Significance of mangrove–mudflat boundaries as nursery grounds for the mud crab, Scylla paramamosain

Fisheries managers frequently try to protect juveniles in order to preserve stocks. Juveniles can be protected by either implementing changes designed to avoid catching immature animals (e.g. increasing mesh size or altering fishing techniques) or protecting nursery grounds. To prevent the capture of immature animals, an estimate of size at maturity is required as well as a knowledge of both fishing methods and the exact location of the nursery grounds. Strong demand for juvenile mud crabs to stock aquaculture ponds has resulted in development of fisheries targeting crabs of all sizes from instar 1 to mature individuals. Using five different fishing methods, different stages in the life cycle of Scylla paramamosain were followed for a period of 16 months in an estuarine population in the Mekong Delta, Vietnam. Mangrove habitat utilisation begins when crabs settle out from the plankton at instar 1 [modal internal carapace width (ICW), 0.5 cm] amongst the pneumatophores at the mangrove fringe. Increasingly larger crabs were found deeper into the mangrove but they were still living on the surface (modal ICW size class, 1.5 cm). As their size increases, the crabs either dig burrows (modal ICW size class, 4.5 cm) or they live in the sub-tidal zone, migrating into the mangrove with each tide to feed (modal ICW size class, 4.5 cm). Larger crabs were caught offshore (modal ICW size class, 12.5 cm) where females accounted for 60% of the catch although of these, only 63% were mature. Recruitment of early instars was continuous but peaked in December to February. Subsequent peaks in the catch rates of larger size classes indicated the development of a single cohort with an estimated growth rate of 2.0 cm ICW per month. On the basis of abdominal width, females were estimated to mature at 10.2 cm ICW although at 9.7 cm ICW, 50% of females had disengaged abdomens. Abdominal disengagement occurred in males at the slightly smaller size of 9.1 cm ICW. Allometric relationships between chela height and carapace width suggested 50% of males acquire mature chelae at 10.2 cm ICW. These results demonstrate the close linkage between early life stages of S. paramamosain and certain specific niches within mangrove habitats, with the main adult population found to be living sub-tidally at some distance from the mouth of the estuary. The study also highlights the special importance of the mangrove fringe, the border between the mangrove forest and the sea, an area which is particularly vulnerable to physical and anthropogenic impacts.     

Leaf consumption by <Emphasis Type="Italic">Sesarma plicata</Emphasis> in a mangrove forest at Jiulongjiang Estuary,China

The feeding ecology of Sesarma plicata (Grapsidae: Sesarminae), the most abundant crab species in a mangrove forest dominated by Kandelia candel at Jiulongjiang Estuary, China, was investigated through field and laboratory experiments. Feeding preference and consumption rates were determined on mature, senescent and decomposed leaves of Kandelia candel, Bruguiera gymnorrhiza and Aegiceras corniculatum. In the laboratory, S. plicata preferred leaves of K. candel over those of B. gymnorrhiza and A. corniculatum, and consumed significantly more decomposed leaves than mature and senescent ones, irrespective of crab size. Field experiments with limited power failed to reveal detectable species preferences despite more consumption of K. candel, but decomposed leaves of each species were again preferred. Leaf characteristics associated with preference changed with plant species and leaf state. Low tannins and high water content characterized the preference for decomposed state of leaves. Species preference was significantly and negatively related to crude fibers and C:N ratios for mature leaves, and crude fiber for senescent leaves, but significantly and positively related to water content for decomposed leaves. Leaf consumption rates averaged for all leaf categories from laboratory no-choice feeding experiments were 0.101, 0.055 and 0.017 gDW ind⁻¹ d⁻¹ for large, medium and small crabs, respectively. In this forest, mean density of S. plicata was 20.5 ind m⁻² as assessed by a manual catching method. Leaf litter removal rate during neap tides by sesarmid crabs was about 1.33 gDW m⁻² d⁻¹ in April 2006. The leaves removed by crabs were grazed on the sediment surface or taken into crab burrows, shredded and stored before being eaten.     

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 dynamics and growth of juveniles of the velvet swimming crab Necora puber (Decapoda: Portunidae)

Recruitment variability plays a critical role in determining local population densities of benthic organisms, but extreme vulnerability at the onset of juvenile life is a trait that is largely responsible for population survivorship trends. The aim of the present study was to determine the role of juvenile recruitment in the population structure of Necora puber. Juveniles of N. puber were collected from the lower intertidal of rocky shores of Plymouth Sound (southwest coast of the UK) and monthly size–frequency distribution were used to determine the dynamics and the growth of the population. The parameters of the von Bertalanffy growth function were estimated (K=0.281 year⁻¹; t ₀=0.043; C=0.103; and t _s=0.268) assuming a L _∞=105 mm. Growth was markedly seasonal and present results indicated a slower juvenile growth rate than described previously for N. puber. The recruitment period was extensive and was two times higher in 2001 than in 2000 at the start of the 1+ year, but levelled off at the end of the 1+ year class on three of the four shores studied. Instantaneous mortality as high as 5.1 year⁻¹(99.4% year⁻¹) was observed during the higher recruitment year. Early juvenile mortality appears to be density dependent and a demographic bottleneck appears to limit the number of juveniles on some shores.     

Aerobic metabolic rates of swimming juvenile mako sharks, <Emphasis Type="Italic">Isurus oxyrinchus</Emphasis>

The shortfin mako shark, Isurus oxyrinchus, is a highly streamlined epipelagic predator that has several anatomical and physiological specializations hypothesized to increase aerobic swimming performance. A large swim-tunnel respirometer was used to measure oxygen consumption (MO₂) in juvenile mako sharks (swimming under controlled temperature and flow conditions) to test the hypothesis that the mako shark has an elevated maintenance metabolism when compared to other sharks of similar size swimming at the same water temperature. Specimen collections were conducted off the coast of southern California, USA (32.94°N and 117.37°W) in 2001-2002 at sea-surface temperatures of 16.0–21.0°C. Swimming MO₂ and tail beat frequency (TBF) were measured for nine mako sharks [77–107 cm in total length (TL) and 4.4 to 9.5 kg body mass] at speeds from 28 to 54 cm s⁻¹ (0.27–0.65 TL s⁻¹) and water temperatures of 16.5–19.5°C. Standard metabolic rate (SMR) was estimated from the extrapolation to 0-velocity of the linear regression through the LogMO₂ and swimming speed data. The estimated LogSMR (±SE) for the pooled data was 2.0937 ± 0.058 or 124 mg O₂ kg⁻¹ h⁻¹. The routine metabolic rate (RMR) calculated from seventeen MO₂ measurements from all specimens, at all test speeds was (mean ± SE) 344 ± 22 mg O₂ kg⁻¹h⁻¹ at 0.44 ± 0.03 TL s⁻¹. The maximum metabolic rate (MMR) measured for any one shark in this study was 541 mg O₂ kg⁻¹h⁻¹ at 54 cm s⁻¹ (0.65 TL s⁻¹). The mean (±SE) TBF for 39 observations of steady swimming at all test speeds was 1.00 ± 0.01 Hz, which agrees with field observations of 1.03 ± 0.03 Hz in four undisturbed free-swimming mako sharks observed during the same time period. These findings suggest that the estimate of SMR for juvenile makos is comparable to that recorded for other similar-sized, ram-ventilating shark species (when corrected for differences in experimental temperature). However, the mako RMR and MMR are apparently among the highest measured for any shark species.     

Population ecology of the green/black turtle (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) in Bahía Magdalena,Mexico

The mangrove channels of Bahía Magdalena, Mexico, are important developmental areas for juvenile green, or black turtles (Chelonia mydas), but incidental bycatch and illegal hunting threaten population persistence. We studied size distribution, condition index (CI), growth rates, and mortality of black turtles in Estero Banderitas, the largest mangrove channel in Bahía Magdalena, to supply information for the development of effective conservation strategies. A total of 213 black turtles (including 88 recaptures) were caught in entanglement nets between July 2000 and July 2003. Average yearly catch per unit of effort (CPUE, 1 unit: 100 m of net fishing for 12 h) dropped during the study from 2.19 to 0.76. About 97% of all turtles were considered juveniles, average size was 54.6 ± 9.5 cm. Turtles were significantly smaller at the head of Estero Banderitas than in the central part of the Estero and in the open bay, indicating size-based habitat segregation. Average growth rate was 1.62 cm/year and declined with increasing size. Growth was seasonal and three times higher in summer (0.28 cm/month) than in winter (0.09 cm/month), body CI was also significantly higher during the summer months. A seasonalized von Bertalanffy growth function (VBGF) was used to model growth for the size range studied (43–73 cm SCL), with the parameters: L _∞ = 101 cm SCL; K = 0.04 year⁻¹; t ₀ = 0; C = 0.4 and t _s = 0.75. Growth data indicate that black turtles may spend up to 20 years in Bahía Magdalena before they reach maturity at about 77 cm SCL. The total mortality estimate (Z) from the length converted catch curve was 0.16, corresponding to a yearly survival probability of 0.85.     

Cadmium accumulation in the female shore crab Carcinus maenas during the moult cycle and ovarian maturation

 The influence of moulting and ovarian maturation on cadmium accumulation in the tissues of female shore crabs Carcinus maenas exposed to 1 mg Cd l⁻¹ in the water was investigated. Cadmium accumulation in all tissues examined was markedly increased in crabs in the postmoult stages (A and B) compared to crabs in all other moult stages. During the moult cycle, average cadmium accumulation in the midgut gland ranged from 29 μg Cd g⁻¹ dw at premoult stage (D₂) to 589 μg Cd g⁻¹ dw at postmoult stage (A). Average cadmium concentrations in the haemolymph ranged from 0.56 μg Cd ml⁻¹ at intermoult stage (C₄) to 4.6 μg Cd ml⁻¹ at postmoult stage (A), while the gills accumulated from 103 μg Cd g⁻¹ dw in intermoult stage (C₃) to 352 μg Cd g⁻¹ dw in postmoult stage (A). Cadmium concentration in gills and haemolymph was also significantly higher in crabs in late premoult stage (D₃) compared to C₄-crabs, while midgut gland cadmium concentration remained elevated in C₁- and C₃- intermoult stages relative to C₄. During ovarian maturation the cadmium accumulation in midgut gland, gills, ovaries and haemolymph decreased. Average cadmium concentration in the midgut gland decreased from 63 μg g⁻¹ dw in ovarian Stage I to 19 μg g⁻¹ dw in ovarian Stage VI. The same pattern was observed for gills, haemolymph and ovaries. The present study demonstrates that cadmium accumulation in the female shore crab strongly depends on the physiological status of the animal. A possible association between physiological calcium requirements and cadmium accumulation during moulting is discussed. Received: 20 January 2000 / Accepted: 20 July 2000     

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     

Population dynamics and growth of <Emphasis Type="Italic">Nassarius reticulatus </Emphasis>(Gastropoda: Nassariidae) in Rhosneigr (Anglesey,UK)

Seasonal changes in catch rate, growth and mortality of Nassarius reticulatus from an intertidal lagoon and a wave-exposed beach at Rhosneigr (Anglesey, North Wales, UK) are described. The number of N. reticulatus caught in baited traps from the lagoon was significantly higher (>125 individuals trap⁻¹) during the summer (>18°C), than at <12°C (<65 individuals trap⁻¹), and the numbers caught in the lagoon were an order of magnitude greater than on the beach, >13 individuals trap⁻¹ in July (>16°C), and <5 individuals trap⁻¹ between December and April (<9.5°C). Predictions of shell growth attained by N. reticulatus annually in the lagoon using graphical modal progression analysis (MPA) of length frequency data, were similar to the growth of marked and recaptured lagoon N. reticulatus. Predictions of shell growth using computerised length frequency distribution analysis (LFDA), however, did not reflect the growth as accurately as MPA. Modal progression analysis demonstrated that N. reticulatus from the lagoon achieved a higher asymptotic maximum shell length (L _∞) and a lower growth constant (K) than animals from the beach. Shell growth was seasonal with growth of the lagoon individuals slowing down towards the end of September and resuming in early April, about a month later than the beach individuals. Mortality of N. reticulatus was greater during the summer, and survival was lower in the lagoon than on the beach. Recruitment patterns were similar in the lagoon and on the beach, and MPA and LFDA predicted that larval N. reticulatus settled between late summer and early autumn, with juveniles (7–8.9 mm) appearing in the population the following year, between February and April. Growth of male and female N. reticulatus in the laboratory was similar and was temperature and size dependent. The different growth patterns between N. reticulatus from the two habitats, predicted using MPA, were maintained when individuals were reared under laboratory conditions for ∼6 months; N. reticulatus <21 mm from the beach grew faster than individuals from the lagoon, although N. reticulatus >21 mm from the lagoon grew faster and attained a larger length (26 mm) than individuals from the beach (24 mm). Low food availability did not affect N. reticulatus survival in the laboratory but significantly suppressed shell growth.     

Population structure,mortality and growth of <Emphasis Type="Italic">Pinna nobilis</Emphasis> Linnaeus, 1758 (Mollusca,Bivalvia) at different depths in Moraira bay (Alicante,Western Mediterranean)

An investigation to characterize the causes of Pinna nobilis population structure in Moraira bay (Western Mediterranean) was developed. Individuals of two areas of the same Posidonia meadow, located at different depths (A1, −13 and A2, −6 m), were inventoried, tagged, their positions accurately recorded and monitored from July 1997 to July 2002. On each area, different aspects of population demography were studied (i.e. spatial distribution, size structure, displacement evidences, mortality, growth and shell orientation). A comparison between both groups of individuals was carried out, finding important differences between them. In A1, the individuals were more aggregated and mean and maximum size were higher (A1, 10.3 and A2, 6 individuals/100 m²; A1, x = 47.2 ± 9.9; A2, x = 29.8 ± 7.4 cm, P < 0.001, respectively). In A2, growth rate and mortality were higher, the latter concentrated on the largest individuals, in contrast to A1, where the smallest individuals had the higher mortality rate [A1, L = 56.03(1 − e^−0.17t ); A2, L = 37.59(1 − e^−0.40t ), P < 0.001; mean annual mortality A1: 32 dead individuals out of 135, 23.7% and A2: 16 dead individuals out of 36, 44.4%, and total mortality coefficients (z), z _A1(−30) = 0.28, z _A1(31–45) = 0.05, z _A1(46−) = 0.08; z _A2(−30) = 0.15, z _A2(31–45) = 0.25]. A common shell orientation N–S, coincident with the maximum shore exposure, was observed in A2. Spatial distribution in both areas showed not enough evidence to discard a random distribution of the individuals, despite the greater aggregation on the deeper area (A1) (A1, χ ² = 0.41, df = 3, P > 0.5, A2, χ ² = 0.98, df = 2 and 0.3 < P < 0.5). The obtained results have demonstrated that the depth-related size segregation usually shown by P. nobilis is mainly caused by differences in mortality and growth among individuals located at different depths, rather than by the active displacement of individuals previously reported in the literature. Furthermore, dwarf individuals are observed in shallower levels and as a consequence, the relationship between size and age are not comparable even among groups of individuals inhabiting the same meadow at different depths. The final causes of the differences on mortality and growth are also discussed.     

Influence of cadmium accumulation and dietary status on fatty acid composition in two colour forms of shore crabs, Carcinus maenas

 The effects of cadmium exposure and dietary status on cadmium accumulation, fatty acid (FA) content and profiles were investigated in two colour forms of the shore crab Carcinus maenas. Groups of shore crabs were either starved or fed with blue mussels, Mytilus edulis, during a 40 d exposure period to 2 or 6 μM Cd²⁺ (as CdCl₂). Starved green individuals accumulated more cadmium in haemolymph and hepatopancreas than did red crabs and green crabs fed during the experiments. In the red colour form, no difference in cadmium accumulation was observed between starved and fed individuals. In both colour forms, hepatopancreas contained more FA than gills and muscle. The FAs often present in the largest amounts in the tissues were 16:0, 16:1ω7, 18:1ω7, 18:1ω9, 20:4ω6, 20:5ω3 and 22:6ω3. However, saturated (SAFA) and mono-unsaturated fatty acids (MUFAs) were dominant in hepatopancreas, whereas poly-unsaturated fatty acids (PUFAs) were dominant in gills and muscles. At the beginning of the experiment, the total FA content in the hepatopancreas was 111.6 mg g⁻¹ (dry weight) for red crabs and 78.4 mg g⁻¹ for green shore crabs. During the experiment, however, the FA content decreased in red crabs. This decrease was more pronounced for starved individuals than for fed individuals. Also, the decrease in FA content was more pronounced in crabs exposed to 6 μM cadmium compared to crabs exposed to 2 μM or crabs not exposed to cadmium. No change in FA content was observed in green shore crabs, irrespective of diet and cadmium exposure. For both colour forms, no change in FA content was observed for gills and muscle. In red crabs, a decrease was observed for all FAs in the hepatopancreas. This decrease, however, was more pronounced for SAFAs and MUFAs than for PUFAs, indicating that the metabolism of FAs during starvation and cadmium exposure is selective. The experiments indicate that green colour forms of shore crabs are more tolerant of natural stress such as starvation and anthropogenic stress, e.g. cadmium exposure, than are red colour forms of shore crabs. Received: 23 September 1999 / Accepted: 29 April 2000     

Feeding ecology of the mangrove crab <Emphasis Type="Italic">Ucides cordatus</Emphasis> (Ocypodidae): food choice,food quality and assimilation efficiency

Food preferences, consumption rates and dietary assimilation related to food quality were investigated for the large semi-terrestrial and litter-consuming mangrove crab Ucides cordatus cordatus (Ocypodidae, L. 1763) in northern Brazil. Stomach contents were composed of mangrove leaves (61.2%), unidentified plant material and detritus (28.0%), roots (4.9%), sediment (3.3%), bark (2.5%), and animal material (0.1%). U. cordatus prefers Rhizophora mangle over Avicennia germinans leaves despite a higher nitrogen content, lower carbon to nitrogen (C/N) ratio and lower tannin content of the latter. Also, assimilation rates for senescent R. mangle leaves (C: 79.3%, N: 45.4%) were higher than for A. germinans leaves (C: 40.6%, N: 9.1%). Faeces composition indicates that A. germinans leaves were more difficult to masticate and digest mechanically. The leaf-ageing hypothesis, according to which crabs let leaves age in burrows to gain a more palatable and nutritive food, was rejected for U. cordatus since N content, C/N ratio and the abundance of microorganisms did not differ significantly between senescent leaves and leaves taken from burrows. The low microbial biomass on leaf surfaces and in the sediment indicates its minor importance for the nutrition of U. cordatus. It is concluded that high ingestion and assimilation rates of a R. mangle diet together with the consumption of algae allow for a high intake of C, N, and energy. The data suggest that the digestibility of mangrove leaves by U. cordatus is not hampered by tannins. This may have provided a competitive advantage over other leaf-consuming invertebrates unable to digest mangrove litter with high tannin concentrations. Due to the large stock biomass of U. cordatus in the study area, a great amount of finely fragmented faeces is produced (about 7.1 ton dry matter ha⁻¹ year⁻¹ in a R. mangle forest) which is enriched in C, N and bacterial biomass compared to the sediment. The decomposition of mangrove litter, and thus nutrient and energy transfer into the sediment, is greatly enhanced due to litter processing by U. cordatus.     

Age and growth of the naticid gastropod <Emphasis Type="Italic">Polinices pulchellus</Emphasis> (Gastropoda: Naticidae) based on length frequency analysis and statolith growth rings

In Red Wharf Bay, UK the naticid gastropod, Polinices pulchellus, was more abundant and more highly aggregated during the summer months (June–August 2001) than during the winter (December 2000). Whilst small numbers of juvenile P. pulchellus (4–6 mm shell length) were present throughout the year the population consisted mainly of individuals of 12–14 mm shell length. Juvenile snails grew rapidly in size during the winter and early spring; growth then virtually ceased between May and June, following which there was a further period of rapid growth between August and February. Densities ranged between 57 and 4,073 ha⁻¹ and the largest individual collected during this investigation measured 16.2 mm in shell length. Statoliths from adult P. pulchellus revealed the presence of a settlement ring and two prominent growth rings (rings 1 and 2). A curvilinear relationship exists between statolith diameter and shell length in snails up to 16 mm in length. Settlement rings ranged in diameter from 19.7 to 45.2 μm (mean 29.8 μm; SE=0.41) giving an estimated shell length of the settled juvenile of 1.1 mm. The diameter of ring 1 and ring 2 were significantly correlated indicating that rapid growth during the first year is maintained during year 2. Shell lengths estimated from the diameters of the prominent statolith rings and those obtained from length frequency data analysis (LFDA), were broadly congruent strongly suggesting an annual periodicity to the statolith rings. The largest snails (>15 mm) present within this population were estimated to be between 2 and 3 years old. Von Bertallanfy seasonal growth curves obtained from the LFDA predicted values of L∞, K and t ₀ of 14.32 mm, 1.54 and −0.14 years, respectively, suggesting that P. pulchellus rapidly attains its maximum asymptotic size.     

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     

UV- and salinity-induced oxidative effects in the marine diatom <Emphasis Type="Italic">Cylindrotheca closterium</Emphasis> during simulated emersion

The diatom Cylindrotheca closterium was exposed to transient light- and osmotic conditions as occur during its tidal emersion. The objective was to analyze how this simulated emersion contributes to the production of active oxygen species (AOS) and via this, to oxidative cell damage. Light- and salinity conditions were varied in factorial combination: low light (no UVB) or high light (unweighted UVB-dose rates of respectively 0.01; 0.07; 0.24; 1.03 W m⁻²) at normal (30 psu) or high salinity (60 psu). UVB (0.01–0.24 W m⁻²) and high salinity had a significant, negative effect on the photosynthetic efficiencies ΔF/F _m’ (steady-state quantum yield) and F _v/F _m (maximum yield). UVB at 1.03 W m⁻² (15 kJ m⁻² d⁻¹) almost arrested electron transport. At ecologically relevant UVB levels, i.e. below 0.24 W m⁻² (≈3.4 kJ m⁻² d⁻¹) with UVB:PAR<0.4:100 (PAR photosynthetically active radiation) only dynamic photoinhibition was observed (protection via heat dissipation). Non-photochemical quenching was positively correlated with the de-epoxidation of diadinoxanthin (DD) to diatoxanthin (DT). A decreasing ratio DT/(DD+DT) after 4 h of UVB at >0.07 W m⁻² and at 60 psu indicated a reversal of the diatom xanthophyll cycle (diminished photoprotection) which may be caused by an enhanced AOS production. Oxidative stress and -damage to C. closterium cells were assessed applying fluorescent indicator dyes, via confocal microscopy and quantitative image analysis. AOS production rates (cellular DCF fluorescence) were stimulated by UV, and were ~50% higher at 60 psu. AOS production decreased with an increasing pre-exposure (0–4 h) to normal UVB (0.24 W m⁻²), which indicated a stimulation of the antioxidative defence. Non-protein thiols (indicator CMF) and glutathione pools (HPLC-analyzed) decreased with UVB-dose rates (0.01–0.24 W m⁻²), most likely due to AOS-mediated thiol oxidation. Hypersalinity (60 psu) and UVB (0.01–0.24 W m⁻²) caused membrane depolarization (dye DIBAC₄(3)) and phospholipid hydrolysis (phospholipase A₂ dye: bis-BODIPY FL-C₁₁-PC). AOS production may have diminished the membrane polarity, and peroxidized the membrane lipids (HPLC-analyzed malondialdehyde) which enhanced PLA₂ activity. The dyes indicated an increased oxidative (lipid) damage at a 15% inhibition of photosynthesis in this diatom, at UVB levels and salinities that can be expected in situ during its periodic tidal emersion.     

Fecundity,brood loss and egg development through embryogenesis of <Emphasis Type="Italic">Armases cinereum</Emphasis> (Decapoda: Grapsidae)

The present work is a comprehensive study of reproduction and embryonic development of Armases cinereum. Ovigerous A. cinereum (Bosc, 1802) females from Sebastian Inlet, Florida (9.88–19.4 mm CW) lay 2,000–12,000 eggs per brood, depending on their CW (mm): fecundity = 24.662 CW^1.9432. A. cinereum displayed significant brood loss through development (ca. 500 eggs per brood) independently from their CW (no senescence). However, since smaller females lay fewer eggs than larger ones, the percentage of eggs lost during embryonic development is greater in smaller females. The number of eggs carried on a later stage of development (potential fertility = 5.5593 CW^2.4417) is a more accurate estimate of the reproductive output and subsequent recruitment. Egg volume increased during development (64%, 0.025–0.041 mm³ or 0.36–0.43 mm of diameter, N = 270) and was strongly correlated with egg water content increase (19.21%, r = 0.89). Lipids, particularly fatty acids, seem to be the major energy source for embryonic development, decreasing 56.31 and 37.08% (respectively) during embryonic development; both are negatively correlated with egg volume (r = −0.90). The utilization of fatty acids through the different developmental stages of A. cinereum is presented. The most consumed fatty acids are the monounsatured (43.33 μg mg⁻¹ dw), followed by the saturated (29.91 μg mg⁻¹ dw) and polyunsaturated (24.03 μg mg⁻¹). Palmitic (16:0) and linoleic (18:2n-6) acids are preferentially consumed (19.5 and 17.9 μg mg⁻¹ dw, respectively). The high proportion of essential polyunsaturated fatty acids of C₁₈ and C₂₀ reflects the consumption of primary producers such as mangrove leaves. EPA/DHA ratio (2.85–3.84) and low DHA content indicated that this species appears in a medium-low level of the trophic chain. The low ratio of 18:1n-7/18:1n-9 and high percentage of 18:1n-9 (marker of carnivory) may be a sign of the consumption of juvenile invertebrates. The high percentage of odd-numbered FA indicated the occurrence of detritivores/scavenger behaviours. The fatty acid composition of the eggs reflects adult feeding ecology (omnivorous) and habitat.     

Swimming behavior of juvenile anchovies (<Emphasis Type="Italic">Anchoa</Emphasis> spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O₂ l⁻¹, except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O₂ l⁻¹. We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s⁻¹ that were near the theoretical optimum speeds between 7.0 and 8.0 bl s⁻¹. An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s⁻¹, expected = 6.4 bl s⁻¹). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.     

Energy balance of <Emphasis Type="Italic">Octopus maya</Emphasis> fed crab or an artificial diet

The present study was designed to evaluate the effect of a natural prey (the crab Callinectes sp.) and an artificial diet (pellet with squid paste and offered as a paste) on the survival and assimilation efficiency of subadult octopuses with 486 g of initial live weight. In order to reach this goal, the effects of the type of diet on energetic balance were assessed by recording ingestion rate (C), respiratory rate (R = R routine, R _rout + R apparent heat increment, R _AHI), ammonia production rate (U = U routine, U _rout + U post-prandial, U _PP) and biomass production (P) of Octopus maya during its growing process. Energy lost from faeces (H) was calculated as H=C−(U+R+P) and assimilated energy (As) as R + P. Octopuses fed an artificial diet had almost five times higher ingestion rate compared to that observed in octopuses fed crab. However, growth rate and production (P) were high in octopuses fed crab in comparison to octopuses fed artificial diet. An inverse relation between faeces (H) and type of food was observed, indicating that animals lost 77% of the ingested energy when fed artificial diet and only 5% when fed crab. A higher assimilation and production efficiency were obtained in octopuses fed crab (P/As: 61%) than in animals fed the artificial diet (P/As: −5%). The routine O : N ratio for animals in fasting was 9.1 and 2.3 for octopuses being fed crabs and the artificial diet, respectively. The post-alimentary O : N ratio was 3.6 and 2.2 for animals fed crabs and the artificial diet, respectively. This indicates that animals fed on both diets rely almost exclusively on protein. Based on energy balance data, a value of 472 kJ week⁻¹ kg⁻¹ of live octopus was estimated as the energy needed to obtain a growth rate near 9 g day⁻¹ (2.8% BW day⁻¹) for O. maya subadults. The total crab biomass needed to obtain 1 kg of fed O. maya biomass was calculated. A comparison with other different energy balance measurements made in other octopus species indicates that O. maya and Enteroctopus megalocyathus (Pérez et al. 2006) tend to be more efficient by channelling more ingested energy to biomass production (P = 69.5% of C) than O. vulgaris (P = 23% of C; Petza et al. 2006) or Paraledone charcoti (P = 4% of C; Daly and Peck 2000).     

Seasonal variations in hepatopancreas fatty acid profiles of two colour forms of shore crabs, Carcinus maenas

 The relationship between intermoult duration (coloration), sex, size and seasonal variations in fatty acid (FA) profiles was studied in a population of shore crabs, Carcinus maenas, inhabiting the Isefjord, Denmark. For male shore crabs, the total hepatopancreas FA content was high in July and December (12.7 to 16.0 mg g⁻¹ dry weight, dw) but lower in May and September (7.3 to 10.0 mg g⁻¹). This indicates that male shore crabs are in relatively good condition before winter, when the crabs migrate off shore, but in relatively poor condition when they return to shallow waters during spring. The hepatopancreas FA content also decreased over the mating season. After the mating season the hepatopancreas FA content of males had decreased to approximately 60% of that prior to the mating season. Female shore crabs had significantly higher hepatopancreas FA levels than males in May (11.7 mg g⁻¹ dw), September (12.6 mg g⁻¹ dw) and December (17.9 mg g⁻¹ dw) but lower levels in July (9.5 mg g⁻¹ dw). This indicates that the spawning season is the most energy-demanding part of the female reproductive cycle. For all seasons, the hepatopancreas FA content of green shore crabs was significantly higher than that of red shore crabs. For both colour forms, the amount of polyunsaturated fatty acids (PUFAs) was significantly higher than that of saturated fatty acids (SAFAs) and monounsaturated fatty acids (MUFAs), with the relative proportion of PUFAs increasing when the total hepatopancreas FA content decreased. For both genders and colour forms, the most dominating SAFA was palmitic acid (16:0). Palmitoleic acid (16:1ω7), vaccenic acid (18:1ω7) and oleic acid (18:1ω9) were the three MUFAs found in highest concentrations. The most dominating PUFA was eicosapentaenoic acid (EPA, 20:5ω3). Docosahexaenoic acid (DHA, 22:6ω3) and arachidonic acid (AA, 20:4ω6) were also abundant in all samples. The results demonstrated that season, sex, size and intermoult duration all influence the amount of FAs present in the hepatopancreas of shore crabs. Received: 23 September 1999 / Accepted: 29 April 2000     

Seasonal variations of ATPase activity and antioxidant defenses in gills of the mud crab <Emphasis Type="Italic">Scylla serrata</Emphasis> (Crustacea,Decapoda)

The mud crab Scylla serrata is an important commercial crustacean being widely distributed along the southeastern coast of China. This crab and particularly its gills are subjected to seasonal fluctuations of environmental factors due to direct exposure to seawater. The investigation of seasonal variations of ATPase and antioxidant defenses in gills could be helpful for the understanding of physiological regulation mechanism of seasonal adaptations. In this study, mud crabs were collected from the subtropical waters near Xiamen island, Southeast China (24°26′46′′N, 118°04′04′′E), in August and November, 2002, and February and May, 2003, respectively, being considered as specimens from summer, autumn, winter and spring, respectively. Only healthy intermoult male crabs without carapace or appendage damage were used, having a carapace width of 7.05 ± 0.52 cm and a wet weight of 130 ± 20 g. The activities of four ATPases and three antioxidation enzymes (superoxide dismutase, catalase, glutathione peroxidase) and the content of malondialdehyde in gills were measured. The results showed that the activities of Na⁺, K⁺-ATPase, Mg²⁺-ATPase, Ca²⁺-ATPase and Ca²⁺, Mg²⁺-ATPase reached maximum levels similarly in summer and decreased in winter. The activities of superoxide dismutase and catalase were higher in summer and lower in winter, with a significant decrease in winter compared to the other seasons (p < 0.01). Glutathione peroxidase activity was higher in summer, and lower in autumn, being significantly higher in summer or winter than in spring or autumn (p < 0.01). Malondialdehyde content was higher in summer and lower in spring with significant differences among the different seasons (p < 0.01). In summary, the obvious seasonal activity variations of four ATPases and antioxidant enzymes and the content of malondialdehyde reflect a seasonal regulation of the physiological metabolising enzyme and the antioxidant capacity to cope with seasonal alterations of environment factors such as fluctuating salinities and temperatures.