This project evaluated the feasibility of salvaging zebra mussel-infested freshwater mussels from their native habitat by determining 1) how zebra mussel infestation affects unionid feeding and body condition, 2) how starvation in quarantine affects body condition of unionids, and 3) what feeding regime maintain unionid condition in quarantine. The effects of zebra mussel infestation on two mussel species were evaluated through glycogen analyses of mantle tissue and gut content analysis. Specimens of Amblema p. plicata (Say, 1817) and Quadrula p. pustulosa (I. Lea, 1831), collected from a heavily infested reach of the Ohio River in 1996, had significantly lower (p<0.05) glycogen levels than specimens collected from a low-infestation reach upstream. In 1996 and 1997, heavily infested Amblema p. plicata and Quadrula p. pustulosa had significantly less (p<0.01) organic matter and fewer algal cells in their guts than lightly infested specimens. In addition, gut contents of individual A. p. plicata contained significantly less (p<0.05) organic matter and fewer algal cells than the combined gut contents of all zebra mussels (18-33 mm in length) attached to their shells. Gut analyses also revealed significant diet overlap between native unionids and infesting zebra mussels. Thus, competitive interactions or interference by zebra mussels likely reduced unionid ingestion and consequently reduced glycogen stores. During quarantine, unionids salvaged from a lightly infested area and starved for 30 days had glycogen levels that declined dramatically. After 30 days without supplemental feeding, mean glycogen levels of A. p. plicata declined 85%, and mean glycogen levels of Q. p. pustulosa declined 70%. Thus, feeding of unionids is necessary to maintain their condition during lengthy quarantine. To determine the best feeding regime for unionids in quarantine, assimilation efficiencies and carbon budgets were established for the rainbow mussel, Villosa iris (Lea, 1829), using radio-labelled cultures of Neochloris oleoabundans (Chantanachat and Bold 1962) at three cell concentrations. Assimilation efficiencies for Villosa iris at 100,000cells/ml, 10,000 cells/ml, and 1,000 cells/ml were similar (45-56%); however, regardless of these similarities, assimilation efficiencies from this study indicate that Neochloris oleoabundans is readily assimilated (~50% AE) by Villosa iris . In addition, total assimilation was maximized at 100,000cells/ml, which indicates that Villosa iris has the greatest amount of energy available for growth, reproduction, and maintenance of condition in captivity at this cell concentration. During a second quarantine experiment, specimens were provided with 100,000cells/ml of N. oleoabundans twice per day. Initial mean glycogen levels for Amblema p. plicata (9.4 + 2.4 mg/g) and Quadrula p. pustulosa collected from ORM 175.5 in July 1997 were not significantly different (p>0.3) than the mean glycogen levels of A. p. plicata and Q. p. pustulosa collected from the same site in July 1996. Glycogen stores of unionids entering quarantine, therefore, were similar in both the starvation and controlled feeding experiments. After 7, 14, and 30 days of controlled feeding in quarantine, mean glycogen levels of A. p. plicata and Q. p. pustulosa did not change significantly (p>0.1). Thus, quarantine protocol for salvaged native mussels should include the feeding of algae to captive specimens to sustain glycogen levels prior to relocation. / Master of Science

The main objective of the diploma thesis was defined as the consideration of the hypothesis that food intake structure of adult roach changes significantly during the spawning period. The practical experiment was conducted to reach the defined aim. The experiment was performed on the Brno and Hamry Reservoirs in 2008 and 2009. The acquired data were analysed by standard tools used for gut contents analyses (Indirect Method, Frequency of Occurrence, Index of Fullness, Index of Preponderance) related to the Gonadosomatic Index.

Black Carp (Mylopharyngodon piceus) is an invasive fish species native to Asia that has become increasingly abundant within the Mississippi River Basin during the past decade. Originally introduced to control snails that are an intermediate host for trematode parasites of fishes in aquaculture ponds, Black Carp are now present in several rivers in the U.S. and represent a potential threat to threatened and endangered mussel populations. Black Carp have historically been classified as molluscivores; however, a recent study that examined gut contents of Black Carp collected from the Mississippi River Basin indicated that Black Carp are opportunistic consumers that prey upon a wide variety of invertebrates and are flexible in their feeding modes. Despite the potential for Black Carp to compete with native riverine fish species for invertebrate prey, only one published study has compared Black Carp trophic position with that of native fishes in a small portion of the Black Carp’s invaded range. Therefore, the objectives of this study were to assess trophic overlap between Black Carp and two fish species native to the Mississippi River Basin using isotopic niche analysis and gut contents analysis. Dorsal muscle tissue samples were collected from Black Carp, Freshwater Drum (Aplodinotus grunniens), and Blue Catfish (Ictalurus furcatus) and analyzed for δ13C and δ15N to assess each species’ isotopic niche. Freshwater Drum and Blue Catfish gut contents were also removed and analyzed and compared to published Black Carp stomach contents data. Gut contents analysis indicated differences in diet composition between Black Carp and the two native fish species. Chironomidae had the highest frequency of occurrence (67%) and percent of taxa by number (47%) for Freshwater Drum. Trichoptera had the highest frequency of occurrence (58%) and percent of taxa by number (30%) for Blue Catfish, and Gastropoda had the highest frequency of occurrence (16.5%) of any specific prey taxa for Black Carp. Black Carp showed low isotopic niche overlap (≤47%) with both native species when muscle tissue δ13C and δ15N data from all fish collection locations were combined and when assessment of isotopic niches was restricted to the subset of locations where all three species were collected. Isotopic niche overlap was also low (10-48%) between Black Carp and both native species when isotopic niches were compared at individual collection locations. Intraspecific isotopic niche overlap among fish collection locations was highly variable (0-69%) within each of the three species, highlighting the need to assess interspecific isotopic niche overlap by collection location. Broad isotopic niches exhibited by Black Carp in the Mississippi River and tributaries are indicative of substantial trophic diversity among individuals and use of multiple basal energy sources, consistent with a recently published study which found that Black Carp diet composition differed among individuals and that Black Carp consumed a variety of invertebrates, including non-benthic taxa.

Originalmente, a teoria de nicho ecológico fundamenta-se em indivíduos de uma espécie sendo ecologicamente equivalentes por utilizarem os recursos de forma similar. Portanto, o nicho de uma espécie é definido em termos do uso médio de recursos. Porém, a qualidade e abundância de recursos consumidos, sexo, idade ou morfotipo do consumidor influenciam o comportamento alimentar individual. Estudos recentes sobre forrageamento ótimo tem foco de interesse na variação interindividual no uso de recursos. Ao longo da sua área de distribuição são reconhecidas populações da toninha Pontoporia blainvillei (Cetacea) e no litoral paulista, possivelmente hajam três subpopulações dessa espécie: norte, centro e sul. Meu primeiro objetivo foi investigar se há variação na dieta entre essas três subpopulações de toninha. Adicionalmente, investiguei quais fatores entre sazonalidade, sexo e idade dos indivíduos poderiam estruturar a dieta em uma dessas subpopulações (central). Utilizei 58 indivíduos provenientes de captura acidental no litoral paulista, com representantes das regiões norte, centro e sul. A amostra incluiu juvenis e adultos de ambos os sexos, capturados em todas as estações do ano. Identifiquei as espécies nos conteúdos estomacais usando os otólitos de peixes e os bicos de lulas. Estimei o tamanho das presas utilizando regressões com as medidas dessas estruturas e investiguei a estruturação da dieta usando um índice de variação da dieta. Houve uma clara variação na dieta ente as subpopulações paulistas, possivelmente devido à diferença espacial na disponibilidade de presas. Para a subpopulação central não houve variação na dieta decorrente da sazonalidade ou do sexo. A ausência de variação sazonal pode ser devido à pouca alteração na abundância da principal presa, P. harroweri, no ambiente ao longo do ano. A ausência de variação decorrente do sexo pode ser devido a seleção de presa estar mais relacionada a características de corpo mole e fácil digestão e não ao tamanho da presa, sendo essa seleção comum para ambos os sexos. Indivíduos de idades diferentes possuem dietas distintas, com juvenis consumindo mais espécies que os adultos. Essa mudança de nicho alimentar pode ser devida ao período de aprendizado. Finalmente, mesmo descontando os efeitos da disponibilidade temporal e espacial de presas, do sexo e da idade, houve variação interindividual da dieta na subpopulação central. Somente estudos com marcadores isotópicos poderão investigar se essa variação é devida à especialização individual em toninhas / Ecological niche theory is originally based on the assumption that individuals of a species use similar resources and therefore are ecologically equivalents. Under this framework, the niche of the species can be defined in terms of average resource use. However, factors such as quality and quantity of prey resources in the environment, consumer gender, age or morphotype may influence the individual feeding behavior. Considering that possible interindividual variation, recently studies under optimal foraging theory have variation among individuals as a focus of interest. The species P. blainvillei is partionated in populations along its range and probably, subpopulations can be found in the Sao Paulo state coast: northern, central and southern. Hence, my first goal was to investigate if there is diet variation among the three franciscana subpopulations from Sao Paulo coast. Additionally, I searched for which factors would be the diet structure defined within one of the subpopulations (the central one). I have considered the seasonality, individual gender and sex as possible factors influencing the diet structure within the central subpopulation. I had 58 franciscana specimens obtained from bycatch in the Sao Paulo coast, being them from northern, central and southern regions. They were juveniles and adults of both genders and bycaught in all the seasons. I identified the prey species from the gut contents with fish otholits and squid beaks. I estimated prey length and weight using regressions with that structures measures and I investigated the diet structure using a diet variation index. Within the central subpopulation I found no diet variation based on seasonality or individual gender, instead I found diet variation for individuals from different ages. Probably, most of the diet variation among subpopulations is due to prey availability spatial difference. The lack of diet variation based on seasonality may be due to little seasonal variation in the abundance of the main prey, P. harroweri. The lack of variation due to gender is probably related to the prey selection on soft body and easy digestion instead of prey size, being this type of selection common for both genders. Juveniles franciscana preyed on more species than the adults and that niche shift as an age effect can be consequence of forraging skills development. Enclosing, even discarding the effects of the spacial and temporal prey availability, individual gender and age, I found interindividual-level diet variation within the central subpopulation, which can points out to franciscana individual specialization, but isotopic studies are required to infer that specialization

Originalmente, a teoria de nicho ecológico fundamenta-se em indivíduos de uma espécie sendo ecologicamente equivalentes por utilizarem os recursos de forma similar. Portanto, o nicho de uma espécie é definido em termos do uso médio de recursos. Porém, a qualidade e abundância de recursos consumidos, sexo, idade ou morfotipo do consumidor influenciam o comportamento alimentar individual. Estudos recentes sobre forrageamento ótimo tem foco de interesse na variação interindividual no uso de recursos. Ao longo da sua área de distribuição são reconhecidas populações da toninha Pontoporia blainvillei (Cetacea) e no litoral paulista, possivelmente hajam três subpopulações dessa espécie: norte, centro e sul. Meu primeiro objetivo foi investigar se há variação na dieta entre essas três subpopulações de toninha. Adicionalmente, investiguei quais fatores entre sazonalidade, sexo e idade dos indivíduos poderiam estruturar a dieta em uma dessas subpopulações (central). Utilizei 58 indivíduos provenientes de captura acidental no litoral paulista, com representantes das regiões norte, centro e sul. A amostra incluiu juvenis e adultos de ambos os sexos, capturados em todas as estações do ano. Identifiquei as espécies nos conteúdos estomacais usando os otólitos de peixes e os bicos de lulas. Estimei o tamanho das presas utilizando regressões com as medidas dessas estruturas e investiguei a estruturação da dieta usando um índice de variação da dieta. Houve uma clara variação na dieta ente as subpopulações paulistas, possivelmente devido à diferença espacial na disponibilidade de presas. Para a subpopulação central não houve variação na dieta decorrente da sazonalidade ou do sexo. A ausência de variação sazonal pode ser devido à pouca alteração na abundância da principal presa, P. harroweri, no ambiente ao longo do ano. A ausência de variação decorrente do sexo pode ser devido a seleção de presa estar mais relacionada a características de corpo mole e fácil digestão e não ao tamanho da presa, sendo essa seleção comum para ambos os sexos. Indivíduos de idades diferentes possuem dietas distintas, com juvenis consumindo mais espécies que os adultos. Essa mudança de nicho alimentar pode ser devida ao período de aprendizado. Finalmente, mesmo descontando os efeitos da disponibilidade temporal e espacial de presas, do sexo e da idade, houve variação interindividual da dieta na subpopulação central. Somente estudos com marcadores isotópicos poderão investigar se essa variação é devida à especialização individual em toninhas / Ecological niche theory is originally based on the assumption that individuals of a species use similar resources and therefore are ecologically equivalents. Under this framework, the niche of the species can be defined in terms of average resource use. However, factors such as quality and quantity of prey resources in the environment, consumer gender, age or morphotype may influence the individual feeding behavior. Considering that possible interindividual variation, recently studies under optimal foraging theory have variation among individuals as a focus of interest. The species P. blainvillei is partionated in populations along its range and probably, subpopulations can be found in the Sao Paulo state coast: northern, central and southern. Hence, my first goal was to investigate if there is diet variation among the three franciscana subpopulations from Sao Paulo coast. Additionally, I searched for which factors would be the diet structure defined within one of the subpopulations (the central one). I have considered the seasonality, individual gender and sex as possible factors influencing the diet structure within the central subpopulation. I had 58 franciscana specimens obtained from bycatch in the Sao Paulo coast, being them from northern, central and southern regions. They were juveniles and adults of both genders and bycaught in all the seasons. I identified the prey species from the gut contents with fish otholits and squid beaks. I estimated prey length and weight using regressions with that structures measures and I investigated the diet structure using a diet variation index. Within the central subpopulation I found no diet variation based on seasonality or individual gender, instead I found diet variation for individuals from different ages. Probably, most of the diet variation among subpopulations is due to prey availability spatial difference. The lack of diet variation based on seasonality may be due to little seasonal variation in the abundance of the main prey, P. harroweri. The lack of variation due to gender is probably related to the prey selection on soft body and easy digestion instead of prey size, being this type of selection common for both genders. Juveniles franciscana preyed on more species than the adults and that niche shift as an age effect can be consequence of forraging skills development. Enclosing, even discarding the effects of the spacial and temporal prey availability, individual gender and age, I found interindividual-level diet variation within the central subpopulation, which can points out to franciscana individual specialization, but isotopic studies are required to infer that specialization

Les poissons herbivores jouent un rôle écologique majeur face au phénomène de « coral-algal phase-shift », qui sévit depuis le début des années 80 dans les récifs coralliens de la Caraïbe. Dans cette étude, les habitudes alimentaires des principales espèces de poissons herbivores ont été étudiées afin de comprendre comment ces espèces utilisent et se partagent les ressources trophiques en milieu récifal et dans les herbiers. Les niches trophiques de ces espèces ont été décrites par deux méthodes complémentaires: les analyses des contenus des tractus digestifs et les analyses des isotopes stables du carbone C3C/ 12C) et de l'azoteCSN/ '4N). La qualité nutritionnelle des sources alimentaires a été évaluée par leurs concentrations de macronutriments (protéines, lipides, glucides solubles et insolubles) afin d'expliquer en partie le choix des ressources par les herbivores. Parmi les 14 espèces de poissons étudiées, appartenant aux familles des Pomacentridae, des Acanthuridae et des Scaridae, huit types de niches trophiques ont été décrits, indiquant une diversité fonctionnelle de ces espèces. Les niches trophiques décrites sont occupées par une ou plusieurs espèces, indépendamment de leur morphologie et de leurs affinités taxinomiques. Les résultats de cette étude suggèrent une complémentarité entre ces espèces, en terme d'utilisation des ressources, et soulèvent la question de leur conservation afin de préserver leur rôle écologique dans les écosystèmes côtiers de la Caraïbe. / Herbivorous fishes play a major ecological role against the « coral-algal phase-shift » phenomenon in the coral reefs of the Caribbean. In this study, the feeding patterns of the most important species of herbivorous fishes have been studied to understand how these species use and share the trophic resources on reefs and seagrass beds . Trophic niches of these species have been described by two complementary methods: the analyses oftheir digestive contents and the analyses of stable isotopes of carbon C3C/ J2C) and nitrogen (lsN/14N). The nutritional quality of the sources has been evaluated by their concentrations of macronutrients (proteins, lipids, soluble and insoluble carbohydrates) to explain partially the choice of resources by herbivorous fishes . Among the 14 studied species of herbivorous fishes (Pomacentridae, Acanthuridae and Scaridae), eight types of trophic niches have been described, indicating a functional diversity of these species . The trophic niches are occupied by one or several species, independently oftheir morphology or their taxonomie affinities. The results of this study suggest a complementarity among these species and raise the question of their conservation to preserve their ecological role on marine ecosystems of the Caribbean.

Field and laboratory studies were used to elucidate the factors affecting temporal and spatial variations of species abundance and structure of macroalgal assemblage and environmental variables between fish farming (FFA) and non-fish farming (NFFA) areas in Hsiao-Lu-Chiao island, a coral island in southwestern Taiwan. Four experiments have been approached: 1. field surveys of macroalgal assemblage structure on 5-m and 10-m depth at 3 sampling sites at FFA (FFA1, FFA2 and FFA3) and 1 sampling site at NFFA from September 2004, January 2005 and April 2005; 2.the relationship between abiotic (monthly maximum air temperature, monthly minimum air temperature, monthly mean air temperature, monthly cumulative precipitation, monthly cumulative irradiance, seawater temperature, light extinction coefficient, water motion, and nutrient (NO3-, NO2-, NH4+, SRP, DON, and DOP) and biotic (seaurchin density) factors and spatio-temporal variations in macroalgal structures analyzed by non-parametric multivariate model; 3. Factors affecting macroalgal abundance and structure: (1). Comparison of growth temperature ranges in different species to field temperature fluctuation; (2).Comparison of growth irradiance ranges in different species to field irradiance fluctuation; 4.Herbivore pressure: (1). Spatio-temporal variations of sea urchin abundance and structure of assemblage; (2). Gut contents and food preference of sea urchin experiment; (3). Herbivore exclusion experiment. Macroalgal %cover, biomass, species richness, diversity (H¡¦) and evenness (J¡¦) showed temporal and spatial variations, low values in January 2005 and also low values in the 5 m- and 10 m-depth areas of FFA1 and the 10 m-depth areas of FFA2. The data of k-dominance curve, hierarchical cluster and ANOSIM tests indicate that macroalgal assemblage is different between 4 sampling sites, between 2 depths and between 3 seasons. Ceratodictyon spongiosum is the most important species that separates September and January assemblages from April assemblage and separates the FFA1 and FFA2 assemblages from the FFA3 and NFFA assemblages. BVSTEP analysis shows that nutrients (NO3-, DON, DOP), temperature, monthly cumulative precipitation, and sea urchin density are the factors corresponding to variations of macroalgal assemblages, this correlation is more significant for 5 m-depth assemblage. Fish farming area FFA1and FFA2 assemblage are affect by sea urchin density, temperature and DON. Sea urchin influnces macroalgal abundance and assemblage structure in FFA1 and FFA2. Macroalgal %cover in 5 m-depth area shows a reversal relationship with sea urchin density; however, this relationship is not observed for 10 m-depth area. FFA1 and FFA2 are belong to high grazing pressure sites as indicated by high sea urchin density and exclusion experiment. Sea urchin gut contents and feeding preference test show that sea urchin has strong food selectivity with Hypnea charoides and Gracilaria coronopifolia as the most preferred species. Herbivore exclusion experiment shows that Hypnea charoides and Gracilaria coronopifolia are the species recruited in the cages. Ceratodictyon spongiosum had high biomass in FFA1 and FFA2 in January, which was ¡¥low DOP/high DON¡¦. The coindicence of temporal variations in FFA3 assemblage structure with a change from ¡¦Halimeda opuntia and Boodlea compostia¡¦ ¡÷ ¡¦Amphiroa fragilissima, Corallina phhulifera and Galaxaura oblongata¡¦ ¡÷¡¦Halimeda opuntia and Boodlea compostia¡¦ with low nitrogen/ high phosphorous¡¦ ¡÷¡¦ high nitrogen/ low phosphorous¡¦ ¡÷ ¡¦low nitrogen/ high phosphorous¡¦ suggest a role of ¡¦low nitrogen (NO3-)/high phosphorous (DOP)¡¦ for FFA3 structure modification. NFFA assemblage is controlled by temperature and monthly cumulative precipitation. Monthly cumulative precipitation in September was higher than January and April, in which Boodlea compostia and Gracilaria coronopifolia were dominant algae in September. The temperature growth responses of algae using the continuous-flow outdoor laboratory tank culture system fit their seasonal growth, reflecting the temperature-dependent manner of seasonal variations in abundance. It could be concluded from the present investigation that the structure of benthic macroalgal assemblage in Hsiao-Lu-Chiao island in southwestern Taiwan is affected by predicted natural and pulse disturbances. Temperature fluctuations involve in overall temporal variations in structure. Sea urchin herbivory and nutrient as pulse nutrient modulate the structure in fish farming area while monthly cumulative precipitation is associated with algal structure in non-fish farming area.

Recruitment of pelagic fish populations is believed to be regulated during the planktonic larval stage due to high rates of mortality during the early life stages. Starvation is thought to be one of the main sources of mortality, despite the fact that there is rarely a strong correlation between the feeding success of larval fish and food availability as measured in the field. This lack of relationship may be caused in part by (i) inadequate sampling of larval fish prey and (ii) the use of total zooplankton abundance or biomass as proxies for larval food availability. Many feeding studies rely on measures of average prey abundance which do not adequately capture the variability, or patchiness, of the prey field as experienced by larval fish. Previous studies have shown that larvae may rely on these patches to increase their feeding success. I assess the variability in the availability of larval fish prey over a range of scales and model the small-scale distribution of prey in Conception Bay, Newfoundland. I show that the greatest variability in zooplankton abundance existed at the meter scale, and that larval fish prey were not randomly distributed within the upper mixed layer. This will impact both how well we can model the stochastic nature of larval fish cohorts, as well as how well we can study larval fish feeding from gut content analyses. Expanding on six years of previous lab and field studies on larval Radiated Shanny (Ulvaria subbifurcata) from Conception Bay, Newfoundland, I assess the feeding success, niche breadth (S) and weight-specific feeding rates (SPC, d-1) of the larvae to determine whether there are size-based patterns evident across the years. I found that both the amount of food in the guts and the niche breadth of larvae increased with larval size. There was a shift from low to high SPC with increasing larval size, suggesting that foraging success increases as the larvae grow. My results suggest that efforts should be made to estimate the variability of prey abundance at scales relevant to larval fish foraging rather than using large-scale average abundance estimates, since small-scale prey patchiness likely plays a role in larval fish feeding dynamics. In addition, the characteristics of zooplankton (density, size and behaviour) should be assessed as not all zooplankton are preyed upon equally by all sizes of larval fish. Overall, this thesis demonstrates that indices based on averages fail to account for the variability in the environment and in individual larval fish, which may be confounding the relationship between food availability and larval growth.

Recruitment of pelagic fish populations is believed to be regulated during the planktonic larval stage due to high rates of mortality during the early life stages. Starvation is thought to be one of the main sources of mortality, despite the fact that there is rarely a strong correlation between the feeding success of larval fish and food availability as measured in the field. This lack of relationship may be caused in part by (i) inadequate sampling of larval fish prey and (ii) the use of total zooplankton abundance or biomass as proxies for larval food availability. Many feeding studies rely on measures of average prey abundance which do not adequately capture the variability, or patchiness, of the prey field as experienced by larval fish. Previous studies have shown that larvae may rely on these patches to increase their feeding success. I assess the variability in the availability of larval fish prey over a range of scales and model the small-scale distribution of prey in Conception Bay, Newfoundland. I show that the greatest variability in zooplankton abundance existed at the meter scale, and that larval fish prey were not randomly distributed within the upper mixed layer. This will impact both how well we can model the stochastic nature of larval fish cohorts, as well as how well we can study larval fish feeding from gut content analyses. Expanding on six years of previous lab and field studies on larval Radiated Shanny (Ulvaria subbifurcata) from Conception Bay, Newfoundland, I assess the feeding success, niche breadth (S) and weight-specific feeding rates (SPC, d-1) of the larvae to determine whether there are size-based patterns evident across the years. I found that both the amount of food in the guts and the niche breadth of larvae increased with larval size. There was a shift from low to high SPC with increasing larval size, suggesting that foraging success increases as the larvae grow. My results suggest that efforts should be made to estimate the variability of prey abundance at scales relevant to larval fish foraging rather than using large-scale average abundance estimates, since small-scale prey patchiness likely plays a role in larval fish feeding dynamics. In addition, the characteristics of zooplankton (density, size and behaviour) should be assessed as not all zooplankton are preyed upon equally by all sizes of larval fish. Overall, this thesis demonstrates that indices based on averages fail to account for the variability in the environment and in individual larval fish, which may be confounding the relationship between food availability and larval growth.