The trophic status of herbivorous fishes on coral reefs

Dietary analyses of 17 species of nominally herbivorous fishes on the northern Great Barrier Reef were carried out to resolve the questions to what extent diet in nominally herbivorous reef fishes is dominated by living plant material and whether short-chain fatty acid profiles in the alimentary tract predict diet. The fishes included members of the families Acanthuridae, Scaridae and Kyphosidae. The analyses revealed consumption of a wide range of food resources including macroscopic algae, turfing algae, pelagic and planktonic animal matter, organic detritus and sediments. In only 7 of the 17 species was living algae the dominant dietary item. Gastrointestinal short-chain fatty acids (SCFA) profiles reflecting the fermentative activities of gut symbionts were used as a framework for the initial analysis of dietary patterns. The feeding patterns implied by the SCFA profiles were validated by direct observation of food material through gut content analysis. We identified four main trophic groupings differentiated by fermentation profiles and diet. These were: (1) species with low levels of SCFA in gut samples (mean mM 9.2ǂ.8) but a high proportion of isovalerate (mean 15.4%dž.7): Acanthurus nigricauda, A. olivaceus, Ctenochaetus striatus, Chlorurus microrhinos, C.sordidus, Scarus schlegeli; (2) species that fed on turfing, filamentous and small thallate algae and displayed moderate levels of SCFA (mean mM 21.9DŽ.9): Acanthurus nigricans, A.lineatus, Naso tuberosus; (3) species that fed on large thallate (macroscopic) algae with high levels of SCFA (mean mM 41.2Dž.5) with very low proportions of isovalerate (mean 0.5%ǂ.1): Kyphosus cinerascens, K.vaigiensis, Naso unicornis, Zebrasoma scopas; and (4) species with diets dominated by planktonic animal material with moderate levels of SCFA (mean mM 21.3ǃ.5): Naso annulatus, N. brevirostris, N. hexacanthus, N. vlamingii. We conclude that a wide range of dietary items are taken by the 17 nominal herbivores, and that dietary groupings do not reflect taxonomic relationships. There was evidence of convergence in feeding modes and diet between phylogenetically distinct taxa and divergence within particular lineages.     

Comparison of herbivory in the closely-related marine fish genera Girella and Kyphosus

  Specimens of the nominally herbivorous, closely-related, fish genera Girella and Kyphosus were collected from Australian waters in 1994 and 1995. The diet of three Girella species (G. cyanea, G. elevata, and G. tricuspidata) consisted mainly of chlorophytes and rhodophytes, with an animal component of␣15.9 ± 4.2% in G. tricuspidata. The diet of four species of Kyphosus (K. bigibbus, K. cinerascens, K. sydneyanus, and K. vaigiensis) included phaeophytes, chlorophytes and rhodophytes, and almost no animal material. Concentration of total short-chain fatty acids in the posterior intestine was <11.4 mM in the Girella spp. and >39.2 mM in the Kyphosus spp. These results suggest that microbial fermentation plays a role in algal digestion in Kyphosus spp., but not in Girella spp. Girellids and kyphosids appear to function quite differently as herbivores. Girellids should be considered as omnivores that complement readily-available energy from algae with protein from invertebrates. Kyphosids appear to be strict herbivores that can derive adequate nutrition from algae poor in easily assimilable energy, through microbial fermentation in the hindgut. Received: 8 July 1996 / Accepted: 2 August 1996     

Chemoautotrophic symbiosis in the tropical clamSolemya occidentalis (Bivalvia: Protobranchia): ultrastructural and phylogenetic analysis

Patterns of algal seasonality, and their effect on the diet and feeding preferences of the herbivorous crab Grapsus albolineatus were investigated over an 18-mo period from March 1993 to August 1994 on an exposed tropical rocky shore (Hok Tsui Peninsula at Cape d' Aguilar, Hong Kong). Algal cover was greatest in the winter months, and lowest in the summer. Foliose algae such as Ulva fasciata, Porphyra suborbiculata, and Dermonema frappieri were dominant in the winter, but died off in the summer. During the hot summer months, perennial encrusting algae e.g., Ralfsia expansa, Hildenbrandia rubra, H. occidentalis, coralline crusts and the encrusting cyanobacteria Kyrtuthrix maculans, were the dominant algal species. Seasonal variation in algal abundance influenced the dietary selectivity of the herbivorous crab G. albolineatus. In the winter, the crab fed selectively on filamentous algae (e.g. Hincksia spp., Cladophora spp., Enteromorpha spp., and the cyanobacteria Lyngbya sp.). Foliose algae (e.g. U. fasciata, P. suborbiculata, Pterocladia tenuis) formed a small part of the diet, despite being the dominant species on the shore. Foliose and filamentous algae were virtually absent from the shore in the summer, and the crabs switched to feeding solely on encrusting algae. Electivity indices revealed preferences for green and brown turf species, and avoidance of foliose algae. Faecal analysis revealed that a greater proportion of the food is digested in the winter, suggesting that G. albolineatus is able to digest filamentous algae more efficiently than encrusting algae. Feeding preferences of G. albolineatus appear to be influenced by a number of factors, including the availability, digestibility and morphology of algae. The foraging behaviour and cheliped morphology of the crab also affect food choice. The monsoonal nature of Hong Kong's climate controls the diversity and abundance of intertidal algae and, therefore, indirectly influences the diet and subsequent growth and reproductive success of the herbivorous crab G. albolineatus.     

Scale-dependent patterns of variability of a grazing parrotfish (<Emphasis Type="Italic">Leptoscarus vaigiensis</Emphasis>) in a tropical seagrass-dominated seascape

Although herbivorous fish form critical linkages between primary producers and higher trophic levels, the major factors regulating their spatial structure in seagrass systems remain poorly understood. The present study examined the parrotfish Leptoscarus vaigiensis in seagrass meadows of a tropical embayment in the western Indian Ocean. Stomach content analysis and direct field observations showed that L. vaigiensis is an efficient grazer, feeding almost exclusively on seagrass leaves. Seagrass shoot density was highly correlated to all density variables (total, juvenile and subadult) and juvenile biomass of L. vaigiensis, while subadult biomass was predicted by distance to neighbouring coral habitat. Moreover, density and biomass of predatory fish (piscivores) were predicted by seagrass canopy height and the distribution patterns of predators followed those of L. vaigiensis. Hence, factors at local (seagrass structural complexity and feeding mode) and landscape scale levels (seascape context and distribution of piscivores) likely mutually structure herbivorous fish communities. The findings underscore the importance of incorporating multiple scale-dependent factors when managing coastal seagrass ecosystems and their associated key species.     

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.     

Food habits, gut morphology and pH, and assimilation efficiency of the zebraperch Hermosilla azurea, an herbivorous kyphosid fish of temperate marine waters

Adult zebraperch, Hermosilla azurea, were found to be functional herbivores in that animal matter constituted <0.01% of the total dry weight of stomach contents of fish collected off Santa Catalina Island in southern California waters. The diet of these fish consisted mainly of red algae (88.2% by dry wt) and also small amounts of brown (7.8%) and green (4.0%) algae. The most important dietary item, the filamentous red algae Polysiphonia spp., was found in >78% of the stomachs and comprised >60% of the contents by dry weight. The digestive tract was long, on average 4.0 times the standard length of the fish, and was composed of the stomach, pyloric caeca, intestine, hindgut chamber with a blind caecum, and rectum. The mean pH of the cardiac stomach was acidic (3.9), whereas that of the intestine was nearly neutral (6.9) and that of the hindgut and blind caecum slightly acidic (6.3 and 6.6, respectively). Algal foods are apparently digested by acid lysis in the stomach and by microbial fermentation in the hindgut. Zebraperch assimilated nutritional constituents from six species of algae with varying degrees of efficiency: carbon (73.7 to 89.7%), nitrogen (72.4 to 84.5%), and protein (71.9 to 94.9%). The fish assimilated these constituents as efficiently or more efficiently from three species of nondietary brown algae as from three species of dietary red and green algae. These results show that zebraperch, like their tropical and subtropical relatives (members of the genus Kyphosus), can digest a wide variety of algae including brown algae containing defensive secondary compounds. Received: 3 November 1997 / Accepted: 19 June 1998     

Trophic ecology of coral reef gobies: interspecific,ontogenetic, and seasonal comparison of diet and feeding intensity

In fishes, a small body size may facilitate cost-effective exploitation of various primary and secondary food resources, but may pose difficulties associated with digestion of plant material and finding sufficient food in a foraging area potentially restricted by a high risk of predation. We examined the trophic ecology of five common, small-bodied coral reef fish from the family Gobiidae. For each species, we determined diet composition, feeding bite rate, foraging substrate, and feeding behaviour, and examined whether diet composition and feeding bite rate changed ontogenetically and seasonally. The five species showed a diverse range of trophic modes: Amblygobius bynoensis and Amblygobius phalaena were herbivores, Valenciennea muralis was a carnivore, Asterropteryx semipunctatus a detritivore, and Istigobius goldmanni an omnivore. Both the herbivores and detritivore supplemented their diet with animal material. The consumption of a wide range of dietary resources by the two smallest species with the most restricted mobility (A. semipunctatus and I. goldmanni) may ensure energy requirements are met within a restricted foraging area. There was a significant difference in mean feeding bite rate among species, with carnivore > herbivore > omnivore > detritivore. None of the species exhibited an ontogenetic shift in diet composition or increase in feeding bite rate, indicating that (1) postmaturation growth is not facilitated by a higher quality diet or increased feeding intensity following maturation, and (2) their small body size does not preclude herbivory. The herbivores had the highest gut:fish length ratio, which may facilitate plant digestion. While diet did not change seasonally, the mean feeding bite rate was significantly lower in winter than summer for four of the study species.     

Atypical plant–herbivore association of algal food and a kleptoplastic sea slug (Elysia clarki) revealed by DNA barcoding and field surveys

The identity of food sources and feeding preferences of specialist herbivores have been commonly inferred from spatial associations between consumer and food items. However, such basic information for well-known marine herbivores, sacoglossans (sea slugs), and their algal diets remains disappointingly lacking, especially from field studies. The sacoglossan, Elysia clarki (Pierce et al. in Molluscan Res 26:23–38, 2006), is kleptoplastic and sequesters chloroplasts from algal food to photosynthesize, so DNA identification of sequestered chloroplasts was employed to verify the algal species fed upon by the slug across its geographic range. The molecular information on the algae consumed by E. clarki was combined with field surveys of slugs and algae in slug habitats in the Florida Keys in July and August of 2008 in order to evaluate whether the diet of this herbivore could be predicted based on its spatial association with algae in the field. A considerable mismatch between food availability and kleptoplast identity was recorded. E. clarki commonly occupied areas devoid of potential food and often contained symbiotic plastids from algal species different from those most frequently found in the surveyed habitats. In three of the four study sites, algal species present were poor predictors of slug diet. These findings suggest that the photosynthetic capability of E. clarki may release the slug from the constraint of requiring proximity to its food sources and may allow for the potential lack of spatial coupling between this herbivore and its algal food. This combination of field surveys and DNA barcoding provided critical and previously unavailable information on herbivore feeding in this marine system.     

Comparing macroalgal food and habitat choice in sympatric, tube-building amphipods, Ampithoe lacertosa and Peramphithoe humeralis

For small tube-building amphipods that live on the algae they consume, food and habitat are tightly linked. This study compared two closely related amphipods to determine whether the species’ algal preferences are based on the food value of the algae or on some other aspect of their algal habitat. Ampithoe lacertosa and Peramphithoe humeralis are both abundant on Shannon Point beach (Anacortes, Washington, USA; 48°30.542′ N, 122°41.070′ W) but specialize on different algae. In observations and laboratory experiments conducted July–September 1997, 2007, and 2008, the two species exhibited markedly different choices of food and habitat when offered six common macroalgae. Ampithoe lacertosa ate all algae offered, but preferentially built tubes on the green alga Ulva lactuca. Survival was relatively low among juveniles maintained on single species diets, except when they were fed Mazzaella splendens. Conversely, P. humeralis consumed primarily the brown kelp Saccharina latissima, Alaria marginata, and Desmarestia ligulata and preferred those species for tube building. Juvenile P. humeralis could not survive on a diet of U. lactuca or M. splendens. While A. lacertosa builds simple, temporary tubes and relocates frequently, P. humeralis is a highly thigmotactic species that builds long-term, complex tubes on the alga it prefers to eat. Feeding and habitat preferences of the two species were not clearly linked to nitrogen content of the algae, C:N ratio, or toughness of the algal tissue. Instead, preferences of the species may be related to their mobility and the permanence of the tubes they build. Ampithoe lacertosa and P. humeralis also use different feeding strategies; the former appears to mix algae to produce a high-quality diet, while the latter is more selective and has a capacity for compensatory feeding. The species are abundant on the same protected rocky shores, but specialize on different algae for habitat and food. Results suggest that the nutritional requirements of these sympatric mesograzers differ considerably and even closely related species can exhibit divergent behavioral strategies.     

Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

Sources of variation in herbivore preference: among-individual and past diet effects on amphipod host choice

Understanding which factors affect the feeding preferences of herbivores is essential for predicting the effects of herbivores on plant assemblages and the evolution of plant–herbivore interactions. Most studies of marine herbivory have focussed on the plant traits that determine preferences (especially secondary metabolites), while few studies have considered how preferences may vary among individual herbivores due to genetic or environmental sources of variation. Such intraspecific variation is essential for evolutionary change in preference behaviour and may alter the outcome of plant–herbivore interactions. In an abundant marine herbivore, we determined the relative importance of among-individual and environmental effects on preferences for three host algae of varying quality. Repeated preference assays were conducted with the amphipod Peramphithoe parmerong and three of its brown algal hosts: Sargassum linearifolium, S. vestitum and Padina crassa. We found no evidence that preference varied among individuals, thus constraining the ability of natural selection to promote increased specialisation on high-quality S. linearifolium. Most of the variation in preference occurred within individuals, with amphipod preferences strongly influenced by past diet. The increased tendency for amphipods to select alternate hosts to that on which they had been recently feeding indicates that amphipods are actively seeking mixed diets. Such a feeding strategy provides an explanation for the persistence of this herbivore on hosts in the field that support poor growth and survival if consumed alone. The effects of past diet indicate that herbivore preferences are a function of herbivore history in addition to plant traits and are likely to vary with the availability of algae in space and time.     

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.     

Increases in surgeonfish populations after mass mortality of the sea urchinDiadema antillarum in Panamá indicate food limitation

In 1983/1984,Diadema antillarum suffered mass mortalities throughout its West Atlantic range. Its populations were reduced by 95% and subsequently have failed to recover. These die-offs led to sustained increases in the abundance of soft algae, including types eaten by herbivorous reef fishes. I monitored adult populations of three herbivorous surgeonfishes (Acanthurus coeruleus, A. chirurugus andA. bahianus) between 1978 and 1990, and the recruitment of their pelagic juveniles between 1979 and 1989, on six patch reefs in Panamá. Adult populations ofA. coeruleus andA. chirurgus, which largely restrict their feeding to reef substrata, increased by averages of 250 and 160%, respectively, after the die-off ofD. antillarum in 1983. No increases occurred in the adult populations ofA. bahianus, which often feeds in off-reef habitats unaffected byD. antillarum. Average annual levels of juvenile recruitment of all three surgeonfishes did not differ before and after the die-off. These results support the hypothesis that adult populations of two herbivorous fishes that are strongly reliant on reef algae for food previously were limited by competition withD. antillarum.     

Fish guts as chemical reactors: a model of the alimentary canals of marine herbivorous fishes

The alimentary canals of marine herbivorous fishes are variously composed of a few basic structures strung together in series. We model the structures where digestive processes occur as chemical reactors following the model of Penry and Jumars: the stomach as a batch or continuous-flow stirred-tank (CSTR), the intestine as a plug-flow reactor (PFR), and the hindgut caecum as a CSTR. Other structures, where food is mechanically processed-gill rakers, pharyngeal mills, and muscular stomachs-are classified as gates. The optimality criterion for the model is the digestion of the most nutrient in the least amount of time. With the model we are able to predict gut configuration as a function of nutrient concentration and hypothesize that the guts of herbivorous fishes always have a PFR component and may or may not have a CSTR component. The Penry-Jumars model appears to provide a consistent theoretical framework for four main types of digestive mechanisms in marine herbivorous fishes and offers specific testable hypotheses on the feeding ecology and digestive physiology of four representative species of fish as well as other, still unstudied, herbivorous fishes.Contribution No. 68 from the Ocean Studies Institute, Long Beach, California 90840, USA     

Foraging by the herbivorous parrotfish Sparisoma radians

The foraging behavior of the bucktooth parrotfish Sparisoma radians was studied in seagrass beds off St. Croix, US Virgin Islands and in laboratory preference tests. Thalassia testudinum was the dominant item in the field diet with the epiphytized distal portion of the blades most favored. Other seagrasses, Syringodium filiforme and Halodule wrightii, were taken in relation to their abundance. Several algae were also eaten, particularly Halimeda spp. and Penicillus spp. Although abundances of the algae varied, the fish maintained a steady mixture of plant species in their diet. Laboratory feeding tests for various plants presented in pairs showed clear preferences in nearly all cases. The preference hierarchy was (1) T. testudinum with epiphytes, (2) H. wrightii, (3) T. testudinum without epiphytes, (4) S. filiforme, and the algae, (5) Dictyota divaricata, (6) Enteromorpha flexuosa, (7) Caulerpa mexicana, (8) Halimeda incrassata and (9) Penicillis pyriformis. Preference did not vary significantly with satiation. The catch per unit effort (kilocalories absorbed per bite) was calculated for each plant, using (a) absorption values calculated for S. radians fed the different plants, (b) the calorific values for tested plants, and (c) an estimate of the amount of material taken per bite. The ranking of catch per unit effort closely paralleled the preference hierarchy with the exception of C. mexicana which has a toxin. Fish fed diets of single plants, mixed plant diet, and starved controls showed differential survival which paralleled the preference hierarchy, the most preferred plants leading to longest survival. Comparison of laboratory results with field feeding behavior shows that inclusion of plants in the diet is not related directly to preference rank, availability or survival value, but that the fish deliberately eat a variety of plants presumably which maintains balanced diet. This result indicates that models of optimal foraging for herbivores should include nutrient constraints and avoidance of toxins in order to predict accurately the behavior of an animal in the field.     

Effect of food concentration and type of diet on <Emphasis Type="Italic">Acartia </Emphasis>survival and naupliar development

We have performed life table experiments to investigate the effects of different food types and concentrations on the larval development and survival up to adulthood of Acartia tonsa. The food species offered comprised a wide taxonomic spectrum: the pigmented flagellates Isochrysis galbana, Emiliania huxleyi, Rhodomonas sp., Prorocentrum minimum, the diatom Thalassiosira weissflogii, grown on medium offering enriched macronutrient concentrations and the ciliate Euplotes sp. initially cultured on Rhodomonas. For the ciliate species, also the functional response was studied. In order to avoid limitation by mineral nutrients, food algae have been taken from the exponential growth phase of the nutrient replete cultures. The suitability of Rhodomonas as a food source throughout the entire life cycle was not a surprise. However, in contrast to much of the recent literature about the inadequacy or even toxicity of diatoms, we found that also Thalassiosira could support Acartia-development through the entire life cycle. On the other hand, Acartia could not complete its life cycle when fed with the other food items, Prorocentrum having adverse effects even when mixed with Rhodomonas and Thalassiosira. Isochrysis well supported naupliar survival and development, but was insufficient to support further development until reproduction. With Emiliania and Euplotes, nauplii died off before most of them could reach the first copepodite stages. Acartia-nauplii showed a behavioral preference for Euplotes-feeding over diatom feeding, but nevertheless Euplotes was an insufficient diet to complete development beyond the naupliar stages.     

Providing a common diet to different marine decapods does not standardize the fatty acid profiles of their larvae: a warning sign for experimentation using invertebrate larvae produced in captivity

Larval decapods are commonly produced in captivity and employed in experiments to evaluate interspecific physiological and biochemical differences. Currently, it is still unknown if different decapod species provided a common diet and exposed to identical abiotic conditions produce newly hatched larvae (NHL) with similar fatty acid (FA) profiles. This study analyzed the FA composition of NHL from five marine shrimp species (Lysmata amboinensis, L. boggessi, L. debelius, L. seticaudata and Rhynchocinetes durbanensis) fed a common diet and stocked at constant temperature. FA profiles of NHL differed significantly within and among genera. NHL from species unable to molt from zoea I to zoea II in the absence of food (L. amboinensis, L. debelius and R. durbanensis) displayed the lowest FA contents. Researchers must be aware that providing a common diet to different species, even if closely related, may not standardize the FA profile of NHL produced in captivity.     

Strain-related physiological and behavioral effects of <Emphasis Type="Italic">Skeletonema marinoi</Emphasis> on three common planktonic copepods

Three strains of the chain-forming diatom Skeletonema marinoi, differing in their production of polyunsaturated aldehydes (PUA) and nutritional food components, were used in experiments on feeding, egg production, hatching success, pellet production, and behavior of three common planktonic copepods: Acartia tonsa, Pseudocalanus elongatus, and Temora longicornis. The three different diatom strains (9B, 1G, and 7J) induced widely different effects on Acartia tonsa physiology, and the 9B strain induced different effects for the three copepods. In contrast, different strains induced no or small alterations in the distribution, swimming behavior, and turning frequency of the copepods. 22:6(n-3) fatty acid (DHA) and sterol content of the diet typically showed a positive effect on either egg production (A. tonsa) or hatching success (P. elongatus), while other measured compounds (PUA, other long-chain polyunsaturated fatty acids) of the algae had no obvious effects. Our results demonstrate that differences between strains of a given diatom species can generate effects on copepod physiology, which are as large as those induced by different algae species or groups. This emphasizes the need to identify the specific characteristics of local diatoms together with the interacting effects of different mineral, biochemical, and toxic compounds and their potential implications on different copepod species.     

Effects of various diets on the nutritional value of Artemia sp. as food for the prawn Penaeus monodon

The nutritional value of Artemia sp. as food for marine fish and crustacean larvae has been linked to the level of its polyunsaturated fatty acid (PUFA) content. Experiments in August 1984 were conducted to determine the effects of various artificial diets and algae on fatty acid composition of PUFA-deficient Artemia sp. (Utah GSL strain) and their resulting value as food for postlarvae of the prawn Penaeus monodon (Fabricius). Nauplii of the brine shrimp were grown on extracts of corn, copra, soybean and rice bran containing precursors (C18) to long-chain PUFA and also on algal species containing different levels of long-chain PUFA (C20). The nauplii were then used as food for P. monodon postlarvae. The results revealed that absence of C20 polyunsaturates from the feeds and their presence in the algae were reflected in the polyunsaturated fatty acid content of the tissues of Artemia sp. When fed with brine shrimp fed on algae, P. monodon displayed better postlarval survival and significantly higher growth; related to the content of polyunsaturated fatty acids in Artemia sp. A practical feeding approach in prawn hatcheries would be to grow Artemia sp. on a cheap diet such as rice bran, and then to enhance its nutritional value with a diet high in PUFA prior to harvesting, in order to improve hatchery production.     

Relation of food preference to fitness for the green sea urchin,Strongylocentrotus droebachiensis

We maintained 30 to 35 mm sea urchins (Strongylocentrotus droebachiensis) on 17 different macroalgal diets in the field over a 130-d period and correlated the resulting changes in somatic and gonadal tissues with measures of food preferences. The varions algae fell into there distinct categories according to their ability to support growth. The most striking effect of diet was on gonadal mass, there being a 14-fold increase for the best algal diet, compared to initial gonadal mass, and nearly a threefold decrease for the worst diet. The relation of food preference to somatic growth was nonlinear. A sharp increase in the growth of test diameter, total mass and test mass occurred between the nonpreferred and intermediately preferred algae, but growth rates were similar for intermediate and preferred algae. Gonadal growth tended to increase exponentially with an important measure of food preference, feeding rate (g ingested d^–1). These observations suggested that urchins only channel food resources into somatic growth up to a given point, additional reserves being used for producing gametes. The positive relationship of food preference to growth, and especially the strong correlation with gonadal production, indicates that the highly selective feeding of the urchin contributes to ils fitness.