Annual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus

Interannual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus (De Haan) was studied for a 7-yr period (1981 to 1987) in an intertidal mudflat in the estuary of the Tatara-Umi River, Fukuoka, Japan. Density of juveniles fluctuated greatly, and maximum annual density was low in 1981 to 1984 (22.0 to 44.0 m^-2) and very high in 1985 to 1987 (98.5 to 188.6 m^-2). In 1980, recruits may have been very few, though there is no data on or before 1980, because no crab of the 1980 cohort was found during the study period. Density of adult crabs was almost stable (11.0 to 32.6 m^-2), perhaps because of density-dependent mortality. Under the high density conditions prevailing from 1985 to 1987 crabs grew slowly and longevity was shorter. Male chela and female abdomen were dwarfish, and females produced few and small broods. The changes of life-history traits under high density conditions were thought to result from strong intraspecific competition for space.     

Depth as a factor in abundance and size of juvenile penaeid shrimps in the absence of estuaries and marshes

Triweekly trawling around low tide during daylight at 1, 2, 3, and 4 m depths along north Kuwait Bay's extensive intertidal mudflats from 1 April 1985 through 9 December 1986 showed the catchability ofMetapenaeus affinis (H. Milne-Edwards, 1837),M. stebbingi Nobili, 1904,Parapenaeopsis stylifera (H. Milne-Edwards, 1837), andPenaeus semisulcatus De Haan, 1844 differed significantly with depth and season. All species were significantly more catchable in 1 and 2 m depths than in 3 and 4 m depths.M. affinis showed the clearest and most consistent relationship with depth: catchability in 1 m depths was significantly greater than that in 2 m depths (24.9 vs 5.4 per 5-min tow), and catchability in 1 and 2 m depths was significantly greater than that in 3 and 4 m depths (15.2 vs 1.4 per 5-min tow).M. affinis also showed the clearest relationship of increasing size with depth. Shrimp captured in 1 m depths were significantly smaller (~ 14 mm carapace length, CL) than those captured in 2 m depths in the spring (~23 mm CL) and summer (~ 18 mm CL). In the absence of low-salinity waters and intertidal vegetation, it is believed that the edge of the advancing and receding tide over the mudflats, i.e. the shallowest waters, provides juvenile shrimp, particularlyM. affinis, protection from predation.     

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     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Distribution of Juvenile,subadult and adult Scylla serrata (Crustacea: Portunidae) on tidal flats in Australia

In Deception Bay, northern Australia, during 1979–1981, a study was made of the distribution of Scylla serrata (Forskal) in an area having a broad intertidal zone. Juveniles (20 to 99 mm carapace width) were resident in the mangrove zone, remaining there during low tide. The majority of subadult crabs (100 to 149 mm) migrated into the intertidal zone to feed at high tide and retreated to subtidal waters at low tide. Adults (150 mm and larger) were caught mainly subtidally and only small numbers were captured in the intertidal at high tide. Few crabs were captured in the coolest months (May to August). Adults were captured on the flats mainly in the warmest months (January to April), but subadults could be captured over the entire summer (September to March). Juveniles were found in the upper intertidal throughout the year.     

Frequency and timing of extrapair fertilisation in the polyandrous red phalarope (Phalaropus fulicarius)

In sequentially polyandrous bird species, where females mate with different males in succession during a single breeding season, sperm stored by females can occasionally lead to extrapair fertilisations (EPFs) in clutches cared for by the second and subsequent males. Thus, we predicted that in red phalaropes (Phalaropus fulicarius) – a sex-role-reversed, sequentially polyandrous, arctic breeding shorebird – EPFs would occur more frequently in clutches laid later in the breeding season. We used multilocus DNA profiling to examine the frequency and timing of EPFs in a population of red phalaropes breeding in the Canadian high arctic. Using a technique to determine parentage without maternal DNA, we inferred that 6 of 70 chicks in 18 broods resulted from EPFs – one extrapair chick in each of 6 broods. These results were supported by a further analysis using microsatellite DNA. As predicted, broods containing EPFs hatched from clutches laid significantly later in the season than did broods containing no EPFs. The difference in median hatch dates between broods with and without EPFs was 9.0 days, or 38% of the entire egg-laying period in that season. For the whole breeding season, we estimated that 6.5% of chicks were sired by extrapair males, which is similar to extrapair paternity estimates for other sex-role-reversed birds, but relatively low compared to the majority of socially monogamous species studied so far. Received: 24 August 1998 / Received in revised form: 18 January 1999 / Accepted: 24 January 1999     

Competition for inorganic carbon between photosynthesis and calcification in the symbiont-bearing foraminifer Amphistegina lobifera

The feeding behaviour of the subtropical, intertidal crab Gaetice depressus (He Haan) has been investigated. This species was found to be an omnivorous scavenger capable of ingesting both animal and plant tissues which are passed to the mouth using the chelipeds. Furthermore, the mouth-parts themselves are modified and bear elongated setae, notably on the third maxillae. The net-like structures which the setae give rise to can be repeatedly flailed through the seawater in unison enabling the crabs to feed on suspended material. Beating frequency decreased with crab size but rose with increasing suspension concentration. Experiments demonstrated that suspension feeding can contribute significantly to nutrient uptake and direct observations confirmed that the crabs use this method in situ. The crabs are also able to use the setose appendages to sweep deposited material from the underlying substrate towards the mouth, presumably for nutritive purposes. The significance of these different feeding methods is discussed.     

Life-history patterns in two forms of Macrophthalmus japonicus (Crustacea: Brachyura)

Two behaviorally distinct forms — courtship signal — (Form L and V) of Macrophthalmus japonicus (De Haan) are known. Monthly samples of each form were collected from two areas in Japan: Form L from Amakusa, western Kyushu, between April 1985 and February 1986; Form V from Fukuoka, northern Kyushu, between November 1982 and December 1983. Life-history traits were compared between two populations and discussed. Individuals from the Amakusa population grew more slowly and matured at a smaller size than those from the Fukuoka population. Maturation period and longevity did not differ significantly. In the first year of reproduction, females in the Fukuoka population produced more, and larger broods than those in the Amakusa population and crab mortality was higher in every age group. Egg size did not differ significantly. The major environmental factors — air-temperature, nitrogen content of the surface mud and period of aerial exposure were similar in both areas, and it is suggested that the differences in life-history exhibited by these crabs are genotypic, and perhaps related to their different form.     

Feeding and burrowing ecology of two East African mangrove crabs

The behavior and ecology of two mangrove crabs,Sesarma meinerti De Man, 1887 andCardisoma carnifex (Herbst, 1794) were investigated at the beginning of the rainy season (October–November 1988) at Mida Creek, Kenya. Both species occupy upper intertidal levels, above mean high-water neap, and completely overlap in their zonation. Each burrow lasts ca. 3 wk, with no significant difference between the species. Both are more active around dusk and dawn and also follow a similar trend in their foraging activity, but differ in that burrowing is mainly diurnal inC. carnifex and nocturnal inS. meinerti. A hierarchy of food preference, established by offeringC. carnifex andS. meinerti leaves of five mangrove species, proved similar for both, withBruguiera gymnorhiza ranking first andAvicennia marina last, but significant only forC. carnifex. A rough estimate of the amount of litter consumed by these two species and of the soil mixed up by their burrowing activity indicates that they play a role of primary importance in the ecology of East African mangroves.     

Reproductive biology of the common shrimp Crangon crangon (Decapoda: Crangonidae) in the central Irish Sea

Aspects of the reproductive biology of the common shrimp Crangon crangon (L.) were studied in Port Erin Bay, Isle of Man, Irish Sea. Size at sexual maturity was determined from the proportions of ovigerous females and of females with maturing ovaries. The size at which 50% of females are mature is estimated (±95% confidence intervals) as 12.5±0.48 mm carapace length. Based on the proportions of ovigerous females and of mature females, the main breeding season was from January to June. Mean ovarian dry weights indicated two broods (winter and summer), with females bearing winter broods (WB) having higher gonad indices than those with summer broods (SB). WB females with non-eyed eggs and with eyed eggs differed in the regression of ovarian dry weight on carapace length, indicating preparation for laying a second brood. In both broods the moult stages of berried females were related to egg stage. Moulting will occur following the release of the brood. During embryonic development, mean egg length and egg volume were larger in all stages, and the mean dry weight of individual eggs of all stages heavier, in WB than in SB; there was no difference in egg number, however. Consequently, reproductive investment, the proportion of female weight devoted to egg production, was 67% higher in WB (0.20±0.04) than in SB (0.12±0.03). There is a significant effect of egg volume on brood weight, but not on egg number. In both broods, egg number was a negatively allometric function of female body size in non-eyed eggs and an isometric function of female body size in eyed eggs. Brood mortality during incubation was higher in SB (17%) than WB (10%). Differences in the reproductive variables and investment between the two broods of C. crangon are discussed in the light of reproductive strategies and life history.Communicated by J.P. Thorpe, Port Erin     

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.     

Population structure and growth in captivity of the spiny lobsterPanulirus penicillatus from Dahab,Gulf of Aqaba,Red Sea

Panulirus penicillatus (Olivier, 1791) (Decapoda: Palinuridae) is the most common spiny lobster in the Red Sea and is widely distributed in the Indian and Pacific Oceans. Lobsters (n=377) were collected on four occasions during 1986 on the coral reef at Dahab, Sinai, Egypt. Average size of the collected individuals was 70.5±24.6 mm carapace length (CL) for males and 63.2±15.9 mm CL for females. The sex ratio was 1:1.64 males to females. Length increment per molt was inversely correlated with size and ranged from 2.1 mm per molt in the 40 to 50 mm CL size class to less than 1 mm in the 60 to 70 mm CL size class. Average intermolt period was ca. 136 d for all size classes. The relationship between carapace length and body weight was expressed by the equation:W _b=6.43 × 10^–4 × (CL)^2.89.P. penicillatus from Dahab differ in size, sex ratio and growth rate compared to other palinurid populations throughout their range. This might represent the effect of isolation and location at the edge of the geographical range for this species. It may also indicate an adaptation to their unique habitat in the coral reef in comparison to other palinurid species.     

Phenotype-environment matching in the shore crab (Carcinus maenas)

The shore crab (Carcinus maenas) exhibits a range of carapace pattern polymorphisms, but little is known regarding their function or maintenance. If patterns represent some form of crypsis, then associations between carapace colouration and substrate are expected; to determine whether such relationships exist, frequency of crab morphs and quantity of substrate type were measured from fifteen 10×40 m² quadrats at each of three sites along the southern shore of the Firth of Forth, Scotland. Five thousand one hundred and thirty-seven crabs and 3.6 km of line intercept transect data were collected during a 9-week period. Crab abundance, relative frequency of morphs and substrate type varied significantly among the three sites. Plain crabs were strongly associated with macro-algal substrates whereas patterned crabs were associated with mussel beds. This pronounced phenotype-environment matching, as well as various characteristics of the carapace patterns themselves, suggests that patterned crabs are cryptic on polychromatic backgrounds. The frequency of patterned crabs and the percentage of white pigment on the carapace both declined significantly with carapace width. The loss of pattern coincides with an ontogenetic shift in habitat use and we present evidence to suggest that individual crabs lose their pigment, rather than larger patterned crabs being preferentially removed from the population by predators. Throughout their life history, shore crabs encounter high variation in predation, food supply, and physical habitat; to survive they have evolved a strategy that includes elements of pattern polymorphism, crypsis, ontogenetic shifts, and plastic responses.     

The annual reproductive cycles of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) from the South-west coast of India

Employing the gonad index method, the reproductive cycles of three decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards) have been studied. In these crustaceans breeding is not continuous all the year round, but extends over several months of the year with distinct peak periods of gonadal activity. The male and female reproductive cycles are not concurrent. The peak of the reproductive cycle of males occurs slightly earlier in the breeding season than that of females. These studies indicate the possibility of production of successive broods of eggs during the same breeding season. In these species, the low saline conditions of the monsoon period are unfavourable for breeding. The medium and high saline conditions during the post-monsoon and pre-monsoon months, respectively, with plenty of planktonic food for the larvae, seem to be the favourable periods for breeding activity.     

Variation in the carapace pattern of juvenile Carcinus maenas

Juvenile shore crabs (Carcinus meanas L.) are highly variable with respect to carapace pattern; young crabs often have conspicuous patches of white or other pigment. Adult crabs, in contrast, are drab and usually very similar in pattern. It is probable that the patterns serve to make young crabs cryptic on certain backgrounds. Juvenile crabs of carapace widths lying between 5 and 15 mm were collected from a range of British shores (totalling 48). Pattern incidence was found to relate to two factors: the percentage of weed cover (mainly of fucoids) on the shore, and the amount of mud present. Shores of high weed cover, or with much mud present, had low frequencies of conspicuously patterned crabs. The presence of patterned crabs among the juvenile population, and their absence among adults, could result from differential visual predation removing patterned crabs progressively so that only unpatterned crabs would survive to adulthood. Alternatively, individual crabs could lose their conspicuous patterns with age. Evidence is presented in support of the latter hypothesis.     

Influence of predation by a crab on the distribution of the size-groups of three intertidal gastropods in South Australia

Two South Australian rocky intertidal platforms were sampled in April 1981 and December 1982 (Marino Rocks) and in December 1982 (Lady Bay), respectively. Three snail species, Nerita atramentosa Reeve, 1855, Bembicium nanum (Lamarck, 1822) and Austrocochlea concamerata (Wood, 1828), exhibited shore-level size-gradients, with smaller individuals occupying the higher intertidal levels. The reef crab (Ozius truncatus M.-Edwards, 1834), whose distribution overlaps that of the gastropods, is an important predator of these gastropods. The shore-level distribution of shell sizes can be explained by differential selection against smaller individuals by reef crabs. The decreased density of small snails at lower intertidal levels may be the result of either their consumption by reef crabs and/or their movement to higher shore levels where crab predation is less intense.     

Measurement of chlorophyll fluorescence reveals mechanisms for habitat niche separation of the intertidal seagrasses <Emphasis Type="Italic">Thalassia hemprichii</Emphasis> and <Emphasis Type="Italic">Halodule uninervis</Emphasis>

In Taiwan, Thalassia hemprichii dominates the upper intertidal zone, whereas Halodule uninervis occupies the lower intertidal zone. We tested the hypothesis that T. hemprichii is better adapted to high irradiance and more resistant to air exposure than H. uninervis. The photosynthetic efficiency, damage, and extent of recovery were determined by measuring chlorophyll fluorescence using pulse amplitude modulated fluorometry. Both species growing in tidal pools, in response to high irradiance alone, revealed a small depression in maximal quantum yield of photosystem II (Fv/Fm) at noon. The second experiment compared the effect of air exposure alone and the combined effect of air exposure with high irradiance by interposing a shading screen on both species, growing in the intertidal zone over a diurnal cycle. Values of Fv/Fm of both the shaded and irradiated T. hemprichii remained high at low tide. However, H. uninervis exhibited a marked depression following air exposure and a synergistic depression under the combined effect. The experimental manipulations of exposure time demonstrated that the tolerance of T. hemprichii to the combined effect was longer and the recovery from air exposure following re-submersion was better than those of H. uninervis. Both species were more susceptible to the combined effect in the dry season than in the wet season. Our results suggest that air exposure is more important than high irradiance in constraining the distribution of H. uninervis in the upper intertidal zone. This was confirmed by transplantation experiments in which a rapid decline of H. uninervis was observed after transplantation into the upper intertidal zone. In the lower intertidal zone, measurements of the response of the photosynthetic electron transport rate to irradiance demonstrated that the transplanted T. hemprichii exhibited a sun-type response and H. uninervis a shade-type response.     

Age-specific filial cannibalism in a paternal mouthbrooding fish

Entire-brood cannibalism by mouthbrooding males of the cardinal fish Apogon doederleini was investigated in temperate waters of southern Japan during two breeding seasons. The rate of cannibalism was 17–18% in each season and did not differ among age-groups. However, the seasonal pattern of cannibalism differed markedly among age-groups: young (1- and 2-year-old) males frequently cannibalized early broods, especially the first brood, of the season, whereas cannibalism by middle-aged (3- and 4-year-old) and old (5- and 6-year-old) males mainly occurred late in the breeding season. We explain this difference in terms of trade-offs between current and future reproduction. Young males, whose future reproductive success is enhanced by the growth increment, may allocate more time and energy to growth by cannibalizing early broods. In contrast, for older males who have had more breeding cycles and grow little, cannibalism could be a way to reverse the deterioration in their somatic condition that occurs as the breeding season progresses. It is also likely that the current reproductive loss entailed by the cannibalism is effectively compensated by quick re-mating with another female. Received: 24 February 1997 / Accepted after revision: 20 July 1997     

Body shape and burial behavior of the sand crab Emerita analoga (Stimpson 1857) in a reflective to intermediate morphodynamic range of sandy beaches

Sand crabs belonging to the genus Emerita are the dominant species on sandy beaches of the Pacific and Atlantic coasts of America. The success of this genus has been attributed to the great phenotypic plasticity of life history traits that enables them to cope with swash climate variability from beach to beach. The burrowing rate and variation in carapace shape of Emerita analoga from south-central Chile were studied on a series of five sandy beaches with contrasting reflective-intermediate swash conditions (harsher swash to benign swash). Environmental sampling was carried out monthly from September 2009 to November 2010, while biological experiments were conducted in September and October 2010. A combination of field experiments, geometric morphometric tools and multivariate analysis was used to assess the relationship between burrowing behavior and carapace shape with variables of swash climate. Our results showed that (1) body shape of E. analoga is directly affected by the frequency of effluent line crossing, (2) burrowing rate of E. analoga is affected directly by shape variability, (3) burrowing rate is affected directly and indirectly by the swash climate variables: frequency of effluent line crossing and mean grain size of sediment. Thus, the physical conditions in the intertidal zone may be the main environmental constraints determining cephalothorax shape, with a wider cephalothorax on beaches with finer sand and more benign physical dynamics, and a narrower shape on beaches with coarser sand and harsher physical dynamics. The latter phenotype (narrower shape) probably allows compensating the high difficulty of burying in coarser sediments.     

The breeding biology of two sympatric species of tropical intertidal hermit crabs,Clibanarius chapini and C. senegalensis

A one-year study of breeding in the two intertidal hermit crabs Clibanarius chapini Schmitt and C. senegalensis Chevreux and Bouvier showed that both species maintain a high reproductive activity throughout the year. Brief low reproductive activities, found in both species, occurred in different months: C. chapini in November, C. senegalensis in May. While these drops cannot be attributed to any recognisable environmental factor, it is suggested that such pattern may result from competition between the two species. Sex-ratio studies indicated slightly more females than males in both species. The maximal size attained by males of both species is considerably greater than that of the females; in C. senegalensis this is very marked. It is suggested that it may be selectively advantageous for a female to attain a smaller maximal size since her brood must also be accomodated within the shell. As expected for continuously breeding species, recruitment of young into the population is also continuous.