Effects of Group Membership and Size Distribution within a Group on Growth Rates of Juvenile Sablefish Anoplopoma fimbria

Group membership can confer both advantages and disadvantages to growth in juvenile fishes. The balance between costs and benefits of social interactions can shift depending on such factors as the composition of the group (density and size disparity) and the availability of food. We examined the effect of these factors on absolute growth and growth depensation in juvenile sablefish, Anoplopoma fimbria. Increasing density and increasing size disparity had little influence on absolute growth rates of juvenile sablefish and the effects of these social factors were not modified by ration level. In experiments testing density effects, absolute growth did not differ among groups of 1, 3, or 10 fish held at high rations, but at low rations single fish exhibited a different pattern of size-dependent growth compared to fish in groups. In experiments testing disparity effects, absolute growth did not differ between groups with an even size distribution and groups with a mixed size distribution. The relative size of an individual within a group, i.e., small, medium, or large, also did not modify growth, despite evidence of higher chasing behavior in mixed size distributions. Although the growth of small fish was not diminished in the presence of large fish, negative impacts of size disparity were expressed in high levels of cannibalism, which occurred in 42% of groups with a mixed size distribution. Significant growth depensation over time occurred in the density experiment, but not in the size disparity experiment, possibly due to the shorter duration of the latter experiment. We suggest that growth depensation was generated by individual variability in growth capacity rather than social effects on growth rates. Schooling behavior, measured by group cohesion indices, increased with fish size and was higher in groups with an even vs. a mixed size distribution. These results for sablefish are consistent with other schooling species in which growth variability is determined by exploitative competition and/or genetic variability in growth capacity rather than interference competition.     

Hierarchical and interactive habitat selection in response to abiotic and biotic factors: The effect of hypoxia on habitat selection of juvenile estuarine fishes

Habitat selection is a shared process among animals where individuals choose areas that differ in biotic and abiotic characteristics to maximize individual fitness. We used manipulative laboratory mesocosm choice experiments to examine hierarchical and interactive relationships influencing this habitat selection process of estuarine fishes. We assessed selection among substrate, dissolved oxygen (DO) concentration, food availability, and predation risk using two common juvenile estuarine fish species, pinfish (Lagodon rhomboides) and Atlantic croaker (Micropogonias undulatus). For both species oxygen concentration greatly influenced selection patterns; fishes strongly avoided low DO conditions, while in higher levels of DO factors such as substrate or food influenced selection patterns. However, both species strongly avoided predators even when alternative habitat was severely oxygen limited. These results suggest that predation risk may be the greatest determinant of habitat selection of the factors considered. Expansion of low DO areas in the world’s oceans is a major anthropogenic disturbance and is rapidly increasing. Assessing impacts of hypoxia on habitat usage of mobile organisms is critical as changes in environmental metrics including predator distribution and DO levels may alter habitat selection patterns disrupting critical ecosystem processes and trophic interactions. Our results indicate that juvenile fishes may forgo emigration from hypoxia due to predation risk. If similar patterns occur for juvenile fishes in estuaries they may potentially suffer from reduced growth, reproductive output, and survivorship.     

Utilization of the exotic cladoceran Daphnia lumholtzi by juvenile fishes in an Illinois River floodplain lake

Daphnia lumholtzi comprises a substantial component of the zooplankton community during mid‐ to late‐summer in Lake Chautauqua, a floodplain lake along the Illinois River near Havana, Illinois. In order to quantify the utilization of D. lumholtzi by juvenile fishes, diet analyses were conducted for seven juvenile fish species collected from Lake Chautauqua during the 2001 annual drawdown period. Freshwater drum Aplodinotus grunniens and emerald shiner Notropis atherinoides demonstrated negative selectivity for D. lumholtzi relative to native zooplankton species whereas four species of fish (bluegill Lepomis macrochirus, white bass Morone chrysops, white crappie Pomoxis annularis and black crappie Pomoxis nigromaculatus) consumed substantial amounts of D. lumholtzi. Although selectivity values for D. lumholtzi varied among these fish species, positive selection for D. lumholtzi increased similarly among larger size classes of each fish species, and corresponded with ontogenetic shifts in diet. Mean body length of D. lumholtzi consumed by 20–69 mm L_T juvenile fishes ranged from 0·75 to 0·99 mm with a calculated total length range of 2·0–2·6 mm. Results from this study provide evidence that high abundances of D. lumholtzi in mid‐ to late‐summer provide an additional food source for several juvenile fish species during a time when abundances of large native cladoceran species (i.e. Daphnia) are low, and juvenile fishes are searching for larger prey associated with ontogenetic shifts from zooplankton to macroinvertebrates and fishes. Because zooplankton production is typically lower in rivers than in lakes, survivorship of juvenile fishes produced in floodplain lakes may be higher in riverine systems if they are not reliant on zooplankton as a primary food resource. Therefore, high abundances of D. lumholtzi may benefit juvenile fishes in managed floodplain lakes, such as Lake Chautauqua, by increasing growth and facilitating the transition from zooplanktivory to insectivory or piscivory.     

The influence of shelter availability on recruitment and early juvenile survivorship of Lythrypnus dalli Gilbert (Pisces: Gobiidae)

The bluebanded goby, Lythrypnus dalli Gilbert, is a common inhabitant of rocky subtidal reefs along the coast of southern California. Experiments utilizing artificial habitats indicate that recruitment and early juvenile survivorship are limited by the presence of adult conspecifics and predators. However, the level of effect is strongly influenced by shelter availability. Both adult and predator effects on limiting juvenile density are significantly reduced as the availability of shelters is increased. Susceptibility to predation is greatest among smaller size classes of fish and is likely to result from displacement from shelters by larger fish. Existing population structures probably reflect the availability of adequate shelter sizes and relative competitive abilities among fish.     

Comparison of microridges in juvenile and adult sunfish,Lepomis gibbosus

Microridges are F-actin-based surface protrusions of the superficial layer cells of fish epidermis. Microridge patterns progress in complexity during fish embryogenesis, often transitioning from abundant surface microvilli to the classical fingerprint arrangement. This progression suggests pattern changes may also occur during later stages of fish development. Fluorescent labelling of F-actin and morphometric analysis were therefore used to assess changes in epidermal microridge patterns in juvenile and adult sunfish (Lepomis gibbosus). The microridge patterns found in adult pumpkinseed were similar to that described for many fishes, consisting of whorls or complex multi-branched ridges. The microridge patterns of the scales from three different-sized groups of juvenile pumpkinseed were distinctly different from that of adult, however, and were present mainly as unbranched concentric or nearly concentric rings in the two larger juvenile groups. In the smallest juveniles, microridges were often apparent as fragmented ridges with abundant actin puncta. Larger juveniles sometimes displayed mixed patterns, with some microridges similar to that of both adult and juvenile patterns. The results show a transition from simple microridge patterns in juvenile pumpkinseeds to distinctly different, diverse and more complex patterns in adults. The different pattern types may reflect particular microridge functions relevant to fish size and age.     

Otolith increment width responses of juvenile Australian smelt Retropinna semoni to sudden changes in food levels: the importance of feeding history

To aid the interpretation of otolith microstructure in wild fishes, the present study assessed responses in daily otolith increment widths of early juvenile Australian smelt Retropinna semoni to sudden changes in feeding conditions. There was an almost immediate response in otolith increment widths (which can be interpreted as growth in length) to sudden changes in feeding conditions, with such changes being detected statistically after c. 4 days. Fish displayed compensatory growth when food supplies were increased following a period of limited food, indicating the magnitude of the response in growth appears highly dependant on feeding history. Additionally, fish size also appeared to influence increment widths suggesting that both fish size and feeding history are important factors that must be considered when interpreting otolith microstructure for the species.     

Juvenile growth of a macrohabitat generalist tied to hydrological character of large‐river system

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Causes of Growth Depensation in Red Drum, Sciaenops Ocellatus, Larvae

Individual growth rates, and thus sizes at a given age, vary within cohorts of fishes reared in the laboratory, field enclosures, and the wild, but the causes of this variability are poorly understood. We examined effects of three variables (food availability, a diet shift, and fish density) on growth depensation in red drum, Sciaenops ocellatus, larvae in laboratory experiments. Absolute variance in length of larvae was greater with higher food availability, and relative variance (CV) for a cohort at 14d posthatching was positively correlated with its CV at 7d posthatching, over which time CV doubled. Supplementation of a rotifer diet with Artemia nauplii (diet shift) increased growth rate and length variance, but not CV. Individually-reared and group-reared larvae had similar growth rates, but group-reared larvae exhibited higher CVs than individually-reared larvae. The observation that food availability and diet shifts affected absolute size variance, while fish density affected relative size variance of red drum larvae, indicates that both absolute and relative size variance are important to understand how environmental factors affect growth depensation.     

Density‐dependent polyphenism and geographic variation in size among two populations of lubber grasshoppers (Romalea microptera)

1. Density‐dependent phase polyphenism occurs when changes in density during the juvenile stages result in a developmental shift from one phenotype to another. Density‐dependent phase polyphenism is common among locusts (Orthoptera: Acrididae). 2. Previously, we demonstrated a longitudinal geographic cline in adult body size (western populations = small adults; eastern populations = large adults) in the eastern lubber grasshopper (Romalea microptera) in south Florida. As lubbers are confamilial with locusts, we hypothesised that the longitudinal size cline was partly due to density‐dependent phase polyphenism. 3. We tested the effect of density, population, and density×population interaction on life‐history traits (pronotum length, mass, cumulative development time, growth rate) of, and proportion surviving to, each of the five instars and the adult stage in a 2 × 3 factorial laboratory experiment with two lubber populations, each reared from hatchling to adult at three different densities. 4. The effect of density on life history and survival was independent of the effects of population on life history and survival. Higher densities led to larger adult sizes (pronotum, mass) and lower survivorship. The western population had smaller adult masses, fewer cumulative days to the adult stage, and higher survivorship than the eastern population. 5. Our data suggest that lubber grasshoppers exhibit density‐dependent phase polyphenism initiated by the physical presence of conspecifics. However, the plastic response of adult size to density observed in the laboratory is not consistent with the relationship between phenotypes and adult density in the field. Genetic differences between populations observed in the laboratory could contribute to size and life‐history differences among lubber populations in the field.     

Predictability of littoral-zone fish communities through ontogeny in Lake Texoma,Oklahoma-Texas,USA

Synopsis We sampled larval, juvenile and adult fishes from littoral-zone areas of a large reservoir (Lake Texoma, Oklahoma-Texas) (1) to characterize environmental factors that influenced fish community structure, (2) to examine how consistent fish–environment relationships were through ontogeny (i.e., larval vs. juvenile and adult), and (3) to measure the concordance of larval communities sampled during spring to juvenile and adult communities sampled at the same sites later in the year. Larval, juvenile and adult fish communities were dominated by Atherinidae (mainly inland silverside, Menidia beryllina) and Moronidae (mainly juvenile striped bass, Morone saxatilis) and were consistently structured along a gradient of site exposure to prevailing winds and waves. Larval, juvenile and adult communities along this gradient varied from atherinids and moronids at highly exposed sites to mostly centrarchids (primarily Lepomis and Micropterus spp.) at protected sites. Secondarily, zooplankton densities, water clarity, and land-use characteristics were related to fish community structure. Rank correlation analyses and Mantel tests indicated that the spatial consistency and predictability of fish communities was high as larval fishes sampled during spring were concordant with juvenile and adult fishes sampled at the same sites during summer and fall in terms of abundance, richness, and community structure. We propose that the high predictability and spatial consistency of littoral-zone fishes in Lake Texoma was a function of relatively simple communities (dominated by 1–2 species) that were structured by factors, such as site exposure to winds and waves, that varied little through time.     

SEX RATIO,PARENTAL INVESTMENT,AND INTERPARENT VARIABILITY IN NESTING SUCCESS IN A SOLITARY BEE

Observations of uniquely marked females of the solitary, twig-nesting bee, Osmia bruneri, were conducted under greenhouse conditions to test several predictions of sex-ratio and parental-investment theory. In support of Fisher's (1958) theory, we found that the observed sex-ratio of progeny in this dimorphic species did not differ from that expected on the basis of average male and female weights. Investment patterns also exhibited a seasonal component: female parents produced more female than male offspring early in the nesting season but reversed this pattern later. Interfemale variability was large for all nesting parameters examined. Neither female-parent size nor the rate at which females completed cells was significantly related to several estimates of parent fitness. Parent-offspring heritability for size was also low. We found no evidence to support the hypothesis that progeny sex-ratios are influenced by maternal condition. Variance in progeny sex-ratios was large, but the population sex-ratio probably departs frequently from the equilibrium value. The results marginally support Kolman's (1960) prediction of large variance in progeny sex-ratios in large panmictic populations. We conclude that variability among females in investment patterns and variability in size among progenies are probably maintained by such factors as resource heterogeneity and the shape of the adult survivorship curve.     

Habitat complexity and sex-dependent predation of mosquito larvae in containers

Studies in aquatic systems have shown that habitat complexity may provide refuge or reduce the number of encounters prey have with actively searching predators. For ambush predators, habitat complexity may enhance or have no effect on predation rates because it conceals predators, reduces prey detection by predators, or visually impairs both predators and prey. We investigated the effects of habitat complexity and predation by the ambush predators Toxorhynchites rutilus and Corethrella appendiculata on their mosquito prey Aedes albopictus and Ochlerotatus triseriatus in container analogs of treeholes. As in other ambush predator-prey systems, habitat complexity did not alter the effects of T. rutilus or C. appendiculata whose presence decreased prey survivorship, shortened development time, and increased adult size compared to treatments where predators were absent. Faster growth and larger size were due to predator-mediated release from competition among surviving prey. Male and female prey survivorship were similar in the absence of predators, however when predators were present, survivorship of both prey species was skewed in favor of males. We conclude that habitat complexity is relatively unimportant in shaping predator-prey interactions in this treehole community, where predation risk differs between prey sexes.     

The effects of tail autotomy on survivorship and body growth of Uta stansburiana under conditions of high mortality

We examined the effects of tail autotomy on survivorship and body growth of both adult and juvenile Uta stansburiana by directly manipulating tail condition. Tail loss decreased neither survivorship nor rate of body growth for individuals in two natural populations. Lack of an influence of tail loss on survivorship in these two populations may be the result of high mortality. Under high mortality any differential effects of tail loss will be lower than in populations facing lower mortality. Growth experiments in the laboratory demonstrated that, under conditions of minimal environmental variation and social interactions, there is no tradeoff between body growth and tail regeneration as has been suggested for other species of lizards. One possible reason for this difference is that U. stansburiana does not use the tail as a storage organ for lipids. The original and regenerated tails are composed mainly of protein. In general, any differential body growth between tailed and tailless individuals may be due to social interactions and not a diversion of limited energy into tail regeneration.     

LIFE-CYCLE COMPONENTS OF SELECTION IN ERIGERON ANNUUS: II. GENETIC VARIATION

Genetic variation for seedling and adult fitness components was measured under natural conditions to determine the relative importance of the seedling stage for lifetime fitness in Erigeron annuus. Variation in lifetime reproductive success can result from both the persistent effects of genetic variation expressed among seedlings and from variation in adult fitness components. Analysis of covariance was used to separate the stage specific from the cumulative effects of genetic variance expressed earlier in the life cycle. E. annuus produces seeds through apomixis, which allowed measurement of the fitness of replicate genotypes from germination through the entire life cycle. There were significant differences among genotypes for date of emergence, seedling size, survivorship and fecundity, but heritabilities were low, indicating slow response to selection. For all characters, environmental components of variance were one to two orders of magnitude larger than genetic variance components, resulting in broad sense heritabilities less than 0.1. For seedling size and fecundity, all of the genetic variance was in the form of genotype-environment interactions, often with large negative genetic correlations across environments. In contrast, genotypes differed in mean survivorship through one year, but there were no genotype-environment interactions for viability. Genetic differences in viability were primarily expressed as differences in overwinter survivorship. Genotype × environment interactions among sites and blocks were generated early in the life cycle while the genotype × environment interactions in response to competitive environment (open, annual cover, perennial cover) first appeared in adult fecundity. Genetic variation in lifetime fitness was not significant, despite a fourfold difference in mean fitness among genotypes.     

Reef-based micropredators reduce the growth of post-settlement damselfish in captivity

Despite the ubiquity of micropredators and parasites on coral reefs, their effects on the survival and growth of juvenile fishes are virtually unstudied. Caging and laboratory experiments were used to investigate whether reef based micropredators fed on recently metamorphosed damselfish, the time of day that micropredation occurred, and whether micropredation affected fish growth and survival. Caged juveniles of the damselfish, Pomacentrus moluccensis, were held on the reef over four consecutive time periods. Micropredators (gnathiid and cirolanid isopods) were found associated with caged fish at night only, and cirolanids were observed attacking and killing some caged fish. In order to test the effect of micropredation on growth and survival without the influence of predatory fishes, groups of five P. moluccensis were caged for 2 weeks in one of three treatments: micropredators excluded, mesh control, or micropredators present. There were no significant differences in survival among the treatments, but fish were larger in cages with fewer survivors suggesting that competition for food was intense. Fish exposed to micropredators were larger than fish in the other two treatments, however, micropredator exclusion also excluded plankton; thus, differences in food availability among treatments during crepuscular periods likely confounded the treatment effect on fish growth. A laboratory growth experiment was performed to better control food availability and minimise handling stress, using a validated host-micropredator model. Individual juvenile damselfish, Dischistodus perspicillatus, were exposed to 0, 1 or 2 micropredators (Gnathia falcipenis) each evening and fed equally for 8 days. Mortalities only occurred in fish exposed to micropredators on the first evening of the experiment, and fish exposed to two micropredators each evening were significantly smaller than unexposed fish. These results suggest that repeated gnathiid infections can reduce fish growth in the first week after settlement. Consequently, micropredation may affect the ecology of damselfish after settling on coral reefs.     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Experimental test of the influence of the size of shoals and density of fish on parasite infections

There is an increasing interest in the relationship between adult coral reef fishes and their parasites; however, there is almost no information concerning post larvae (i.e. newly recruited) fish and their parasites. Newly recruited and juvenile fish are highly sensitive to their environment and they have a very low survival rate. Fish grouping behavior has been studied in several ways and many studies have shown that increasing group size might be a way to share a cost linked to parasitism, via a dilution effect. To my knowledge, this hypothesis has never been tested for juveniles of fish. The potential influence of school size and density of a juvenile coral reef fish, Haemulon flavolineatum (Haemulidae), on the success of a monogenean fish parasite, Haliotrema sp., was studied. First, the fish acquired their monogeneans when they were more than 40 mm long, size corresponding also to a change in their foraging behavior. Second, there was a positive effect of fish density on the success of the establishment of the parasite on the host population. Finally, although the parasite success was not directly related to fish group size, the parasite abundance was higher for intermediate shoal size. These results are discussed in terms of the consequence parasitism may have on a fish's shoaling and to a larger extent on the fish population.     

Evolutionary divergence in life history traits among populations of the Lake Malawi cichlid fish Astatotilapia calliptera

During the early stages of adaptive radiation, populations diverge in life history traits such as egg size and growth rates, in addition to eco‐morphological and behavioral characteristics. However, there are few studies of life history divergence within ongoing adaptive radiations. Here, we studied Astatotilapia calliptera, a maternal mouthbrooding cichlid fish within the Lake Malawi haplochromine radiation. This species occupies a rich diversity of habitats, including the main body of Lake Malawi, as well as peripheral rivers and shallow lakes. We used common garden experiments to test for life history divergence among populations, focussing on clutch size, duration of incubation, egg mass, offspring size, and growth rates. In a first experiment, we found significant differences among populations in average clutch size and egg mass, and larger clutches were associated with smaller eggs. In a second experiment, we found significant differences among populations in brood size, duration of incubation, juvenile length when released, and growth rates. Larger broods were associated with smaller juveniles when released and shorter incubation times. Although juvenile growth rates differed between populations, these were not strongly related to initial size on release. Overall, differences in life history characters among populations were not predicted by major habitat classifications (Lake Malawi or peripheral habitats) or population genetic divergence (microsatellite‐based F_ST). We suggest that the observed patterns are consistent with local selective forces driving the observed patterns of trait divergence. The results provide strong evidence of evolutionary divergence and covariance of life history traits among populations within a radiating cichlid species, highlighting opportunities for further work to identify the processes driving the observed divergence.     

Interpopulation variation in growth and life-history traits of the introduced sunfish, pumpkinseed Lepomis gibbosus, in southern England

We examined attributes of growth and reproduction in 19 populations of pumpkinseed (Lepomis gibbosus) introduced into southern England in order to: (i) assess variability of these traits in a northern European climate; (ii) assess inter‐relationships among these variables; and (iii) compare these attributes with populations from other parts of Europe where pumpkinseeds have been introduced. Growth rates varied considerably among populations, but juvenile growth rates and adult body sizes were generally among the lowest in Europe. Mean age at maturity ranged from 2.0 to 3.9, and was strongly predicted by the juvenile growth rate (earlier maturity with faster juvenile growth). Other population parameters that also displayed significant negative associations with mean age at maturity were gonadosomatic index, body condition, and adult body size (total length, TL at age 5). Mean TL at maturity and the adult growth increment showed no significant associations with any of the other growth or life‐history variables. Pumpkinseed populations in England matured significantly later than those introduced into warmer, more southerly areas of the continental Europe. All of these data suggest that a combination of cool summer temperatures and resource limitation is the cause of slow growth, small adult body size and delayed maturity relative to introduced populations on the European mainland.     

Patterns and propensities in reproduction and growth of marine fishes

A number of strong regularities characterize certain very basic biological parameters in marine fishes. For example, the ovulated eggs of fish usually measure approximately 1 mm in diameter. The small, relatively uniform size of the eggs means that almost all fish larvae experience environmental variability at very similar scales, which itself establishes strong constraints for, and links between reproduction and recruitment. Additional constraints emerge from seawater being a poor medium for respiration, which establishes further linkages between growth and mortality. These constraints have produced strongly convergent features, and thence the patterns in reproduction and growth of marine fishes that are presented.