Entrainment of the circadian rhythm in larval release of the crab Dyspanopeus sayi by temperature cycles

Marine and estuarine crabs brood attached eggs, which hatch synchronously releasing larvae at precise times relative to environmental cycles. The subtidal crab Dyspanopeus sayi has a circadian rhythm, in which larvae are released within the 4-h interval after the time of ambient sunset. Previous studies demonstrated that the rhythm can be entrained by the light:dark cycle. Since subtidal crabs are also exposed to temperature fluctuations, an unstudied question was whether the circadian rhythm could be entrained by the diel temperature cycle. To answer this question, ovigerous D. sayi were entrained in darkness to 2.5, 5, and 10 °C temperature cycles that were reverse in phase from the ambient temperature cycle. After entrainment, larval release times were monitored in constant conditions of temperature and darkness with a time-lapse video system. The effectiveness of a temperature cycle to shift the timing of larval release increased as the magnitude of the temperature cycle increased and as crabs were exposed to increasing numbers of entrainment cycles. However, entrainment to a 10 °C cycle only lasted 2 days in constant conditions. When crabs were entrained to a light:dark vs. a 10 °C temperature cycle, the light:dark cycle was dominant for entrainment. Nevertheless, ovigerous crabs do sense temperature cycles and in areas where daylight is too low for entrainment, temperature cycles can be used to regulate the time of larval release.     

Predatory behavior and preference of a successful invader, the mud crab Dyspanopeus sayi (Panopeidae), on its bivalve prey

Predator-prey relationships between the panopeid crab, Dyspanopeus sayi, and the mytilid, Musculista senhousia, were investigated. Through laboratory experiments, prey-handling behavior, prey size selection, predator foraging behavior and preferences for two types of prey (M. senhousia and the Manila clam Ruditapes philippinarum) were assessed. Handling time differed significantly with respect to the three prey sizes offered (small: 15.0-20.0 mm shell length, SL; medium: 20.1-25.0 mm SL; and large: 25.1-30.0 mm SL); mud crabs were more efficient in predating medium-small than large prey. Although differences in prey profitability were not evident, D. sayi exhibited a marked reluctance to feed on larger-sized prey whilst smaller, more easily predated mussels were available. Size selection may be the result of a mechanical process in which encountered prey are attacked but rejected if they remain unbroken after a certain number of opening attempts. D. sayi exhibited inverse density-dependent foraging. A significant higher mortality of prey was evident at low prey density. Thus, at low predator density, the D. sayi-M. senhousia interaction was a destabilizing type II functional response. Interference responses affected the magnitude of predation intensity by D. sayi on M. senhousia, since as the density of foraging crabs increased, their foraging success fell. At high density (4 crabs tank⁻¹), crabs engaged in a high amount of agonistic activity when encountering a conspecific specimen, greatly diminished prey mortality. Finally, presenting two types of prey, Manila clam juveniles were poorly predated by mud crabs, which focused their predation mostly on M. senhousia. It is hypothesized that, when more accessible prey is available, mud crabs will have a minimal predatory impact on commercial R. philippinarum juvenile stocks.     

Herbivory by crabs and the control of algal epibionts on Caribbean host corals

Summary A short-term experiment was conducted to examine the relationships among the branching coral Porites porites, algal epibionts, and a facultative crab associate Mithrax sculptus in Belize, Central America. Initial field observations suggested that coral colonies supporting resident crabs generally had lower algal cover than colonies without crabs. The hypothesis was tested that Mithrax significantly depresses host coral algal cover and thereby indirectly affects host survivorship and growth. Crab accessibility to an array of coral colonies, similarly covered with algal epibionts, was manipulated in three treatments. Results strongly support the hypothesis, with significant differences in algal cover (primarily Dictyota spp.) noted among treatments after only one month. Caged heads with crabs included and uncaged natural controls allowing crabs free access averaged less than 10% cover, whereas mean algal cover exceeded 75% where crabs were excluded. The uncaged treatment, in which crabs were allowed free access to Porites heads was not significantly different from the crab inclusion treatment. Collectively, these results demonstrate that under natural conditions, crabs can have pronounced effects on host corals by reducing fouling algal epibionts. Furthermore, these facultative coral associates may have more important, albeit localized effects on Caribbean corals than has been suggested previously.     

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

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

Male-produced aggregation pheromone of Carpophilus sayi, a nitidulid vector of oak wilt disease, and pheromonal comparison with Carpophilus lugubris

Abstract 1 Carpophilus sayi, a nitidulid beetle vector of the oak wilt fungus, Ceratocystis fagacearum, was shown to have a male‐produced aggregation pheromone. 2 Six male‐specific chemicals were identified from collections of volatiles. The two major compounds were (2E,4E,6E,8E)‐3,5‐dimethyl‐7‐ethyl‐2,4,6,8‐undecatetraene and (2E,4E,6E,8E)‐3,5,7‐trimethyl‐2,4,6,8‐undecatetraene, in a ratio of 100 : 18. These compounds, in a similar ratio, were previously reported to be the pheromone of Carpophilus lugubris, a closely related species. The four minor C. sayi compounds (less than 4% as abundant as the first) were also alkyl‐branched hydrocarbons and consisted of two additional tetraenes and two trienes. 3 The pheromone of C. lugubris was re‐examined to refine the comparison with C. sayi, and C. lugubris was found to have the same additional, minor tetraenes as C. sayi, but not the trienes. 4 A synthetic mixture of the two major compounds was behaviourally active for both sexes of C. sayi in oak woodlands in Minnesota. The pheromone was tested in combination with fermenting whole wheat bread dough (a potent synergist of nitidulid pheromones). The combination of the 500‐µg pheromone dose and dough attracted at least 30‐fold more C. sayi than either pheromone or dough by itself. The synergized pheromone has potential as a tool for monitoring insect vector activity in an integrated management program for oak wilt. 5 Although C. lugubris was not present at the Minnesota test sites, two other Carpophilus species, Carpophilus brachypterus and Carpophilus corticinus, were clearly cross‐attracted to the synergized pheromone of C. sayi.     

Effect of gastropod shell characteristics and hermit crabs on shell Epifauna

The epifauna on gastropod shells occupied by the hermit crabs Pagurus pollicaris (Say) and P. longicarpus (Say) was examined, as was the utilization of shells by these two hermit crabs. In the study area in Tampa Bay, Florida, shells were not a limiting factor to the hermit crab population, and there apparently was little competition for shells. Interspecific competition for shells was limited because the two hermit crab species differed in size and hence occupied shells of different sizes. The total number and density of most epifaunal species were higher on shells occupied by hermit crabs than on unoccupied shells, possibly because hermit crabs prevent their shells from being buried and hence lengthen the time the epifaunal community can grow and develop. The hermit crab species also appeared to affect the epifaunal community, for the total number and density of most epifaunal species were larger on shells occupied by P. pollicaris than P. longicarpus. With increasing shell size, the populations of most epifaunal species, also were larger but not their density. Least influential in affecting the epifaunal community was the species of shells.     

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

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

Host location and selection by the symbiotic sargassum crab <Emphasis Type="Italic">Portunus sayi</Emphasis>: the role of chemical,visual and tactile cues

The Sargassum community consists of a unique and diverse assemblage of symbiotic fauna critical to pelagic food chains. Associated symbionts presumably have adaptations to assist in finding Sargassum. In situ scattered Sargassum patches accumulate as they are pushed toward the shoreline (via wind, waves, currents or tides) and are frequently less than 1 m apart and in depths of 10 cm or less as the patches approach the shoreline Crabs, and other symbiotic fauna, must relocate to another patch that is seaward in direction or likely perish as their current patch will likely become beached. This study investigated sensory cues used for host location and selection by the Sargassum crab, Portunus sayi. Chemical detection trials were conducted with a two-chamber choice apparatus with Sargassum spp. and Thalassia testudinum as habitat source odors. Visual detection trials (devoid of chemical cues) and habitat selection trials were conducted in which crabs were given a choice between hosts. Results showed that P. sayi responded to chemicals from Sargassum spp. Crabs visually located host habitats but did not visually distinguish between different hosts. In host selection trials, crabs selected Sargassum spp. over artificial Sargassum and T. testudinum. These results suggest that crabs isolated from Sargassum likely use chemoreception; within visual proximity of a potential patch, crabs likely use both chemical and visual information.     

Chromatic Patterns of the Hermit Crab Calcinus tubularis Related to the Occupied Shell

Crustacea are known to develop different chromatic patterns due to many factors. Regarding decapods, chromatism was mainly studied in crabs, while very little is known about chromatic patterns in hermit crabs. Calcinus tubularis is a typical infralittoral rocky bottom hermit crab, studied for different aspects of its biology except chromatic variations. This paper aims at describing the different colour morphologies of C. tubularis, discussing hypothesis of why they develop, and testing if in nature the crab prefers a shell with a chromatic pattern similar to that of its body. One hundred and forty crabs were observed and filmed in the laboratory. They were subdivided into two groups, according to their chromatic pattern: 1) light and 2) dark crabs; the shells they occupied were also subdivided into the two groups of 1) light and 2) dark shells on the basis of the epibionts encrusting them. Observations of 129 crabs suggest that the colour depends neither on depth nor on size, intermoult period, diet, reproductive period but it might be connected to genetic factors and might help crab to camouflage. Camouflage is suggested by the fact that 79.3% of the total examined specimens occupy shells with a chromatic pattern resembling that of the crabs themselves. This phenomenon is significantly more recurrent in females than in males and could help the crabs to be cryptic, first with the occupied shell and secondly with the habitat (rocks encrusted by photophylous algae).     

Shell occupancy of the intertidal hermit crab Clibanarius erythropus (Decapoda,Diogenidae) on São Miguel (Azores)

The intertidal hermit crab Clibanarius erythropus was collected at three sites on São Miguel (Azores) during low spring tides. Shells occupied were identified and measured. Crab sizes ranged from 1.78 to 13.67 mm (cephalothoracic shield length), with an average size of 4.40 ± 1.44 mm. Of the 19 different shells utilised, the most frequent were Littorina striata (23.8%), Nassarius incrassatus (22.5%) and Mitra sp. (22.0%). At Fenais da Luz, L. striata was most frequently occupied, while at Água de Alto it was N. incrassatus and, at Caloura, Mitra sp. shells were most frequently used. Shell selection appears to be determined by respective sizes of hermit crab and shell species. Small size-class crabs occupy more shell species than larger crabs. The smallest crab was found at Fenais da Luz occupying a small Bittium sp., whereas the largest crab was found at Caloura inhabiting Stramonita haemastoma.     

Positive abundance and negative distribution effects of a gastropod on an intertidal hermit crab

Summary Field experiments were used to determine the effect of a common intertidal snail (Nerita funiculata) on the use of space for foraging by the hermit crab Clibanarius digueti. Removals of Nerita resulted in an increased density of foraging Clibanarius, while additions of the gastropod had the opposite effect. The observed negative effect of the gastropod on individual hermit crabs appears to be food-related. Field surveys, however, suggested that the hermit crab population is limited by shell number, rather than food. Because Nerita contributes to the shell resource, its effect on the hermit crab population is positive. Nerita, therefore, has a negative effect on the distribution of foraging hermit crabs, but a positive effect on their abundance. Such decouplings of distribution and abundance effects are rare.     

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

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

Influence of prior experience on shell selection by the white spotwrist hermit crab Pagurus criniticornis (Crustacea: Paguridae)

Individuals of Pagurus criniticornis in a free-choice situation were experimentally tested under different laboratory conditions. In order to assess the effect of recently occupied shells on the size- and type-preference by hermit crabs, individuals were held for 30 days under one of the following two conditions: (1) excess of shells and (2) absence of shells. The crabs were then allowed to select shells from a wide array of empty gastropod shells of the two most-occupied species, as observed previously in the field: Cerithium atratum and Morula nodulosa. Preferred shell type (species) and size (shell aperture width and length) were correlated with hermit-crab size. The crabs showed a strong (100%) preference for C. atratum shells, demonstrating that recent and past experience did not influence either shell-type or shell-size preferences in this pagurid. Handling editor: K. Martens     

Oviposition site preference inNeacoryphus bicrucis (Hemiptera: Lygaeidae): Responses to the density and dispersion of a single host-plant species

Caged females of Neacoryphus bicrucis(Hemiptera: Lygaeidae) were permitted to settle upon and oviposit within a small or large ragwort patch, each of which contained both maturing (= adult food/water source) and dehisced (=oviposition substrate/resource for nymphs) flower heads. The proportion of females settling in the large patch was significantly greater than for the small patch but not significantly greater than expected based on the proportion of maturing flower heads contained in each patch. Also, the number of egg masses laid in each patch was proportional to the number of females that settled there. However, egg mass size was significantly skewed toward larger numbers of eggs in the larger patch. Within the large patch egg mass size was significantly larger where the number of dehisced flower heads was greater and accounts for the between-patch difference in number of eggs per mass. Thus, female settlement does not demonstrate a resource concentration effect but females do appear to evaluate patches after settlement with regard to suitability for oviposition.     

A forced association between the slippersnail Crepidula convexa and the hermit crab Pagurus longicarpus?—possible influence from a third party

The suspension-feeding slippersnail Crepidula convexa is commonly associated with hermit crabs (Pagurus longicarpus) living in periwinkle shells (Littorina littorea) at our study site in Nahant, MA, USA. In 15 field surveys conducted at Nahant in 2000, 2001 and 2003, we found that (1) more than 61.8% of individuals of C. convexa resided on shells occupied by hermit crabs, as opposed to the shells of live periwinkles, empty periwinkle shells or other solid substrates; (2) an average of 8.3% of hermit crabs carried at least one individual of C. convexa; and (3) 39.1-75.0% of hermit crabs carrying C. convexa were carrying “large” individuals (snails with wet weight >10% of the weight of the periwinkle shells they occupied). However, it is unlikely that individuals of C. convexa seek out shells occupied by hermit crabs to colonize, and they showed no preference for empty periwinkle shells over other solid substrates in the laboratory. Moreover, in the laboratory the hermit crabs preferentially occupied intact shells bearing individuals of C. convexa only when the alternatives were shells that had been drilled by naticid snails. Thus, neither party preferentially associates with the other: rather, extensive predation by naticid snails on periwinkles at Nahant appears to limit the availability of suitable shells for the hermit crabs, forcing them to inhabit shells bearing “large” individuals of C. convexa. Individuals of C. convexa may benefit from this inadvertent association with hermit crabs: by facilitating snail dispersal, transport by hermit crabs should reduce the potential for inbreeding, an important consideration for a species that lacks free-living larvae in its life history.     

Are the preference and selection patterns of hermit crabs for gastropod shells species- or site-specific?

Experimental analyses of hermit crabs and their preferences for shells are essential to understand the intrinsic relationship of the crabs' dependence on shells, and may be useful to explain their shell use pattern in nature. The aim of this study was to evaluate the effect of crab species and site on the pattern of shell use, selection, and preference in the south-western Atlantic hermit crabs Pagurus brevidactylus and Pagurus criniticornis, comparing sympatric and allopatric populations. Differently from the traditional approach to evaluate shell preference by simply determining the shell selection pattern (i.e., the number of shells of each type selected), preference was defined (according to [Liszka, D., Underwood, A.J., 1990. An experimental design to determine preferences for gastropod shells by a hermit-crab. J. Exp. Mar. Biol. Ecol., 137(1), 47–62]) by the comparison of the number of crabs changing for a particular shell type when three options were given (Cerithium atratum, Morula nodulosa, and Tegula viridula) with the number of crabs changing for this same type when only this type was offered. The effect of crab species was tested at Cabelo Gordo Beach, where P. brevidactylus was found occupying shells of C. atratum, M. nodulosa, and T. viridula in similar frequencies, whereas P. criniticornis occupied predominantly shells of C. atratum. In laboratory experiments the selection patterns of the two hermit-crab species for these three gastropods were different, with P. criniticornis selecting mainly shells of C. atratum, and P. brevidactylus selecting more shells of M. nodulosa. The shell preference was also dependent on crab species, with P. criniticornis showing a clear preference for shells of C. atratum, whereas P. brevidactylus did not show a preference for any of the tested shells. The effect of site was tested for the two species comparing data from Cabelo Gordo to Preta (P. brevidactylus) and Araçá beaches (P. criniticornis). The pattern of shell use, selection, and preference was demonstrated to be dependent on site only for P. brevidactylus. The results also showed that the shell use pattern of P. criniticornis can be explained by its preference at both sites, whereas for P. brevidactylus it occurred only at Cabelo Gordo, where the absence of preference was correlated with the similar use of the three gastropod species studied. Finally, the results showed that the shell selection pattern cannot be considered as a measure of shell preference, since it overestimates crab selectivity.     

A test of the competitive ability–cold tolerance trade-off hypothesis in seasonally breeding beetles

1. Closely related species that use similar resources often differ in their seasonal patterns of activity, but the factors that limit their distributions across seasons are unknown for most species. One hypothesis to explain seasonal variation in the distributions of species involves a trade-off between competitive ability and cold tolerance, where tolerance to the cold compromises competitive ability in warmer (benign) temperatures, either at the level of the individual or population.
2. We tested both individual-level and population-level mechanisms of this hypothesis in two co-occurring species of temperate burying beetles (Silphidae: Nicrophorus sayi, N. orbicollis) that differ in their seasonal patterns of activity.
3. We measured cold tolerance, breeding activity as a function of temperature, and competitive ability as a function of temperature and season.
4. Consistent with our hypothesis, the mid-season N. orbicollis was less able to function at the cold temperatures that characterise early spring, when the early-season N. sayi is most active. The larger beetle, however, always won one-on-one competitive trials at warm temperatures, regardless of species, inconsistent with an individual-level trade-off. N. orbicollis was usually larger and successful when competing for the same carrion later in the season, mostly because of its larger population size, consistent with a trade-off between competitive ability, and cold tolerance acting at the population level.
5. Our findings suggest that cold temperatures limit the mid-season N. orbicollis from earlier spring emergence, while competitive pressure from the more abundant, larger N. orbicollis constrains the early-season N. sayi from remaining active through the summer.

     

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.     

Use of Hyperspectral Imagery to Assess Cryptic Color Matching in Sargassum Associated Crabs

Mats of the pelagic macroalgae Sargassum represent a complex environment for the study of marine camouflage at the air-sea interface. Endemic organisms have convergently evolved similar colors and patterns, but quantitative assessments of camouflage strategies are lacking. Here, spectral camouflage of two crab species (Portunus sayi and Planes minutus) was assessed using hyperspectral imagery (HSI). Crabs matched Sargassum reflectance across blue and green wavelengths (400–550 nm) and diverged at longer wavelengths. Maximum discrepancy was observed in the far-red (i.e., 675 nm) where Chlorophyll a absorption occurred in Sargassum and not the crabs. In a quantum catch color model, both crabs showed effective color matching against blue/green sensitive dichromat fish, but were still discernible to tetrachromat bird predators that have visual sensitivity to far red wavelengths. The two species showed opposing trends in background matching with relation to body size. Variation in model parameters revealed that discrimination of crab and background was impacted by distance from the predator, and the ratio of cone cell types for bird predators. This is one of the first studies to detail background color matching in this unique, challenging ecosystem at the air-sea interface.     

Food partitioning and spatial subsidy in shelter‐limited fishes inhabiting patchy reefs of Patagonia

The diets of the most conspicuous reef‐fish species from northern Patagonia, the carnivorous species Pseudopercis semifasciata, Acanthistius patachonicus, Pinguipes brasilianus and Sebastes oculatus were studied. Pinguipes brasilianus had the narrowest diet and most specialized feeding strategy, preying mostly on reef‐dwelling organisms such as sea urchins, limpets, bivalves, crabs and polychaetes. The diet of A. patachonicus was characterized by the presence of reef and soft‐bottom benthic organisms, mainly polychaetes, crabs and fishes. Pseudopercis semifasciata showed the broadest spectrum of prey items, preying upon reef, soft‐bottom and transient organism (mainly fishes, cephalopods and crabs). All S. oculatus guts were empty, but stable‐isotope analyses suggested that this species consumed small fishes and crabs. In general, P. brasilianus depended on local prey populations and ate different reef‐dwelling prey than the other species. Pseudopercis semifasciata, A. patachonicus and probably S. oculatus, however, had overlapping trophic niches and consumed resources from adjacent environments. The latter probably reduces the importance of food as a limiting resource for these reef‐fish populations, facilitating their coexistence in spite of their high trophic overlap.