Relative importance of benthic microalgae,phytoplankton, and mangroves as sources of nutrition for penaeid prawns and other coastal invertebrates from Malaysia

Carbon, sulfur, and nitrogen stable isotope ratio techniques were used in 1989 and 1990 to evaluate the relative importance of algae and of mangrove detritus in the nutrition of two penaeid prawn species on the west coast of Peninsular Malaysia. Mangrove detritus was found to contribute to the nutrition of juvenile Penaeus merguiensis de Man living within tidal creeks, but not to adult P. merguiensis and juvenile and adult Parapenaeopsis sculptilis (Heller) captured offshore. Results from radiotracer feeding studies, with refractory ¹⁴C mangrove lignocellulose as the food source, indicated that juvenile P. merguiensis from tidal creeks assimilated mangrove carbon with an efficiency of 13.4%. This did not differ significantly (P=0.05) from the assimilation efficiencies of juvenile and adult P. sculptilis living offshore (10.0 and 10.9%, respectively); these values were significantly higher (P<0.001) than for adult P. merguiensis (2.1%). Thus, the differential contribution of mangrove material to the nutrition of prawns in the tidal creeks and offshore was not related to differences in the prawn's ability to utilize detritus. Instead, our analysis of C stable isotopes in sediments indicated that mangrove detritus was generally more abundant within the tidal creeks than offshore. Juvenile prawns in the tidal creeks may also utilize mangrove material indirectly by feeding on small detritivorous invertebrates. Stable isotope analyses suggest that benthic microalgae constitute the other major dietary component for prawns living in tidal creeks. Prawns offshore were utilizing mainly phytoplankton-based material.     

Diverticulum sand in a miniature sand dollar Sinaechinocyamus mai (Echinodermata: Echinoidea)

Sinaechinocyamus mai (Wang) were collected from the low intertidal zone at Tunghsiao, western Taiwan, periodically from July 1990 to July 1991 and May 1993. The mean body length was 7.3±1.01 mm from 146 juveniles and adults. All individuals contained sand grains in their intestinal diverticula. The densities of sand dollars ranged from 1.2 to 2.5 mg cm^-3 and had an inverse linear relationship to the body length. The percentage of sand weight in the diverticulum to the wet body weight is inversely correlated to the body length, indicating that juveniles contain a higher proportion of sand than adults. The ratio of sand weight to body length did not change significantly during the study period. 74% of the diverticulum sand was 0.075 to 0.125 mm in diameter, whereas only 0.1% of the sand in the substrate was this size. The sand in both the diverticulum and the substratum consisted of feldspars and quartz (88.4%) and iron oxide (11.6%). The sand grains of the diverticulum were eliminated from the anus.     

Influence of subtle substrate differences on feeding by shorebirds on intertidal mudflats

Shallow-feeding shorebirds, dowitchers (Limnodromus griseus and L. scolopaceus), western sandpipers (Calidris mauri), dunlin (C. alpina) and American avocets (Recurvirostra americana), reduced the density of their prey in mudflats with little sand but not in mudflats with a moderate admixture of sand. An experiment in Upper Newport Bay, Southern California, during October and November 1979 to explain the difference in density is described. The effect of sand on shorebird feeding was compared in the field by measuring the times spent feeding in plots where sand had or had not been added, respectively. Increasing the sand content to 14% from 2% in the top centimeter decreased the time spent in treated plots by all species compared to adjacent untreated plots. The prey species were small polychaete and oligochaete worms (0.25 to 1.25 mm wide) similar in diameter to sand grains (0.5 to 1.0 mm). In plots where sand had been added, avocets fed by pecking at the surface in addition to scything, the more common method of feeding on muddy substrates. The results suggest that sand interferes with the detection and or capture of prey that are similar in diameter to small sand grains and explains the differences in the effects of predation by these birds seen on mudflats with a moderate admixture of sand compared to the effects on mudflats with little sand. Differential success in prey capture between one microhabitat and the nest (rather than a reduction in competition, as suggested by some authors) might explain the different use of such habitats.     

Microphytobenthos of the Dogger Bank: a comparison between shallow and deep areas using phytopigment composition of the sediment

The surface sediment characteristics related to benthic microalgae primary production were studied at the Dogger Bank, North Sea, in order to evaluate the potential role of microphytobenthos as a food source for the macrobenthic fauna. Twenty-one stations were sampled in July 2001 and May 2002, with water depth ranging from 16.3 to 68.5 m. High-performance liquid chromatography pigment analyses revealed that concentrations of chlorophyll a, chlorophyll c and fucoxanthin are mainly associated with benthic diatom flora at most parts of the Dogger Bank. High percentage of phytopigments (>50%) was firmly attached to sand grains at the stations shallower than 40 m water depth. The deeper stations were characterized by a phytopigment composition originating from pelagic phytoplankton settled on the sea floor. Qualitative microscopy showed that the benthic microflora on top of the Bank mainly consists of small diatoms (5–10 μm), such as e.g., Diploneis spp., living attached to the sand grains. The results are discussed concerning possible implications for ecology and biogeochemistry of the Dogger Bank area.     

Impact of laboratory-imposed physical disturbance on the abundance of four isolates of marine gymnamoebae

Four species of gymnamoebae isolated from a sandy shore were subjected to physical disturbance within sand microcosms containing either 250 or 125 μm sand. Firstly, the effect of two different strengths of physical disturbance on amoebae populations within sand was investigated. Secondly, whether the size of sand grains occupied by amoebae populations changed their response to disturbance events was assessed. Highest abundances of amoebae were recorded in control flasks containing 125 μm sand. Increasing the strength of disturbance decreased the abundance of all four species. Abundances of the two smallest isolates (Platyamoeba sp. and a limax amoeba) were least affected by increasing disturbance. The greatest decrease in abundance was observed in populations of two larger (>15 μm) Mayorella sp. and Vannella sp. The greatest impact of physical disturbance was seen on populations within 250 μm sand. This study shows that physical disturbance may be an important community structuring force within natural marine gymnamoebae communities.     

Natural diet of larval Penaeus merguiensis (Decapoda: Penaeidae) and its effect on survival

The relationship between Penaeus merguiensis protozoea larvae and their phytoplankton diet was examined using seasonal plankton surveys and in situ rearing experiments. Larval abundance, phytoplankton community structure, and chlorophyll a concentration in Albatross Bay, Gulf of Carpentaria, were monitored monthly for 2 yr. Larval abundance peaked in November (spring) and March (autumn), at which times diatoms were the most abundant group in net samples of phytoplankton and in the guts of larvae. During November 1989 and March 1990, larvae were reared in nylon mesh enclosures positioned throughout the water column at three depths: 0 to 3 m, 3 to 6 m and 6 to 9 m. Overall, larval survival and gut fullness were both higher in November than in March. In both months, larval survival was lower at the surface than at other depths. This correlated with lower chlorophyll a concentrations, but lower total cell densities were not detected. During the in situ experiments, diatoms were the most abundant phytoplankton group in the water column and in the guts of larvae and, therefore, appeared to be the principal diet of larvae. Pigment analysis demonstrated that while gut contents generally reflected the composition of the phytoplankton community, the larvae were not feeding exclusively on diatoms. They also ingested green algae and possibly seagrass detritus. The in situ experiments demonstrated that the predominantly diatom flora in Albatross Bay can provide a nutritionally adequate environment for prawn larvae even at seasonally low levels. It is unlikely, therefore, that starvation is a major cause of mortality of P. merguiensis larvae during either of the biannual peaks in their abundance in Albatross Bay, Gulf of Carpentaria.     

Structure and function of the third maxillipeds of the banana prawn Penaeus merguiensis

The functions of the third maxilliped of Penaeus merguiensis de Man are described. It has a 6-segmented endopodite which is used in feeding, grooming and possible pheromone reception. In feeding, large pieces of food are gripped by the stout spines on inner borders of the ischia which hold them to the mouth so that particles can be torn off and ingested. All the distal segments have long setae used in grooming chelate pereiopods and the antennules. In mature males there is a tuft of about 300 slender setae at the end of the propus, but its function remains obscure.     

Diets of piscivorous fishes in a tropical Australian estuary,with special reference to predation on penaeid prawns

The diets of fish from the tropical Embley Estuary in the eastern Gulf of Carpentaria, Australia, were analysed with particular reference to piscivory and predation on juveniles of commercially important penaeid prawns from October 1986 until July 1988. Of the 77 species caught, 52 were piscivorous, and of these 37 ate penaeid prawns. The most numerous piscivores wereScomberoides commersonianus, Arius proximus, Lates calcarifer, Polydactylus sheridani andRhizoprionodon acutus, the first four of which accounted for over 90% of all prawns eaten. Twenty species are commercially important species of prawns. The proportions of penaeids in the diets varied seasonally, according to the density of penaeids in the estuary. The proportion was highest during the pre-wet period (November) and lowest during the dry period (July–August). Predation on prawns was highest in the lower and middle reaches of the river. Estimates of the rates of predation on the two most common juvenile commercial prawns,Penaeus merguiensis andP. semisulcatus, are presented and discussed.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Sand stress as a non-determinant of habitat segregation of indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels in South Africa

Periodical sand inundation influences diversity and distribution of intertidal species throughout the world. This study investigates the effect of sand stress on survival and on habitat segregation of the two dominant mussel species living in South Africa, the invasive Mytilus galloprovincialis and the indigenous Perna perna. P. perna occupies a lower intertidal zone which, monthly surveys over 1.5 years showed, is covered by sand for longer periods than the higher M. galloprovincialis zone. Despite this, when buried under sand, P. perna mortality rates were significantly higher than those of M. galloprovincialis in both laboratory and in field experiments. Under anoxic condition, P. perna mortality rates were still significantly higher than those for M. galloprovincialis, but both species died later than when exposed to sand burial, underlining the importance of the physical action of sand on mussel internal organs. When buried, both species accumulate sediments within the shell valves while still alive, but the quantities are much greater for P. perna. This suggests that P. perna gills are more severely damaged by sand abrasion and could explain its higher mortality rates. M. galloprovincialis has longer labial palps than P. perna, indicating a higher particle sorting ability and consequently explaining its lower mortality rates when exposed to sand in suspension. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. Although sand stress strongly affects the survival of the two species, it does not explain their vertical zonation. Contrary to our expectations, the species that is less well adapted to cope with sand stress maintains dominance in a habitat where such stress is high. GI Zardi, KR Nicastro contributed equally to the work     

Dehydrogenases as tools in the study of marine sediments

A method is outlined for the measurement of NADH₂-dependent dehydrogenase activity in whole samples as well as in fractions of carbonate sediments. The method is based on extraction of enzyme activity with phosphate buffer containing 0.12% Triton X-100, dialysis, and a photometric assay which permits initial velocity determinations of the enzyme reaction under controlled conditions of pH, temperature, and substrate concentration. The dehydrogenases extracted from carbonate sediment of a subtropical beach at Bermuda are characterized by low K _m values (0.03 to 0.07 mM NADH₂/1), pH maxima around 8.5, and temperature maxima between 35° and 40°C. Their vertical distribution in the sediment agrees with what is known about the distribution of biomass on beaches of this type, maximum activity occurring in the topmost centimetre. Approximately 80 to 90% of the total electron transport activity is contained in the grain fraction, the remainder in the interstitial fraction of a sediment sample. From this it can be inferred that the interstitial fauna contributes only little to the total energy budget of this type of sediment, which is dominated by the Aufwuchs on the sand grains. The electron transport activity (ETA) measured by means of the dehydrogenase assay is taken to represent the maximum transport capacity (ETC) of all the organisms living in the sand ecosystem. From published evidence it is inferred that oxygen consumption of sediment samples and of individual organisms measured in vivo usually accounts for 10 to 20% of the maximum electron transport capacity measured in vitro with saturating concentrations of substrates and with Triton X-100 as part of the enzyme assay. It is suggested further that the ratios of dehydrogenase activities in different fractions or zones of sediment, or of different organisms, may represent one of the best indicators, so far available, of energy relationships in marine sediments.Contribution No. 652 from the Bermuda Biological Station for Research.     

Settlement cues in successive developmental stages of the ghost shrimps Callichirus major and C. islagrande (Crustacea: Decapoda: Thalassinidea)

Larvae of Callichirus major (Say) and C. islagrande (Schmitt) were given unconditioned water with sand, water conditioned by previous exposure to adults (AW) without sand, or the combination of both AW and sand to determine if these factors served as settlement cues. Shrimp were first given stimuli upon reaching the last zoeal stage (ZIV), the decapodid stage (D), or the first juvenile stage (J1). Once initiated, exposure of shrimp to stimuli was maintained until termination of␣the experiment. Stage duration, burrowing activity of decapodids and early juveniles and location of exuviae were observed daily. Exposure to sand and AW did not decrease duration of the decapodid stage in either species. However duration of ZIV in C. major appeared to decrease when these stimuli were offered at this stage; this suggests that molting to Stage D may be delayed without such cues. When stimuli were examined separately, AW was found to have a significant effect on the duration of ZIV, while the effect of sand with unconditioned water and effects of simultaneously offered stimuli were not significant. Shrimp that received stimuli at ZIV subsequently burrowed more at D and J1 than individuals that did not receive stimuli at ZIV. Location of exuviae was recorded as on top of sand (not burrowed during molt) or within sand (burrowed during molt) for each shrimp's molt from Stages D to J1 (D molt), and from J1 to J2 (J1 molt). In C. major, the percentage of D exuviae and J1 exuviae found within sand as opposed to on top of sand was significantly higher in individuals which received stimuli at ZIV than for those that first received them at D or J1. When stimuli were examined separately in C. major, the effect of sand on burrowing activity was highly significant, while AW and the stimuli in combination had weaker effects. For C. major, AW appeared to accelerate development to D, while sand triggered settlement. Neither stimulus altered stage duration or burrowing activity of C. islagrande. Received: 7 December 1997 / Accepted: 24 July 1998     

Larval settlement and metamorphosis in the sand crab Emerita talpoida (Crustacea: Decapoda: Anomura)

The effects of various potential cues on the survival and time to metamorphosis of larvae of the sand crab Emerita talpoida (Say) were examined. Zoeal duration, which ranged from 25 to 43 d, was not correlated with subsequent megalopal survival but had a weak, negative correlation with the duration of the megalopal stage. Neither food, sand nor water from containers with adults directly affected megalopal survival or duration. In a second experiment, the presence of sand both increased the probability of survival and shortened the duration of late zoeal stages. This is the first experimental evidence for delayed zoeal metamorphosis in a decapod. Settlement of E. talpoida appears to occur primarily during the zoeal phase, not at the megalopal stage.     

Effects of Trichodesmium spp. blooms on penaeid prawn larvae

The effects of blooms of the cyanobacterium Trichodesmium spp. on penaeid prawn larvae were examined using in situ and laboratory rearing experiments and plankton surveys in Albatross Bay, Gulf of Carpentaria, Australia. The in situ experiments demonstrated that, during a bloom of Trichodesmium spp., larvae of the prawn Penaeus merguiensis did not develop beyond the first protozoea stage, and survival was low compared with times when diatoms were dominant in the same study area. Laboratory experiments confirmed the in situ results. None of the prawn larvae fed Trichodesmium sp. in laboratory experiments developed beyond the first protozoeal stage. In contrast, 94% of prawn larvae fed the green flagellate Tetraselmis suecica successfully developed to the second protozoea stage. Electron microscopy of larvae gut-contents revealed that Trichodesmium spp. were ingested by larvae but were of no nutritional value, resulting in starvation. A 7 yr plankton survey, from 1985 to 1992, showed that minimum abundance of prawn larvae occurs during the annual summer blooms of Trichodesmium spp. and that maximum abundance of prawn larvae generally occurs just after the bloom. There was a negative correlation between the abundance of larvae and the abundance of Trichodesmium at individual sites, one offshore and one inshore, indicating that the blooms affect the survival of larvae. We conclude that variations in both timing and magnitude of Trichodesmium blooms are important determinants of prawn larvae abundance in Albatross Bay. Received: 28 April 1997 / Accepted: 2 April 1998     

A preliminary account of two sandy beaches in South West India

Observations were made on 2 beaches during the premonsoon period (January to March) when conditions were relatively stable. The beach below the berm at Cochin is 14 m wide with a slope of 1 in 8.5, and at Shertallai 40 m wide with a slope of 1 in 16. The well-sorted sand has a mean particle size of 275 at Cochin and 175 at Shertallai. The amount of chlorophyll attached to sand grains is very small. Chlorophyll in the water varies between 0.9 and 13.0 g/l. Carbon in the sand amounts to 210 to 630 g/g, and in the water to 355 to 3660 g/l. Throughout observations, the water temperature remained about 31°C, but on the dry sand surface it reached 50°C. Fauna zonation on the 2 beaches is similar to that of other tropical beaches; crabs of the genus Ocypode and the isopod Eurydice sp. occur at high water mark, the polychaetes of the genera Glycera, Onuphis, Scoloplos and Lumbrinereis in the mid-intertidal region and below; the tidal migrants Bullia melanoides (Gastropoda), Donax incarnatus and D. spiculum (Bivalvia) and Emerita holthuisi (Crustacea) inhabit the wash zone of the surf. The average biomass is 0.73 g carbon/m² at Shertallai and 0.85 g C/m² at Cochin, where fewer species are present. This compares with a value of 0.63 g C/m² for a sandy beach on the west coast of Scotland.     

Variations in benthic community structure in a sub-tropical estuary

Abundances of macroinfaunal invertebrates in sand dollar (Dendraster excentricus) beds and in adjacent areas free of sand dollars were studied at 10 intertidal sites in the Pacific Northwest region of the USA and Canada. Each site was sampled once in late summer of 1977. There were no significant differences in overall diversity of species between the beds and adjacent areas; nor did cluster analysis indicate a sand dollar bed community. However, a polychaete (Armandia brevis) was significantly more abundant inside the beds than just outside them. Conversely, in comparison to the beds, adjacent regions contained significantly more specimens of the following species: a bivalve (Transennella tantilla), several tubicolous crustaceans (Corophium spp., Ampelisca agassizi, and Leptochelia savignyi) and two polychaetes (Glycinde polygnatha and Malacoceros arenicola). Sediment characteristics at each study site were not correlated with the presence or absence of sand dollars; moreover, a comparison among the 10 sites revealed no sediment characteristics typical of sand dollar beds. At most of the sites, significantly fewer tube-building invertebrates occurred inside sand dollar beds than outside. The discussion considers the possibility that mechanical disturbance by the sand dollars tends to exclude some macroinvertebrates (especially tubicolous ones) from sand dollar beds.     

Additional notes on Tubiluchus corallicola (Priapulida), based on scanning electron microscope observations

Cuticular differentiations on the body surface and the pharyngeal armature of adult Tubiluchus corallicola van der Land were examined with the scanning electron microscope. The morphology of abdominal tumuli, flosculi and tubuli, and of pectinate teeth and fimbrillae on the pharynx is described, and an attempt is made to interpret their functional properties. Taking into account the structural details, relative size, and topographic arrangement in relevance to interaction with the interstitial environment, we assume that tumuli are protective, flosculi chemoreceptive, and tubuli adhesive, and that the pharyngeal armature is a scraping mechanism for feeding on epigrowth on sand grains.Supported in part by NSF Grant GB-7952 to the first author.     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.