An investigation of genetic population structure in blue crabs,Callinectes sapidus,using nuclear gene sequences

Spatial and temporal genetic heterogeneity in the sequences of nuclear protein-coding genes were investigated in populations of the blue crab, Callinectes sapidus. Haplotype frequencies of these novel markers were determined for adult crabs collected along 300 km in the northern Gulf of Mexico in two different years (2010 and 2011), as well as for megalopal recruits collected in 2010. Tests of genetic differentiation among all locations and between locations spanning known genetic breaks in other species were conducted. In addition, samples from distant locations within the range of C. sapidus were used to assess genetic divergence on a broader geographic scale. Significant between-year differences were found for adults at one location and near significant spatial differentiation was found across northern Gulf of Mexico locations in 2010. These results suggest that although the large population sizes and meroplanktonic life history of blue crabs promote widespread gene flow on a regional scale, genetic composition can change over just one year. Substantial divergence between the northern Gulf of Mexico and Venezuela suggests the possibility that temporal shifts in haplotype frequencies could result from variation in the rate of immigration from genetically distinct source populations. The possible effects of the Deepwater Horizon Oil Spill and attendant fisheries closures during the sampling phase of this study are also considered.     

Comparative vulnerability to predators,and induced defense responses,of eastern oysters Crassostrea virginica and non-native Crassostrea ariakensis oysters in Chesapeake Bay

Management agencies are considering introducing the Suminoe oyster Crassostrea ariakensis into Chesapeake Bay, USA. It is unknown if the growth of feral populations of this non-native oyster would be regulated by the same predators that once controlled the abundance of the native eastern oyster C. virginica. In laboratory studies, we compared the relative susceptibility of juvenile diploids (shell height < 25 mm) of both oyster species to invertebrate predators of eastern oyster juveniles. Predators included four species of mud crabs [Rhithropanopeus harrisii (carapace width 7–11 mm), Eurypanopeus depressus (6–21 mm), Dyspanopeus sayi (8–20 mm), and Panopeus herbstii (9–29 mm)], the blue crab Callinectes sapidus (35–65 mm), and two sizes of polyclad flatworms (Stylochus ellipticus and possibly Euplana gracilis; planar area ≲5 mm² and ∼14 to 88 mm²). All four species of mud crab and the blue crab preyed significantly (ANOVA, P ≤ 0.05) more on C. ariakensis than on C. virginica, but predation by flatworms of both sizes did not differ significantly between oyster species. The greater susceptibility of C. ariakensis to crab predation was likely due to its shell compression strength being 64% lower than that of C. virginica (P = 0.005). To test for predator-induced enhancement of shell strength, we held oysters of both species for 54 days in the presence of, but protected from, C. sapidus and R. harrisii. Crabs were fed congeneric oysters twice weekly within each aquarium. Compared to controls, shell strength of C. virginica exposed to R. harrisii increased significantly (P < 0.043), as did shell strength of both oyster species exposed to C. sapidus (P < 0.01). Despite the changes in shell strength by both oyster species in the presence of C. sapidus, the shell of C. ariakensis remained 57% weaker than C. virginica. We conclude that, because C. ariakensis exposed to predators continued to have a weaker shell relative to C. virginica, the natural suite of crab and flatworm predators in Chesapeake Bay will likely serve to control the abundance of feral C. ariakensis. We caution that the situation in the natural environment may be sufficiently different in some locations that C. ariakensis may be able to compensate for its greater vulnerability to crab predation and hence become a nuisance species.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Harvest in a fluctuating environment and conservative harvest for the Fox surplus production model

Relations between optimal yield and abundance in a fluctuating environment and conditions for a conservative level of harvest were obtained for the Fox surplus production model and compared with those for the logistic surplus production model. Environmental variation was included in the optimization of harvest with the Fox surplus production model to obtain a relation in which the maximum sustainable yield (MSY) and biomass at the MSY varied as the environment varied. The relation can be applied for management of fisheries at the optimum levels in a fluctuating environment. For both models there is only one maximum sustainable yield under equilibrium conditions, but in a variable environment the maximum sustainable yield and optimum biomass and effort vary as the environment varies. The results were applied to the blue crab (Callinectes sapidus) fishery of the Chesapeake Bay. Although several numerical results for the logistic and Fox models were similar, the parameter estimates were different and the Fox model predicted a much larger decrease in population abundance at the MSY. Harvesting at a conservative level with either the Fox model or the logistic model could increase blue crab abundance substantially with little decrease in harvest. At a conservative level of harvest, there is a 20% increase in biomass with a 6% decrease in yield for the logistic model and a 37% increase in biomass with a 9% decrease in yield for the Fox model. Both the Fox and the logistic surplus production models indicate that the blue crab fishery has been consistently over harvested.     

Flood tide transport of blue crab, Callinectes sapidus, postlarvae: behavioral responses to salinity and turbulence

Blue crab (Callinectes sapidus) postlarvae (megalopae) use flood tide transport to move upstream in estuaries during nocturnal flood tides. The megalopae have a endogenous diel rhythm in activity that is inconsistent with this tidally timed behavior. Thus, it is hypothesized that this behavior is regulated by behavioral changes in response to exogenous cues associated with tidal currents. In a laboratory flow tank, blue crab megalopae were exposed to simultaneous changes in salinity and turbulence to simulate tides in an estuary. On simulated flood tides, megalopae ascended upon exposure to a salinity increase, remained swimming during times of high turbulence, and descended at times of low turbulence. Turbulence stimulated swimming for several hours, approximating the duration of tidal currents in estuaries. Swimming was inhibited by decreasing salinity on simulated ebb tides. These results support a model for regulation of flood tide transport by blue crab megalopae as follows: (1) blue crab megalopae are stimulated to swim into the water column by increasing salinity associated with flood tide; (2) megalopae remain swimming during flood tide in response to high levels of turbulence; (3) megalopae descend at the end of flood tide, when current speed and turbulence decline to low levels; and (4) megalopae are inhibited from swimming on ebb tides by the associated salinity decrease. This is the first model for regulation of flood tide transport in a species lacking a tidal rhythm in activity.     

Abundance,breeding and growth of the crab Scylla serrata in two South African estuaries

Populations of the crab Scylla serrata (Forskal) were studied in two South African estuaries from 1971 until 1974. One estuary was open to the sea, the other was closed by a sandbar. Population density in the closed estuary was estimated by means of mark-release-recapture data from tagged crabs. The population of S. serrata was estimated as 1 crab/124m² and the production at 3.4 g/m²/annum. Catch per unit effort data from the closed estuary indicated a natural mortality of 41% in the crabs' second year and 60% in the third. Growth was studied in both estuaries by means of size-frequency analysis and from tagging returns. Growth was rapid in the first 12 to 15 months, when the crabs attained a carapace width of 80 to 160 mm. Thereafter growth slowed, and after 3 years crabs were between 140 and 180 mm in carapace width. Females mated in summer at a carapace width of 103 to 148 mm, males at 141 to 166 mm. After mating, females migrated out of both estuaries into the sea.     

Use of Fully Biodegradable Panels to Reduce Derelict Pot Threats to Marine Fauna

Fishing pots (i.e., traps) are designed to catch fish or crustaceans and are used globally. Lost pots are a concern for a variety of fisheries, and there are reports that 10 – 70% of deployed pots are lost annually. Derelict fishing pots can be a source of mortality for target and bycatch species for several years. Because continual removal of derelict gear can be impractical over large spatial extents, modifications are needed to disarm gear once it is lost. We tested a fully biodegradable panel with a cull or escape ring designed for placement on the sides of a crab pot that completely degrades into environmentally neutral constituents after approximately 1 year. This panel is relatively inexpensive, easy to install, and can be used in multiple fisheries. We used the blue crab (Callinectes sapidus) fishery as a test case because it is a large pot fishery and blue crab pots are similar to traps used in other pot fisheries. We had commercial fishers deploy pots with panels alongside standard pots in Chesapeake Bay (U.S.A.) to assess potential effects of our experimental pots on blue crab catch. We compared the number, biomass, and size of crabs captured between standard and experimental pots and evaluated differences in catch over a crabbing season (March–November) at five locations. There was no evidence that biodegradable panels adversely affected catch. In all locations and time periods, legal catches were comparable in abundance, biomass, and size between experimental and standard pots. Properly designed biodegradable panels appear to be a viable solution to mitigate adverse effects of derelict pots. Uso de Paneles Completamente Biodegradables para Reducir las Amenazas de Vasijas Abandonadas para la Fauna Marina     

Genotypic variation in the parasitic dinoflagellate Hematodinium perezi along the Delmarva Peninsula,Virginia

Hematodinium perezi (genotype III) is a parasitic dinoflagellate that infects blue crabs along the eastern seaboard and Gulf of Mexico, USA. In order to examine the intra-specific genetic variation of this parasite, eleven microsatellite markers from H. perezi (III) were amplified from 227 infected blue crabs collected during 2008–2009 from six sites in Virginia. Simultaneous infections with multiple genetic types in a single-host individual were common and observed in 42 % of the samples. The remaining 58 % of samples had a single allele per locus at all eight polymorphic loci suggesting that the life history stages of the parasite in the host hemolymph are likely haploid. The composition and distribution of multi-locus genotypes (MLG) from samples with infections of a single genetic type indicated high genotypic variation along the Delmarva Peninsula, Virginia, with no evidence of population structure. The lack of linkage disequilibrium combined with the large number of unique MLGs (84 %) is strong evidence for recombination in the life cycle, but the sexual stages remain undetermined. This is the first evidence of ploidy level, infections by multiple genetic types in an individual host animal, high levels of genotypic variation, and sexual reproduction for any species of Hematodinium.     

Natural food,foregut clearance-rate and activity of the crab Scylla serrata

The natural diet, rate of foregut clearance and diurnal activity of the crab Scylla serrata were determined. The gut volume is related to size of crab as gut volume (ml)=0.07e^0.033x , where x=carapace width in millimetres. Fifty per cent of crabs collected in Australia and South Africa contained molluscan remains and about 21% contained crustacean remains — chiefly grapsid crabs. Fish remains were rarely found, and it was concluded that S. serrata does not normally catch mobile forms such as fish and penaeid prawns. Gut clearance of organic tissue was rapid and almost complete after 12 h. Fish bone was retained for a mean time of 2 to 3 days, and shell for 5 to 6 days. Time-lapse photography, using infra-red light, was used to record activity. Visible light flashes reduced activity. S. serrata remained buried during the day, emerging at sunset to spend the night feeding, which occurred intermittently even when unlimited food was available. If no food was present the amount of time spent on the substrate surface was halved.     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Some effects of temperature on the growth and metabolic rate of juvenile blue crabs,Callinectes sapidus,in the laboratory

Juvenile blue crabs, Callinectes sapidus Rathbun, were grown in the laboratory at different temperatures, and metabolic-rate determinations were made. Growth is shown to be dependent upon temperature. Crabs kept at high temperatures (34° and 27°C) grow faster than those kept at lower temperatures (13°, 15°, and 20°C). Increase in size per molt is less at higher temperatures than at lower ones. Mortality is directly proportional to temperature between 13° and 34°C and is very high during ecdysis at elevated temperatures. Metabolic rate increases with temperature, but various degrees of acclimation are seen after 4 weeks exposure. No acclimation of general activity to temperature was found. The findings are applied theoretically to crabs living in the region of heated discharge canals of electrical generators: the motile blue crab could extend its growing season without decreasing size at maturity by active selection of thermal surroundings.In part based on a thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science at the University of Florida, USA.     

Endogenous swimming rhythms in estuarine crab megalopae: implications for flood-tide transport

Up-estuary migration of crab larvae to adult habitats is thought to be accomplished by selective tidal transport in which late-stage larvae enter the water column on flood tides and remain on or near the bottom on ebb tides. This study measured endogenous rhythms in swimming by the last larval stage (megalopa) of blue crabs Callinectes sapidus and fiddler crabs Uca spp. Previous field studies found that megalopae of both species were only abundant in the estuarine water column on nocturnal rising tides. Megalopae were collected from the Newport River Estuary, North Carolina (34°41N; 76°40W) during August–September 1992 and swimming activity was recorded for 4.5 to 7 d under constant conditions with a video system. Rhythms exhibited by both genera in the laboratory were not identical to those recorded in the field. Uca spp. displayed a circatidal rhythm, with maximum swimming occurring near the time of high tide in the field. Rhythm amplitude increased when crushed oyster shells were present, which suggested that megalopae bury or cling to the substrate during quiescent periods. In contrast, C. sapidus had a circadian rhythm in which maximum swimming coincided with the day phase in the field. In most trials, the activity of blue crab megalopae was unrelated to the expected tidal cycle. It was concluded that a tidal rhythm in swimming was the behavioral basis of flood-tide transport for fiddler crab larvae. The endogenous rhythm in blue crabs does not participate in transport, which probably results from behavioral responses to environmental cues associated with flood tide.     

Detection and avoidance of hypoxic water by juvenile Callinectes sapidus and C. similis

Detection and avoidance of hypoxic water by juvenile blue crabs, Callinectes sapidus Rathbun and C. similis Williams were observed under laboratory conditions. Hypoxia avoidance was quantified within an avoidance chamber using response time (time in minutes before a crab's initial entry into the normoxic side of the avoidance chamber after the introduction of hypoxic water into the chamber), total time (amount of time spent by each crab at the hypoxic end of the chamber during the final 50 min of each 60 min treatment), activity (total distance travelled by a crab during a treatment), percent avoidance and preferred oxygen tension as indices of measurement. The mean preferred oxygen tension for C. sapidus was 112 torr oxygen (range = 98 to 125 torr) and for C. similis was 108 torr oxygen (range = 82 to 121 torr). Both species were able to detect hypoxic water and remain at an optimum oxygen tension. When percent avoidance, response time and total time were used as measures of avoidance, C. similis was found to detect and avoid hypoxic water at 0, 25 and 50 torr oxygen tension but not at any higher levels of hypoxia. C. sapidus, on the other hand, did not initiate any significant avoidance behavior even when the water was completely anoxic. Both species of crabs were more active at the higher oxygen tensions than in the more hypoxic levels. Behavioral responses associated with short-term hypoxic exposure included increased activity with the introduction of hypoxic water into the chamber, frequent movement of the eye-stalk, avoidance of hypoxic water by slowly crawling out of the hypoxic zone, restless and erratic movements, and rapid movements of the crabs' antennae. We conclude that C. similis can better detect and avoid hypoxia than C. sapidus.     

Synergistic effects of cadmium and salinity combined with constant and cycling temperatures on the larval development of two estuarine crab species

The developmental stages from megalopa to third crab of the blue crab Callinectes sapidus Rathbun were tested in 12 combinations of cadmium (0, 50, and 150 ppb) and salinity (10, 20, 30, and 40) at 25°C. A reduction in survival and a significant delay in development from megalopa to third crab occurred within each salinity regime in 50 ppb compared with the control. Comparison of the delay in development within each salinity regime revealed that the sublethal effect of cadmium was most pronounced in the salinities normally preferred by C. sapidus. A similar comparison within each cadmium concentration, however, showed that the developmental time from megalopa to third crab was approximately the same irrespective of salinity. The developmental stages from hatch to first crab of the mud-crab Rhithropanopeus harrisii (Gould) were examined in 63 combinations of cadmium (0, 50, and 150 ppb), salinity (10, 20, and 30), constant temperature (20°, 25°, 30°, and 35°C) and cycling temperature (20° to 25°C, 25° to 30°C, and 30° to 35°C). The results indicated that cycling temperatures may have a stimulating effect on survival of the larvae compared to constant temperatures, both in the presence and in the absence of cadmium. Effects of cadmium and salinity and their interaction on the survival of the larvae from zoeae to megalopa were documented at most of the temperatures by analyses of variance. The zoeal larvae were more susceptible to cadmium than the megalopa. Effects of different combinations of cadmium and salinity on the duration of larval development were assessed by a t-test.     

“Natural” agglutinins against sea urchin sperm in the hemolymph of the crab Cardisoma guanhumi

“Natural” agglutinins against sperm have been considered to be broadly specific. However, the clumping of sperm from one species of sea urchin in hemolymph from the crab Cardisoma guanhumi, revealed a narrowly specific agglutinin. Additional testing could establish the taxonomic distribution of the reacting antigen.     

A Contemporary,Sex‐Limited Change in Body Size of an Estuarine Turtle in Response to Commercial Fishing

Abstract: Juvenile growth rate and adult body size are important components of life‐history strategies because of their direct impact on fitness. The diamondback terrapin (Malaclemys terrapin) is a sexually dimorphic, long‐lived turtle inhabiting brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of its range, terrapins face anthropogenically imposed mortality: juveniles of both sexes inadvertently enter commercial crab traps and drown. For adult females, the carapace eventually grows large enough that they cannot enter traps, whereas males almost never reach that critical size. We compared age structure, carapace dimensions, growth curves, and indices of sexual dimorphism for a Chesapeake Bay population of terrapins (where mortality of turtles is high due to crab traps) with contemporary terrapins from Long Island Sound and museum specimens from Chesapeake Bay (neither group subject to commercial crab traps). We also calculated the allochronic and synchronic rates of evolutionary change (haldanes) for males and females to measure the rate of trait change in a population or between populations, respectively. We found a dramatic shift to a younger male age structure, a decrease in the length of time to terminal female carapace size, a 15% increase in female carapace width, and an increase in sexual dimorphism in Chesapeake Bay. In a new twist, our results implicate a fishery in the selective increase in size of a reptilian bycatch species. These sex‐specific changes in life history and demography have implications for population viability that need to be considered when addressing conservation of this threatened turtle.     

Cover Caption

Cover : Diamondback terrapins (Malaclemys terrapin) inhabit brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of the turtle's range, juveniles (pictured) and adult males that inadvertently enter commercial crab traps typically drown, whereas the larger carapace of adult females excludes them from traps. On pages 1268‐1277, Wolak et al. document sex‐speci. c changes in carapace length and width in populations that co‐occur with commercial crab traps. Th ese changes have not been observed in contemporary and historic populations in areas without crab trapping.     

Effect of acanthocephalan parasites on the behaviour and coloration of the mud crab Macrophthalmus hirtipes (Brachyura: Ocypodidae)

In the field, the numbers of cystacanths of the parasitic acanthocephalan Profilicollis spp. harboured by crabs are relatively high and correlate with carapace width. In a field experiment, the responses of crabs to the simulated approach of a bird predator (the parasite's definitive host) was not influenced by the number of acanthocephalans they harboured. Crabs that were exposed at the surface of the sediments during receding high tide, however, tended to harbour more parasites than nearby crabs hidden in burrows. An analysis of colour patterns on the carapace of crabs showed that infection levels did not influence carapace pigmentation, and thus did not affect the conspicuousness of a crab relative to the background environment. However, the likelihood of a male crab winning a ritualized fight against a conspecific in the field was associated with its infection level, but in a way that suggests that this finding is a consequence of pathology rather than an adaptation of the parasite to increase its transmission rate. Although only weak evidence was found indicating that Profilicollis manipulates the behaviour or colour of its host to its own benefit, the high infection levels observed suggest that the crab population acts as a major reservoir for larval stages of this parasite that are infective to birds.     

Relationships between sponges and crabs: patterns of epibiosis on Inachus aguiarii (Decapoda: Majidae)

The association between sponges and the crab Inachus aguiarii Brito Capello, 1876 was studied by analysing the relationships between sponge distributional patterns on the crab carapaces and several morphological and biological crab parameters. Juveniles, mature females and mature males were differentiated on the basis of sex dimorphism and terminal pubertary moult. All three groups were fouled to different degrees by sponges. Percent sponge cover was related to carapace size only in mature males whereas mature females, all but one of which were ovigerous, were extensively covered regard-less of their size. It is proposed that some behavioural patterns unique to females, such as long resting periods in sponge-rich microhabitats, are responsible for these high sponge covers in females. Sponges showed two trends in the colonization of the carapace, leading to either a monopolizing or a sharing of the available carapace surface. The sponge species found on the carapaces studied are not obligatory epibionts of crabs, but are believed to reflect the sponge population characterizing the crab home range.     

Settlement times of blue crab (Callinectes sapidus) megalopae during flood-tide transport

Settlement by blue crab (Callinectes sapidus Rathbun) megalopae on artificial settlement substrates was monitored relative to tidal currents throughout ten nights from July to September 1997 in which the phase relationship between tides and the light dark cycle differed. Most megalopae were in intermolt, and the total number settling to collectors sampled at hourly intervals was greater than totals on collectors immersed all night. Maximum settlement occurred at slack water before ebb tide (SBE), with a smaller peak at slack water before flood tide (SBF). These results support the hypothesis that during flood-tide transport (FTT) blue crab megalopae remain swimming during flood tide at night in response to water turbulence and settle in response to the decline in turbulence occurring near SBE. Settlement peaks near SBF can be explained by a behavioral response of megalopae to increasing salinity at the beginning of flood tide, which results in an ascent response lasting only a few minutes. Depth maintenance in the water column is not maintained at SBF because of low water turbulence. Since light inhibits swimming and upward movement into the water column, settlement, and, presumably, transport were reduced when SBE occurred near the times of sunrise and sunset. Collectively, these results suggest that the phase relationship between the tide and light: dark cycles affects FTT, the timing of settlement, and behaviors associated with habitat selection. Published online: 9 August 2002