The spatial scale of density-dependent growth and implications for dispersal from nests in juvenile Atlantic salmon

By dispersing from localized aggregations of recruits, individuals may obtain energetic benefits due to reduced experienced density. However, this will depend on the spatial scale over which individuals compete. Here, we quantify this scale for juvenile Atlantic salmon (Salmo salar) following emergence and dispersal from nests. A single nest was placed in each of ten replicate streams during winter, and information on the individual positions (±1 m) and the body sizes of the resulting young-of-the-year (YOY) juveniles was obtained by sampling during the summer. In six of the ten streams, model comparisons suggested that individual body size was most closely related to the density within a mean distance of 11 m (range 2–26 m). A link between body size and density on such a restricted spatial scale suggests that dispersal from nests confers energetic benefits that can counterbalance any survival costs. For the four remaining streams, which had a high abundance of trout and older salmon cohorts, no single spatial scale could best describe the relation between YOY density and body size. Energetic benefits of dispersal associated with reduced local density therefore appear to depend on the abundance of competing cohorts or species, which have spatial distributions that are less predictable in terms of distance from nests. Thus, given a trade-off between costs and benefits associated with dispersal, and variation in benefits among environments, we predict an evolving and/or phenotypically plastic growth rate threshold which determines when an individual decides to disperse from areas of high local density.     

Effects of age,size, and density on natural survival for an important coral reef fishery species,yellow tang, <Emphasis Type="Italic">Zebrasoma flavescens</Emphasis>

Hundreds of thousands of juvenile yellow tang, Zebrasoma flavescens, are caught each year in the state of Hawai’i (USA) for the live aquarium trade. As part of an extensive adaptive management strategy built around a network of protected areas, an emphasis was placed on understanding this important species’ life history. Multiple capture-mark-recapture techniques and a model selection approach to data analysis in Program MARK were used to estimate the effects of individual age and conspecific density on natural per-capita daily survival probabilities of yellow tang recruits (recently settled individuals, 30–50 mm total length) and the effects of body size and site on natural per-capita monthly survival probabilities for juveniles (58–127 mm total length). The models of recruit survival that included additive effects of density and age were best supported by the data and indicated an increase of survival with age and decrease of survival with increased conspecific density. At 1 day post-settlement, the model averaged daily per-capita survival probability ranged from 0.963 (95% CI: 0.932–0.981) at a low density of 0.1 recruits m⁻² to 0.848 (95% CI: 0.752–0.911) at a high density of 1.3 recruits m⁻². The best supported model of juvenile survival had no effect of fish length or site, with a constant monthly per-capita survival of 0.939 (95% CI: 0.925–0.950). Only about 1% of recruits may survive to adulthood when protected from fishing. These results can be used to better analyze and interpret data from protected area monitoring surveys and refine management practices. Continued long-term monitoring, combined with targeted life history studies and demographic modeling, is needed to further investigate the population level effects of fishing yellow tang juveniles.     

Fish recruitment, dispersal, and trophic interactions in a heterogeneous lotic environment

I examine how dispersal of juvenile creek chubs (Semotilusatromaculatus) from beaver ponds into adjacent stream environments interacts with temporal abiotic variability to influence fish foraging, growth, and long-term persistence in the lotic ecosystem. Minnow trapping in upstream and downstream beaver ponds, along with weir traps used to monitor directional movement, indicated that most chubs colonized the stream from the downstream beaver pond. Large annual fluctuations in density of age 0 creek chubs occurred in the stream over a 10-year sampling period. Multiple regression analysis indicated that stream temperature, precipitation, and the density of reproductive creek chubs were not correlated with summer density of age 0 chubs in the stream. The factor most strongly associated with increased density of age 0 creek chubs was creation of the downstream beaver pond during the 6th–7th years of the study, suggesting dispersal from the pond was the primary factor determining age 0 fish density in the stream. Most individuals in the strong year classes neither persisted in the stream through their first winter nor resulted in an increased abundance of older age classes in later years. Comparison of age 0 fish density in summer to the proportion of fish surviving to age 1 in spring suggested that overwinter mortality was density dependent. Furthermore, a comparison of the size structure for age 0 individuals in summer to age 1 individuals the following spring indicated that winter mortality was size dependent. Experiments in an artificial stream adjacent to the natural channel revealed that fish growth was strongly density dependent, decreasing as fish density increased across both spring and summer, and elevated and low discharge. The decline in invertebrate prey captured by the fish and the subsequent decline in fish growth appeared to be particularly pronounced under low discharge in summer. Changes in juvenile creek chub density had no significant effect on benthic insect or crustacean abundance, suggesting that exploitative competition for limited invertebrate drift resources was a more important cause of density- dependent growth than depressed local benthic invertebrate abundance. These results suggest that lotic regions adjacent to beaver ponds act as potential reproductive “sinks” for dispersing juveniles confronting seasonal and flow-mediated restrictions on resource acquisition and growth, and the occurrence of seasonal bottlenecks to their survival, especially harsh winter conditions. Received: 9 September 1996 / Accepted: 8 August 1997     

Density-dependent growth rate in an age-structured population: a field study on stream-dwelling brown trout Salmo trutta

A field experiment during autumn, winter and spring was performed in a small stream on the west coast of Sweden, aiming to examine the direct and indirect consequences of density-dependent intercohort competition in brown trout Salmo trutta . Individual growth rate, recapture rate and site fidelity were used as response variables in the young-of-the-year (YOY) age class, experiencing two different treatments: presence or absence of yearlings and over-yearlings (age ≥ 1+ year individuals). YOY individuals in stream sections with reduced density of age ≥ 1+ year individuals grew significantly faster than individuals experiencing natural cohort structure. In the latter, growth rate was negatively correlated with density and biomass of age ≥ 1+ year individuals, which may induce indirect effects on year-class strength through, for example, reduced fecundity and survival. Movement of YOY individuals and turnover rate ( i.e. proportion of untagged individuals) were used to demonstrate potential effects of intercohort competition on site fidelity. While YOY movement was remarkably restricted (83% recaptured within 50 m from the release points), turnover rate was higher in sections with reduced density of age ≥1+ year individuals, suggesting that reduced density of age ≥1+ year individuals may have released favourable microhabitats.     

Behavioral signature of intraspecific competition and density dependence in colony‐breeding marine predators

In populations of colony‐breeding marine animals, foraging around colonies can lead to intraspecific competition. This competition affects individual foraging behavior and can cause density‐dependent population growth. Where behavioral data are available, it may be possible to infer the mechanism of intraspecific competition. If these mechanics are understood, they can be used to predict the population‐level functional response resulting from the competition. Using satellite relocation and dive data, we studied the use of space and foraging behavior of juvenile and adult gray seals (Halichoerus grypus) from a large (over 200,000) and growing population breeding at Sable Island, Nova Scotia (44.0 ^oN 60.0 ^oW). These data were first analyzed using a behaviorally switching state‐space model to infer foraging areas followed by randomization analysis of foraging region overlap of competing age classes. Patterns of habitat use and behavioral time budgets indicate that young‐of‐year juveniles (YOY) were likely displaced from foraging areas near (<10 km) the breeding colony by adult females. This displacement was most pronounced in the summer. Additionally, our data suggest that YOY are less capable divers than adults and this limits the habitat available to them. However, other segregating mechanisms cannot be ruled out, and we discuss several alternate hypotheses. Mark–resight data indicate juveniles born between 1998 and 2002 have much reduced survivorship compared with cohorts born in the late 1980s, while adult survivorship has remained steady. Combined with behavioral observations, our data suggest YOY are losing an intraspecific competition between adults and juveniles, resulting in the currently observed decelerating logistic population growth. Competition theory predicts that intraspecific competition resulting in a clear losing competitor should cause compensatory population regulation. This functional response produces a smooth logistic growth curve as carrying capacity is approached, and is consistent with census data collected from this population over the past 50 years. The competitive mechanism causing compensatory regulation likely stems from the capital‐breeding life‐history strategy employed by gray seals. This strategy decouples reproductive success from resources available around breeding colonies and prevents females from competing with each other while young are dependent.     

The social interactions of juvenile Japanese monkeys on Koshima Islet

This study reports on the social interactions of juvenile Japanese monkeys (1–5 years old) living in their natural habitat. Juvenile males preferred to associate and play with males of the same age. Male-male interactions initiated by a subordinate of the same age or by an older frequently involved play. Juvenile females tended to interact with younger individuals, and to avoid older males. Older females (4–5 years old) tended to avoid one another. Sibling pairs had high frequencies of interaction, although the occurrences of particular types of interactions varied from pair to pair. Patterns of social relations among adults are discussed in relation to juvenile behavior.     

Combined effects of chemical and visual information in eliciting antipredator behaviour in juvenile Atlantic salmon Salmo salar

Under natural conditions, both young-of-the-year (YOY; 0+ year) and parr (1+ year) Atlantic salmon Salmo salar exhibited strong antipredator behaviour ( e.g. increase in latency to resume foraging) following the exposure to damage-released chemical alarm cues relative to a stream water control. Subsequent exposure to a novel visual stimulus had contrasting results. Parr increased their reactive distance to the visual stimulus if they had been previously exposed to a chemical alarm cue, whereas YOY did not. On the other hand, both YOY and parr took significantly longer to resume foraging when exposed to a visual stimulus if they had been previously exposed to a chemical alarm cue than control groups. While YOY and parr differed in the type and intensity of antipredator responses to both chemical and visual stimuli, perhaps due to differential costs and benefits associated with age, both used the chemical and the visual information in a combined manner.     

Juvenile Anadromous Atlantic Salmon of Three Lakes of Newfoundland

Anadromous Atlantic salmon parr (Salmo salar) were captured from three Newfoundland lakes. Yearling and 2 year parr were captured most often in shallow water <2 m deep around lake perimeters and were not abundant at greater depths. The 3+ and 4+ parr age groups were rarely found inshore (<2 m) but were captured in deeper offshore areas. Though the parr population in one deep lake was too small to be estimated by mark-recapture, estimates of the yearlings plus 2+ age groups in two shallower lakes were 55 and 63 parr ha⁻¹. respectively. No under yearlings were found in lake habitats. Parr density in the three study lakes varied inversely with mean lake depth. Lacustrine parr growth rate was greater than or equal to that of stream dwelling parr. It is concluded that some shallow lakes of Newfoundland provide major rearing areas for juvenile anadromous Atlantic salmon.     

Temporal and spatial variation in growth of juvenile Atlantic salmon

Spatial and temporal variation in length‐at‐age and environmental factors affecting variation in growth rate of juvenile Atlantic salmon Salmo salar were studied using data from a long‐term study in the River Stjørdalselva, central Norway. Mean annual instantaneous growth rate among 1+ and 2+year juvenile Atlantic salmon varied between 0·59 and 1·50 g g⁻¹ year⁻¹ and mean instantaneous daily growth rate of young‐of‐the‐year (YOY) varied between 0·013 and 0·033 g g⁻¹ day⁻¹. Between year variation in growth was larger than the within year intra‐watercourse spatial variation. For YOY and 1+year Atlantic salmon, a major part of the observed between year variation in growth rates was explained by variation in mean daily water discharge and spring temperature. For 2+year juvenile Atlantic salmon, mean daily water discharge and cohort density were the only variables to significantly explain variation in growth rates. A large part of the within water‐course spatial variation could not be explained by temperature variations and juvenile Atlantic salmon in the uppermost areas of the river, experiencing the lowest ambient temperatures during the growth period, displayed the highest growth rates. Within the baselines set by temperature, biotic and abiotic factors connected to water flow regime and variation in food availability are suggested to be a major determinants of the temporal and spatial variation in juvenile Atlantic salmon growth rates.     

Growth of larval and juvenile perch: the importance of diet and fish density

In August, growth rate of young–of–the–year (YOY) Perch In lake enclosures could be explained by both YOY density and mean cladoceran biomass, suggesting that in a lake where YOY perch are dominant, growth may be density dependent in late summer and mediated through top–down control on daphnid biomass. In June, growth rate of YOY perch could not be fully explained by YOY density or by mean cladoceran biomass, suggesting that growth and survival during the first part of the summer is negatively affected by a diet of Bosmina and cyclopoid copepods only. The experiments also suggest why YOY perch have a slow growth and a low abundance in eutrophic lakes where small zooplankton dominate. The June experiment also indicated that growth of late larval or early juvenile perch improved when a larger cladoceran became available and was included in the diet.     

Life histories of Atlantic salmon altered by winter temperature and summer rearing in fresh- or sea-water

Increased growth during winter increased the incidence of age 1+ Salmo salar smolts in spring. High condition factor in spring and good growth in summer was associated with a high incidence of sexually mature males in autumn. In two experiments, groups (n=160–237 per group) of individually identified parr, either ungraded (lower and upper modal groups: LMG, UMG) or size-graded (LMG only), were reared at either 10, 6 or 3 °C overwinter (Nov to May). At 10 °C, up to 51% of parr originally in the LMG became smolts in spring at age 1+. In contrast, at either 6 or 3 °C (control), < 6% of LMG parr became smolts. The probability of being recruited into the UMG overwinter was positively related to initial body size, and was increased by size-grading. Smolt recruitment was two-fold higher among females compared to males; a proportion of males by age 0+ had already opted to mature at age 1+ rather than smolt at age 1+. In contrast, smolting at age 1+ was not inhibited in males previously mature at age 0+. During summer (May to Nov), all experimental groups were reared at ambient temperature, each subdivided between fresh water (max 21 °C) or seawater (max 15 °C). Good growth in seawater of winter recruits into the UMG confirmed they had completed smolting. Mortality in seawater among parr was 41–83%, and among smolts was 10–22%. Specific growth rate during summer was inversely related to winter rearing temperature. The incidence of sexual maturity in autumn at age 1+ among male parr was positively related to winter rearing temperature, fork length and condition factor in May, but there was large variation among individuals with respect to body size and maturity. Summer rearing in seawater reduced growth and the incidence of maturation. Parr and post-smolt maturity was 84–99% and 100% in fresh water respectively, 21–58% and 0% in seawater.     

Ontogenetic constraints and diet shifts in Perch (Perca fluviatilis): mechanisms and consequences for intra‐cohort cannibalism

1. In many populations, sufficient size variation to allow for cannibalism may develop not only among age cohorts but also within them. Here, we used data on resource dynamics, consumer body size distribution and gape size limitation to unravel mechanisms promoting cannibalism within cohorts of young‐of‐the‐year (YOY) perch (Perca fluviatilis). 2. Perch are strongly gape limited when feeding on large zooplankton during early ontogeny. As a consequence, only initially large fish were able to shift to feeding on abundant large invertebrates, necessary to sustain fast growth. 3. We suggest that a combination of high initial size variation and exclusive access to resources for individuals with an initial size advantage is a prerequisite for the development of a size distribution sufficient for intra‐cohort cannibalism to occur. 4. During the time when cannibalism was observed, growth of the largest individuals in YOY perch cohorts was faster than that of smaller individuals. However, the energy gain from cannibalism did not increase growth rate enough to reach a size necessary to feed on more abundant size classes of victims, and therefore, the effect of cannibalism on overall cohort density was minor. 5. In addition to a high energy gain from cannibalism allowing for fast growth, strong resource limitation and slow growth rates of small individuals (i.e. potential victims) are a prerequisite not only for the development of intra‐cohort cannibalism but also for its persistence.     

Patchy recruitment patterns in marine invertebrates: a spatial test of the density-dependent hypothesis in the bivalve Spisula ovalis

Density-dependent and density-independent processes have been shown to influence the population dynamics of marine invertebrates, especially recruitment. However, their relative importance has not been evaluated in natural populations. High adult densities have been suggested to inhibit recruitment, especially in suspension-feeders which may ingest incoming larvae. Age structure and juvenile abundance were investigated in the bivalve Spisula ovalis in order to evaluate the importance of density dependence in generating spatial patterns. Age structure is readily established in this species owing to annual shell lines. An extensive sample (from about 100 sites a few hundred meters apart over 4 consecutive years) was analyzed in the statistical framework of spatial analyses, avoiding spurious correlations due to non-independence between neighboring sites. The area studied supports about ten annual cohorts, though only a few occur at each site. The overall picture is a mosaic of kilometer-scale patches of contrasted age structures, as revealed by highly significant spatial autocorrelations. To our knowledge, such large-scale spatial patterns in age structure have not previously been described in benthic invertebrates. Strong patterns are detected even for juveniles, and are independent of the adult biomass present before settlement. Therefore, patchy patterns of age structure mainly reflect density-independent effects, such as spatial variations in larval supply, passive transport of juveniles, or predation on recruits. In the absence of detailed spatial analyses, such patterns have been misinterpreted previously as negative effects of adult density on settlement success. Received: 21 November 1996 / Accepted: 20 February 1997     

Influence of intraspecific interaction and substrate type on initial growth and establishment of Hydrilla verticillata

Both substrate type and plant–plant interaction can greatly influence the growth and establishment of plants. In order to assist the re-vegetation of submerged macrophytes, the growth of Hydrilla verticillata with increasing equi-distance neighboring plant density on two substrate types (sediment and sand, representing high- and low-nutrient level, respectively) was assessed in monoculture stands. The results showed that substrate type greatly changed the biomass allocation patterns of the target plants, with a smaller root mass ratio on sediment compared to sand (0.70 vs. 3.11%). However, interaction between substrate type and neighboring density was observed. At low density, growth on sediment greatly increased plant height (43.90 vs. 22.10 cm), leaf biomass (216.63 vs. 68.41 mg), and total biomass (298.39 vs. 121.77 mg) when compared to growth on sand. However, at high density, no significant effect of the substrate type was found in those parameters. On sediment, high neighboring density greatly decreased the height, root number, total root length, root mass, and total biomass, implying large intraspecific plant–plant competition. However, such competition can be greatly reduced in infertile environments. Therefore, when the plants were grown on sand, neighboring density showed little effect on the height (22.10–26.53 cm), total root length (21.34–40.50 cm), and root biomass (3.14–6.27 mg). Total biomass and root number significantly increased by 50% and 115%, respectively, at high density compared to low density on sand, suggesting that facilitation rather than competition was occurring. Therefore, plant–plant interaction can vary from competition in fertile environments to facilitation in infertile environments. In summary, neighboring density should be manipulated according to the environmental nutrient level, so as to reduce intraspecific competition or increase intraspecific facilitation, and finally enhance the initial growth and establishment of H. verticillata in re-vegetation activities.     

Refugees or ravenous predators: detecting predation on new recruits to tropical estuarine nurseries

Many of the most abundant small and juvenile fishes within shallow water estuarine nursery habitats consume other fish to some degree but have rarely been considered as potentially important predators in the functioning of these systems because of the low (<50%) average occurrence of fish in their diets. Predation by abundant minor piscivores on new recruits when they first enter the nursery may make a significant contribution to the predation mortality of this critical life-history stage. To determine the potential importance of minor piscivores as predators on new recruits, temporal patterns in the diets of 15 common species of minor piscivores were examined and related to the abundance of new recruits (≤20 mm FL) in biweekly seine samples over 13 months in shallow (<1.5 m) sandy habitats in the Ross River estuary in north-eastern Queensland, Australia. The high spatial patchiness of new recruits made it difficult to correlate their abundance with their consumption by minor piscivores, and there was no relationship detected between the abundance of new recruits and the occurrence of fish in the diets of minor piscivores. To gain broader insight into spatio-temporal patterns in the consumption of fish prey by minor piscivores, we utilised a collection of fishes sampled during various studies over 6 years from 17 estuaries in the region to examine the diets of >3500 individuals from 20 spp. of minor piscivores. Patterns in the consumption of fish prey by these minor piscivores, especially the highly abundant sparids, sillaginids and ambassids, revealed that the low average occurrence of fish in their diet greatly underestimated the predation pressure imposed by these on fish prey at particular locations and times. For most sampling occasions and locations few minor piscivores consumed fish prey (consumed by 0% of individuals examined), while occasionally a large proportion of individuals within a taxon did so (50–100% of individuals consumed fish prey). Often at such times/locations multiple species of minor piscivores simultaneously preyed heavily on fish. When minor piscivores consumed fish, they preyed mainly on small new recruits. Because many of these minor piscivores are relatively recent recruits, many of the small and juvenile fishes believed to gain refuge in shallow estuarine nurseries may themselves be important predators on fish subsequently recruiting to these habitats, and so potentially play a significant role in structuring estuarine fish faunas and the functioning of shallow water nurseries.     

Size-dependent foraging capacities and intercohort competition in an ontogenetic omnivore (Arctic char)

Intraspecific competition for resources is strongly influenced by the size of competitors. In this study, we estimated the size‐scaling of the foraging capacities on zooplankton and benthic macroinvertebrates in Arctic char (Salvelinus alpinus) to link size‐dependent performance to effects from competition. The competitive interactions between two size‐classes (YOY and 1‐y) of char were then studied in a large‐scale pond experiment and in two small subarctic lakes. The attack rate function on zooplankton was hump‐shaped whereas the attack rate on benthic chironomids increased monotonically with size. The size‐scaling exponent's for zooplankton and chironomids were 0.65 and 0.30, respectively, leading to that critical resource density (CRD) and maximum growth resource density (G_MRD) increases with size, suggesting an exploitative competitive advantage of small individuals over large. Correspondingly, large (1‐y) char growth was negatively affected by cohort competition whereas small (YOY) char growth was not. Diets of both size classes were dominated by macroinvertebrates with large overlap in prey size suggesting only small gape size advantages for large char. Small char fed to a larger extend on cladocerans which, due to the hump‐shaped foraging efficiency function on zooplankton, were a relatively more profitable resource for small than large char. Estimates of CRD and G_mRD were in correspondence with observed growth responses and resource estimates for zooplankton, whereas for macroinvertebrates only qualitatively correspondence with foraging estimates and char performance was found. Although we were able to explain our results with exploitative competition only, we suggest a general need for size‐dependent foraging estimates on prey in more complex habitats in order to quantitatively link performance and resource abundances. Interference and size‐dependent resource use as mechanisms for observed stable population dynamics in char was not supported by this study and instead a low per capita fecundity and early cannibalism on recruits are suggested to be potential mechanisms that may stabilize char dynamics.     

Unexplained sexual segregation in polygamous ungulates: a defense of an ontogenetic approach

Two processes are generally mixed when considering ungulate sexual segregation: the social segregation which appears to be a rule in polygamous ungulate species and the spatial segregation which is facultative. Early in life, there is evidence that males and females exhibit different levels of activity and patterns of interaction which could lead to behavioural incompatibility. It is proposed that juvenile females, differently motivated than males to interact socially, may soon avoid the pseudo-sexual and agonistic male behavioural components. as body and behavioural dimorphism increase with age, the behavioural incompatibility would be intensified. Since adults often interact agonistically with or avoid them, juveniles will attend to interact with same-sex peers of similar status. Because social experiences are occurring since early stages of development, i.e., during sensitive periods, a social auto-segregation by sex/age classes is expected. Then population would be structured in coherent socio-spatial sets of compatible individuals, sharing similar constraints and properties. Around the birth period, females become asocial promoting a temporal social segregation. Costs linked to gestation and lactation may make females more dependent on water and rich food resources. Female with offspring are also likely to select areas where the risks of predation are reduced, sometimes at detriment to energy intake, while other animals exhibit opportunistic feeding behaviour, resulting in seasonally ecological segregation. Body growth channels a lot of energy for males which are expected to use the best feeding areas. However, indirect competition for the same resources may result in a spatial segregation between the sexes. Sex- and age-specific activity budgets are suspected to favour social and spatial segregations. The social segregation among males and females use exclusive ranges. Whether spatial segregation results in ecological segregation would however depend on habitat heterogeneity, population density, local ecological conditions and the biological cycle periods. It is suggested that spatial segregation will occur not only between the sexes but also among males. If populations are structured in coherent socio-spatial units, spatial segregation among individuals of age and sex classes may result from indirect interactions between sets of individuals (of same age and sex class), that may be translated as an indirect competition between individuals. Studies on ethogenesis are considered as fundamental as they allow to know on what basis social and spatial segregations are set up. In particular, studies on behavioural development could highlight how social segregation between adults may result from a process that originates much earlier in the life of animals. By considering the role of behaviour, we defend the idea that social and spatial segregations may be logically distinguished.     

The partitioning of density-dependent dispersal, growth and survival throughout ontogeny in a highly fecund organism

The way in which density‐dependent effects are partitioned amongst survival, growth and dispersal are key in determining the temporal and spatial dynamics of populations. Here we propose a mechanistic approach to understanding how the relative importance of these sources of density dependence can change over ontogeny through changes in dispersal abilities, energy stores and mortality risks. Whereas the potential for active dispersal typically increases over ontogeny as a function of body size, susceptibility to starvation and predation decreases. The joint effect of these mechanisms suggests a general model for the ontogenetic sequence of how density dependence is manifested, with density dependence early in ontogeny being primarily expressed as mortality on local spatial scales, whereas later stages respond to local density in terms of dispersal and potentially growth. Here we test this model by manipulating the densities of juvenile Atlantic salmon (Salmo salar L.) at two life‐history stages in the wild. Density‐dependent mortality during the early juvenile stage (i.e. fry at onset of exogenous feeding) was accompanied by no effects on body size and weak effects on dispersal. In contrast, dispersal of older juveniles (i.e. parr 2–3 months after onset of feeding) was strongly density‐dependent, with more individuals emigrating from high‐density release sites, and with no effect of initial density on mortality. This dispersal, however, appeared insufficient to produce an ideal free distribution within the study stream, as indicated by the effect of spatial variation in density on body size by the end of the first growth season. These results demonstrate that the way density‐dependent effects are partitioned amongst survival, growth and dispersal changes throughout ontogeny. Furthermore, these changes occur in correlation with changes in individual mortality risks and dispersal abilities, and suggest a general paradigm for the way in which juvenile density‐dependence is manifest in spatially structured populations of highly fecund organisms.     

Size-asymmetric competition and size-asymmetric growth in a spatially explicit zone-of-influence model of plant competition

Size-asymmetric competition among plants is usually defined as resource pre-emption by larger individuals, but it is usually observed and measured as a disproportionate size advantage in the growth of larger individuals in crowded populations (“size-asymmetric growth”). We investigated the relationship between size-asymmetric competition and size-asymmetric growth in a spatially explicit, individual-based plant competition model based on overlapping zones of influence (ZOI). The ZOI of each plant is modeled as a circle, growing in two dimensions. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. We grew simulated populations with different degrees of size-asymmetric competition and at different densities and analyzed the size dependency of individual growth by fitting coupled growth functions to individuals. The relationship between size and growth within the populations was summarized with a parameter that measures the size asymmetry of growth. Complete competitive symmetry (equal division of contested resources) at the local level results in a very slight size asymmetry in growth. This slight size asymmetry of growth did not increase with increasing density. Increased density resulted in increased growth asymmetry when resource competition at the local level was size asymmetric to any degree. Size-asymmetric growth can be strong evidence that competitive mechanisms are at least partially size asymmetric, but the degree of size-asymmetric growth is influenced by the intensity as well as the mode of competition. Intuitive concepts of size-asymmetric competition among individuals in spatial and nonspatial contexts are very different.     

A biological sampling problem illustrated by the population structure and growth patterns of Sardinella aurita at Tripoli,Libya

Synopsis Sardinella aurita were sampled from catches of the lampara fishery at Tripoli, Libya during October 1979 through to September 1980. Landings consisted mainly of adult fish with large (mode 21–26 cm total length), relatively fast growing 3–5-group individuals in winter and spring and smaller (mode 14–20 cm total length), slow growing 2- and 3-group fish in summer. Gonad growth commenced in April, when the fish were feeding on zooplankton, and continued at the expense of mesenteric fat until July. It is suggested that the older and bigger fish spawn first, probably after having moved westwards, and that this results in spatial and temporal differences in length at age within the population. There was no consistent difference in growth rate between the sexes, but because first maturity in males tends to occur earlier than in females, there were length distribution differences between the sexes in the youngest (2+) age group as sampled.