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.     

Herbivory and detritivory among gammaridean amphipods from a Florida seagrass community

The gammaridean amphipods Cymadusa compta (Smith), Gammarus mucronatus Say, Melita nitida Smith and Grandidierella bonnieroides Stephensen from a seagrass community in the Indian River estuary of Florida (USA) fed variously upon large drift algae, small algae epiphytic on seagrasses and seagrass leaf debris and detritus. Consumption was measured in the laboratory using an index (CI) equivalent to mg ingested mg^-1 amphipod day^-1. Observations revealed that the amphipods fed by macrophagy, an attack upon large algae and seagrass debris, and by microphagy, small particle detritus feeding and scraping of plant surfaces for diatoms and other epiphytic algae. C. compta was a macrophagous feeder with a generalized diet of algae and seagrass debris, preferring epiphytic algae and drift algae at mean rates of 1.10 and 0.87 CI, respectively. Gammarus mucronatus fed upon epiphytes and seagrass debris equally at mean rates of 0.90 and 0.97 CI, respectively. The diet of M. nitida condisted primarily of epiphytes, consumed at an average rate of 1.05 CI. Grandidierella bonnieroides fed in a specialized microphagous manner, grooming plant surfaces for small particle detritus and diatoms at an approximate CI rate of 1.45. Assimilation of plants ingested, as reflected by carbon-14 uptake, varied similarly among the 4 amphipods. Epiphytic algae appeared to be most useful as food, providing means of 41 to 75% carbon-14 uptake as ingesta. Drift algae and seagrass debris were of less value, with means varying between 11 and 24 % of carbon-14 uptake by the amphipods. The data show a pattern of feeding which resembles resource partitioning of food both by size and kind. Other evidence, however, including population limitation by predators and an apparent overabundance of food, indicate that resource partitioning as seen may be an artifact, and one which has no co-evolutionary basis among the present species.Contribution No. 102 of Harbor Branch Foundation, Inc.     

Experiments on resuspension of natural microphytobenthos populations

Laboratory experiments on resuspension of benthic diatoms were performed in 1988 on intact cores of sediment obtained from two transects in the Ems estuary (Dutch Wadden Sea, NW Europe). Diatoms were most abundant in sediment from sheltered stations. In the experiments, which were performed in a small carousel system, the percentage of resuspended diatoms increased concomitantly with angular velocity of the agitating paddles. Maximum microphytobenthos resuspension in terms of cell numbers was up to 45% of the total population present in the 0.5-cm top layer, and up to 11% in terms of chlorophylla. Maximum sediment resuspension was 6%. The results show that the presence of a diatom film (numerous cells on the sediment surface) increases sediment stability, thereby suppressing resuspension of sediment and diatoms. However, if the sediment contains many fine particles and much detritus, sediment stability decreases, leading to increased resuspension.These results support the contention that low detritus content and large numbers stabilize the surface layer of tidal flats, whereas high detritus content and fewer cells (<2 × 10⁵ cm^–2) do not. The stabilization is thought to be dependent on a combination of differences in sediment consolidation and in bottom roughness caused by the presence or absence of diatom films.     

Epizoic diatoms on the euphausiidNyctiphanes australis: Consequences for gut-pigment analyses of whole krill

We report the first record of infestation by the epizoic pennate diatoms (Licmophora sp.) of the euphausiidNyctiphanes australis (G. O. Sars, 1883) collected from Otago Harbour and coastal waters, New Zealand (45°50S; 170°37E) between October and April 1986–1989: at times up to 50 to 70% of the krill examined were infested. Infestation by epizoic diatoms introduces a significant source of error into determinations of egestion rates using gut fluorescence of whole individuals, and prevents measurement of background fluorescence in krill. Diatom infestation can lead to overestimation of ingestion rates by a factor of 2 to 5. We suggest that investigators using the gut fluorescence method to measure grazing rates should determine the occurrence of epizoic diatoms in samples prior to analysis. If diatoms are present, it is necessary to hold krill in the laboratory until diatoms are shed at moult before makingin vitro gut-pigment measurements. Alternatively, the stomachs must be dissected for analyses, negating the benefits of rapid sample preparation and quantitative recovery of pigments that are the advantage of the technique. Epizoic diatoms can significantly contribute to variability of apparentin situ ingestion rates inN. australis.     

Diet of copepods (Scopalatum vorax) associated with mesopelagic detritus (giant larvacean houses) in Monterey Bay,California

The feeding structures or houses of the giant larvacean Bathochordaeus sp. serve as both habitat and food for the calanoid copepod Scopalatum vorax. Gut contents of S. vorax include both microbial and metazoan associates of larvacean houses, and possibly the house-mucus matrix itself. Copepods were observed and collected from larvacean houses between 100 and 500 m in Monterey Bay, California, using a submersible ROV (remotely operated vehicle) from the Monterey Bay Aquarium Research Institute. Gut contents were compared to potential food items on the houses and in the open water (not associated with the house). Copepods were generalist feeders, with amorphous detritus, diatoms, and copepods or other crustacean parts dominating gut contents. Protozoans and algae other than diatoms were rarer in guts. Houses contained a diverse assemblage of microplankton and metazoans, both intact specimens and detrital remains of these. Numbers of diatoms and fecal pellets were enriched by 1 to 3 orders of magnitude on houses compared to numbers in surrounding water. Many of the abundant species of diatoms and copepods on houses occurred in S. vorax guts. This observation coupled with S. vorax feeding habits observed in situ and in the laboratory provide evidence for feeding on houses. S. vorax appears to possess special adaptations to living in a resource-limited environment, such as gorging as a feeding adaptation, chemosensory structures to help locate houses, and the ability to change feeding modes. Consumption of detritus at depth by S. vorax provides evidence that metazoans contribute to remineralization of particulate organic carbon in the mesopelagic zone.     

Organic detritus of a tropical estuary

Organic detritus of the Cochin Backwater (India) is largely composed of fine silt and sand particles around which organic matter adheres and forms aggregates. In addition to plankton, large quantities of benthic algae, rooted plants, animal matter, suspended soft mud and the material brought down by the rivers and land runoff constitute the main sources of detritus. Detritus sedimentation in the estuary attains its maximum from April to June. Detritus forms a major portion of seston, while phytoplankton productivity constitutes 0.1 to 1.0% of settled detritus. Detrital pigments include degraded chlorophyll (phaeophytin), which has a marked seasonal variation; detrital carbon, phosphorus and nitrogen show little change during the year. The caloric value of detritus, calculated from protein, carbohydrate and lipid fractions and also from total carbon, indicates that sedimented detritus does not have a high nutritional value. However, being a readily available material, its entry into the food chain seems to increase the efficiency of energy transfer from one trophic level to another.     

Natural diets of vertically migrating zooplankton in the Sargasso Sea

The feeding preferences of three common diel vertically migrating zooplankton were investigated from December 1999 to October 2000 at the U.S. JGOFS Bermuda Atlantic Time-Series Study (BATS) station in the Sargasso Sea. Gut content analysis of the copepods Pleuromamma xiphias (Giesbrecht) and Euchirella messinensis (Claus) and of the euphausiid Thysanopoda aequalis (Hansen) indicated that all three species fed on a wide variety of phytoplankton, zooplankton, and detrital material. Diet changes generally reflected seasonal trends in phytoplankton community structure. However, species-specific feeding preferences and differences in feeding selectivity among the three species were evident, and in general agreement with feeding habits predicted from the analysis of mouthpart morphology. The euphausiid T. aequalis fed equally on more different food types compared to both copepod species. The copepod P. xiphias consumed a diverse assemblage of phytoplankton from late winter through the summer (78-93% of gut items, by number, were phytoplankton) and based its diet more strongly on carnivorous feeding in autumn and early winter (31% and 61% of gut items were phytoplankton, respectively). E. messinensis showed the greatest feeding specialization, with a strong preference for pennate diatoms in winter and spring and for coccolithophorids during late summer and fall (constituting 67-93% of gut items by number). All three species consumed diatoms more than other phytoplankton taxa, even though diatoms form only a small fraction of the phytoplankton biomass in the Sargasso Sea. Although the majority of gut items identified were phytoplankton cells, the relative biomass contribution of these small cells may be lower than that of zooplankton and detritus. Zooplankton on which the three species primarily preyed were protozoans and crustaceans, but also included other metazoans such as chaetognaths and cnidarians. Marine snow was also an important component of the diet in all three species, with typically >50% and rarely <20% of the gut content being olive-green debris. Marine snow from larvacean houses was found in the guts of all three species, while E. messinenis appeared to selectively consume marine snow aggregates enriched with bicapitate Nitzschia spp. Large cyanobacteria (>4 µm in diameter) found in guts were also likely consumed with marine snow. The species-specific differences in the diets of these three migrating species suggest that an individual species approach is important in determining how feeding habits affect the structure of pelagic food webs and carbon cycling in the sea. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0815-8.     

Trace metal ingestion and assimilation by the green mussel <Emphasis Type="Italic">Perna viridis </Emphasis>in a phytoplankton and sediment mixture

We examined trace metal ingestion and assimilation (Cd, Se, Zn) by the green mussel Perna viridis in a mixture of diatoms and sediment at concentrations below and above the pseudofeces production levels. Dual gamma radiotracers (¹⁰⁹Cd, ⁶⁵Zn) were used to investigate particle selection on marine diatoms and sediments. The diatom (Thalassiosira weissflogii) was radiolabeled with ¹⁰⁹Cd, and the natural sediment was radiolabeled with ⁶⁵Zn. By comparing the ratios of ¹⁰⁹Cd:⁶⁵Zn in the particle mixture, mussel tissues and pseudofeces within a short-term exposure period (35 min), the results demonstrated that the green mussels were able to selectively ingest the diatom particles at a high particle load. Efficiency of selection for nutritious particles (e.g. diatoms) increased with increasing ratio of sediments in the particle mixture. Pseudofeces contained a higher ratio of sediments relative to that in the feeding suspension. No major particle selection was observed at concentrations below the level for pseudofeces production. The assimilation of Cd, Se and Zn by the green mussels was quantified using a pulse-chase feeding technique. The assimilation of Se and Zn by the green mussels from ingested diatoms was reduced with the presence of sediment within the mussel gut, presumably due to the resorption of metals onto the sediment, leading to a quicker passage of metals through the digestive tract and a lower proportion of metals subjected to intensive digestion. In contrast, the presence of diatoms did not significantly affect metal assimilation from ingested sediment. A significant correlation between metal assimilation efficiency and metal gut passage time was also observed. Metal assimilation by the green mussels appeared to be little dependent on the particle concentration in seawater. Our study suggests that particle selection may potentially alter metal influx from ingested food sources, particularly at high particle concentrations. Selective feeding of nutritious particles, coupled with a high assimilation efficiency from these ingested particles, may increase metal influx into mussels from the dietary phase.     

Food resources of postlarval brown shrimp (Penaeus aztecus) in a Texas salt marsh

Field and laboratory experiments were conducted to identify the sources of food in the natural diet of postlarval brown shrimp (Penaeus aztecus Ives). A series of enclosures placed in East Lagoon (29°20N; 94°45W) on Galveston Island, Texas, USA, in May 1985, were used to evaluate the individual and combined contribution of Spartina alterniflora detritus, epiphytes of S. alterniflora, plankton, and demersal fauna in terms of differences in shrimp growth and carbon assimilation (stable carbon-isotope analysis). Demersal fauna (harpacticoid copepods, amphipods, tanaids and polychaete annelids), and plankton (>0.095 mm) accounted for approximately 53 and 47% of the growth of the postlarvae (11 to 22 mm rostrum-telson length), respectively, while the autochthonous plant substrates, S. alterniflora detritus and epiphytes, contributed little. Laboratory experiments confirm that a mixed diet consisting of both animal protein and phytoplankton promotes maximum growth. Our results indicate that plankton may be an important allochthonous source of carbon contributing to the growth and development of shrimp in the salt marsh.     

A non-motorized dye ejector for visualization of flow in situ and its use with coral reef crinoids

We describe a portable, non-motorized device for delivering a tracer dye into seawater under field conditions. Dye is ejected at a constant flow rate over a period of tens of minutes. The ejector works in a wide range of ambient pressures without external energy requirements. The flow rate is adjusted simply by varying the length of the delivery tube. The dye streams permitted observations of the upcurrent and downcurrent flow regimes for a filter-feeding crinoid (Comanthus bennetti) living at a depth of 8 m on a coral reef. The results indicate that the crinoid may enhance the rate of particle capture by changing the scale of turbulence in the water passing through the mesh of the filtration fan.     

An assessment of seaweed decomposition within a southern Strait of Georgia seaweed community

An assessment of litter and detritus decomposition and nitrogen content of decomposing litter is presented for ten important seaweeds within a southern Strait of Georgia (British Columbia, Canada) seaweed community sampled from August 1975 until October 1976. Litter decomposition rates varied among species with the time required for litter to disappear from litter bags ranging from 6 d for the lamina of Nereocystis luetkeana to about 70 d for Fucus distichus. Decomposition was characterized by an accelerating increase in the nitrogen: dry weight ratio of remnant litter as decomposition proceeded. Iridaea cordata detritus decomposed most rapidly, at 5.7% d^-1, while rates for Gigartina papillata, N. luetkeana, Laminaria saccharina and Laminaria groenlandica ranged from 1.8 to 3.6% d^-1. The remaining species decomposed more slowly. There was a tendency toward more rapid decomposition with decreasing crude fibre content and detritus particle size; however, it appears that morphology, habitat and growth rate are also correlated with relative decomposition rates. Of 43 taxa identified within quantitative litter collections, F. distichus (41%), I. cordata (26%), N. luetkeana (27%) and Laminaria spp. (4%) accounted for 98% of total deposition with mean peak accumulation occurring in August and September from a low near zero in January and February. Litter distribution was patchy, with most litter decomposing near its place of deposition. The application of litter decomposition rates to measured litter accumulation in a mathematical simulation of decomposition predicted the rate of seaweed litter decomposition to peak at about 1.1 g AFDW (ash-free dry weight) m^-2 d^-1 in early September from a mid-winter low near zero. In total, 56±4% of decomposing litter formed detritus, with the remainder being released as soluble matter. The annual contribution of seaweed litter biomass to detrital pathways from our study site was calculated to be 152 g AFDW m^-2.     

Incorporation of organic aggregates by marine mussels

Two marine mussels, Geukensia demissa (Dillwyn) and Mytilus edulis (L.) collected in 1990 in Old Silver Beach, Falmouth, Massachusetts, incorporated nitrogen when fed ¹⁵N-labelled organic aggregates produced from dissolved organic nitrogen released by the brown sea-weed Fucus vesiculosis. Uptake of ¹⁵N on the aggregate diet was linear over the course of 24 h, and unincorporated ¹⁵N was eliminated from the gut after 48 h. Both species of mussels incorporated approximately five times more N when they were fed organic aggregates than when they were fed either ¹⁵N-labelled dissolved organic material (DOM) or particulate detritus, both of which were also derived from the seaweed. Nitrogen uptake was greatest in controls fed the diatom Thalassiosira weissflogii; mussels fed phytoplankton incorporated seven times more nitrogen than those fed aggregates. However, aggregates could supply an estimated 7 to 14% of the N requirements for both mussels, whereas DOM or particulate detritus could only supply 1 to 3%. These data provide evidence that a food web pathway exists from primary producer to released dissolved organic nitrogen to microbial organic aggregate to metazoan consumer, and, further, that it can be more important in a detrital food web than either particulate detritus or DOM.     

Diel vertical migration and feeding patterns of Mysis mixta (Crustacea,Mysidacea) in the Baltic Sea

The extent of the nocturnal vertical migration of Mysis mixta Lilljeborg varied between early July and late October (of 1985 and 1986) in a coastal area of the Baltic Sea. Migration was more restricted in early July and late October. Seasonal changes in surface light levels and transparency were sufficient to explain the observed differences. Mysids avoided light levels above 10^-4 lux throughout the study period. Smaller juveniles migrated higher up than larger juveniles and adults. A two-layered distribution with part of the population close to the bottom was observed at night. Zooplankton were more abundant in water layers above the main concentration of mysids. M. mixta fed on phytoplankton, detritus, copepods, cladocerans, rotifers and tintinnids. Diel changes in gut fluorescence indicated a higher intake of phytoplankton at night, but levels were low compared to primarily herbivorous zooplankton. Comparisons of stomach contents of mysids caught at the bottom in the evening and in the water column at night showed a higher ingestion of zooplankton at night and of detritus during the day. Mysids caught at the bottom at night had an intermediate diet. Copepods and cladocerans constituted between 90 and 100% of ingested material by weight in all mysid groups.     

Selective feeding of littoral harpacticoids on diatom algae: hungry gourmands?

We studied benthic harpacticoid grazing on diatom algae from two sites on the White Sea intertidal sandflat. Diatoms from sediments and from harpacticoid gut contents were sampled in situ, identified and counted, and grazing rates were calculated by two ways: (1) using potential daily ration estimations and (2) from the gut content and gut-residence time data. Paraleptastacus kliei did not contain any diatoms in the guts and presumably fed on other objects (bacteria or flagellates). Two other dominating species studied, Heterolaophonte minuta and Huntemannia jadensis, contained an average of 604 and 222 diatom cells per specimen. Diet composition differed significantly from the natural algal community. Two diatoms of intermediate cell size (Nitzschia palea var. debilis and Navicula sp.) contributed 92–97% of gut content for H. minuta and 58–81% for Hn. jadensis, whereas these diatoms amounted to only about 10% of biomass in native community. Mean consumption rates were estimated as 50–200 μg of wet biomass/day/cm², so the harpacticoids grazed only between 3 and 11% of the total microalgae biomass per day. The grazing impact on the two preferred diatom populations, however, was much more intensive, 10–30% per day for Navicula sp. and 55–228% for N. palea. Therefore, native harpacticoid populations demonstrate highly selective feeding and could be strongly limited by their food in spite of seemingly plentiful total abundance of microphytobenthos. This disproportionally high grazing pressure upon some species apparently could affect the structure of microalgae communities resulting in low relative abundance of mid-sized forms. We hypothesize that a very dynamic spatio-temporal distribution of epibenthic harpacticoids (short-living micropatches) may be the possible adaptation to such local food limitation.     

Functional morphology of the podia and ambulacral grooves of the comatulid crinoid Antedon bifida (Echinodermata)

Feeding units — viz. triplets of unequally-sized podia associated with protective lappets — occur all along the pinnules of adult Antedon bifida (Pennant). Small food particles are trapped by direct mucus impingement to the wall of primary and secondary podia (there are no mucus net or mucus thread helping in this process). Large particles are caught by primary and secondary podia which partly curl over them. Small particles accumulate on the collecting podia before being transferred to the groove, while large particles are transferred one by one. Transfer of particles to the groove occurs by wiping the collecting podia on the ciliary tracts against the ciliary current. When active, tertiary podia always paddle against the ciliary current. They serve in bolus formation (mucus embedding of food particles), but do not participate in bolus compaction or propulsion. Elimination of unwanted particles occur through the action of secondary podia whose movements may disrupt the lappet's palisade, thus creating a sideward current that moves particles from the groove to the outside. Typical podial triplets do not occur along brachial and calycinal grooves. Both brachial and calycinal podia function mostly in guiding and regulating particle flow. The feeding structures of early pentacrinoid larvae of Antedon bifida recall to mind those of pinnules of adult individuals. They consist of twenty-five podia arranged in five radial triplets alternating with five interradial pairs. They are similar to the pinnular feeding structures of adults in that they have both collecting (radial) and paddling (interradial) podia.     

A note on the feeding of Calanus helgolandicus on detritus

Experiments on the feeding of Calanus helgolandicus females on detrital material were carried out. Natural detritus from the ocean was never ingested, whereas dead diatoms were readily eaten. Furthermore, it was shown that C. helgolandicus females ingested large amounts of fecal material produced by the same species in other vessels. These results indicate that the copepod C. helgolandicus feeds not only on living organisms, but also on dead particles.     

On detritus as a food source for pelagic filter-feeders

Data on the amount of organic detritus within the particle size fraction 1 to 150 are presented for the Western Kiel Bight. Grouped into a mixed surface layer and a stratified lower layer in accordance with the main hydrographic features of the Western Baltic Sea, the rounded-off values show a seasonal variation between 100 and 600 mg m^-3 expressed as dry weight of organic matter. The overall average for both layers is about 200 mg m^-3. Organic detritus thus comprises more than 40% of total organic matter in the above size class, which is the class most easily accessible to the relatively small filter-feeders in this area. Attempting to trace the origin of organic detritus, a positive correlation to phytoplankton standing stock was found in some cases, suggesting the predominance of autochthonous detritus. Proceeding from the assumption that pelagic filter-feeders select their food mainly by size and not by taste, it is concluded that organic detritus plays an important role as a supplementary food source, being ingested together with phytoplankton and small nonmotile heterotrophs. The nutritive value of detritus is increased by the adsorption of dissolved organic matter and above all through the subsequent colonization by bacteria, which utilize the dissolved substances. Detritus particles serving as a substratum for bacteria thus form a means whereby dissolved organic substances reenter the food chain. The ingestion of detritus by filter-feeders is, therefore, thought to be instrumental in increasing the effectivity of energy transfer from the primary to the secondary food-chain level.     

Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients

Growth rates of juvenile Capitella sp. I were determined on different rations of six food types: Gerber's mixed cereal, TetraMin fish flakes, benthic diatoms, Ulva sp., spring detritus, and summer detritus. A simple growth model estimated maximum growth rate and maintenance ration for each food. There were differences in the growth response among foods relative to nitrogen content. As juveniles increased in size, differences in growth between foods became more pronounced. For all juveniles, growth rates were correlated with levels of the amino acids histidine, phenylalanine, threonine, and valine, and the polyunsaturated fatty acid 20:5w3; correlations with histidine and phenylalanine levels were the most significant. Regressions of growth rates as a function of these two amino acids suggest a daily maintenance ration of 300 pg histidine and phenylalanine mg^-1 nitrogen biomass. Juvenile worms grew on spring detritus but not on summer detritus, indicating the probable importance of micronutrients (polyunsaturated fatty acids, amino acids) for growth of juvenile Capitella sp. I in the field.     

Relationship between the habitat and diet of three species of atherinids and three species of gobies in a temperate Australian estuary

The distributions and diets of the six most abundant species of teleost in the shallows of a large south-western Australian estuary were examined from samples collected between March 1988 and February 1989. Fish were collected monthly by seine net from over bare sand and from within patchy and dense areas of the aquatic macrophyte Ruppia megacarpa. Their gut contents were compared with samples of the benthos and plankton collected from each of these three habitat types. The densities of the atherinids Leptatherina wallacei and Atherinosoma elongata and of the goby Favonigobius suppositus were greatest in dense R. megacarpa, whereas those of the atherinid L. presbyteroides and the goby F. lateralis were greatest over bare sand. The density of the goby Pseudogobius olorum was greater in patchy R. megacarpa than in the other two habitat types. The gut contents of each of the six species was dominated by crustaceans and/or polychaetes, with detritus also making a major contribution to the diet of P. olorum and F. lateralis. The relative proportions of prey items in the guts of fish in a particular habitat corresponded to the preys' relative occurrence in the environment. This indicates that the fish had been feeding predominantly in one particular habitat prior to capture. Within each habitat type, the six species partitioned the available food, the major components of the diet of each species being different. The gobiid species fed mainly on the benthos and the atherinids typically fed higher in the water column; A. elongata and P. olorum tended to be less selective as to where they foraged. There were no consistent differences in either the dietary breadth or the fullness of the guts of any species among habitat types.     

Food composition of crinoids (Crinoidea: Echinodermata) in relation to stalk length and fan density: their paleoecological implications

Crinoids have been diverse organisms in marine epifaunal filter feeding communities at any level of tiering above the substrate since they appeared in the Ordovician. Feeding is regarded as the most important factor in producing the crinoid tiering, which is primarily defined by stalk length. The gut contents of five sympatric crinoid species (three isocrines and two comatulids) were observed, and these were compared with the stalk length and the fan density. We have classified these crinoid species into four groups based on the stalk length and fan density, e.g., long stalk with low fan density, long stalk with high fan density, short stalk with low fan density, and short stalk with high fan density. In the gut contents, diatom crusts were found mainly from species with longer stalks, and chlorophyll-like fluorescent material were only detected from the groups with a shorter or no stalk. The group with lower fan density contained more inorganic particles than the group with higher fan density. Therefore, the gut contents and their amounts depend on their stalk lengths and their fan densities. The results imply that diversified morphologies in the crinoids have evolved through adaptations to different ecological factors such as difference in their diets.