The trophic status of herbivorous fishes on coral reefs

Estimates of feeding rates, alimentary tract structure and temporal patterns of food processing obtained from twelve species of nominally herbivorous fishes on the northern Great Barrier Reef were compared. These included members of the families Acanthuridae, Scaridae and Kyphosidae. Based on an analysis of diet and short-chain fatty acid (SCFA) profiles from a previous study we initially partitioned the twelve species into four dietary categories, as follows: (a) Category 1: herbivores with a diet of macroscopic brown algae and high SCFA profiles in the hindgut region (Naso unicornis, Kyphosus vaigiensis); (b) Category 2: herbivores feeding on turfing and filamentous red and green algae with moderate SCFA profiles in the hindgut region (N. tonganus, K. cinerascens, Zebrasoma scopas, Acanthurus lineatus); (c) Category 3: zooplankton feeders with moderate SCFA profiles (N. vlamingii, N. brevirostris); (d) Category 4: species feeding on detrital and sedimentary materials with low levels of SCFA (Chlorurus microrhinos, Scarus schlegeli, Ctenochaetus striatus, A. olivaceus). The purpose of this comparison was to determine whether measures of feeding activity, alimentary tract structure, and food processing were concordant with diet. A dichotomy in feeding rates was observed. Species with a diet of algae and zooplankton (categories 1–3) had slower feeding rates than those feeding on detrital aggregates and sediment (category 4). The pattern of food processing also followed the same dichotomy with species of categories 1–3 retaining food in the alimentary tract overnight and commencing the feeding day with substantial amounts of food in the intestine and hindgut. Category-4 species commenced the feeding day with empty alimentary tracts suggesting a rapid turnover of gut contents. Within the herbivorous and zooplankton-feeding species neither alimentary tract structure nor food processing mode were predicted by diet or SCFA profiles. A hindgut fermentation chamber was present in K. vaigiensis but not in N. unicornis, a species with high levels of SCFA in the hindgut region and a diet of brown macroscopic algae. In contrast N. vlamingii, with a diet dominated by animal matter, retained large amounts of food material in a hindgut chamber over the entire feeding cycle. In tropical perciform fishes, herbivory and fermentation are not associated with the alimentary tract structures that characterise herbivorous terrestrial vertebrates. Estimates of the abundance of the different groupings of nominally herbivorous fishes indicated that the dominant elements in the reef grazing and browsing fauna were consumers of detrital and sedimentary materials. These could not be classified as herbivores. Members of this group were dominant in all habitats investigated. Explicitly herbivorous taxa were a minority component in all habitats investigated.Communicated by G.F. Humphrey, Sydney     

A re-evaluation of the diel feeding hypothesis for marine herbivorous fishes

We re-evaluated the "diel feeding hypothesis" by measuring diel variation in starch, protein, and floridoside in three algal "types" collected from a fringing coral reef at Lizard Island, Great Barrier Reef, Australia. Samples of two species of rhodophyte algae, Gracilaria arcuata and Acanthophora spicifera, and the turf assemblage from the territories of the herbivorous pomacentrid Stegastes nigricans were collected at four time periods through the day: 0630-0730, 1000-1100, 1330-1430, and 1630-1730 hours. We also measured the ability of several species of marine fish (the herbivores Acanthurus nigricans, A. lineatus, A. olivaceus, and Parma alboscapularis and the detritivore Ctenochaetus striatus) to hydrolyse floridoside by estimating !-galactosidase activity in tissue from the anterior intestine. We detected no diel pattern in protein content of the algae but found a significant steady increase in starch content throughout the day. Floridoside content increased in the morning and decreased in the afternoon, a pattern that may be driven by midday photoinhibition of the algae. All the fishes tested could utilise floridoside. Our results support the diel feeding hypothesis. Although floridoside content decreased in the afternoon, our results suggest floridoside was used during the day by the algae to synthesise starch. Thus the algae increased in nutritional value until photoinhibition occurred at midday then subsequently maintained their nutritional value during the afternoon. This pattern of algal nutrients increasing to a midday peak and remaining relatively constant throughout the afternoon correlates well with the diel feeding pattern in many species of marine herbivorous fish.     

Genetic evolution among selected members of the genus Naso (Nasinae), "unicornfishes" from Guam

A species-level phylogeny of selected surgeonfishes in the genus Naso (Acanthuridae: subfamily Nasinae) was constructed using allozyme data and found to be concordant with previous inferences drawn from morphological data. Naso spp. examined from waters surrounding Guam between 1992 and 1994 were: N. caesius (Randall and Bell, 1992), N. hexacanthus (Bleeker, 1855), N. lituratus (Schneider, 1801), N. thynnoides (Valenciennes, 1835), and N. unicornis (Forskål, 1775). Acanthurus lineatus (Linnaeus, 1758) (Acanthuridae: subfamily Acanthurinae) and Zanclus cornutus (Linnaeus, 1758) (Zanclidae) were included for the purpose of outgroup comparison. Analyses of the allozyme data set generated from 16 presumptive loci using both cladistic and phenetic tree-constructing techniques identified the sibling species N. caesius and N. hexacanthus as the most genetically similar taxa among the Naso species surveyed. The pair shared the most synapomorphic alleles and the lowest degree of genetic divergence with respect to other species studied. The unweighted pair-group method using the arithmetic mean (UPGMA) and the neighbor-joining (NJ) topologies, along with two synapomorphic alleles (Aat-2 and Gpi-A), characterize N. caesius, N. hexacanthus, N. lituratus, and N. unicornis as a monophyletic group. Many allelic autapomorphies were detected in N. thynnoides, a member of the subgenus Axinurus. This supports the morphological data, which suggest that one-plated unicornfishes form a monophyletic group. The proposed partial phylogeny inferred from the allozyme data is concordant with the morphological differences (i.e. dentition, osteology, and myology) that have been used to support the partition of the genus Naso into the subgenera Axinurus and Naso.     

Gut characteristics and assimilation efficiencies in two species of herbivorous damselfishes (Pomacentridae: Stegastes dorsopunicans and S. planifrons)

The morphological and physiological mechanisms by which marine herbivores assimilate energy and nutrients from primary producers and transfer them to higher trophic levels of reef ecosystems are poorly understood. Two wide-ranging Caribbean fishes, the dusky damselfish, Stegastes dorsopunicans, and the threespot damselfish, S. planifrons, defend territories on patch reefs in the Archipelago de San Blas, Republic of Panama. We examined how relative intestine length and retention time influence digestion and absorption of energy and nutrients in these fishes. The dusky damselfish has a relative intestine length (RIL=intestine length/standard length) of 1.2 and a Zihler index {ZI=intestine length (mm)/10[mass(g)^1/3]} of 3.4. These values are significantly lower (P_RIL=P_ZI<0.0001) than those for the threespot damselfish (3.0 and 8.2, respectively). Both RIL and ZI for both species fall well below previously published values for other herbivorous pomacentrids, and may reflect their primary food resource at San Blas (diatoms). Energy-rich diatoms may be easier to digest than refractory macroalgae characteristic of diets of many herbivorous fishes (RIL range: 2-20). Despite differences in RIL and ZI between these two species, gut retention time is the same (P>0.05) for both dusky (6.6 h) and threespot damselfish (6.5 h). Thus, food travels the length of the threespot damselfish intestine ~2.5 times faster than it does in the dusky damselfish intestine. Levels of protein, carbohydrate, and lipid are significantly (0.003<P<0.030) higher in the feces of dusky damselfish than in the feces of threespot damselfish, when both species were fed a natural diet of benthic diatoms collected from damselfish territories. This indicates threespot damselfish have a greater nutrient-specific and total assimilation efficiency than do dusky damselfish. Furthermore, when fed an artificial pellet diet, protein absorption efficiency differed significantly (P=0.014) between species; threespot damselfish absorbed 98.3% of dietary protein, whereas dusky damselfish absorbed 96.4% of dietary protein.     

Demographic plasticity in tropical reef fishes

We use age-based analyses to demonstrate consistent differences in growth, mortality, and longevity of coral reef fishes from similar habitats (exposed reef crests) 20 km apart. On outer-shelf reef crests of the northern Great Barrier Reef (GBR), size in four taxa of reef fishes (Chlorurus sordidus, Scarus frenatus, and S. niger and the acanthurid Acanthurus lineatus) was systematically and significantly smaller when compared with the same taxa on adjacent mid-shelf reef crests. Differences in size could be attributed to differences in growth between habitats (shelf positions). On outer reef crests the species examined had consistently lower size at age profiles and also reduced life spans compared with populations from mid-shelf reefs. To confirm this relationship, two of the most abundant species (C. sordidus and S. frenatus) were selected for more detailed spatial analysis of demographic patterns. Sampling adults of both taxa from reef crests on three mid- and three outer-shelf reefs revealed that most of the variation in growth was explained by shelf position, although C. sordidus also displayed differences in growth among mid-shelf reefs. We conclude that differences in body sizes across the continental shelf of the northern GBR are primarily determined by these trends in growth. Strong spatial patterns also existed in the mean ages of populations and longevity estimates for C. sordidus and S. frenatus between shelf positions. Both species on outer-shelf reefs displayed less variable cohort sizes, significantly reduced mean ages, and foreshortened longevity compared with populations on mid-shelf reefs. Furthermore, differences in these parameters were rare among replicate reefs within mid- and outer-continental-shelf positions. Age-based catch curves suggested that rates of S. frenatus natural mortality on the outer shelf were nearly twice as high as on the mid shelf. Visual surveys indicated that total scarid densities on outer-shelf reef crests are on average fourfold higher than for equivalent mid-shelf habitats. This fact, coupled with significantly reduced growth rates, reduced mean ages, and increased mortality rates, suggests that density-dependent processes may be responsible for observed differences among localities.     

In situ feeding rates and grazing impact of zooplankton in a South African temporarily open estuary

Recent studies in temporarily open estuaries of South Africa have shown that phytoplankton biomass is at times low, when compared to the high standing stock of the grazers. In situ grazing rates of the dominant zooplankton species were estimated at the Mpenjati Estuary once during the winter closed phase, in August 1999, and once during the summer open phase, in February 2000. The study aimed at determining what proportion of the energetic demands of the dominant grazers of the estuary is met by the available phytoplankton. Results show that the gut of all species exhibited higher pigment concentrations during the night than during the day, both in winter and summer. Gut pigment contents ranged from 0.27 to 5.38 ng pigm individual^-1 in the mysid Gastrosaccus brevifissura, from 0.16 to 1.63 ng pigm individual^-1 in the copepod Pseudodiaptomus hessei, from 0.12 to 0.45 ng pigm individual^-1 in the copepod Acartia natalensis, and from 0.8 to 5.44 ng pigm individual^-1 in the caridean Palaemon sp. [where pigm is the sum of chlorophyll-a (chl-a) and phaeopigments]. During the winter closed phase, gut evacuation rates for G. brevifissura, P. hessei, and A. natalensis were 0.62, 0.42, and 0.46 h^-1, respectively. In summer, gut evacuation rates were 0.68, 0.48, and 0.46 h^-1 for G. brevifissura, P. hessei, and Palaemon sp., respectively. The rate of gut pigment destruction for G. brevifissura was 99.6% of the total ingested, one of the highest values ever recorded for any crustacean. A gut pigment destruction of 79.0% was measured for Palaemon sp., 95.7% for P. hessei, and 93.8% for A. natalensis. During winter the total grazing impact of the dominant zooplankton species ranged from 5.05 to 22.7 mg chl-a m^-2 day^-1 and accounted for 34-69% of the available chl-a in the water column. During summer, the grazing impact ranged between 0.45 and 0.65 mg chl-a m^-2 day^-1, accounting for 17-41% of the available chl-a in the water column. This shows that the dominant zooplankton species of the Mpenjati have a very high grazing impact on algal cells. At times this may exceed 100% of the available phytoplankton production, suggesting that the zooplankton community may often resort to other food sources to meet all its energetic demands.     

Feeding ecology of five fishes from the mid-slope micronekton community off southern Tasmania, Australia

We examine the feeding ecology of characteristic micronekton species inhabiting the mid-slope region (~700 to 1,500 m depths) off southern Tasmania. Five fishes, Diaphus danae, Hygophum hanseni, Lampanyctus australis (Myctophidae), Phosichthys argenteus (Phosichthyidae) and Chauliodus sloani (Chauliodontidae), were sampled by depth-stratified midwater trawling on a diel and seasonal basis. Overall, 74 prey taxa were identified from 2,132 stomachs. Euphausiids (mostly Euphausia spinifera and E. similis) and calanoid copepods (mostly Pleuromamma species) were the main prey of the three myctophids; P. argenteus ate fishes and decapods in addition to the euphausiids, while C. sloani ate only fishes. Copepods were less important in the diets of larger D. danae, L. australis and P. argenteus and were replaced by euphausiids in the myctophids and by fishes in P. argenteus. In autumn, when euphausiid biomass increased six-fold, all three myctophids and P. argenteus fed most intensively and consumed a high proportion of euphausiids. The three myctophids appeared to feed nocturnally. Differences in the timing and duration of feeding corresponded to differences in their spatio-temporal overlap with key prey. Daily rations of H. hanseni (1.93% dry-weight biomass) and L. australis (1.43%), estimated from data on stomach fullness, were typical for temperate myctophids and higher than that of the non-migratory P. argenteus (0.43%). The vertical flux of near-surface plankton production to the mesopelagic food web is based primarily on diel feeding in the upper water column (<500 m) rather than consumption of species that migrate seasonally into the deeper mesopelagic zone. Because species such as P. argenteus and C. sloani feed above the third trophic level, their predators, including the commercially important orange roughy, are feeding between levels four and five.     

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.     

Algal products as naturally occurring substrates for p-glycoprotein in Mytilus californianus

Mytilus californianus is a filter feeder that removes seaweed particulates, phytoplankton, and their byproducts from the water. The gills of this animal express high multixenobiotic resistance (MXR) and multixenobiotic transport activity that is related to the mammalian p-glycoprotein (p-gp). The high p-gp observed in mussel gills may provide the mussel protection from natural toxins in the diet. To test this hypothesis, extracts of various seaweeds and phytoplankton found in and around the mussel habitat were examined for presence of p-gp substrates by competition assays with a fluorescent dye substrate, rhodamine B. Accumulation of rhodamine was increased whereas dye efflux was slowed in the presence of algal extracts, indicating that p-gp substrates are present in the seaweeds Macrocystis pyrifera, Egregia menziesii, and Phyllospadix scouleri and the phytoplankton Alexandrium catanella and Pseudonitzchia australis. Initial fractionation of E. menziesii extracts by HPLC showed several distinct peaks of moderate hydrophobicity with p-gp-modulating ability. Additionally, Egregia extract showed potential as a chemosensitizer in tests with mussel (Mytilus edulis) and sea urchin (Lytechinus pictus) embryos. These data indicate that marine algae contain compounds that are substrates and/or chemosensitizers for the p-gp transporter in marine bivalves, thus providing evidence that MXR may have evolved in response to dietary pressures.     

A proteomic approach to the study of the marine mussels Mytilus edulis and M. galloprovincialis

In this study, a proteomic approach was applied for the generation of reference maps and subsequently to detect, quantify and compare the global protein expression between two related species of marine mussels, Mytilus edulis and Mytilus galloprovincialis, growing in their own geographical habitats. A comparative study of the protein profiles generated from analytical two-dimensional electrophoresis gels was performed, and changes in protein expression were analysed quantitatively by computer analysis. An average of 1,278 spots per gel was detected in 16 individuals (8 M. edulis and 8 M. galloprovincialis); however, not all spots were included in the study. Expression of 420 spots was compared, and significant differences in the intensity levels were detected in 37 protein spots (8.8%). Fifteen proteins showed higher expression in M. edulis, and 22 proteins, in M. galloprovincialis. The technique of peptide mass fingerprinting using MALDI-TOF (matrix-assisted laser desorption ionisation/time-of-flight) and/or nanoelectrospray double subfragmentation mass spectrometry enabled the unambiguous identification of 15 of these 37 differentially expressed proteins. Most of the identified proteins can be grouped basically into four broad functional classes: cytoskeletal and myofibrillar proteins, proteins associated with stress response, proteins associated with the storage or production of energy, and proteins related to rearrangement in the synthesis of native structures. These results expand our understanding of the molecular differentiation of the two mussel taxa and serve as a useful base for future ecological, physiological and genetic studies.     

Gut length and mass in herbivorous and carnivorous prickleback fishes (Teleostei: Stichaeidae): ontogenetic, dietary, and phylogenetic effects

Relative gut length, Zihler’s index, and relative gut mass were measured in four species of prickleback fishes and the effects of ontogeny, diet, and phylogeny on these gut dimensions were determined. Of the four species, Cebidichthys violaceus and Xiphister mucosus shift to herbivory with growth (>45 mm SL), whereas X. atropurpureus and Anoplarchus purpurescens remain carnivores. A. purpurescens belongs to a carnivorous clade, and the three other species belong to an adjacent, herbivorous clade. Gut dimensions were compared in three feeding categories of the four species: (1) small, wild-caught juveniles representing the carnivorous condition before two species shift to herbivory; (2) larger, wild-caught juveniles representing the natural diet condition of the two carnivores and the two species that have shifted to herbivory; and (3) larger, laboratory-raised juveniles produced by feeding a high-protein artificial diet to small juveniles until they have reached the size of the larger, wild-caught juveniles. Comparisons of gut dimensions in categories (1) versus (2) tested for an ontogenetic effect, in (2) versus (3) for a dietary effect, and within each category for a phylogenetic effect. C. violaceus and X. mucosus increased gut dimensions with increase in body size and did not change ontogenetic trajectory in gut dimensions on the high-protein artificial diet, suggesting that they are genetically programmed to develop relatively large guts associated with herbivory. X. atropurpureus increased its gut dimensions with increase in size similar to its sister taxon, X. mucosus, suggesting a phylogenetic influence, but decreased gut dimensions on the high-protein artificial diet, suggesting phenotypic plasticity. Nevertheless, X. atropurpureus displayed a larger gut than A. purpurescens, further evidence that it evolved in an herbivorous clade. A. purpurescens possessed a relatively small gut that was little affected by ontogeny or diet. Ontogeny and phylogeny more than diet appear to influence gut dimensions in the four species, thus favoring genetic adaptation over phenotypic plasticity as the major force acting on digestive system features in the two prickleback clades.     

Under-ice amphipods in the Greenland Sea and Fram Strait (Arctic): environmental controls and seasonal patterns below the pack ice

During three "Polarstern" cruises to the ice-covered Greenland Sea (spring 1997, summer 1994, autumn 1995) studies on the under-ice habitat (morphology, hydrography, ice-algal biomass) and on the macrofaunal, autochthonous under-ice amphipods (species diversity, abundance) were carried out in order to describe environmental controls and seasonal patterns in this community. In spring, the ice underside was rather smooth and whitish, while in summer melting structures and sloughed-off ice-algal threads were observed, in autumn detritus clumps accumulated in depressions at the ice underside. Only in summer, a thin layer of warm (up to -0.6°C) and less saline (as low as S=6.3) water was found at the ice-water interface above Polar Water. Integrated ice-algal biomass was highest during autumn (2.6 mg chl a m^-2) and lowest during summer (1.2 mg chl a m^-2). Four species of under-ice amphipods occurred in spring and summer (Apherusa glacialis, Onisimus glacialis, O. nanseni, Gammarus wilkitzkii), but only the last species was observed at the ice underside in autumn. A. glacialis and G. wilkitzkii were equally abundant in spring; A. glacialis dominated in summer. The highest total abundance of amphipods occurred during summer (31.9 ind. m^-2), compared to lower abundances in spring and autumn (5.3 and 1.1 ind. m^-2, respectively). A factor analysis revealed seasonal patterns in the data set, which mainly influenced A. glacialis, and species-specific relations between several environmental factors and the distribution of under-ice amphipods. Abundance of A. glacialis was closely related to the under-ice hydrography and ice-algal biomass, whereas the other amphipod species were more influenced by the under-ice morphology. It is therefore stated that the observed thinning of the Arctic sea ice and the resulting increased meltwater input and change in morphology of floes will have a profoundly adverse effect on the under-ice amphipods.     

Aphids are unaffected by the elemental defence of the nickel hyperaccumulator Streptanthus polygaloides (Brassicaceae)

Summary. Nickel hyperaccumulation, resulting in plant Ni contents of >1000 mg kg^у dry mass, has been shown to defend plants against folivorous herbivores. We determined whether this elemental defence tactic protected hyperaccumulating plants from attack by a phloem-feeding herbivore. We used the pea aphid, Acyrthosiphon pisum, and the Ni-hyperaccumulating plant Streptanthus polygaloides. Aphids were allowed to colonize mixed arrays of S. polygaloides in which plants either were hyperaccumulating Ni, not hyperaccumulating Ni and treated with a systemic insecticide, or not hyperaccumulating Ni. Aphid numbers g^у dry mass of plant biomass were lowest for the insecticide treatment, intermediate for low-Ni plants, and highest for plants hyperaccumulating Ni. Artificial liquid aphid diet, amended with varying levels of Ni, resulted in decreased aphid survival at 2500 mg kg^у Ni dry mass (or 5.03 mM Ni). We concluded that Ni levels in the phloem of hyperaccumulating plants of S. polygaloides were < 5.03 mM and, as a result, were not effective in defending plants against aphid attack.     

Assimilation of dietary free amino acids,peptides and protein in post-larval Atlantic halibut (<Emphasis Type="Italic">Hippoglossus</Emphasis><Emphasis Type="Italic">hippoglossus</Emphasis>)

With the aim of improving the formulation of diets for early stages of marine fish, post-larval Hippoglossus hippoglossus (56 days after first feeding; 23 mg dry weight) were tube-fed liquid diets of radiolabelled L-(¹⁴C) free amino acids (FAA diet), peptides (PEPT diet) and protein (PROT diet), in order to compare the kinetics of transfer from gut into the liver and carcass. The test diets were prepared using equal concentrations of amino acids (AA) and fed as a single pulse (0.3 µl, 2.2 µg AA). The PROT diet consisted of methylated-¹⁴C-labelled (mainly lysine) bovine serum albumin (BSA), the PEPT diet was the PROT diet digested with trypsin and the FAA diet consisted of an amino acid mixture based on BSA composition with added ¹⁴C-lysine. The common labelling with ¹⁴C-lysine permitted a comparison of the three diets. Based on dissection and distribution of the tracer in each compartment, the FAA diet was almost completely absorbed 1 h after the tube-feeding; PEPT and PROT diets required more time before they were processed by the digestive tract. The FAA diet was absorbed about eight times as fast as protein and about six times as fast as peptides from the gut. A low retention of label in the liver for all diets at all sampling times suggests that the liver does not store the AA at these absorption rates. The FAA diet had a higher assimilation efficiency (about 90%) than the PEPT diet (about 12%) and PROT diet (about 32%). FAAs, thus, appear to be superior to protein and peptides as a dietary source of amino acids in post-larval Atlantic halibut.     

Cross-shelf variation in calanoid copepod production during summer 1996 off the Oregon coast, USA

The fecundity of nine species of adult female calanoid copepods, and molting rates for copepodite stages of Calanus marshallae were measured in 24 h shipboard incubations from samples taken during the upwelling season off the Oregon coast. Hydrographic and chlorophyll measurements were made at approximately 300 stations, and living zooplankton were collected at 36 stations on the continental shelf (<150 m depth) and 37 stations offshore of the shelf (>150 m depth) for experimental work. In our experiments, maximum egg production rates (EPR) were observed only for Calanus pacificus and Pseudocalanus mimus, 65.7 and 3.9 eggs fem^-1 day^-1 respectively, about 95% of the maximum rates known from published laboratory observations. EPR of all other copepod species (e.g., C. marshallae, Acartia longiremis and Eucalanus californicus) ranged from 3% to 65% of maximum published rates. Fecundity was not significantly related to body weight or temperature, but was significantly correlated with chlorophyll a concentration for all species except Paracalanus parvus and A. longiremis. Copepod biomass and production in on-shelf waters was dominated by female P. mimus and C. marshallae, accounting for 93% of the adult biomass (3.1 mg C m^-3) and 81% of the adult production (0.19 mg C m^-3 day^-1). Biomass in the off-shelf environment was dominated by female E. californicus, P. mimus, and C. pacificus, accounting for 95% of the adult biomass (2.2 mg C m^-3) and 95% of the adult production (0.08 mg C m^-3 day^-1). Copepodite (C1-C5) production was estimated to be 2.1 mg C m^-3 day^-1 (on-shelf waters) and 1.2 mg C m^-3 day^-1 (off-shelf water). Total adult + juvenile production averaged 2.3 mg C m^-3 day^-1 (on-shelf waters) and 1.3 mg C m^-3 day^-1 (off-shelf waters). We compared our measured female weight-specific growth rates to those predicted from the empirical models of copepod growth rates of Huntley and Lopez [Am Nat (1992) 140:201-242] and Hirst and Lampitt [Mar Biol (1998) 132:247-257]. Most of our measured values were lower than those predicted from the equation of Huntley and Lopez. We found good agreement with Hirst and Lampitt for growth rates <0.10 day^-1 but found that their empirical equations underestimated growth at rates >0.10 day^-1. The mismatch with Hirst and Lampitt resulted because some of our species were growing at maximum rates whereas their composite empirical equations predict "global" averages that do not represent maximum growth rates.     

Fragmented seagrass habitats on the Mediterranean coast, and distribution and abundance of mysid assemblages

Four different habitats were sampled in order to determine the spatial and temporal distribution and abundance of mysids in a seagrass meadow ecosystem. These habitats included meadows of Posidonia oceanica and Cymodocea nodosa, the edge of P. oceanica meadow and sandy substrates. Three random sites for each habitat were sampled along the coast of Alicante at different times, using a hand-net technique (6 m²). Seven species of mysids were identified: Leptomysis posidoniae, Leptomysis buergii, Paramysis helleri, Siriella clausii, Mesopodopsis slabberi, Siriella armata and Mysidopsis gibbosa. Total abundance of mysids was highest at the edge of the P. oceanica meadow, followed by the C. nodosa meadow, sandy substrate and P. oceanica meadow, respectively (P<0.05); this pattern was found during all sampling periods except one. Three species showed persistent spatial segregation by habitat (S. clausii in Posidonia; L. posidoniae at the edge; S. armata in Cymodocea), whereas the other four did not show any clear segregation by habitat (ANOVA, P<0.01). On a site scale (km), spatial variability was very high. Multivariate analyses confirm the existence of differences in the assemblage structure among habitats, with a gradual change from the Posidonia meadow to sand. Posidonia and Cymodocea showed different mysid assemblages. Fragmented, shallow, vegetated habitat on the Mediterranean coast seems to support a higher abundance and species richness of mysids, through habitat diversification, in comparison with homogeneous habitats.     

Characterizing the resident, fermentative microbial consortium in the hindgut of the temperate-zone herbivorous fish, Hermosilla azurea (Teleostei: Kyphosidae)

The zebraperch, Hermosilla azurea Jenkins and Evermann, a warm-temperate marine fish species with a strictly macroalgal diet, has a relatively long digestive tract with an enlarged hindgut and an associated blind caecum (HC). In zebraperch sampled off Santa Catalina Island, California (33°19′42′′N; 118°18′37′′W) in years 1995 through 2001, direct cell counts, gut epithelium assessment of bacterial attachment, and short-chain fatty acid (SCFA) analyses verified that the zebraperch HC possesses a dense and morphologically diverse, fermentative microbiota. Bacterial cell counts and morphological diversity were significantly higher in HC contents compared to anterior gut regions, suggesting that microbial populations were growing along the digestive tract. Similarly, electron micrographs of the HC epithelium revealed attached microbes, further supporting the possibility that these organisms constitute resident microbiota. Five different SCFAs were detected in all three regions of the digestive tract, but levels were up to three times greater in HC contents. Acetate was consistently the prevailing SCFA in all gut regions. Sequence analysis of bacterial 16S rDNA was used to identify predominant bacterial groups in HC contents. Of the seven main bacterial types identified, Enterovibrio spp. were the dominant bacteria in HC contents followed by species of Bacteroides,Faecalibacterium, and Desulfovibrio. Taken together, our findings show that the zebraperch HC harbors a consortium of microbes that appears to assist in the breakdown of algal polysaccharides in the herbivorous diet of the fish.     

Dietary selectivity in the field and food preferences in the laboratory for two herbivorous fishes (Cebidichthys violaceus and Xiphister mucosus) from a temperate intertidal zone

Diets and food selectivity of two stichaeid fishes (Cebidichthys violaceus and Xiphister mucosus) from the rocky intertidal zone of the central California coast (USA) were studied at each season of the year by gut content analysis and abundance measurements of potential macrophyte food items. Both fishes, after reaching a standard length of about 44 mm, were almost exclusively herbivorous. The bulk of the diet consisted of 8 to 10 species of chlorophytes and rhodophytes. These main dietary components were chiefly annual seaweeds with high surfaceto-weight ratios (sheetlike forms or small, highly branched forms). Perennial seaweeds were eaten in relatively large amounts only during the winter. Macrophytes eaten in only trace amounts included about 20 species of chlorophytes, chrysophytes, phaeophytes, rhodophytes and a spermatophyte. The small amount of animal material in the diet (never more than 2% by weight) could well have been ingested incidentally while the fishes were feeding on seaweeds. Food preference tests with up to 19 macrophyte species in the laboratory revealed that both fishes chose to eat three annual rhodophytes (Smithora naiadum, Porphyra perforata and Microcladia coulteri) in preference to Ulva lobata, an annual chlorophyte that was more abundant in the diets of field-caught specimens.     

Seasonal fluctuations in the photosynthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef-building coral <Emphasis Type="Italic">Montastraea</Emphasis>

The photosynthetic capacity of photosystem II (PS II) in symbiotic dinoflagellates (Symbiodinium sp.), as measured by analysis of chlorophyll fluorescence, was investigated in the primary Caribbean reef-building corals, Montastraea annularis and Montastraea faveolata, for 5 years and Montastraea franksi over 2 years in the Bahamas. Significant seasonal fluctuations in the quantum yield of charge separation (F_v/F_m) of PS II were found in all species at all depths, with the highest photosynthetic capacity consistently recorded between mid-winter and early spring and the lowest photosynthetic capacity occurring in the mid to late summer. Corals residing in shallow depths of 1-2 m showed the greatest fluctuations in F_v/F_m, whereas deeper corals (3-4 and 14 m depths) had consistently higher values of F_v/F_m. Densities of symbiotic dinoflagellates and photosynthetic pigments followed a similar pattern. Fluctuations of photosynthetic capacity showed a strong correlation with seasonal patterns of water temperature and light. Such seasonal shifts in photosynthetic capacity are most likely due to several biochemical processes in the algae that lead to alterations of both photoprotection and photodamage. While symbiont density changed significantly on a seasonal basis, visual signs of coral bleaching were noted only in the fall of 1995 and the spring and summer of 1998. Comparisons of photosynthetic capacity and the decrease in the number of symbionts and their subsequent recovery indicated that symbiont populations in this study had the ability to recover quickly following bleaching events, as long as continued physical perturbation (e.g. thermal stress) did not shorten the recovery phase. Large-scale bleaching events are best viewed as the end points of seasonal physiological variation in which photosynthetic capacity and density of symbiotic dinoflagellates are reduced to a lower level than during "non-bleaching" years.     

Distribution of the alien ctenophore <Emphasis Type="Italic">Mnemiopsis</Emphasis><Emphasis Type="Italic">leidyi</Emphasis> in the Caspian Sea in August 2001

In this study, spatial and vertical distributions of the invasive ctenophore Mnemiopsis leidyi in the Caspian Sea were evaluated by using data collected at 41 stations during the August 2001 cruise. A comparison of data from different depths revealed that M. leidyi were generally confined to surface waters. The maximum size of the ctenophore was only 41-45 mm, and the bulk of individuals (85.5%) were <10 mm in length. The average and maximum biomasses of M. leidyi were calculated as 120 and 351 g wet weight m^-2, respectively. Whilst highest biomasses were observed in the western and central Middle Caspian Sea, hot spot areas of reproduction were present along the coasts of the western Caspian Sea, with abundance values of up to 2285 ind. m^-2. The impact of such high densities of M. leidyi is expected to be significant for the pelagic ecosystem of the Caspian Sea.