Community assembly in Nothobranchius annual fishes: Nested patterns,environmental niche and biogeographic history

The assembly of local communities from regional species pools is shaped by historical aspects of distribution, environmental conditions, and biotic interactions. We studied local community assembly patterns in African annual killifishes of the genus Nothobranchius (Cyprinodontiformes), investigating data from 168 communities across the entire range of regionally co‐existing species. Nothobranchius are small fishes associated with annually desiccating pools. We detected a nested pattern of local communities in one region (Southern Mozambique, with Nothobranchius furzeri as the core and dominant species), but no nestedness was found in the second region (Central Mozambique, with Nothobranchius orthonotus being the dominant species). A checkerboard pattern of local Nothobranchius community assembly was demonstrated in both regions. Multivariate environmental niche modeling revealed moderate differences in environmental niche occupancy between three monophyletic clades that largely co‐occurred geographically and greater differences between strictly allopatric species within the clades. Most variation among species was observed along an altitudinal gradient; N. furzeri and Nothobranchius kadleci were absent from coastal plains, Nothobranchius pienaari, Nothobranchius rachovii, and Nothobranchius krysanovi were associated with lower altitude and N. orthonotus was intermediate and geographically most widespread species. We discuss implications for ecological and evolutionary research in this taxon.     

Asymmetric reproductive isolation between two sympatric annual killifish with extremely short lifespans

Background

Interspecific reproductive isolation is typically achieved by a combination of intrinsic and extrinsic barriers. Behavioural isolating barriers between sympatric, closely related species are often of primary importance and frequently aided by extrinsic factors causing spatial and temporal interspecific separation. Study systems with a severely limited role of extrinsic factors on reproductive isolation may provide valuable insights into how reproductive isolation between sympatric species is maintained. We used no-choice experimental set-up to study reproductive barriers between two closely related sympatric African killifish species, Nothobranchius furzeri and Nothobranchius orthonotus. These fish live in small temporary savannah pools and have complete spatial and temporal overlap in reproductive activities and share a similar ecology.

Principal Findings

We found that the two species display largely incomplete and asymmetric reproductive isolation. Mating between N. furzeri males and N. orthonotus females was absent under standard experimental conditions and eggs were not viable when fish were forced to mate in a modified experimental setup. In contrast, male N. orthonotus indiscriminately mated with N. furzeri females, the eggs were viable, and offspring successfully hatched. Most spawnings, however, were achieved by male coercion and egg production and embryo survival were low. Behavioural asymmetry was likely facilitated by mating coercion from larger males of N. orthonotus and at relatively low cost to females. Interestingly, the direction of asymmetry was positively associated with asymmetry in post-mating reproductive barriers.

Significance

We showed that, in fish species with a promiscuous mating system and multiple matings each day, selection for strong mate preferences was relaxed. This effect was likely due to the small proportion of resources allocated to each single mating and the high potential cost to females from mating refusal. We highlight and discuss the fact that males of rarer species may often coercively mate with females of a related, more abundant species.     

Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

Trophic niche partitioning in communities of African annual fish: evidence from stable isotopes

Annual killifish of the genus Nothobranchius often co-occur in temporary savannah pools. Their space- and time-limited environment does not allow for any substantial habitat or temporal segregation. Coexisting species are therefore predicted to have well separated trophic niches to avoid intense food competition. Although in a previous “snapshot” study using stomach content analysis (SCA), the trophic niches of three sympatric species (N. furzeri, N. orthonotus, and N. pienaari) were found to vary among species, the difference was relatively weak and inconsistent across different sites. Here, we used the time-integrative capacity of stable isotope analysis to test whether the trophic niches of sympatric Mozambican Nothobranchius are more distinct over a long-term period. Analysis of carbon and nitrogen stable isotopes separated the trophic niche and trophic position of N. pienaari but failed to find any difference between N. furzeri/N. kadleci and N. orthonotus. No segregation was found at the sites with low prey diversity. In contrast, SCA identified N. orthonotus as the species with the most distinct trophic niche. We discuss the effect of prey diversity and different sensitivities of stomach content and stable isotope analysis in general and conclude that the trophic niches of the three sympatric Nothobranchius species are well separated.     

Evidence for diet partitioning among three coexisting native freshwater fishes in South Africa's Cape Fold Ecoregion

The partitioning of limited resources commonly explains how different species can coexist within the same ecological community. In this 2010 study, the diets of three coexisting freshwater fishes (Cape galaxias Galaxias zebratus, n = 27; Cape kurper Sandelia capensis, n = 60; Breede River redfin Pseudobarbus burchelli, n = 77) were characterised and compared in three headwater streams in South Africa's Cape Fold Ecoregion using gut contents and stable isotope analyses. These data were analysed to ascertain whether the three species exploit distinct trophic niches. Both approaches provided evidence that these species occupy different trophic niches, though with some overlap. However, dietary differences among sites were not consistent and were probably influenced by site-specific factors like resource availability. Pseudobarbus burchelli had a broader niche breadth at Tierkloof Stream than the other two species, but not at Waaihoek or Tierstel Streams. Our results also suggest that P. burchelli consumed a more omnivorous diet than do the other two species, whereas S. capensis occupied a higher trophic position than the other two species and consumed vertebrates. Our findings suggest that these species occupy non-equivalent feeding niches in Cape Fold Ecoregion headwater streams, and that diet partitioning might facilitate their coexistence in these systems.     

Female bias in the adult sex ratio of African annual fishes: interspecific differences,seasonal trends and environmental predictors

Many populations have consistently biased adult sex ratios with important demographic and evolutionary consequences. However, geographical variation, the mechanisms, temporal dynamics and predictors of biased sex ratios are notoriously difficult to explain. We studied 334 wild populations of four species of African annual fish (Nothobranchius furzeri, N. kadleci, N. orthonotus, N. rachovii) across their ranges to compare their adult sex ratio, its seasonal dynamics, interpopulation variation and environmental predictors. Nothobranchius populations comprise a single age cohort and inhabit discrete isolated pools, with wide-ranging environmental conditions (habitat size, water turbidity, structural complexity, predators), making them ideal to study adult sex ratio variation. In captivity adult sex ratios were equal. In natural populations, adult sex ratios were biased 1:2 toward females in three study species while N. kadleci had sex ratios at unity. There was a decline in the proportion of males with age in one species, but not in the other species, implying most severe male mortality early after maturation, declining later perhaps with a decrease in male abundance. In general, the populations at vegetated sites had relatively more males than populations at sites with turbid water and little vegetation. Selective avian predation on brightly coloured male fish likely contributed to female dominance and vegetation cover may have protected males from birds. In addition, an aquatic predator, a large belastomid hemipteran, decreased the proportion of males in populations, possibly due to greater male activity rather than conspicuous colouration. Alternative explanations for a sex ratio bias, stemming from male–male contests for matings, are discussed. We conclude that the effect of environmental conditions on adult sex ratio varies dramatically even in closely related and ecologically similar sympatric species. Therefore, difficulties in explaining the ecological predictors of sex ratio biases are likely due to high stochasticity rather than limited sample size.     

Cannibalism and Intraguild Predation Among Phytoseiid Mites: Are Aggressiveness and Prey Preference Related to Diet Specialization?

We tested whether specialist and generalist phytoseiid mites differ in aggressiveness and prey choice in cannibalism and intraguild predation. Specialists tested were Galendromus occidentalis, Neoseiulus longispinosus, Phytoseiulus persimilis, and P. macropilis; tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, N. fallacis, and Typhlodromus pyri. Aggressiveness of cannibalistic females against larvae was not related to diet specialization except that highly aggressive species were exclusively generalists. Seldom to moderately cannibalizing species occurred in both generalist and specialist phytoseiids. In contrast to aggressiveness in cannibalism, generalists and specialists differed in aggressiveness in intraguild predation. Adult females of specialists were only slightly aggressive against heterospecific larvae, whereas adult females of all generalists except T. pyri were highly aggressive. Adult females of generalists were able to discriminate between con- and heterospecific larvae and preferentially consumed the latter when given a choice. Adult females of specialists except G. occidentalis showed no preference when given a choice between con- and heterospecific larvae. We conclude that aggressiveness in intraguild predation, species recognition and subsequent preferential consumption of heterospecifics when given a choice is common in generalist but not specialist phytoseiids. We discuss the evolutionary pathways that might have led to the difference between specialists and generalists in species discrimination.     

Feeding behavior of Geoffroy's marmoset (Callithrix geoffroyi) in an Atlantic forest fragment of South-eastern Brazil

We report on the diet and feeding behaviour of a group of Geoffroy's marmosets (Callithrix geoffroyi) in an Atlantic forest fragment in south-eastern Brazil, studied during the period February 1993 to Januaray 1994. Major food categories were gums (68.6%) fruits (15%), and small animal prey (invertebrates 14.6% and vertebrates 0.8%). Dietary changes were observed between the wet and dry seasons, although they were not statistically significant. Insects and gums were consumed throughout the year but fruits were more prevalent in the diet in the wet season. Plant species exploited for their gums includedInga stipularis, followed byAcacia paniculata, Paulinia carpopodia, andBauhinia angulosa. Chemical analysis of the gum of the four species most used all presented high values for carbohydrates and crude proteins. Fruits of an unidentified species of Myrtaceae (sp. 2) were consumed the most. Animal prey consisted mainly of insects, arachnids, snails, and in three cases, lizards and frogs, with orthopterans being the most prevalent insect prey. This study demonstrates thatC. geoffroyi efficiently exploits resources typically available in secondary and disturbed forest habitat. The main threat to the species is forest destruction, degradation, and fragmentation, but the management of small forest fragments may be an effective corservation strategy.     

Partitioning of food resources among Sillago japonica, Ditremma temmincki, Tridentiger trigonocephalus, Hippocampus japonicus and Petroscirtes breviceps in an eelgrass, Zostera marina, bed

We carried out dietary analysis on five numerically abundant fishes, Sillago japonica, Ditremma temmincki, Tridentiger trigonocephalus, Hippocampus japonicus and Petroscirtes breviceps in an eelgrass bed in Kwangyang Bay, Korea. Comparisons between species demonstrated that the dietary composition of each fish species was significantly different from that of every other species. Although gammarid amphipods and caprellid amphipods were consumed by all species, their individual contributions to each species' diet varied. Furthermore, polychaetes contributed to the diets of S. japonica and T. trigonocephalusand crab larvae were consumed byD. temmincki. Algae and eelgrass were not consumed by four fish species and made only a minimal contribution to the diet of P. breviceps. The diet of each fish species except H. japonicus underwent size-related changes; smaller fishes consumed gammarid amphipods, mysids and copepods, while larger fishes ate polychaetes, gastropods, isopods and other fishes. Differences in the prey organisms consumed of each individual species could be often related to differences in mouth length and width. S. japonica, D. temmincki, T. trigonocephalus, and P. breviceps underwent also a significant diel changes that could be related to differences in foraging behavior and/or prey availability. Thus, use of vision to detect prey would account for the greater daytime consumption of copepods by S. japonica and of crab larvae by D. temmincki, whereas the nocturnal emergence of gammarid amphipods, polychaetes and isopods from the substrate explained their greater consumption by S. japonica, D. temmincki, T. trigonocephalusand P. breviceps at night. Dietary breadth was greater for species with larger mouth dimensions.     

Metazoan parasites of African annual killifish (Nothobranchiidae): abundance,diversity, and their environmental correlates

Estimates of biodiversity and its global patterns are affected by parasite richness and specificity. Despite this, parasite communities are largely neglected in biodiversity estimates, especially in the tropics. We studied the parasites of annual killifish of the genus Nothobranchius that inhabit annually desiccating pools across the African savannah and survive the dry period as developmentally arrested embryos. Their discontinuous, non‐overlapping generations make them a unique organism in which to study natural parasite fauna. We investigated the relationship between global (climate and altitude) and local (pool size, vegetation, host density and diversity, and diversity of potential intermediate hosts) environmental factors and the community structure of killifish parasites. We examined metazoan parasites from 21 populations of four host species (Nothobranchius orthonotus, N. furzeri, N. kadleci, and N. pienaari) across a gradient of aridity in Mozambique. Seventeen parasite taxa were recorded, with trematode larval stages (metacercariae) being the most abundant taxa. The parasites recorded were both allogenic (life cycle includes non‐aquatic host; predominantly trematodes) and autogenic (cycling only in aquatic hosts; nematodes). The parasite abundance was highest in climatic regions with intermediate aridity, while parasite diversity was associated with local environmental characteristics and positively correlated with fish species diversity and the amount of aquatic vegetation. Our results suggest that parasite communities of sympatric Nothobranchius species are similar and dominated by the larval stages of generalist parasites. Therefore, Nothobranchius serve as important intermediate or paratenic hosts of parasites, with piscivorous birds and predatory fish being their most likely definitive hosts.     

Relative consumption of three aphid species by the lacewing, Chrysoperla rufilabris, and effects on its development and survival

Relative consumption of three aphid species, Aphis gossypii Glover, Myzus persicae (Sulzer) and Lipaphis erysimi (Kaltenbach) (Homoptera: Aphididae), by larvae of the lacewing, Chrysoperla rufilabris (Burmeister) (Neuroptera: Chrysopidae), was determined in the laboratory, together with effects on lacewing development and survival. Percentages of survival of C. rufilabris from first instar to adult eclosion were significantly different among lacewing larvae fed different aphid species. When larvae were fed A. gossypii and M. persicae, all larvae developed to adulthood. All larvae died prematurely when they were fed L. erysimi. Developmental duration of C. rufilabris larvae was significantly shorter when larvae were fed A. gossypii (18.0 d) than when larvae were fed M. persicae (19.2 d). The number of fourth instar aphids consumed during development by C. rufilabris larvae differed significantly among individuals fed different aphid species. Chrysoperla rufilabris consumed an average of 168 M. persicae, followed by 141.6 A. gossypii, and only 26.6 L. erysimi. The percentage of these total number of aphids consumed by each larval stadium of C. rufilabris varied significantly among aphid species. The percentage of A. gossypii consumed by each larval stadium was similar to that for M. persicae, 12.1 and 11.4% by the first instar, 15.7 and 13.1% by the second instar, and 72.2 and 75.5% by the third instar, respectively; whereas in the case of L. erysimi, 23.3% of the total number of aphids were consumed by the first instar, 30.1% by the second instar, and 46.6% by the third instar.     

Trophic positions of omnivores are not always flexible: Evidence from four species of freshwater crayfish

Omnivores are generally believed to be flexible in their diet and trophic position: seasonal, ontogenetic and site‐based differences in trophic position have been observed. We compared consumed and assimilated diet among four species within a group of omnivorous freshwater crayfish, to determine whether species that occur together at a site occupy different trophic positions. Diets of Geocharax falcata, Gramastacus insolitus, Cherax destructor and Euastacus bispinosus (Decapoda: Parastacidae) were compared using stable isotopes (δ¹³C and δ¹⁵N) and gut content analysis across nine sites that varied in their species composition. Gramastacus insolitus consumed mainly plant material across all sites. Geocharax falcata consumed either plants or animals or both at different sites. Its trophic level was consistently similar to G. insolitus, despite differences in gut contents and source for dietary carbon. Cherax destructor consumed animals and had a relatively stable trophic position among sites. Relative trophic position of these three species was consistent across sites and regardless of food consumed, they were positioned as omnivores at a lower trophic level than predators but higher than primary producers and herbivores. Euastacus bispinosus occupied a higher trophic level than other invertebrate species but δ¹³C levels did not differ among sites. Cherax destructor and G. falcata may show flexibility in food sources and in the assimilation of food that determines their trophic position relative to other crayfish species. In contrast, G. insolitus and E. bispinosus are likely to show both a more fixed diet and less flexible trophic position. Therefore, not all omnivores show the flexible diet and trophic position generally reported in the literature. Some species of omnivorous crayfish may maintain a relatively constant trophic position across sites, seasons or changes in food availability regardless of whether their consumed diet alters or not.     

Species‐specific ontogenetic diet shifts among Neotropical Crenicichla: using stable isotopes and tissue stoichiometry

Ontogenetic diet shifts were compared among five sympatric pike cichlids Crenicichla in a subtropical South American stream using stable C and N isotopes and tissue stoichiometry (C:N). Within species, stable N isotopes were positively related to body size while C:N showed negative relationships. Stable C isotopes, however, were not related to body size in any species. By modelling the switch to piscivory using gut content‐isotope‐body size relationships, diet shifts were shown to be species‐specific with regard to both rate and degree of piscivory. Compared to other piscivorous lineages, Crenicichla appear to be unusually small‐bodied (based on maximum body size). Because of their diversity, abundance and dynamic size‐structured functional roles, Crenicichla may exert broad and complex predation pressures on the aquatic community.     

Seasonal Variation in the Diets of Bluefish, Pomatomus saltatrix (Pomatomidae) and Striped Weakfish, Cynoscion guatucupa (Sciaenidae) in Southern Brazil: Implications of Food Partitioning

The stomach contents of 516 Pomatomus saltatrix and 506 Cynoscion guatucupa from southern Brazil were examined. Both species preyed primarily upon teleosts and crustaceans and occasionally squid. Seasonal variation in the diet was related to prey availability in the area, confirming the opportunistic feeding behaviour of both species. During the summer, P. saltatrix preyed upon four prey species (Loligo sanpaulensis, Artemesia longinaris, Mullus argentinae and Paralonchurus brasiliensis), whereas in the autumn and winter Engraulis anchoita was the predominant prey. C. guatucupa similarly preyed upon A. longinaris and P. brasiliensis during the summer, with juvenile C. guatucupa and E. anchoita also consumed. The latter species was an important dietary constituent during other seasons, as were A. longinaris (autumn), Umbrina canosai (winter), Trachurus latami and euphausids (spring). Although P. saltatrix had a broader diet than C. guatucupa during the summer, the latter species had a broader diet overall. Diet breadths were lower from autumn to spring, due to the predominance of E. anchoita in the diet of both species. This restricted diet resulted in significant intra- and inter-specific diet overlap during the winter but, because of the seasonal abundance of E. anchoita, the high levels of diet overlap do not necessarily indicate competition.     

Is resource partitioning the key? The role of intra‐interspecific variation in coexistence among five small endemic fish species (Characidae) in subtropical rivers

This study focused on intra and interspecific variations in the diet of small species in the Pelotas River, upper Uruguay River ecoregion. We tested the hypothesis that five small characids coexist via resource partitioning. Samplings were performed quarterly between August 2013 and May 2014 at 14 sites in the Pelotas River basin, using gillnets of different mesh sizes, seines, cast nets and electrofishing. In this study, the contents of 1,253 stomachs were analysed from Astyanax dissensus, A. paris, A. saguazu, A. xiru and Bryconamericus patriciae. The main food resources consumed were aquatic insects, leaves, seeds and detritus. There were significant seasonal intra and interspecific variations in the use of food resources by the five species. Permutational analysis of multivariate dispersions results indicated significant differences in individual intraspecific variability between the sampled periods, with the highest values found in November and February because of an increase in allochthonous resources (leaves and seeds). Besides this, significant differences in the proportions of consumption of allochthonous and autochthonous were observed. The trophic niche breadth was significantly different among the species, such that in August, the species had narrower niche breadths. In the other periods, the highest niche breadth values were due to an increase in the consumption of allochthonous items. Diet overlap was low (< 0.4) between 62% of the species pairs. The small characid species showed pronounced changes in resource use and intra and interspecific variations indicating resource partitioning. These factors appear to be essential in species interactions and for coexistence.     

Diet of brown-nosed coatis and crab-eating raccoons from a mosaic landscape with exotic plantations in southern Brazil

We described the diets of two procyonids, the brown-nosed coati Nasua nasua and the crab-eating raccoon Procyon cancrivorus, through analysis of stomach contents of road-killed specimens in southern Brazil. We compared them with previously published dietary information for another syntopic mesopredator, the crab-eating fox Cerdocyon thous. The landscape of the study area includes native grasslands, forests, exotic tree plantations, and other crops. Food items were represented by frequency of occurrence (FO) and relative volume (RV). Stomach contents of 23 coatis were analyzed. Animal and plant items were equally frequent, although the volume of plants was greater. Exotic plant species were consumed more than native plants. Among prey items, invertebrates were more important than vertebrates, mainly because of the frequency of coleopterans and annelids and the volume of necrophagous dipteran larvae. Five specimens of raccoons were analyzed, in which animal items had the highest FO and plant items had the highest RV. Both invertebrate and vertebrate prey had the same FO, although vertebrates had a higher RV. Our data suggest that these procyonids are opportunistic hypocarnivores, utilizing anthropic sources, with diet overlap. The coatis also overlapped their diet substantially with the foxes. The stomach-contents analysis revealed the importance of fruit pulp biomass, soft-bodied animals such as larvae, and also soil, which might indicate feeding habits such as scavenging and geophagy.     

Diets of deepwater oreos (Oreosomatidae) and orange roughy Hoplostethus atlanticus

The diets of black oreo Allocyttus niger, smooth oreo Pseudocyttus maculatus, spiky oreo Neocyttus rhomboidalis and orange roughy Hoplostethus atlanticus were determined from examination of contents of 240, 311, 76 and 415 non‐empty stomachs, from fishes sampled by bottom trawl on Chatham Rise to the east of South Island, New Zealand. Hoplostethus atlanticus had an opportunistic predatory strategy with a broad diet dominated by prawns and mesopelagic teleosts, but with substantial components of mysids and cephalopods. Pseudocyttus maculatus was strongly specialized on gelatinous zooplankton (jellyfish and salps). Allocyttus niger consumed mainly salps and hyperiid amphipods, and to a lesser extent fishes, prawns, mysids and copepods. Neocyttus rhomboidalis primarily consumed salps, along with mysids, euphausiids and fishes. Only P. maculatus did not exhibit significant ontogenetic variation in diet. The diets were also influenced by year and bottom depth. Differences in the distributions and diets of the four species probably reduce conflicts in resource use.     

Pollen consumed by the solitary bee Tetrapedia diversipes (Apidae: Tetrapediini) in a tropical agroecosystem

Pollen analysis of the larval food supply is an important tool for identifying the plants that provide the floral resources used by bees. The present study documents the pollen sources consumed by larvae of the solitary bee Tetrapedia diversipes in a tropical agroecosystem. A total of 60 pollen types were recorded with three families being the most important. Euphorbiaceae (60.5%), Malpighiaceae (16.8%) and Asteraceae (12.2%) pollen had the greatest representation in the samples examined. The pollen of Dalechampia dioscoreifolia predominated in the diet of the larvae of T. diversipes (RF?=?56.35%) and indicates the importance of this plant in maintaining populations of this solitary bee.     

Food consumption and growth of larvae and juveniles of three cyprinid species at different food levels

Synopsis The relationships between food availability, consumption and growth were analyzed from the onset of feeding to an age of 90 days in three cyprinid species. Fish were held at 20 ± 0.5° C and given two (three) constant rations of approximately 30, (40) or 100% dry body weight (dbw) ind^-1 day^-1. Food consisted of living zooplankton, the size of which correlated with fish size. At high food densities consumption rates decreased rapidly with fish size in all three species. At reduced rations, fish consumed most of the food offered until they were larger than 10 mg dbw. In all species and at each feeding level daily rations consumed increased allometrically with body size. Respiration rate, expressed as routine metabolic rate differed insignificantly between the three species. At high ration levels, growth rates of small bleak, Alburnus alburnus, were distinctly lower than those of roach, Rutilus rutilus, and blue bream, Abramis ballerus. At low food supply all three species grew at similar rates. Assimilation efficiency at low food conditions was approximately twice that of the well-fed groups. If the caloric equivalents of oxygen consumption as measured in well-fed fish are applied to fish fed at low rations their energy budgets do not balance. This indicates the limitations of fish larvae in the partitioning of energy for growth or activity at such conditions.