Feeding and reproductive behaviour and their effect on catchability of the spanner crab Ranina ranina

The behaviour of 16 adult (8 male and 8 female) spanner crabs (Ranina ranina), collected off southern Queensland, Australia, was monitored continuously in captivity by closed-circuit television for fifteen months from September 1982. Spanner crabs spent most of the time buried in the substrate, emerging mainly only when food became available. They remained emerged for twice as long on feeding days as on days without food. Females responded significantly faster than males to the presence of food. Response was slowest in October and November. There was no correlation between temperature and response time. The average feeding time was 2.0 min (SE=0.12), with no significant difference between males and females. Crabs without food were aggressive towards crabs with food, which sometimes led to fighting and wounding. These interactions also could result in food being transferred from one individual to another. When males interacted, food was transferred more often than when either females, or males and females interacted. Around moulting, male crabs did not feed for 52 d (SE=9.0) and females for 22 d (SE=2.2). This habit would reduce the frequency with which newly moulted crabs are caught in the baited tangle nets used by commercial fishermen. In mating interactions, copulation was always initiated by the males. Males dug up other crabs but, apparently unable to distinguish the sex of these individuals, attempted copulation with either sex. The majority of copulations occurred between midday and midnight and in the period August to December. The frequency of copulations with a female increased 10 d before she extruded eggs, after which it dropped to zero for the following 41 to 50 d. Eggs were carried for 39 to 44 d in the period September to November. The females remained emerged for long periods before extruding their eggs, but the period shortened immediately afterwards. This behaviour would lead to low catchability of ovigerous females. Because females respond more rapidly than males to a food stimulus, they may be more catchable in baited nets than are males. It is concluded that seasonal changes in the behaviour of spanner crabs could affect the number caught by baited tangle nets and may influence the sex ratio in catches.     

Juvenile length-at-age data reveal that spanner crabs (<Emphasis Type="Italic">Ranina ranina</Emphasis>) grow slowly

The spanner crab (Ranina ranina) is a widespread and abundant brachyuran in offshore sand substrata of the Indo-Pacific region. Little is known of this species biology, population dynamics and ecology, despite it being the target of commercial fishing operations in many areas. Previous studies of R. ranina growth using length-frequency analysis of samples collected with commercial fishing gear have derived widely divergent estimates of growth parameters. The estimated time taken to reach 100 mm rostral carapace length (minimum legally exploited size in Queensland, Australia) in those studies has ranged from 1.75 to 8.83 years for females and from 1.08 to 3.58 years for males. Our data show that the commercial fishing apparatus used in those studies is size selective and catches only adult crabs. The resulting size bias in samples collected using that apparatus precludes the application of length-frequency-based techniques to estimate growth parameters from those samples. We devised a new dredge to collect samples of juvenile R. ranina and to calculate juvenile growth rates from modal progression in those samples. We combined those data with estimated mean maximum lengths (L) of 121.7 mm for females and 155.9 mm for males from commercial catch data to model other von Bertalanffy growth parameters using bootstrap methods. Those modelled parameters (K=0.29, T₀=–0.24 for females; K=0.23, T₀=–0.25 for males) indicate that R. ranina grows more slowly than most previous estimates suggest, with females requiring an average of 6.35 years and males 4.31 years to reach 100 mm rostral carapace length. This slow growth is consistent with the slow metabolism of R. ranina, and indicates that this species would be likely to recover slowly from overexploitation.Communicated by G.F. Humphrey, Sydney     

Demography of <Emphasis Type="Italic">Nebalia</Emphasis> sp. (Crustacea: Leptostraca) determined by carrion bait trapping in Lobster Bay,Cape d‘Aguilar Marine Reserve,Hong Kong

The feeding ecology of leptostracans is not well-known. In the present study, species of Nebalia (Crustacea: Leptostraca) were captured every month for 1 year from Lobster Bay, Hong Kong, using baited traps. Captured individuals fell into a size range of 0.36–3.64 mm and 0.95–5.52 mm in carapace and total lengths, respectively. They were divided into different reproductive stages based on the morphology of the second pair of antennae and thoracopods. Sexual dimorphism becomes apparent between 0.7–1.5 mm (CL) and 1.4–3.2 mm (TL). Most individuals collected were sexually differentiated, but immature (79.7% of the captured individuals), followed by post-ovigerous (11.4%) and ovigerous (8.9%) females. Sexually undifferentiated juveniles were rare (3.9%). The sex ratio of the immature individuals approached 1:1 with a propensity towards male dominance. Not one mature male was captured. Although females with setose thoracopod endopods, indicating that they were ovigerous, were captured frequently, only two were carrying eggs. The two brooding females had clutch sizes of 11 and 17 eggs. Alternate peaks in relative abundance of ovigerous and post-ovigerous females were identified, the latter following the former. Reproductive activities occurred mainly in April–May and October–December, indicating that two cohorts are recruited. Catches using baited traps could be explained by the reproductive season of Nebalia sp. High catches were usually associated with the months of highest reproductive activity, low catches with peaks in post-ovigerous females. It is therefore concluded that not all individuals at various reproductive stages are attracted to carrion and hence adopt a scavenging feeding mode. Catches of Nebalia sp. in Hong Kong possibly reflect their annual cycle of reproductive activity     

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.     

The biology of the blue manna crab Portunus pelagicus in an Australian estuary

Various aspects of the biology of the blue manna crab Portunus pelagicus have been investigated in the large Peel-Harvey estuarine system of Western Australia, using samples collected regularly by beach seine, gill net and otter trawl between February 1980 and July 1981. Whereas crabs were widely dispersed throughout Peel Inlet, Harvey Estuary and the saline regions of tributary rivers during the summer and autumn, they were found mainly near the estuary mouth in the winter and spring. Since our data suggest that P. pelagicus has a preference for salinities of 30 to 40, the above changes in distribution are apparently related to the marked seasonal variation in salinity which results from the very seasonal pattern of rainfall. The number of ovigerous crabs in the estuary were greatest in January and February. The mean carapace width and number of eggs of ovigerous females were 110 mm (range 85 to 157 mm) and 509 433 (range 270 183 to 847 980), respectively. P. pelagicus started to reach the minimum legal size for capture (carapace width 127 mm) in the summer when they were approximately I yr old, and left the system in large numbers in the following winter when they were 15 to 20 mo old. These features explain why the fishery for P. pelagicus is highly seasonal, with the vast majority of crabs being taken between January and May. As crabs approached the end of their first year of life, the ratio of females began to exceed that of males, apparently as a result of the movement of males out of the system and legislation against the capture of ovigerous females.     

Offshore spawning by the portunid crabScylla serrata (Crustacea: Decapoda)

The portunid crabScylla serrata (Forskål) is shown to migrate offshore to spawn. Records of 447S. serrata caught as bycatch by trawlers in the tropical waters of northern Australia were analysed with respect to area, depth, distance offshore and month of capture as well as the sex and size of the crabs and whether the females were ovigerous. The crabs were caught mainly in three areas that correspond to the tiger prawn trawl fishery, at between 10 and 60 m depth (mean 28.5 m), 3 to 95 km offshore (mean 17.9 km). Most (87%) of the crabs were captured in October and November, which suggests they move offshore in September and October. No crabs were reported from offshore by February. Over 97% of the crabs caught offshore were female, of these 61.5% were ovigerous. The size range of females (100–109 to 200–209 mm carapace-width size classes) and males (120–129 to 200–209 mm carapace-width size classes) caught was similar. The frequent occurrence in coastal waters of females that are larger than the modal size at spawning, as well as mature females with spent ovaries, suggests that many females return to the coast after spawning. Although some species of portunid crab are euryhaline, mature females of estuarine species migrate to the sea to spawn. The migration byS. serrata described here is far more extensive than would be required to reach sea water salinities; it probably provides a dispersal mechanism for larvae to enable the megalopa stage to recruit to habitats distant from those of the parents.     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Biochemical composition of the deep-sea red crab Chaceon quinquedens (Geryonidae): organic reserves of developing embryos and adults

Deep-sea red crabs Chaceon quinquedens (Smith) were collected in traps at depths of 860 and 1043 m in the northern Gulf of Mexico. Ovigerous crabs were maintained in the laboratory and the developing embryos were sampled every 2 wk until hatching. Proximate analysis (lipid, protein, carbohydrate, and ash) of embryos was performed to determine patterns and rates of organic reserve utilization during embryogenesis. Midgut gland, gonads, and clutch (as appropriate) of adult crabs (males, non-ovigerous females and ovigerous females) were analyzed for the same components as the embryos. Red crab embryos exhibited different patterns of yolk deposition and subsequent depletion of yolk components during embryogenesis. There was a range of lipid to protein (L:P) ratios among the different clutches examined, indicating plasticity in the relative proportions of lipid and protein yolk. The energy used for embryogenesis was estimated by converting the amounts of lipid, protein and carbohydrate in the embryos to their caloric equivalents; final values, taken from 9 mo-old embryos whose siblings were hatching as zoeae, were subtracted from the initial values of sibling embryos sampled at the time of collection (2 to 3 mo old). The amount of energy consumed during embryogenesis in the laboratory was relatively constant (0.12 to 0.13 cal egg^-1). There was considerable variability among the concentrations of organic reserves in the midgut gland of adult crabs and in the ovaries of females. Variations in midgut gland L:P ratios and ovaries were related to the reproductive status of the females, but there were no trends related to depth of capture.     

Mound building in Ilyoplax pingi (Crustacea: Brachyura: Ocypodidae)

The ocypodid crab Ilyoplax pingi, observed in Kanghwa I., Korea in 1992, builds a mound at the burrow entrance, by piling mud dug out from the burrow. The mounds were made by both waving and non-waving males, and by both ovigerous and non-ovigerous females. The burrow diameter at the widest part tended to be larger in crabs with mounds than in crabs without mounds, whereas the burrow depth was not different between them. The mound density increased in the late exposure period, when the crabs' surface activities declined. Mound removal and rebuilding experiments revealed that the presence of the mounds has the effect of keeping neighboring crabs away.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

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.     

Activity pattern and spatial strategy in Pachygrapsus marmoratus (Decapoda: Grapsidae) from Mediterranean and Atlantic shores

The spatial strategies and activity patterns of two populations of Pachygrapsus marmoratus (Fabricius), from Italy and Portugal, were studied in the field. The Mediterranean site was a non-tidal splash zone about 30 cm wide, while the Atlantic site was a 30 m wide intertidal belt. Almost all P. marmoratus of both populations were active on the emerged rocks, rarely entering the sea. The Mediterranean crabs actively fed on the algal turf throughout the 24 h, while in Portugal active crabs were more numerous at night. In the Italian population, all crabs were packed together, although large males had larger activity areas than females and smaller males. In Portugal, the large males were more concentrated in the sublittoral fringe, while both small males and females were confined to the eulittoral and littoral fringe. The Italian P. marmoratus performed feeding loops within areas smaller than 9 m². Although most Portuguese crabs showed the same spatial strategy, some of them performed feeding migrations to the sublittoral fringe, covering distances of up to 20 m. Environmental factors (e.g. temperature), physical factors (e.g. refuge availability) and biotic factors (e.g. competitors and predators) of the different shores probably determine the behavioural plasticity of␣P.␣marmoratus. Received: 1 February 1998 / Accepted: 13 November 1998     

Movement within and between different habitats by the portunid crab Scylla serrata

Tagging experiments were carried out on Scylla serrata in three different types of habitat in Moreton Bay, Queensland, Australia, during 1976–1981. A total of 6 233 crabs were tagged and 1 180 recaptured. Two categories of movement were found, a free ranging type and an offshore migration by females. Crabs in a narrow creek with mangrove-covered banks displayed little movement. In areas with large intertidal flats bare of mangroves, crabs underwent more movement and adults (carapace width 150 mm or greater) and subadults (carapace width 100 to 149 mm) moved similar distances (mean 3.9 km). In an area with direct access to the sea, males and females moved equal distances, but in a long channel behind an island, mean female movement (6.6 km) was significantly more than that of males (mean 3.7 km). The distance between tag and recapture site was not greatly affected by the time at liberty over a period of 1 to 36 wk. Tag recaptures showed an exchange between the populations of a mangrove creek and those in the neighbouring bay. There was very limited exchange between the population in an estuary and the adjacent bay and no exchange was found between neighbouring areas separated by a region of habitat unsuitable for S. serrata. None of the more than 3 000 females captured in the study area was carrying eggs, but two tagged ovigerous females were caught at sea after having moved out of the study area. Eight other females were recaptured in other inshore or estuarine areas 20 to 65 km from their release site.     

Vertical migrations in the tree crab Sesarma leptosoma (Decapoda,Grapsidae)

Sesarma leptosoma, somewhat similar to the Atlantic related species, Aratus pisonii, is an East-African mangrove crab which spends its entire life on the roots and branches of mangrove trees (mostly Rhizophora mucronata, Bruguiera gymnorhiza and Ceriops tagal). S. leptosoma never enters the water, nor does it ever venture onto the free mud surface at low tide. Part of the day and night it remains on the lower parts of the mangrove aerial roots, which are often encrusted with a wet algal mat of Bostrichia spp., searching for food and water. Twice a day, from ca. 06:00 to 08:00 hrs in the morning and 16:00 to 18:00 hrs in the afternoon, many of the crabs migrate as far as the leaves at the top of the tree on which they feed. However, they only spend a brief period among the leaves, from ca. 07:00 to 10:00 and 17:00 to 19:00 hrs, after which time they make their way back towards the roots again in two downward migrations. In the morning, the downward migration brings all the crabs back to the roots, but in the evening not all the crabs take part in the downwards migration and some of them pass the whole night in the tree tops. A comparison of the migration time patterns for two different periods of the year (June–July and November) shows that the number of crabs migrating along the tree trunk is modulated by the spring-neap tidal cycle, while the onset of daily migration seems to be controlled mostly by the light level and/or other climatological cues. The adaptive significance of this migratory behaviour is discussed. Observations reported in our study were made in Mida Creek, Dabaso, Kenya in 1991 and 1992.     

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.     

Growth and longevity of the mud snail Batillaria attramentaria

Crab pots were used to sample a population of Scylla serrata (Forskal) in an estuarine area in Queensland, Australia. Pots were laid 100 m apart at fixed positions for 4 d each month for 1 yr (April 1980–June 1981). Data from recapture of tagged crabs showed that males larger than 140 mm carapace width and females larger than 150 mm had a higher capture probability than did smaller crabs; thus size-frequency distributions based on crab-pot captures are biased. Spacing trials showed that pots positioned 50 m apart fished competitively but that there was no difference in catch between those placed 100 and 200 m apart. Catch distribution indicated that the presence of a crab in a pot reduced the probability of further captures. Temperature and incidence of recently moulted crabs accounted for 66% of variation in monthly catches. Catch-per-unit-effort (CPUE) data can be used as a measure of relative abundance of adults if allowance is made for temperature and the incidence of moulting. The capture-mark-release-recapture method for making population estimates was tested but it is concluded that, because of cost and bias in collecting techniques, it cannot be recommended as a technique for studying S. serrata.     

The breeding cycles of two species of grapsid crabs (Crustacea: Decapoda) from the North Coast of Kyushu,Japan

The incidence of ovigerous females in populations of two grapsid crabs, Hemigrapsus penicillatus (de Haan, 1835) and Sesarma (Parasesarma) pictum (de Haan, 1835) were followed from August 1975 to November 1976. H. penicillatus, which inhabits the lower intertidal region near the mouth of Tatara-Umi Estuary, breeds from March to November. S. pictum, which inhibits crevices and abandoned holes of other species and is abundant at and above the upper intertidal region of the estuary, breeds from May to September. H. penicillatus matures when the female reaches 6 to 7 mm carapace width, whereas S. pictum becomes mature when the carapace width is 12 to 13 mm, although the maximum size attained by females of both species is almost the same. H. penicillatus produces 5 to 6 broods, S. pictum 2 to 3 broods, during a breeding season. The size of a newly laid egg of H. penicillatus is smaller than that of S. pictum. The number of eggs produced by a female H. penicillatus is greater than that of S. pictum during a breeding season. In both species the peak of the breeding season is during summer, at which time the ovarian activity is also apparently accelerated. The major environmental factor which controls the breeding in these crabs appears to be temperature. H. penicillatus is submerged at every high tide, and is relatively inactive from late November to February when the ambient water and air temperatures are rather low. S. pictum is dormant in crevices or understones of the splash zone from November to March. The length of the breeding season of these crabs appears to be inversely proportional to the period of their winter dormancy.     

Distribution of Todarodes pacificus (Cephalopoda: Ommastrephidae) paralarvae near the Kuroshio off southern Kyushu, Japan

The distribution and abundance of Japanese common squid [Todarodes pacificus (Steenstrup, 1880)] paralarvae off southern Kyushu Island near the Kuroshio were examined in relation to water types in the region. Surveys were conducted in 1996 and 1997 using paired, 70 cm diameter Bongo nets. Temperature–salinity plots from 0, 50 and 100 m depths were used to assign sampling stations to three water types: inshore, mixing and Kuroshio. In total, 4103 T. pacificus paralarvae were collected at 59 of 72 stations. Catch densities at positive stations ranged between 8 and 4282 ind./50 m² sea surface area. Catches during both years were highest in mixing waters, followed by inshore and Kuroshio waters. Total catches were highest just north of the oceanographic front at the inshore edge of the Kuroshio and declined with increasing distance away from the front. Catches of paralarvae ≤1.0 mm mantle length also peaked near the front. In both years, ≥85% of all paralarvae and ≥69% of those ≤1.0 mm mantle length occurred less than 25 km from the front. We conclude that most spawning off southwest Kyushu occurs near the frontal zone, where Kuroshio and inshore waters meet. Received: 18 November 1998 / Accepted: 11 May 1999     

Reproductive activities in the sesarmid crab Sesarma intermedia in the coastal and estuarine habitats of Hakata,Japan

Aspects of activity pattern, mating, breeding behaviour and reproductive organs were studied in Sesarma intermedia (de Hann) along the Tatara river estuary, Hakata Bay, western Japan. Activity started from early May after a six-month hibernation period. The daily activity pattern was bimodal with one peak after sunset and another before sunrise. Mating started after emerging from hibernation and was ended by July. Practically no mating was seen in August and thereafter, by which time most of the females were carrying eggs. Larger males mated more frequently with any sized adult female. Copulation seemed to take place both in the burrow and on the surface of natural shelter. Females began to carry eggs from mid June on the estuarine habitat, whereas on the coastal habitat ovigerous females could be seen from mid-May. Multiple ovipositions of eggs were realized. Females were able to oviposit three times without any repeated mating. It took about 30 d to incubate one egg mass and less than one week to spawn again. The release of larvae was not sporadic but rather took place within a limited time interval. Sunset seemed to have a great influence on larvae release along the seashore. Energy allocation for reproductive purposes seemed to start before crabs awoke from the hibernation period. Towards the end of the breeding period there was a drastic decline in ovarian development with a remarkable gain in hepatopancreas, a reservoir for energy.