Factors affecting the interval between clutches in the hermit crab <Emphasis Type="Italic">Pagurus nigrivittatus</Emphasis>

Interval between reproductive events is an important factor for iteroparous animals because it determines the number of clutches throughout life. This study examined whether female size, clutch size, shell size and prenuptial molting affected the clutch interval in the hermit crab Pagurus nigrivittatus. Precopulatory guarding pairs of P. nigrivittatus were sampled in the field and kept in the laboratory until the female extruded eggs. The clutch interval of each female was assessed as one of two types of relatively “short” and “long” intervals by checking whether the guarded female had eggs and/or egg cases from the preceding brood or not when the guarding pair was collected. The clutch interval was longer in females with prenuptial molting than those without molting and these females usually grew larger at the prenuptial molt. This suggests that female P. nigrivittatus with a long interval might allocate energy into growth at the expense of the number of clutches during the current reproductive season. The allocation to growth is theoretically predicted to decrease with female size. Gastropod shell size is also known to affect the reproductive activity in hermit crabs. However, female size did not significantly affect the clutch interval in P. nigrivittatus, and the effect of gastropod shell size on clutch interval was not consistent with previous empirical studies. These results may be caused by differences in the gastropod species of shell occupied by the females of P. nigrivittatus.     

Effects of shell utilization on reproductive patterns in tropical hermit crabs

Dittoral and sublittoral hermit crabs (Coenobita clypeatus, Clibanarius tricolor, Calcinus dibicen, Petrochirus diogenes, Paguristes grayi, Pagurus miamensis, Paguristes tortugae, Paguristes cadenati and Pagurus sp.) were studied on Lighthouse Atoll, Belize. The Clibanarius tricolor population differed markedly in mean clutch size from congeneric populations at temperate latitudes and from conspecific populations in the Florida Keys. These differences can be attributed primarily to variation in shell utilization pattern, overlap with competitors, and breeding season. Regression analyses indicate that shell weight but not internal shell volume affects the clutch size of C. tricolor in Belize, probably due to selection for optimum volume resulting in the use of broken and encrusted shells with high weight: volume ratios. Both egg size and clutch size differed significantly among egg developmental stages in C. tricolor, Calcinus tibicen, and Pagurus miamensis.     

Consequences of shell-species preferences for female reproductive success in the hermit crab Pagurus bernhardus

Littoral hermit crabs, Pagurus bernhardus, show a strong preference for Littorina obtusata shells rather than those of Gibbula species. The fitness consequences, in terms of fecundity, for this shell preference is examined for female crabs. Females in the preferred species produced eggs earlier in the season, produced more eggs in the first brood, and produced a second brood more often than did females in the less preferred species. The smaller brood for Gibbula spp. was not a consequence of egg loss from the pleopods due to an unfavourable shape of shell, but rather reflected lower egg production. It is not clear, however, if this differential reproduction is due to direct costs of carrying an unfavourable shell, i.e. the shell impedes reproduction, or whether crabs compete aggressively for favoured shells so that only crabs of low quality inhabit lowquality shells.     

Endogenous swimming rhythms underlying the spawning migration of the blue crab, Callinectes sapidus: ontogeny and variation with ambient tidal regime

Spawning female blue crabs, Callinectes sapidus, use ebb-tide transport (ETT) to migrate seaward. In estuaries with semi-diurnal tides, ETT in ovigerous blue crabs is driven by a circatidal rhythm in vertical swimming in which crabs ascend into the water column during ebb tide. The ontogeny of this rhythm was examined by monitoring swimming behavior of females before the pubertal molt, females that had recently undergone the molt but had not yet produced a clutch of eggs, and ovigerous females from an estuary with strong semi-diurnal tides. To assess variation in swimming rhythms with ambient tidal regime, swimming rhythms of ovigerous females from semi-diurnal (Beaufort, North Carolina), diurnal (St. Andrew Bay, Florida), and non-tidal (South River, North Carolina) estuaries were compared. Experiments were conducted during the summers of 2006–2008. Female crabs prior to oviposition had variable endogenous swimming rhythms (circadian, circatidal, or circalunidian). Ovigerous females from estuaries with semi-diurnal and diurnal tides had pronounced circatidal or circalunidian rhythms with swimming during the time of ambient ebb tide. Swimming rhythms of several ovigerous crabs switched between circatidal and circalunidian during the ~5-day observation period. Ovigerous crabs from a non-tidal estuary had a circadian rhythm with vertical swimming around the time of sunset. These results suggest that, while endogenous swimming rhythms are present in some female blue crabs prior to oviposition, rapid seaward movement via ETT in tidal estuaries begins following oviposition of the first clutch of eggs.     

Effects of costs and benefits of brood care on filial cannibalism in the sand goby

This study tested experimentally whether clutch size and the cost of care affect filial cannibalism in the sand goby, Pomatoschistus minutus. Evolutionary models of filial cannibalism suggest that egg eating has evolved as a way for the male parent to prolong his breeding season. These models assume that eggs function as an alternative energy source for the constrained parent. I manipulated clutch size by allowing males to mate with either one or two females, representing a small and a large clutch, respectively. The addition of a small male shore crab, a common egg predator, increased the cost of care. I quantified fat reserves as a measure of the condition of guarding males. Males who did not build nests had lower fat reserves than males who built nests, suggesting that males with low energy reserves do not start breeding. Males with small clutches lost their nest to the crab more often than males with large clutches. Neither filial cannibalism nor the amount of eggs eaten were affected by the treatments. Males who consumed eggs had a higher fat percentage than males who did not eat eggs. The result that males with small clutches lost their nests to the crabs supports the idea that eggs are defended only if the benefit from continued care will outweigh the cost and that males therefore are sensitive to the trade-off between present and future reproductive success. Received: 15 May 1997 / Accepted after revision: 15 November 1997     

Male mate choice and male–male competition in the hermit crab Pagurus nigrofascia: importance of female quality

Male mate choice has recently been reported in some animals with male–male competition. In the laboratory, we examined whether males choose their mates based on female quality that was indicated by body size and/or days to prenuptial molt, and the effects of female quality on male–male competition in the hermit crab Pagurus nigrofascia. We collected samples from April to May 2009 at an intertidal shore in Hokkaido, Japan (41°N, 140°E). When a male simultaneously encountered two receptive females in the mate choice experiment, males chose females which require less time to molt. When a male guarding a female with less time to molt was challenged by an intruder, the guarding male defended the female for a longer period and was more likely to win the contest. These results indicate that male P. nigrofascia use time to molt to discriminate between females.     

Female sand gobies gain direct benefits by choosing males with eggs in their nests

In some fish species with paternal care, females prefer to spawn with males whose nests already contain eggs. Several hypotheses have been put forward to explain this behaviour, such as reduced risk of predation or cannibalism (the dilution effect), increased parental investment, and mate copying. This experimental study focuses on female mate choice in the sand goby, Pomatoschistus minutus. Females were found to choose males with eggs in their nests. In addition, hatching success increased with clutch size, mainly because males with larger clutches showed less filial cannibalism. Increased egg survival in large clutches may thus be explained by a combination of the dilution effect and higher parental investment. In another experiment, females did not seem to copy the observed mate choice of other females. In conclusion, female preference for males with eggs in their nests is adaptive, and can be explained by direct benefits, as more surviving offspring are produced. Received: 23 December 1995/Accepted after revision: 11 May 1996     

Descriptions and durations of premolt setal stages in female Dungeness crabs,<Emphasis Type="Italic"> Cancer magister</Emphasis>

For crustaceans with a well-defined annual molting season, such as adult female Dungeness crabs (Cancer magister Dana), setal molt staging can, in principle, be used to predict molting destiny of individual crabs. Size-specific predictions of molting probability could, in turn, be useful for estimating mortalities due to molting. Female Dungeness crabs were collected January–March 1997 at depths of 10–30 m off the California coast, USA (41°N). Intermolt stage A₁–C₄ and premolt stages D₀, D₁, D₁, D₁ and D₂ were described based on features of the branchial epipod. Laboratory experiments suggested that stage D₁ was the earliest stage beyond which eventual molting was inevitable so that molting destiny could be determined. Estimated mean number of days from the beginning of stage D₁ to molting was 85. Estimated individual stage durations, based on laboratory experiments, were 24, 51, –4, 18, and 20 days, respectively, for stages D₀, D₁, D₁, D₁, and >D₂. The estimated –4 days for stage D₁ suggests that this stage must be very brief and raises a question of its utility in designation of molt stages. Molt staging of three field samples, 300–600 crabs in each, collected prior to the 1997 annual molting season, indicated an increase in the size and frequency of crabs staged D₁ or later as the time to molt approached, although crabs of 150 mm carapace width and larger showed few signs of molt preparation. Because the duration of the molting season (approximately 120 days) for adult female C. magister in northern California exceeds the estimated maximum duration of reliable prediction of molting destiny (85 days), it does not appear that molt staging can be used to predict molting destiny in this population. However, the procedures that we have used in this paper for application to female C. magister might be used with success for other crustaceans if the duration of stages D₁ to molting exceeds the duration of the molting season, and furthermore may be used for describing temporal molting trends.Communicated by J.P. Grassle, New Brunswick     

Ecdysial rhythms in megalopae and first instars of the Dungeness crab Cancer magister

Diel molting cycles of megalopae and first instar Dungeness crabs Cancer magister Dana captured in the Grays Harbor estuary (46° 55N; 124° 05 W) in May 1991 were studied under laboratory conditions. Sixtyone percent of the megalopae and 76% of the first instar crabs molted during periods of ambient darkness under a normal light — dark diel regime, and molting pattern was not affected by changes in the photoperiod (24 h daylight or 24 h darkness). Time until metamorphic molt increased as conspecific density increased. Habitat type (shell or mud) did not affect time until molt of megalopae and first instar crabs, nor did it affect daily molting rhythm of first instars. We hypothesize that nightly ecdysis of megalopae and first instar Dungeness crabs and density-dependent molting may be an adaptive response to predation and cannibalism among young-of-the-year.Contribution No. 875 from the School of Fisheries, WH-10, University of Washington, Seattle, Washington     

Obligate commensalism of <Emphasis Type="Italic">Curvemysella paula</Emphasis> (Bivalvia: Galeommatidae) with hermit crabs

Curvemysella paula is a markedly crescent-shaped bivalve that lives inside snail shells occupied by hermit crabs. Here, we describe the unique symbiotic life, growth pattern, and reproductive biology of this bivalve, based on specimens collected from the shallow, muddy bottom of the Seto Inland Sea, Japan. C. paula was found attached to columellae in the siphonal canal, mainly of nassariid snail shells occupied by two types of hermit crabs: Diogenes edwardsii (Diogenidae) and Spiropagurus spiriger (Paguridae). The crescent-shaped shell of C. paula is an adaptation to symbiotic life in the narrow interspace between the snail shell and the hermit-crab abdomen. C. paula is a protandric hermaphrodite. In our samples, each host snail shell harbored one (or rarely a few) large female and several males. All the female bivalves settled on the host shells with their anterior end facing outward and benefited from currents created by the hermit crab when feeding. In the muddy bottom, snail shells are a limited resource for both the hermit crabs and symbiotic bivalves. The bivalves benefit from the mobility of the hermit crabs, which prevent the shells from becoming buried in the mud. C. paula represents the only example of obligate commensalism with hermit crabs found in Bivalvia.     

Genetic differentiation between morphotypes of the marine sponge Suberites ficus (Demospongiae: Hadromerida)

Sponges of three morphotypes of Suberites ficus (Johnston, 1842) were collected during February and March 1985 off the south-west of the Isle of Man, and were compared by using spicule size distributions and genetic allele frequencies of isozyme loci. The populations did not show any significant differences of spicule size or type, but could be easily differentiated into three separate species based on isozyme patterns. Samples of pale orange S. ficus growing on gastropod shells inhabited by hermit crabs (Pagurus spp.) were reproductively isolated from the redorange and the pale yellow colour morphs encrusting the bivalve Chlamys opercularis. These latter two colour morphs were genetically similar, but significant differences were observed at two of the 19 gene loci assayed. All the sponges studied were sympatric, and therefore the genetic differences, indicating reproductive isolation, are strong evidence for separate gene pools and, hence, that they are different species. The genetic identity between the two colour morphs of S. ficus on C. opercularis shells was 0.977, whilst between each of these and S. ficus on hermit crabs it was about 0.65. In all three species genetic variability was high, with mean expected and observed heterozygosity values per locus ranging from 0.17 to 0.36.     

Studies on portunid crabs from the Eastern Pacific. II. Significance of the unusual distribution of Euphylax dovii

Euphylax dovii Stimpson (Brachyura: Portunidae: Podophthalminae), a tropical Eastern Pacific swimming crab, has distinctive morphological adaptations for pelagic existence. Crabs in collections from the open ocean had a sex ratio approximating 1:1, with no crabs bearing eggs. Samples from the continental shelf of Colombia contained thousands of females, mostly ovigerous, but no males. Egg attachment has posed a major problem in the evolution of decapod crustaceans, and the two genera of portunid crabs thus far observed cannot attach eggs unless females can bury partly in soft sediments. This suggests that mated E. dovii females must migrate into shallow shelf waters to encounter sediments necessary for spawning. The high energetic cost of swimming while carrying eggs and the presence of abundant food for larvae are factors favoring residence of females in shelf waters until hatching is complete.     

Utilization of partially predated snail shells by the hermit crab Pagurus longicarpus Say, 1817

The hermit crab Pagurus longicarpus was shown to inhabit shells that were partially predated from intertidal areas of Long Island, New York. Among field collections of P. longicarpus, 2.13% of the hermit crabs (46 of 2155) were found with shells with snail tissue present. Over 90% of these partially predated snail shells were occupied by male hermit crabs. Although hermit crabs were in 8 species of snail shells, only Littorina littorea and Nassarius obsoletus were found occupied by hermit crabs and containing snail tissue. In the laboratory, we found that specimens of the spider crab Libinia emarginata were able to pull off the operculum of snails, leaving damage as found in field collections. In contrast, specimens of P. longicarpus were not able to prey on live, healthy snails. When specimens of P. longicarpus were placed in communal tanks, hermit crabs preferred partially predated snail shells to empty and original shells. However, original shells and empty shells were occupied with more frequency than partially predated shells when crabs were isolated. These findings indicate P. longicarpus actively seeks shells soon after attack and abandonment by snail predators, especially in the presence of competitors.     

Cannibalize or care? The role of perceived paternity in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Parental care is a costly part of reproduction. Hence, natural selection should favor males which avoid caring for unrelated young. However, the decision to abandon or reduce care requires cues which are evaluated to give information on potential reproductive value of the offspring. The prediction that male sand gobies, Pomatoschistus minutus, care for foreign eggs as long as they were spawned in their own nest and at least some of such cues are fulfilled was tested. Egg-guarding males that had recently taken part in a spawning event were given a clutch of eggs that was sired either by themselves or another male, in either their own or another male’s aquarium. Males that had not taken part in a spawning event were used as controls and were given eggs sired by another male. We measured the amount of filial cannibalism and nest building. Control group males did not care for eggs and ate them all before rebuilding the nest. In the other treatments, there were no significant effects of paternity, though males moved to another male’s aquarium increased their clutch area threshold and completely consumed larger clutches than males that were not moved. There was no intermediate response in any treatment in the form of increased partial filial cannibalism or less well-constructed nests. Our results suggest that egg-guarding males cannot distinguish between eggs sired by themselves and those sired by other males but are able to react to cues indicating paternity state. Males do not adopt eggs to attract females in P. minutus.     

Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Epibiotic community of the horseshoe crab Tachypleus gigas

Horseshoe crabs act as moving substrata for simple to complex communities of small marine organisms. Amplexed adult pairs migrate for breeding once every 2 weeks from deep waters towards nearshore waters during highest high tide. Female horseshoe crabs bury themselves to the level of the lateral eyes to deposit eggs while the male crabs fertilize them. Subsequently eggs are buried by the female. Tachypleus gigas (Müller) is the most abundant horseshoe crab species above available along the Orissa coast (India). Adults reach terminal anecdysis once sexually mature and live with their carapace for 4 to 9 years. In spite of this, epibiosis is limited. In the current investigation, differences in the epibiotic community (diatoms and macro-epibionts) present on horseshoe crabs, according to gender, were evaluated, and the macro-epibiont population from different regions of the carapace was mapped. In general, female horseshoe crabs harbored fewer epibionts than the males. Among the diatoms, Navicula spp., Nitzschia spp. and Skeletonema sp. were dominant in both sexes. However, the abundance and diversity of diatoms was greater on the carapaces of male crabs. Among the macro-epibionts, the acorn barnacle (Balanus amphitrite Darwin) and encrusting bryozoan (Membranipora sp.) were the most dominant forms. Barnacles and bryozoans were greater in abundance in the “rough” zone (cardiopthalmic region and anterior region of the opisthosoma). Mapping of the macro-epibionts from different regions of the carapace revealed differential distribution in males and females. Such differentiated distribution of the macro-epibionts can be related to factors such as changing habitat by the horseshoe crabs during breeding, mechanical abrasion and surface availability during mating and nesting periods, requirements of epizootic larvae and surface properties of the carapace (wettability and roughness). In the case of females, mechanical abrasion and surface availability played an important role in the epibiotic community structure and distribution patterns. The surface wettability measurements indicated male carapace to be slightly more hydrophobic than the female carapace. Scanning electron microscopy revealed that the male carapace was comparatively rough compared to the smooth carapace of females. A comparison of surface properties of the carapace indicated that the male carapace is more conducive for epibiosis. Received: 23 August 1999 / Accepted: 25 January 2000     

Influence of extracts of adult crabs on molting of fiddler crab megalopae (<Emphasis Type="Italic">Uca pugnax</Emphasis>)

Larvae of many benthic invertebrates metamorphose in response to habitat cues, which include the presence of adult conspecifics. Prior research showed that fiddler crab [Uca pugnax (Smith)] megalopae advance molting to the first crab stage in seawater in which conspecific adult crabs were maintained. In the present study, extracts of adult crabs were prepared and the specificity and protein content were characterized. U. pugnax megalopae were reared in the laboratory to minimize their prior exposure to potential molting cues. Then they were presented with extract solutions (in 1995, 1996, 1997, and 2000) to determine the specificity of the molting response to extracts of several crab species and the effects of the protein concentration of the extract, age of the megalopae at exposure to extract, and the duration of exposure on the molting response. Megalopae of U. pugnax molted sooner in seawater containing extract from adult conspecifics than in filtered seawater without extract or in seawater containing extract from the congener U. minax. Extract from the mud crab Dyspanopeus sayi retarded molting of U. pugnax megalopae. The stimulatory effect of U. pugnax extract on molting of megalopae was dependent on extract concentration. U. pugnax megalopae did not respond to extract until 6 days after molting, and only 1 day of exposure was sufficient to stimulate molting. The boiled extract was effective after being frozen (–15°C) for 2 years. The preparation of stable extract provides a uniform stimulus for multiple experiments examining the specificity of molting cues for brachyuran crustacean larvae and the onset of receptivity to cues.Communicated by J.P. Grassle, New Brunswick     

Brood parasitism and cooperative nesting in the moorhen,Gallinula chloropus

Summary During a two year study of the moorhen, Gallinula chloropus, a quarter of the nests found had two or more females laying in them. This was the result of two features of the moorhens' breeding biology: (1) Cooperative nesting; two (or more) females were paired to the same male. Both laid in the same nest, the second laid a large clutch synchronously to the first, and both cooperated in parental care. (2) Intraspecific brood parasitism; some females (parasites or dumpers) laid a small number of eggs at random in the nests' of neighbours. Cooperatively nesting females were mother and daughter, while parasite and host were not. Parasitic females laid in their own nest as well, and generally did so after completing a dumping sequence. Parasitic eggs were laid at random during the host's laying and incubation period, and were about a quarter as likely to produce an independent chick as were non-parasitic eggs. Parasitic females would probably have reared more chicks by laying in their own nests straight away, instead of dumping any eggs at all, because there was a strong decline in reproductive success with season. Possible reasons why they did not do so are discussed. Parasitised pairs tended to rear fewer own chicks than non-parasitised pairs. There was no evidence that hosts selectively destroyed parasitic eggs. Four hypotheses that explain why hosts accepted dumped eggs are considered. (1) The host pair were unaware of the dump, (2) the host male had copulated with the dumping female, (3) the dumper was related to one of the host pair, and (4) there was a benefit to the host pair because they gained extra helpers. These hypotheses were not supported by the data, but further tests are needed. As the cost of desertion was greater than the cost of acceptance of parasitic eggs, hosts may have been forced to accept.     

Morphological and genetic evidence for vicariance and refugium in Atlantic and Gulf of Mexico populations of the hermit crab Pagurus longicarpus

The number and wide variety of southeastern United States marine taxa with significant differentiation between Gulf of Mexico and Atlantic Ocean populations suggests that these taxa may have experienced major vicariance events, whereby populations were subdivided by geological or ecological barriers. The present study compared variation in morphology, allozymes, and mtDNA in Gulf of Mexico and western Atlantic populations of the longwrist hermit crab Pagurus longicarpus Say collected during 1997 and 1998. Combined Atlantic populations had significantly fewer denticles on the second segment of the third maxilliped than did Gulf of Mexico populations, and the mean ratio of dactyl length to propodus length was significantly greater in the Atlantic crabs than in the Gulf of Mexico crabs. Allozyme allele frequencies at three loci showed genetic differentiation between a Gulf of Mexico population and two Atlantic populations. Analysis of mtDNA sequence data revealed a clear reciprocal monophyly between Gulf and Atlantic populations, with an estimated divergence age of ~0.6 million years ago. This estimated age of divergence is significantly more recent than an age previously estimated for its congener Pagurus pollicaris (~4 million years ago), suggesting that species with a similar genetic break between Gulf and Atlantic populations may not necessarily share an identical history. Surprisingly, there is evidence of geographic subdivision within Atlantic populations of P. longicarpus along the east coast of North America. This differentiation is especially strong between Nova Scotia and southern populations, suggesting that the Nova Scotia population may represent survivors from a northern refugium during the last glacial maximum.