SEASONAL MODULE DYNAMICS IN SARGASSUM SUBREPANDUM (FUCALES,PHAEOPHYTA)1

Module dynamics of the fucoid alga SARGASSUM SUBREPANDUM (Forssk.) C. Agardh was studied in the southern Red Sea. Seasonal variation in thallus density and size was determined, and the initiation, growth, reproduction, and shedding of modules (primary laterals) were ascertained, using a tagging approach. Possible effects of different size‐related parameters on module initiation, growth, reproduction, and shedding were analyzed in the context of contradicting results for other macroalgae, in comparison with terrestrial plants. Thallus density varied little; most of the seasonal variation occurred at the modular level. A restricted period of new module formation early in the cooler season was followed by fast growth and reproduction. Massive shedding of modules occurred toward the end of the cooler season leading to strongly reduced biomass in summer. There was some evidence that high module numbers inhibited new module formation and enhanced the maximum module elongation rate (fastest‐growing module per thallus). On the other hand, elongation rates generally decreased, and apical tissue losses increased with increasing module length. This response was observed over a wide size range, suggesting grazing losses. There was no evidence of suppressed growth in small modules due to intraspecific competition. Elongation rates remained unaffected by reproductive status, indicating that there was no direct trade‐off between growth and reproduction. Module survivorship was independent of module number and size, but fertile modules were more persistent than vegetative ones. We conclude that module dynamics are determined by seasonal changes in the environment, size‐dependent processes, and interactions among the modules.     

SEASONAL CHANGES IN SIZE STRUCTURE OF SARGASSUM AND TURBINARIA POPULATIONS (PHAEOPHYCEAE) ON TROPICAL REEF FLATS IN THE SOUTHERN RED SEA1

Seasonal variation in density, thallus length and biomass, population size structure, and allometric length‐biomass relationships was investigated in populations of Sargassum ilicifolium (Turner) C. Agardh, Sargassum subrepandum (Forssk.) C. Agardh, and Turbinaria triquetra (J. Agardh) Kütz. (Phaeophyceae) on shallow reef flats in the southern Red Sea. Thallus length and biomass varied strongly with season, with the highest values occurring in the cooler months. Thallus densities showed no significant temporal variation. Log‐total biomass versus log‐density relationships were positive throughout the growth season without any decrease in the slope of the relationship. In two populations, biomass‐density combinations approached the interspecific biomass‐density line, but the massive annual shedding of modules occurred before self‐thinning would set in. Allometric length‐biomass relationships varied with season in all populations and were associated with seasonal module initiation, growth, and shedding. Evidence of a strong asymmetric competition was found in two high‐density populations. These populations showed a predominance of small thalli during peak development, asymmetrical Lorenz curves, increasing Gini coefficients, and increasing thallus length relative to biomass during the main growth phase. In two other less crowded populations, small thalli were absent during peak development, Lorenz curves were symmetrical, and Gini coefficients decreased during the main growth phase. In these populations, size equalization appears to be due to responses at the modular level rather than size‐dependent mortality. We conclude that changes in size structure in this highly seasonal environment are determined by module dynamics, modified by asymmetric competition in some populations, with a minor role of recruitment and no regulatory effect of self‐thinning.     

Intraspecific competition and light effect on reproduction of Ligularia virgaurea,an invasive native alpine grassland clonal herb

The relationship between sexual reproduction and clonal growth in clonal plants often shows up at the ramet level. However, only a few studies focus on the relationship at the genet level, which could finally account for evolution. The sexual reproduction and clonal growth of Ligularia virgaurea, a perennial herb widely distributed in the alpine grasslands of the Qinghai‐Tibetan Plateau of China, were studied under different competition intensities and light conditions at the genet level through a potted experiment. The results showed that: (1) sexual reproduction did not depend on density or light, and increasing clonal growth with decreasing density and increasing light intensity indicated that intraspecific competition and light intensity may affect the clonal life history of L. virgaurea; (2) both sexual reproduction and clonal growth show a positive linear relationship with genet size under different densities and light conditions; (3) a threshold size is required for sexual reproduction and no evidence of a threshold size for clonal growth under different densities and light conditions; (4) light level affected the allocation of total biomass to clonal and sexual structures, with less allocation to clonal structures and more allocation to sexual structures in full sunlight than in shade; (5) light determined the onset of sexual reproduction, and the genets in the shade required a smaller threshold size for sexual reproduction to occur than the plants in full sunlight; and (6) no evidence was found of trade‐offs between clonal growth and sexual reproduction under different densities and light conditions at the genet level, and the positive correlation between two reproductive modes indicated that these are two integrated processes. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Accounting for interspecific competition and age structure in demographic analyses of density dependence improves predictions of fluctuations in population size

Understanding species coexistence has long been a major goal of ecology. Coexistence theory for two competing species posits that intraspecific density dependence should be stronger than interspecific density dependence. Great tits and blue tits are two bird species that compete for food resources and nesting cavities. On the basis of long‐term monitoring of these two competing species at sites across Europe, combining observational and manipulative approaches, we show that the strength of density regulation is similar for both species, and that individuals have contrasting abilities to compete depending on their age. For great tits, density regulation is driven mainly by intraspecific competition. In contrast, for blue tits, interspecific competition contributes as much as intraspecific competition, consistent with asymmetric competition between the two species. In addition, including age‐specific effects of intra‐ and interspecific competition in density‐dependence models improves predictions of fluctuations in population size by up to three times.     

Factors affecting growth rates of young tropical floodplain fishes: seasonality and density-dependence

Synopsis Deviations of growth increments from a model describing average growth of 12 common fish species, mostly juveniles, were compared with hydrological variables and biomass density (ranging from 1 gm^-2 to 83 gm^-2 of potentially competing species of similar size ranges (guilds). Seasonal effects on growth were highly significant for omnivores but not for detritivores. Omnivores exhibited faster growth during the rising water season, and higher growth rates were associated with periods of increased rates of flooding during this season, which could be associated with increased availability of food. No species exhibited density-dependent growth when all seasons were considered together. During the rising-water-level season, no inverse relationship between growth and biomass density was found in any of the 11 species tested at p = 0.05. During the short falling-water season, 2 of 8 species tested did exhibit density-dependent growth but a sign test of correlation coefficients from all species was not significant. These two significant results were from four omnivores tested. However, density-dependent growth was not indicated when the data were pooled within either detritivore or omnivore guilds by hydrological season. It was concluded that with the possible exception of juvenile omnivores during the limited falling-water season, density-dependent growth, and by implication interspecific competition, had no effect in regulating the species' populations investigated in the floodplain environment. Conversely, the importance of a seasonal hydrological regime in maintaining growth rates, at least for omnivores, was evident.     

Effects of nutrition and density in Culex pipiens

Mosquito larvae face numerous biotic and abiotic challenges that affect their development and survivorship, as well as adult fitness. We conducted two experiments under semi‐natural conditions to evaluate the effects of intraspecific competition, nutrient limitation and sub‐lethal doses of malathion on individual life history traits in adult Culex pipiens (Diptera: Culicidae). In the first experiment, larvae of Cx. pipiens were reared at different intraspecific densities and exposed to sub‐lethal doses of malathion. In the second experiment, different intraspecific densities of Cx. pipiens larvae were reared under conditions of low or high larval nutrients, and subsequent adults were fed on either water or 10% sucrose solution. Malathion treatment had relatively minor effects compared with density, which had significant negative effects on development rate, survivorship to adulthood, body size (wing length) and longevity. As larval density increased, a sex ratio distortion in survivorship to adulthood emerged, in which a bias towards males was apparent. Nutrient‐rich larval environments alleviated, in part, the effects of increasing density and extended the lifespan of mosquitoes fed on water and 10% sucrose. Density‐dependent alterations in adult longevity attributable to the larval environment are complex and show contrasting results depending on interactions with other environmental factors. This study suggests that larval resource availability and competition influence Cx. pipiens population growth correlates and have lasting effects on traits that relate to a mosquito's ability to vector pathogens.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

Population density and module demography in Trapa natans (Trapaceae), an annual,clonal aquatic macrophyte

The effects of population density on module demography were studied in Trapa natans L., an annual aquatic macrophyte capable of extensive clonal propagation. At low density, the floating plants produced ten times as many ramets (clonal offshoots) as those at high density. Module mortality occurred at three levels: leaf, ramet (shoot), and genet (genetic individual). There was 100-fold variation in the size of nuts containing germinable seeds. In early summer there was a highly significant linear relationship between dry mass of nuts and the total mass of ramets that each had produced. In early summer most (73–83%) of the variation in total plant biomass was attributable to variation in initial nut size. However the significance of initial nut size was diminished later in the season. The great success of the exotic weed T. natans at colonizing and monopolizing an aquatic habitat is a function of its highly productive clonal growth response to low-density conditions, combined with greater proportional allocation of biomass to reproductive structures, resulting in greatly increased nut production at low initial density. The species appears able to develop and maintain a population at extremely high density: plant buoyancy and the production of large, well-protected nuts allow rapid early growth from the sediment each year and early pre-emption of the water surface.     

Seasonal changes in population structure of the tropical deep‐water kelp Laminaria abyssalis

Tropical kelp populations are rare and anomalous relicts of shallow‐water populations that existed during glacial periods of cooler oceanographic climate. The endemic Brazilian tropical kelp, Laminaria abyssalis Joly and Oliveira Filho, occurred at depths below 40 m. The seasonal variations in biological aspects of L. abyssalis sporophyte populations and local variations in seawater nutrients and temperature were evaluated. A population was sampled four times between the austral spring of 2005 to winter 2006. Seasonal variations in the population structure and in the tissue content of nitrogen (N), carbohydrate and pigments were observed. Higher density (6.3 individuals m^?2), biomass (7.3 kg m^?2) and blade area (13 221 cm²) were observed in summer, while the highest percentage of tissue total N (1.6%), carbohydrates (32.9%) and photosynthetic pigments (chlorophyll a = 1.9% and chlorophyll c = 0.4%) were observed in spring. The highest surface area of fertile tissue in L. abyssalis sporophytes (2.36%) was recorded in winter, indicating winter as the season when more investments are made in reproduction. The highest concentrations of total N (6.3 μM) and phosphate (0.6 μM) in seawater were observed in spring and summer, respectively. Seawater temperatures lower than 20°C, which are characteristic of upwelling waters, occurred every month and were most frequent in spring and summer. We show that L. abyssalis invests more in growth in spring and summer and reaches the greatest thallus size and population density in summer. The lower abundance during winter may be related to the lower frequency of temperatures below 20°C and the local seasonal storms that cause turnover of rhodoliths, the main substrate for L. abyssalis.     

Experimental Examination of Intraspecific Density-Dependent Competition during the Breeding Period in Monarch Butterflies (Danaus plexippus)

A central goal of population ecology is to identify the factors that regulate population growth. Monarch butterflies (Danaus plexippus) in eastern North America re-colonize the breeding range over several generations that result in population densities that vary across space and time during the breeding season. We used laboratory experiments to measure the strength of density-dependent intraspecific competition on egg laying rate and larval survival and then applied our results to density estimates of wild monarch populations to model the strength of density dependence during the breeding season. Egg laying rates did not change with density but larvae at high densities were smaller, had lower survival, and weighed less as adults compared to lower densities. Using mean larval densities from field surveys resulted in conservative estimates of density-dependent population reduction that varied between breeding regions and different phases of the breeding season. Our results suggest the highest levels of population reduction due to density-dependent intraspecific competition occur early in the breeding season in the southern portion of the breeding range. However, we also found that the strength of density dependence could be almost five times higher depending on how many life-stages were used as part of field estimates. Our study is the first to link experimental results of a density-dependent reduction in vital rates to observed monarch densities in the wild and show that the effects of density dependent competition in monarchs varies across space and time, providing valuable information for developing robust, year-round population models in this migratory organism.     

Context‐specific influence of water temperature on brook trout growth rates in the field

1. Modelling the effects of climate change on freshwater fishes requires robust field‐based estimates accounting for interactions among multiple factors. 2. We used data from an 8‐year individual‐based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season‐specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third‐order stream in western Massachusetts, U.S.A. 3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context‐dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density. 4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow‐temperature interaction) differed among seasons. 5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures. 6. Our study provided robust, integrative field‐based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold‐water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season‐specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales.     

Reproductive allocation patterns in different density populations of spring wheat

The effects of increased intraspecific competition on size hierarchies （size inequality） and reproductive allocation were investigated in populations of the annual plant, spring wheat （Triticurn aestivurn）. A series of densities （100, 300, 1 000, 3 000 and 10 000 plants/m^2） along a gradient of competition intensity were designed in this experiment. The results showed that average shoot biomass decreased with increased density. Reproductive allocation was negatively correlated to Gini coefficient （R^2 = 0.927）, which suggested that reproductive allocation is inclined to decrease as size inequality increases. These results suggest that both vegetative and reproductive structures were significantly affected by intensive competition. However, results also indicated that there were different relationships between plant size and reproductive allocation pattern in different densities. In the lowest density population, lacking competition （100 plants/m^2）, individual reproductive allocation was size independent but, in high density populations （300, 1 000, 3 000 and 10 000 plants/m^2）, where competition occurred, individual reproductive allocation was size dependent： the small proportion of larger individuals were winners in competition and got higher reproductive allocation （lower marginal reproductive allocation; MRA）, and the larger proportion of smaller individuals were suppressed and got lower reproductive allocation （higher MRA）. In conclusion, our results support the prediction that elevated intraspecific competition would result in higher levels of size inequality and decreased reproductive allocation （with a negative relationship between them）. However, deeper analysis indicated that these frequency- and size-dependent reproductive strategies were not evolutionarily stable strategies.     

Multiple life stages with multiple replicated density levels are required to estimate density dependence for plants

Negative density dependence resulting from intraspecific competition can regulate plant populations by limiting demographic rates (survival, growth, fecundity). However, the strength of intraspecific competition can vary within and among populations due to spatial or temporal environmental heterogeneity, or genetic differences. Quantification of variation under a relatively constant environment is needed to assess the inherent potential for density dependence to vary. This knowledge will help adjust data collection effort required for parameterisation of density dependence. Our review of published plant demographic studies revealed that only half of the studies included the whole life‐cycle in the analysis of density dependence. Approximately half of the studies manipulated density, while the rest examined density dependence from observed densities in the field. Regardless of the design used, density dependence was estimated from a small number of replicates. To investigate inherent variation in density dependence during the life‐cycle, and the effect of low replication on density dependence estimates, we combined an experimental approach with simulations for an invasive herb Senecio madagascariensis. We found significant negative density dependence for five out of six examined demographic rates in a constant environment, with the strength of density dependence increasing during the life‐cycle. An exception was plant growth, in which the direction of density dependence varied from positive to negative depending on the life stage. Simulations showed substantial deviation for density dependence parameterised from a small number of replicates even when environmental variation was minimal. This suggests that data collection procedures currently used to assess the effect of density on plant demographic rates may produce inaccurate estimates, increasing uncertainty in demographic models. Due to variation in the direction and strength of density dependence during the life‐cycle, multiple life stages with multiple replicated density levels are required to parameterise density dependence for demographic rates.     

What shapes local density? The importance of migration rates and local growth for density‐patch size relationships in two Cionus weevils

1. The relative effect of migration and local growth on the spatio‐temporal density‐distribution of two co‐existing herbivorous weevils, Cionus scrophulariae L. and C. tuberculosus Scop., in 32 host plant Scrophularia nodosa L. patches of varying sizes was investigated. 2. Predictions of the temporal development of the slope in the density‐patch size relationships were derived from a basic population model with scale‐dependent migration rates. The model indicated that the slopes in the density‐patch size relationships during the early season should be reflected by the net scaling of immigration and emigration rates, whereas the slopes during the later season should increase as a result of local growth. 3. Emigration rates of the weevils were estimated in a field experiment, were the weevils coexisted in space and time. These results were then combined with a previous estimate of immigration rates in order to determine the net scaling of migration rates. 4. The emigration rate differed between species, caused by different movement rates in small patches, which could explain differences in the general slope of the density‐patch size relationships of the weevils in the natural figwort patches throughout the summer. The slopes in the relationships in the early season were largely predicted by the net scaling of migration rates. The slope also increased in the later season for C. tuberculosus, whereas the slope decreased for C. scrophulariae. 5. It was concluded that the understanding of both inter‐ and intra‐specific variations in density‐patch size relationships of insect herbivores can be improved using population models incorporating scale‐dependent migration and local growth.     

The adaptive significance of population differentiation in offspring size of the least killifish,Heterandria formosa

We tested the hypothesis that density‐dependent competition influences the evolution of offspring size. We studied two populations of the least killifish (Heterandria formosa) that differ dramatically in population density; these populations are genetically differentiated for offspring size, and females from both populations produce larger offspring when they experience higher social densities. To look at the influences of population of origin and relative body size on competitive ability, we held females from the high‐density population at two different densities to create large and small offspring with the same genetic background. We measured the competitive ability of those offspring in mesocosms that contained either pure or mixed population treatments at either high or low density. High density increased competition, which was most evident in greatly reduced individual growth rates. Larger offspring from the high‐density population significantly delayed the onset of maturity of fish from the low‐density population. From our results, we infer that competitive conditions in nature have contributed to the evolution of genetically based interpopulation differences in offspring size as well as plasticity in offspring size in response to conspecific density.     

Using spatially‐explicit capture–recapture models to explain variation in seasonal density patterns of sympatric ursids

Understanding how environmental factors interact to determine the abundance and distribution of animals is a primary goal of ecology, and fundamental to the conservation of wildlife populations. Studies of these relationships, however, often assume static environmental conditions, and rarely consider effects of competition with ecologically similar species. In many parts of their shared ranges, grizzly bears Ursus arctos and American black bears U. americanus have nearly complete dietary overlap and share similar life history traits. We therefore tested the hypothesis that density patterns of both bear species would reflect seasonal variation in available resources, with areas of higher primary productivity supporting higher densities of both species. We also hypothesized that interspecific competition would influence seasonal density patterns. Specifically, we predicted that grizzly bear density would be locally reduced due to the ability of black bears to more efficiently exploit patchy food resources such as seasonally abundant fruits. To test our hypotheses, we used detections of 309 grizzly and 597 black bears from two independent genetic sampling methods in spatially‐explicit capture–recapture (SECR) models. Our results suggest grizzly bear density was lower in areas of high black bear density during spring and summer, although intraspecific densities were also important, particularly during the breeding season. Black bears had lower densities in areas of high grizzly bear density in spring; however, density of black bears in early and late summer was best explained by primary productivity. Our results are consistent with the hypothesis that smaller‐bodied, more abundant black bears may influence the density patterns of behaviorally‐dominant grizzly bears through exploitative competition. We also suggest that seasonal variation in resource availability be considered in efforts to relate environmental conditions to animal density.     

Intraspecific resource partitioning by an opportunistic strategist, inland silverside Menidia beryllina

Seasonal variation in prey consumption and food resource overlap was evident in an inland water body for mature male, mature female and immature inland silverside (Menidia beryllina). During the first growth phase marked by intensive somatic growth by immature inland silverside, few adults were present in the population (28% of total catch), thus minimizing intraspecific competition for food resources between juvenile and adult inland silverside. During the second growth phase by adults, few juvenile inland silverside were present (0% of total catch) in the population, again minimizing intraspecific competition for food resources between juvenile and adult inland silverside. A divergence in food resource overlap was observed when mature male, mature female and immature inland silverside were present in the population. These population‐level demographic responses to energy acquisition are likely necessary to maximize individual growth of mature male, mature female and immature inland silverside.     

Intraspecific competition in larvae of the ant-lion Morter sp. and interspecific interactions with Macroleon quinquemaculatus

Abstract.

• 1 Intraspecific competition was investigated in the pit-digging larvae of the ant-lion Morter sp., which normally occurs at low density in a food-rich habitat.
• 2 Morter larvae interact by displays, challenges, and sand tossing. Challenges/displays affected pit spacing while sand tossing affected pit size. Pit ownership determined the outcome of contests regardless of the relative sizes or feeding condition of the contestants.
• 3 Competition had no effect on the pit size of winners but losers showed a density dependent reduction in pit size. Interference competition caused a density dependent reduction in larval growth rates.
• 4 Macroleon quinquemaculatus (Hagen) always won contests with Morter, even when smaller. Cannibalism occurred in 60% of these contests.
• 5 Little intraspecific competition occurs in Morter in the field. Morter is more willing to incur energy costs than Macroleon.

     

THALLUS DIFFERENTIATION OF PHOTOSYNTHESIS,GROWTH, REPRODUCTION,AND UV‐B SENSITIVITY IN THE GREEN ALGA ULVA PERTUSA (CHLOROPHYCEAE)1

The chlorophyte Ulva is perceived as a simple and uniform algal form, with little functional differentiation within a thallus. We compared morphology, pigmentation, photosynthesis, growth, reproduction, and UV‐B sensitivity between different thallus regions of Ulva pertusa Kjellman. Thallus thickness and cell size were significantly greater, whereas cell number was less in the basal region than in other regions. Photosynthetic pigment contents were lowest in the basal region and increased toward the marginal region. Photosynthetic capacity and photosynthetic efficiency normalized to fresh weight, area, volume, and cell number showed a progressive increase from the basal to marginal parts; however, on a chl basis those values were equal regardless of thallus part. Values of light saturation point were not statistically different between regions. Growth rates increased from marginal to basal and to middle parts of the thallus, whereas sporulation was highest in marginal (100%) followed by middle (30%) and basal parts (0%). Daily observation over 9 days showed that 56% of the basal cells divided once and did not produce spores, whereas every marginal cell went through its first division and 89% of the primary daughter cells also divided, resulting in 100% sporulation. A 7‐day treatment with PAR and PAR + UV‐A caused a significant decrease in the effective quantum yield of all thallus regions, followed by a recovery toward the initial values, whereas PAR + UV‐A + UV‐B irradiation led to greater photoinhibition and less recovery. Marked differences in the UV‐B sensitivity were observed, with marginal parts being more sensitive and basal parts most resistant.     

Ontogenetic variation in density‐dependent growth of brown trout through habitat competition

1. Density‐dependent growth has been widely reported in freshwater fishes, but the ontogenetic evolution of competition and its subsequent effects on growth through a life span remains unclear. 2. Patterns of competition can be described by integrating population abundance data with habitat‐modelling results. Weighted usable area (WUA; m² WUA ha^?1) curves are obtained for each flow value and are then coupled with demographic data to obtain the occupancy rates (trout m^?2 WUA, the density of a given age class related to its suitable habitat) of the WUA for every age class, year and site. 3. We examined a long‐term data series searching for temporal variation in the influence of habitat occupancy rate on the growth of brown trout Salmo trutta. We tested whether (i) mean cohort mass (mean mass of the cohort during the first 3 years of life) is affected by the occupancy rate experienced across a life span; and (ii) the occupancy rate experienced at different ages influenced mean body size. 4. We observed a consistent negative power relationship between average cohort mass and mean occupancy rate through a life span, indicating that stronger cohorts were related to a reduced growth, with likely consequences for individual fitness. 5. The effects of occupancy rate on size‐at‐age were mainly detected in the size attained at the second year of life, but they were because of the competition at different times. Thus, the level of competition varied through ontogeny, in some of the rivers affecting growth since the first year of life, whereas in most of the rivers the main effects on body size resulted from the competition during the second year of life. 6. Occupancy rate appears more appropriate than density for assessing the occurrence of habitat competition in freshwater fishes, since it encompasses the differences in quantity and quality of suitable habitat for each age class. 7. Our study highlights the importance of density‐dependent growth as a key process in the dynamics of brown trout populations, its temporal variation depending on the temporal changes of density and the variation of competition associated with the habitat capacity for each life stage.