport in ebb tidal flows, while postlarvae (megalopae) return to estuaries using a similar flood-phased endogenous rhythm. We also determined if the expression of the rhythm was influenced by the salinity conditions zoeae and megalopae experience as they transition between low-salinity regions of estuaries and high-salinity coastal waters. Stage I zoeae were collected by holding ovigerous female crabs in the lab until hatching. Megalopae were collected from the plankton and identified to species using molecular techniques (PCR-RFLP). Under constant laboratory conditions, both zoeae and megalopae exhibited endogenous circatidal rhythms in swimming that matched the principal harmonic constituent of the local tides (12.39 ± 0.07 h; X¯ ± SE). Upward swimming in Stage I zoeae occurred 2.5-4 h after high tide near the time of expected maximum ebb currents in the field. Rhythmic swimming of megalopae occurred slightly earlier in the tide (2.5 ± 0.09 h after high tide; X¯ ± SE) but was not entirely synchronized with flood currents, as expected. Salinity conditions had no apparent effect on the expression or pattern of the rhythms. Results indicate that this circatidal rhythm forms the behavioral basis of selective tidal-stream transport (STST) in early stage U. minax zoeae, but does not undergo a sufficient phase shift to account for vertical distribution patterns exhibited by megalopae in the field.     

Multiple cues from multiple habitats: Effect on metamorphosis of the Florida stone crab, Menippe mercenaria

The Florida stone crab, Menippe mercenaria, is an economically and ecologically important species that ranges from North Carolina throughout the Caribbean and the southeastern Gulf of Mexico. However, there is little known about its early life history stages as compared to other commercially important species in the region. The goal of this research was to examine effects of putative cues on metamorphosis from the megalopa stage to the first juvenile stage. Our study investigated the effect of water-soluble exudates from four substrata, as well as natural biofilms, and exudates from adult stone crabs. In addition, the influence of natural substrata was compared to that of artificial substrata. Adult exudate had no significant effect on metamorphosis, despite a wide range of tested concentrations. In contrast, there was a significant effect on mean time to metamorphosis in experimental groups exposed to multiple cues associated with the brown alga Sargassum fluitans, rubble from stone crab habitat, the eastern oyster Crassostrea virginica, and biofilms associated with the oyster. Furthermore, we provide evidence for metamorphic responses to water-soluble chemical cues, as well as biochemical and physical cues associated with different substrata. Overall results were coherent with the relevant body of previous work on metamorphosis of brachyuran crab larvae and indicate that both physical and chemical cues are important factors in facilitating the settlement and metamorphosis of M. mercenaria larvae in juvenile nursery habitat.     

Effects of substratum and conspecific adults on the metamorphosis of Chasmagnathus granulata (Dana) (Decapoda: Grapsidae) megalopae

The ability of marine invertebrate larvae to delay their metamorphosis in the absence of adequate environmental cues has been reported for numerous sedentary and sessile species. In the present study, the effect of various substrata and the presence of conspecific adults on the metamorphosis of a mobile species, the crab Chasmagnathus granulata, was evaluated. The duration of the megalopa stage in experiments with six different substrata and in the presence or absence of conspecific adults was compared in a laboratory study. In addition, the influence of natural substrata was compared with that of artificial substrata of similar grain size or texture. In a further experiment, the two most effective cues (natural mud and conspecific adults) were tested as single vs. combined factors. Natural mud and unidentified chemical cues from conspecific adults had the strongest accelerating effects on development duration to metamorphosis. With the exception of nylon threads (artificial filamentous substratum), none of the artificial substrata had a significant effect on the duration of the megalopa stage. Simultaneous exposure to natural mud and water containing chemical cues from conspecific adults accelerated metamorphosis more than each of these factors separately. Megalopae that were reared without a substratum (control) delayed their metamorphosis by 29% (about 3 days) compared with those in simultaneous contact with natural mud and rearing water of adult conspecifics. The results indicate that the metamorphosis of the megalopa of C. granulata is influenced by the presence or absence of environmental stimuli that are associated with the preferred adult habitat.     

Location, location, location: finding a suitable home among the noise

While sound is a useful cue for guiding the onshore orientation of larvae because it travels long distances underwater, it also has the potential to convey valuable information about the quality and type of the habitat at the source. Here, we provide, to our knowledge, the first evidence that settlement-stage coastal crab species can interpret and show a strong settlement and metamorphosis response to habitat-related differences in natural underwater sound. Laboratory- and field-based experiments demonstrated that time to metamorphosis in the settlement-stage larvae of common coastal crab species varied in response to different underwater sound signatures produced by different habitat types. The megalopae of five species of both temperate and tropical crabs showed a significant decrease in time to metamorphosis, when exposed to sound from their optimal settlement habitat type compared with other habitat types. These results indicate that sounds emanating from specific underwater habitats may play a major role in determining spatial patterns of recruitment in coastal crab species.     

Larval settlement and metamorphosis in the slipper limpet Crepidula onyx (Sowerby) in response to conspecific cues and the cues from biofilm

In the marine environment, aggregated distribution in the genus Crepidula is a very common phenomenon. Works from Pechenik's group suggested that this is the result of gregarious settlement of larvae in response to cues associated with conspecific adults. In this study, we investigated the existence of larval metamorphic cues associated with adults of C. onyx, a slipper limpet introduced to Hong Kong from the U.S. in the 1970s, through a series of laboratory bioassays. The results showed that derived cues in adult C. onyx were waterborne and the waterborne cues were not derived from bacteria associated with the shell and soft body of the adult Crepidula. The natural biofilm also induced the larval metamorphosis of C. onyx. The cues from the biofilm were associated with the surface of the biofilm and were not waterborne. The aggregated distribution in nature of adult C. onyx may result from a selective larval settlement process. On a small scale in the water column near the conspecific adults, larvae of C. onyx initially detect the waterborne conspecific cues, which then lead to positive downward swimming or passive sinking. This activity increases the chances for larvae to make contact with the biofilm and to be exposed into the higher concentration of waterborne conspecific cues. This may eventually lead to the enhanced larval settlement pattern on or near the conspecific adults.     

The effects of interspecific competition and prey odor on foraging behavior in the rock crab, Cancer irroratus (Say)

The effects of competitor pressure and prey odor on foraging behavior of the rock crab, Cancer irroratus (Say), were investigated. The Jonah crab, Cancer borealis (Stimpson), was chosen as the interspecific competitor because it shares resources with C. irroratus. Four treatments were tested for their effect on foraging: the presence or absence of a competitor and two types of prey odor; body odor (living mussel) and tissue extract (dead mussel tissue). The presence of Jonah crabs did not influence location time, search time, prey size selected, or handling time of the rock crabs. However, rock crabs responded differently to the presence of body odor and tissue extract cues. The presence of extract odor decreased the time to locate prey while increasing the number of prey manipulated and prey size selected. When prey body odor was present, rock crabs displayed less investigative behaviors than in the presence of extract odor, illustrated by reduced location time. Extract odor provided a stronger and more attractive cue than body odor, but increased prey manipulation and search time. Extract odor induced increases in manipulation and searching for prey but canceled out the benefits of decreased location time, resulting in crabs from both treatments displaying similar search times. These elevated behaviors may be associated with foraging for injured and cracked prey or may indicate an area of conspecific feeding.     

Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides

Homarus americanus (Milne-Edwards), the American lobster, is a predator in New England subtidal communities, feeding on ecologically important grazers (sea urchins), mesopredators (crabs), and basal species (mussels). In this study, we provide the first report of adult American lobsters foraging in rocky intertidal habitats during nocturnal high tides. Censuses by SCUBA divers in the low intertidal (Chondrus crispus Stackhouse) zone showed mean densities of 2.2 lobsters/20 m² on nocturnal high tides, with contrasting low densities of 0.18/20 m² during diurnal high tides. Nocturnal high-tide intertidal densities were 62% of those reported in a previous study of lobsters in nearby subtidal rocky areas (Novak, 2004). The average carapace length of lobsters in the intertidal at night was > 50 mm. These lobsters were actively foraging in the intertidal with collected individuals having a mean stomach fullness of 67%. Prey found in the stomach contents primarily consisted of crabs, mussels and snails. Field experiments showed that lobsters rarely fed on medium to large size individuals of the common intertidal snail, Littorina littorea (L.). In contrast, experiments with local crab species demonstrated that lobsters actively and readily prey on Cancer irroratus (Say) and Carcinus maenas (L.), but were significantly less likely to consume Cancer borealis (Stimpson). The abundance of Carcinus maenas and blue mussels (Mytilus edulis L.) in the intertidal zone may explain the upshore movement of lobsters. Since nocturnal migration of Homarus americanus into the intertidal zone has not been documented before, our understanding of the dynamics of New England intertidal communities needs to be expanded to include this predator.     

Conspecific cues affect stage-specific molting frequency, survival, and claw morphology of early juvenile stages of the shore crab Carcinus maenas

Benthic conspecific cues are used by competent larvae of many marine invertebrates to locate and settle on suitable habitat. However, aggregations of conspecifics can generate strong intraspecific competition and inter-cohort cannibalism. We investigated the effects of adult conspecific cues on general fitness parameters of juvenile Carcinus maenas (stages J1–J5), and used geometric morphometrics to investigate patterns of allometric growth indicative of life-history strategies and resource use potential. Cues induced faster metamorphosis and slightly shortened intermolt time in J2 individuals, at the expense of acute mortality in J1 crabs. These effects are cumulative but compensatory processes nullify differences by the end of the experiment. Allometric carapace change toward the adult standard remained unchanged, but conspecific cues induced first a change in size (J1) and then in shape (J5) of claws. In both control and cued juveniles, heterochely was incipient but apparent in J5 crabs. Independently of body side, conspecific cues triggered a very marked increase of the propodus posterior margin, presumably enhancing general strength. Therefore, early benthic stages may grow slightly faster to a size refuge, and develop stronger claws providing competitive advantage for the use of high-value food items when population density-dependent processes are more probable.     

Marine invasive species: validation of citizen science and implications for national monitoring networks

Approximately 1,000 volunteers assessed the presence of invasive (Carcinus maenas and Hemigrapsus sanguineus) and native crabs within the intertidal zone of seven coastal states of the US, from New Jersey to Maine. Identification of crab species and determination of the gender of the observed crabs was documented at all 52 sites across a 725-km coastal transect. Using quantitative measures of accuracy of data collected by citizen scientists, a significant predictor of a volunteer’s ability was determined and eligibility criteria were set. Students in grade three and seven had the ability to differentiate between species of crabs with over 80% and 95% accuracy, respectively. Determination of gender of the crabs was more challenging and accuracy exceeded 80% for seventh grade students, while 95% accuracy was found for students with at least 2 years of university education. We used the data collected by citizen scientists to create a large-scale standardized database of the distribution and abundance of the native and invasive crabs. Hemigrapsus sanguineus dominated the rocky intertidal zone from Sandy Hook, New Jersey to Boston Harbor, Massachusetts while C. maenas dominated the northern extent of the sampled coastline. A citizen scientist of this monitoring network detected a range expansion of H. sanguineus. We identified obstacles to creating a national monitoring network and proposed recommendations that addressed these issues.     

Potential Impacts of a Western Pacific Grapsid Crab on Intertidal Communities of the Northwestern Atlantic Ocean

Population density and size distribution, salinity tolerance, and feeding activity were examined in a western Pacific grapsid crab, Hemigrapsus sanguineus, that was recently introduced to the mid-Atlantic coast of North America. Seasonal abundance on a boulder/cobble shore (Crane Neck Pt.) in central Long Island Sound, New York, USA, during 1997–1998, ranged from 7 to 10 crabs m⁻² averaged over the entire intertidal zone. Crabs occurred throughout the intertidal during summer and fall, but appeared to move from high to low elevations during winter. In laboratory experiments, H. sanguineus tolerated salinity down to 10 ppt for 7 d, but showed significant preference for 20 or 27 ppt over 10 ppt. The crabs readily consumed juvenile snails (Littorina littorea) and mussels (Mytilus edulis), as well as other common species of macroalgae and invertebrates occurring at Crane Neck Pt. High feeding rates and the ability to consume littorine snails up to 13 mm in height and mussels up to 20 mm in length suggest that this nonindigenous species has the potential to significantly affect the structure of rocky intertidal communities in the northwestern Atlantic Ocean; however, rigorous field studies are needed to accurately determine the impact of this recent introduction. This revised version was published online in July 2006 with corrections to the Cover Date.     

Timing of the water-to-land transition and metamorphosis in the land hermit crab Coenobita compressus H. Milne Edwards: evidence that settlement and metamorphosis are de-coupled

After metamorphosing from the last larval stage to the transitional megalopal stage in the marine plankton, the hermit crab Coenobita compressus moves ashore where it undergoes a second metamorphosis to the first juvenile instar on land. In two experiments using laboratory-reared crabs, I moved megalopae from water to land after different amounts of time at this stage and investigated the impact of this manipulation on the timing of and survival through the second metamorphosis. In the Involuntary Settlement experiment, megalopae were moved to land when they were 3, 6, 9, 12, or 15 days old. None of those moved between the ages of 3 and 6 days survived through metamorphosis, but the majority of 9-day-old megalopae survived, as did most 12- and 15-day-old megalopae. This suggests that developmental changes early in the megalopal stage prepare C. compressus for terrestrial life. Once on land, megalopae that had been moved to land at 9 days spent about nine additional days there before metamorphosing, while 12- and 15-day-old megalopae metamorphosed after spending about 5 and 4 days, respectively, on land. In the Voluntary Settlement experiment, megalopae were given access to land when they were 1, 5, 10, or 15 days old, but were not forced to make the transition. Those given access to land after 1 day voluntarily left their dishes for the first time after an average of 7 days in water. Those given access when they were 5 days old remained in water about 4 days longer, while those given access when they were 10 and 15 days old left after less than a day. In both experiments, the timing of metamorphosis relative to settlement (i.e., transition to land) showed that these events are dissociated to a degree and revealed the presence of a metamorphic clock. I discuss why the dissociation of settlement and metamorphosis may have been favored in the land hermit crab and in another anomuran crab.     

Effects of reduced salinities on metamorphosis of a freshwater-tolerant sesarmid crab, Armases roberti: Is upstream migration in the megalopa stage constrained by increasing osmotic stress?

Numerous species of estuarine and freshwater-tolerant crabs show an “export strategy”, i.e. an early larval downstream transport towards coastal marine waters, later zoeal development at higher salinities, and a return of the last larval stage, the megalopa, into estuaries or rivers. The speed and extent of the upstream migration of the megalopa through strong salinity gradients may be constrained by increasing hypo-osmotic stress. In an experimental laboratory study with Armases roberti, a freshwater-inhabiting sesarmid crab from the Caribbean region, we studied in the megalopa stage (after zoeal rearing at 25‰) the tolerance of reduced salinities.In the first experiment, the larvae were exposed directly to various constant salinities (1-25‰). For the second experiment, they were transferred stepwise to strongly diluted media (within 6 days from 25‰ to ≤ 3‰), simulating differential scenarios of upstream migration into brackish or freshwater habitats.When postmoult megalopae were exposed directly to salinities ≤ 3‰, they all died within 24 h. A slightly higher salt concentration (5‰), however, allowed for considerable survival (46%) through metamorphosis to the first juvenile crab stage. In treatments with continuous exposure to 10-15‰, as well as in a control group (25‰), survival to metamorphosis was significantly higher (83-96%), and the average duration of development was shorter compared to 5‰ (12-13 vs. 16 days). In the second experiment, with stepwise salinity reductions, gradual acclimation to decreasing osmotic pressures permitted a successful development to metamorphosis at ≤ 3‰ and even in freshwater (< 0.2‰).This strong physiological adaptability enables the megalopa of A. roberti to cross during its upstream migration, within a short time (6 days), strong osmotic gradients, so that metamorphosis is possible also in freshwater habitats where the conspecific adult crabs live. The speed of migration appears to be limited by physiological constraints related to changes in the capability for osmoregulation occurring during the course of the moulting cycle.     

Comparison of low salinity tolerance in Callinectes sapidus Rathbun and Callinectes similis Williams postlarvae upon entry into an estuary

Planktonic larvae of estuarine crabs are commonly exported to the continental shelf for development and then return to coastal and estuarine areas as postlarvae (megalopae). Megalopae returning to estuaries must be adapted to survive in brackish water whereas those of coastally distributed species should not need such adaptations. We investigated 1) whether megalopae of the estuarine crab Callinectes sapidus and the coastal crab Callinectes similis undergo changes in salinity tolerance upon entry into an estuary and 2) what factors induce those changes. Megalopae were collected at a coastal site and a nearby estuarine site and exposed to a range of salinities (5, 10, 15, 20 and 30) for 6 h. Percent survival was determined after 24 h reintroduction to the collection site water. We also investigated 1) whether increased salinity tolerance was induced by reduced salinity or estuarine chemical cues, 2) the time to acclimation and 3) the salinity necessary for acclimation. C. sapidus megalopae from the estuarine site were more likely to survive exposure to low salinities than those from the coastal site. C. sapidus megalopae from the coastal site exhibited increased survival after acclimation to salinities of 27 and 23 for 12 h. Estuarine chemical cues had no effect on salinity tolerance. C. similis megalopae were less likely to survive at low salinities and did not exhibit an acclimation response upon exposure to reduced salinities. These results suggest that megalopae of C. sapidus are physiologically adapted to recruit to estuaries whereas megalopae of C. similis are unable to acclimate to low salinity conditions.