Density-dependent Growth Research Articles

Harvest and density‐dependent predation drive long‐term population decline in a northern ungulate

The relative effect of top‐down versus bottom‐up forces in regulating and limiting wildlife populations is an important theme in ecology. Untangling these effects is critical for a basic understanding of trophic dynamics and effective management. We examined the drivers of moose (Alces alces) population growth by integrating two independent sources of observations within a hierarchical Bayesian population model. We used one of the largest existing spatiotemporal data sets on ungulate population dynamics globally. We documented a 20% population decline over the period examined. There was negative density‐dependent population growth of moose. Although we could not determine the mechanisms producing density‐dependent suppression of population growth, the relatively low densities at which we documented moose populations suggested it could be due to density‐dependent predation. Predation primarily limited population growth, except at low density, where it was regulating. After we simulated several harvest scenarios, it appeared that harvest was largely additive and likely contributed to population declines. Our results highlight how population dynamics are context dependent and vary strongly across gradients in climate, forest type, and predator abundance. These results help clarify long‐standing questions in population ecology and highlight the complex relationships between natural and human‐caused mortality in driving ungulate population dynamics.

Life and death in a dynamic environment: Invasive trout, floods, and intraspecific drivers of translocated populations.

Understanding the relative strengths of intrinsic and extrinsic factors regulating populations is a long‐standing focus of ecology and critical to advancing conservation programs for imperiled species. Conservation could benefit from an increased understanding of factors influencing vital rates (somatic growth, recruitment, survival) in small, translocated populations, which is lacking owing to difficulties in long‐term monitoring of rare species. Translocations, here defined as the transfer of wild‐captured individuals from source populations to new habitats, are widely used for species conservation, but outcomes are often minimally monitored, and translocations that are monitored often fail. To improve our understanding of how translocated populations respond to environmental variation, we developed and tested hypotheses related to intrinsic (density dependent) and extrinsic (introduced rainbow trout Oncorhynchus mykiss, stream flow and temperature regime) causes of vital rate variation in endangered humpback chub (Gila cypha) populations translocated to Colorado River tributaries in the Grand Canyon (GC), USA. Using biannual recapture data from translocated populations over 10 years, we tested hypotheses related to seasonal somatic growth, and recruitment and population growth rates with linear mixed‐effects models and temporal symmetry mark–recapture models. We combined data from recaptures and resights of dispersed fish (both physical captures and continuously recorded antenna detections) from throughout GC to test survival hypotheses, while accounting for site fidelity, using joint live‐recapture/live‐resight models. While recruitment only occurred in one site, which also drove population growth (relative to survival), evidence supported hypotheses related to density dependence in growth, survival, and recruitment, and somatic growth and recruitment were further limited by introduced trout. Mixed‐effects models explained between 67% and 86% of the variation in somatic growth, which showed increased growth rates with greater flood‐pulse frequency during monsoon season. Monthly survival was 0.56–0.99 and 0.80–0.99 in the two populations, with lower survival during periods of higher intraspecific abundance and low flood frequency. Our results suggest translocations can contribute toward the recovery of large‐river fishes, but continued suppression of invasive fishes to enhance recruitment may be required to ensure population resilience. Furthermore, we demonstrate the importance of flooding to population demographics in food‐depauperate, dynamic, invaded systems.

Predicting density-dependent somatic growth in Norwegian spring-spawning herring

Abstract Density-dependent growth, which might influence the effects of fisheries on a population, is often ignored when management strategies are evaluated, mainly due to a lack of appropriate models readily available to be implemented. To improve on this, we investigated if somatic growth in Norwegian spring-spawning herring (Clupea harengus) depends on cohort density using a formulation of the von Bertalanffy growth function on cohorts from 1921 to 2014 and found a significant negative correlation between estimated asymptotic length and density. This clearly indicates density-dependent effects on growth, and we propose a model that can be used to predict the size-at-age of Norwegian spring-spawning herring as a function of herring density (the abundance of two successive cohorts) in short-term predictions of catch advice, and in Management strategy evaluations, including estimation of their reference points such as FMSY.

Assessment of the effect of stand density on the height growth of Scots pine using repeated ALS data

• Repeated ALS allows detecting the height increment variation influenced by stand density. • Higher stand density stimulate height growth, particularly on most productive sites. • Repeated ALS data allows developing density-dependent top height growth models. The spread of remote sensing technologies allows for continuous and precise mapping of forest ecosystem attributes over large areas. One of the most important attributes of a forest ecosystem is site productivity, an essential determinant of resource availability. Knowledge of site productivity is crucial for sustainable forest management because it is the basis for predictions of future forest growth and is fundamental to long-term planning in forestry. Models of forest dynamics allow us to better understand the functioning of forest ecosystems and to assess the provision of ecosystem services. However, discussions on the factors affecting forest growth are ongoing. Previous research has not clarified the influence of stand density on the height increment of trees. Consequently, this uncertainty leads to inconsistent predictions for stand growth and nonoptimal forest-management decisions. We aimed to analyse the effect of stand density on top height (TH) growth and the development of the TH growth model for Scots pine based on bitemporal ALS measurements. We applied repeated ALS observations from two acquisitions (2011, 2019) to fit a density-sensitive TH growth model. We found that stand density affected the TH growth of Scots pine. A higher stand density stimulated TH growth, particularly in productive sites showing a TH increment of about 10%. We demonstrated the usefulness of bitemporal ALS measurements for developing a TH growth model for Scots pine considering stand density. We found that the predictive accuracy of the TH increment increased when stand density was considered.

Cell density-dependent antibiotic tolerance to inhibition of the elongation machinery requires fully functional PBP1B

The peptidoglycan (PG) cell wall provides shape and structure to most bacteria. There are two systems to build PG in rod shaped organisms: the elongasome and divisome, which are made up of many proteins including the essential MreB and PBP2, or FtsZ and PBP3, respectively. The elongasome is responsible for PG insertion during cell elongation, while the divisome is responsible for septal PG insertion during division. We found that the main elongasome proteins, MreB and PBP2, can be inhibited without affecting growth rate in a quorum sensing-independent density-dependent manner. Before cells reach a particular cell density, inhibition of the elongasome results in different physiological responses, including intracellular vesicle formation and an increase in cell size. This inhibition of MreB or PBP2 can be compensated for by the presence of the class A penicillin binding protein, PBP1B. Furthermore, we found this density-dependent growth resistance to be specific for elongasome inhibition and was consistent across multiple Gram-negative rods, providing new areas of research into antibiotic treatment.

Interacting effects of density and temperature on fish growth rates in freshwater protected populations.

Despite the demonstrated benefits of marine protected areas, there has been relatively little dialogue about freshwater protected areas (FPAs) even though some have been established to protect freshwater species from recreational and commercial fishers. After populations recover from fishing pressure, abundances and densities of formerly fished species increase, and we should therefore expect changes in demographic traits compared to those in exploited populations. To test this, we used capture-mark-recapture data for 10 Galaxias maculatus populations across a density gradient mediated by different degrees of fishery closure. We examined the extent to which density-dependent (DD) and density-independent (DI) effects interact to affect specific growth rates in post-recruit populations. We found that population density, stream temperature and individual size interact to affect growth rates. When population densities were high, compensatory responses of far slower growth rates were strongest, indicating that DD growth is a key mechanism regulating post-recruit populations of G. maculatus. This study emphasizes the importance of understanding DD and DI processes, their interactions, function and effectiveness for freshwater fisheries management. For FPAs to be effective, the extent and quality of target species' habitats must serve as key criteria for protection to alleviate competition for limited resources that underpins DD processes.

Bifurcation analysis of a predator-prey system with density dependent disease recovery

The center manifold is an invariant manifold that plays a crucial role in the bifurcation analysis of dynamical systems. The center manifold existence theorem assures the local existence of an invariant submanifold of the state space of a dynamical system around a non-hyperbolic equilibrium point. Center manifold theory is essential in the reduction of different bifurcation scenarios to their normal forms. Our study focuses on a predator-prey interactive system with density-dependent growth in predators subject to a contagious disease. The disease is assumed to be horizontally transmitted, and the rate of recovery of the infected predator is assumed to be density-dependent. At the trivial (zero) equilibrium, the center manifold is calculated whose dynamical behaviour is similar to that of the original system. Further, using the center manifolds, the normal form of a Hopf bifurcation point is determined fromwhich the criticality of the system can be deduced. Finally, numerical simulations are performed with biologically plausible parameters to substantiate the analytical findings. Using numerical continuation methods we detect Generalized Hopf and Zero-Hopf bifurcation points. We discuss their normal form coefficients, compute their two-parameter unfoldings and relate these results to the mathematical theory of codimension two bifurcations.

Breeding Season Survival and Reproduction in a High-Density Bobwhite Population: A case study

The demographic behavior of northern bobwhite (Colinus virginianus; hereafter, bobwhite) populations at high densities could provide important insights into why bobwhite populations fluctuate. Therefore, we documented breeding season demographics of bobwhites to understand how prebreeding density influenced reproductive effort and postbreeding density on an intensively managed property in Leon County, Florida, USA, 2002–2006. We estimated prebreeding bobwhite density each April using multi-observer strip-transects and postbreeding densities each November using covey call grid surveys. We radio-tagged 217 bobwhites in March and located bobwhites at least 5 days/week, 15 April–30 September to determine vital rates. Prebreeding density ranged from 1.5–8.6 birds/ha, peaking in 2002, declining through 2005, then increasing in 2006. Breeding season survival was 0.55, 0.17, 0.20, and 0.59, and nesting rate was 0.47, 0.67, 0.80 and 0.89, 2002–2005, respectively. Postbreeding density ranged from 5.2–13.6 birds/ha, also peaking in 2002 and declining through 2004 before increasing beginning in 2005 and 2006. High breeding season survival and nesting success (>0.55) resulted in greater chick production during periods of population growth. Nesting rate was inversely related to prebreeding density. Declines in bobwhite nesting rate at high prebreeding densities appeared to regulate population growth near population peaks. Lower adult survival and nesting success appeared to cause population declines. We suggest density-dependent intraspecific competition limited population growth at high bobwhite densities by reducing nesting rate while predation of adults and nests explained population fluctuations.

Developing integral projection models for ecotoxicology

In many ecosystems, especially aquatic ecosystems, size plays a critical role in the factors that determine an individual's ability to survive and reproduce. In aquatic ecotoxicology, size informs both realized and potential acute and chronic effects of chemical exposure. This paper demonstrates how chemical and nonchemical effects on growth, survival, and reproduction can be linked to population-level dynamics using size-structured integral projection models (IPM). The modeling approach was developed with the goals and constraints of ecological risk assessors in mind, who are tasked with estimating the effects of chemical exposures to wildlife populations in a data-limited environment. The included case study is a collection of daily time-step IPMs parameterized for the life history and annual cycle of fathead minnows (Pimephales promelas), which motivated the development of modeling techniques for seasonal, iteroparous reproduction, density dependent growth effects, and size-dependent over-winter survival. The effects of a time-variable annual chemical exposure were interpreted using a toxicokinetic-toxicodynamic model for acute survival and sub-lethal growth effects model for chronic effects and incorporated into the IPMs. This paper presents a first application of integral projection models to ecotoxicology. Our research demonstrates that size-structured IPMs provide a promising, flexible, framework for synthesizing ecotoxicologically relevant data and theory to explore the effects of chemical and nonchemical stressors and the resulting impacts on exposed populations.

Intra- and inter-specific density dependence of body condition, growth, and habitat temperature in chub mackerel (Scomber japonicus)

Abstract The density dependence of growth and body condition have important impacts on fish population dynamics and fisheries management. Although population density affects habitat selection, which in turn changes habitat temperature, how this affects growth and body condition remains unclear. Here, we investigated annual changes in body condition, habitat temperature, and cohort-specific growth of chub mackerel (Scomber japonicus) in the western North Pacific and examined quarterly changes in the density dependence of body condition. We hypothesized that chub mackerel body condition is affected both directly (e.g. through competition for food) and indirectly (through changes in habitat temperature) by the abundance of both conspecifics (i.e. chub mackerel) and heterospecifics (the Japanese sardine Sardinops melanostictus). Indeed, chub mackerel body condition, habitat temperature, and growth all decreased with increasing conspecific and heterospecific abundance. Mean annual growth rates in chub mackerel were positively corelated with body condition. The final model showed that conspecific and/or heterospecific abundance had strong negative effects on chub mackerel body condition in all seasons, and influenced habitat temperature through habitat selection in some seasons. By contrast, temperature effects on body condition were weak. Therefore, direct effects likely have more impact than indirect effects on density-dependent body condition and growth.

Lifting the Vendace, Coregonus albula, on the Life Table: Survival, Growth and Reproduction in Different Life-Stages during Very High and Low Abundance Regimes

We analyzed the variability in vendace population parameters in the context of a life table. Parameters related to growth, fecundity and survival were estimated for post-recruitment (age > 1 growing season) life stages of Lake Southern Konnevesi vendace in both very high and low population abundance regimes. Pre-recruitment survival producing population stability was then determined. We found a very strong compensatory density dependence in growth, fecundity and survival: during low abundance, a lifetime reproductive output of a female was almost 20 times that of the abundant regime. To maintain the low abundance regime, pre-recruitment survival must counterbalance it by decreasing to a very low level. Potential drivers of high variability in pre-recruit survival and potential for depensation, Allee effect, are discussed, as well as the implications of the results on fisheries management and risk of extinction due to anthropogenic stressors such as global warming or extensive predator stocking.

Evaluating growth, production and economics of a new candidate species Labeo bata in cages: A regional model for table fish production in floodplain wetlands of North East India

Indian minor carp Labeo bata fingerlings were reared in square shaped cages at different numbers to evaluate the effect of stocking density on growth, production and economics. Fifteen cages (5 × 5 × 2 m each) were installed in deeper areas (6–8 m) of Samaguri beel (a seasonally-flooded floodplain wetland) of Assam, India. The cages were stocked with L.bata (mean length 11.93 ± 0.20 cm; mean weight 13.05 ± 0.77 g) at five different stocking densities of 10 (SD10), 20 (SD20), 30 (SD30), 40 (SD40) and 50 (SD50) fingerlings m−3 in triplicates. Fish were fed with floating pellets containing 32% CP at 3–5% body weight twice-a-day for six months. Density-dependent growth was evident from the results as growth of fish decreased with increasing stocking densities; the specific growth rate of the fish at SD10, SD20, SD30, SD40 and SD50 were 1.03 ± 0.02, 0.88 ± 0.01, 0.84 ± 0.02, 0.67 ± 0.02 and 0.52 ± 0.01 with corresponding weight gain percent of 533.3 ± 13.41, 385.8 ± 10.73, 353.3 ± 14.17, 232.2 ± 8.08 and 155.9 ± 5.38, respectively. However, the highest net yield of fish was achieved at SD30 (50.8 kg cage−1), which was significantly higher than other stocking densities. Also, economic analysis demonstrated highest benefit-cost (B:C) ratio at 30 fish m−3 (1.44). Results indicated decreased fish yield at lower densities though the individual growth rate of fish was higher. On the other hand, fingerling stocking more than 30 fish m−3 was uneconomical because of lower individual growth rate. Hence, a stocking density of 30 fish m−3 could be considered as optimum for production of L.bata table fish in cages. The study generated new and critical information on species diversification in cage aquaculture for tropical floodplain wetlands.

Modeling the Allee effects induced by cost of predation fear and its carry-over effects

Predation-driven Allee effects play an important role in the dynamics of a small prey population; however, such effects cannot occur for the model with type I functional response. Predator-driven Allee effects generally occur when a generalist predator targets some specific prey. However, apart from the lethal effects of predation, there are some non-lethal effects in the presence of a predator. Due to the fear of predation, positive density dependence growth of prey may be observed at low population density because of reduced foraging activities. Moreover, such a non-lethal effect can be carried over through generations or seasons. In the present manuscript, we investigate the role of predation fear and its carry-over effects in the prey-predator model. First, we study the single-species model from a global perspective. We have shown that depending on the birth rate; our single-species model describes three types of growth dynamics, namely, strong Allee dynamics, weak Allee dynamics, and logistic dynamics. Then we include the explicit dynamics of the predator, with type I functional response. Basic dynamical properties, as well as the global stability of each equilibrium, have been discussed. From our analysis, we can observe that both the fear and its carry-over effects have a significant role in the stability of the coexistence equilibrium, even for the model with type I functional response. The phenomenon ‘paradox of enrichment’ can be observed in our model, which cannot be observed in the classical prey-predator model with type I functional response. However, we can see that such a phenomenon can be ruled out by choosing suitable non-lethal effect parameters. Therefore, our study shows how non-lethal effects change the dynamics of a prey-predator model and has powerful biological insights, especially for understanding the dynamics of small populations.

Diffusion modeling reveals effects of multiple release sites and human activity on a recolonizing apex predator

BackgroundReintroducing predators is a promising conservation tool to help remedy human-caused ecosystem changes. However, the growth and spread of a reintroduced population is a spatiotemporal process that is driven by a suite of factors, such as habitat change, human activity, and prey availability. Sea otters (Enhydra lutris) are apex predators of nearshore marine ecosystems that had declined nearly to extinction across much of their range by the early 20th century. In Southeast Alaska, which is comprised of a diverse matrix of nearshore habitat and managed areas, reintroduction of 413 individuals in the late 1960s initiated the growth and spread of a population that now exceeds 25,000.MethodsPeriodic aerial surveys in the region provide a time series of spatially-explicit data to investigate factors influencing this successful and ongoing recovery. We integrated an ecological diffusion model that accounted for spatially-variable motility and density-dependent population growth, as well as multiple population epicenters, into a Bayesian hierarchical framework to help understand the factors influencing the success of this recovery.ResultsOur results indicated that sea otters exhibited higher residence time as well as greater equilibrium abundance in Glacier Bay, a protected area, and in areas where there is limited or no commercial fishing. Asymptotic spread rates suggested sea otters colonized Southeast Alaska at rates of 1–8 km/yr with lower rates occurring in areas correlated with higher residence time, which primarily included areas near shore and closed to commercial fishing. Further, we found that the intrinsic growth rate of sea otters may be higher than previous estimates suggested.ConclusionsThis study shows how predator recolonization can occur from multiple population epicenters. Additionally, our results suggest spatial heterogeneity in the physical environment as well as human activity and management can influence recolonization processes, both in terms of movement (or motility) and density dependence.

Virus-Host Interactions Shape Viral Dispersal Giving Rise to Distinct Classes of Traveling Waves in Spatial Expansions

Reaction-diffusion waves have long been used to describe the growth and spread of populations undergoing a spatial range expansion. Such waves are generally classed as either pulled, where the dynamics are driven by the very tip of the front and stochastic fluctuations are high, or pushed, where cooperation in growth or dispersal results in a bulk-driven wave in which fluctuations are suppressed. These concepts have been well studied experimentally in populations where the cooperation leads to a density-dependent growth rate. By contrast, relatively little is known about experimental populations that exhibit density-dependent dispersal. Using bacteriophage T7 as a test organism, we present novel experimental measurements that demonstrate that the diffusion of phage T7, in a lawn of host E. coli, is hindered by steric interactions with host bacteria cells. The coupling between host density, phage dispersal and cell lysis caused by viral infection results in an effective density-dependent diffusion coefficient akin to cooperative behavior. Using a system of reaction-diffusion equations, we show that this effect can result in a transition from a pulled to pushed expansion. Moreover, we find that a second, independent density-dependent effect on phage dispersal spontaneously emerges as a result of the viral incubation period, during which phage is trapped inside the host unable to disperse. Additional stochastic agent-based simulations reveal that lysis time dramatically affects the rate of diversity loss in viral expansions. Taken together, our results indicate both that bacteriophage can be used as a controllable laboratory population to investigate the impact of density-dependent dispersal on evolution, and that the genetic diversity and adaptability of expanding viral populations could be much greater than is currently assumed.

Density-dependent plant growth drives grazer stimulation of aboveground net primary production in Yellowstone grasslands

The mechanisms by which grazing animals influence aboveground net primary production (ANPP) in grasslands have long been an area of active research. The prevailing wisdom is that grazing can increase ANPP by increasing the availability of growth-limiting resources such as nitrogen and water, but recent theory suggests that the density-dependent growth of grassland vegetation can lead to grazer-stimulation of ANPP simply by removing shoot biomass and increasing relative growth rate (RGR). We compared the relative roles of resource availability and density-dependent growth in driving positive responses of ANPP to grazing in Yellowstone National Park. We measured the effects of clipping (50% simulated grazing intensity) and natural grazing on soil nitrogen availability, soil moisture, and shoot growth over 2months in two grassland plant communities (mesic and dry) grazed primarily by bison. Clipping increased RGR by over 100% in both grassland types but had no effect on N availability or soil moisture during the same growth periods. Clipping stimulated ANPP only at mesic grassland, and the magnitude of this effect was strongly related to the initial plant biomass at the time of clipping relative to estimated peak biomass, supporting the density-dependent framework. Bison grazing had qualitatively similar effects on ANPP and RGR to clipping with no accompanying effects on N availability or soil moisture. Our results show how grazing can stimulate ANPP independent of a direct influence on resource availability simply by exploiting the dynamics of density-dependent plant growth.

Elephant population responses to increased density in Kruger National Park

The elephant population in the Kruger National Park (KNP) has been increasing since the cessation of culling in the mid-1990s. This contrasts with recent trends in elephant populations in many parts of Africa where poaching continues to decrease numbers. Logistic growth theory predicts that increased competition for vital resources when densities increase should serve to constrain population growth, implying a negative density-growth correlation. We tested this prediction using 28 years of elephant census data to investigate how the growth of the KNP’s elephant population responds to increasing elephant density from the period 1985 to 2012. We expected a spatially variable population growth pattern in response to the distribution of elephant densities in the park and thus classified the park into zones with low, medium or high long-term (28 years) average, dry-season elephant density. Zones were named ‘peripheral’, ‘semi-peripheral’ and ‘core’ zones, respectively, and represent proxies of resource availability to elephant herds. Using a Stochastic Ricker growth model, we tested for the presence of negative density-dependence in population growth in the core versus peripheral zones. In response, we only detected density-dependent growth in the core zone. Overall the population grew at 4.1% per year, coupled with local recruitment rates that increased over time, particularly in the peripheral zones. These density-dependent trends support previous observations of homogenisation of elephant distribution and density across the KNP landscapes.Conservation implications: Density-dependent changes to elephant growth rates are scale-dependent (local vs. park level). Only core areas with long-term high density show signs of density-dependent growth. Overall, the distributions of elephants are homogenising in the KNP. Conservation authorities should monitor the impact of such homogenisation to landscape heterogeneity. The spatial variation of the negative density-growth correlation, especially between the core and peripheral zones, can be considered when developing effective strategies to manage the KNP elephant population.

GROWTH MODELS IN MICROBIAL ECOSYSTEMS - RESOURCE OR DENSITY DEPENDENCE?

This paper aims at discussing the two main modeling schemes that are used to describe dynamically the growth of microbial ecosystems, that are resource and density-dependent growth functions, respectively. Monod has been the first to hypothesize that this growth is, before all, an increasing saturated function of the main limiting substrate concentration. Contois assumed that the growth is not only a function of the substrate but also of the biomass density-itself, and thus the name « density-dependent ». In re examining their respective experiments (species used, conditions of experiments, mode of reactor functioning, measurement techniques), we try to understand the engines for a density-dependent phenomenon to appear. In particular, we refer to recent experiments where it was shown that density-dependent appeared as soon as the biomass structures into flocs or in the presence of filamentous bacteria even at relatively low concentrations. Based on this historical review of data, it is shown that density-dependent kinetics is not systematically a question of biomass density but rather related to its structure within the medium and to the mobility of microbial cells.

Early life growth is affecting timing of spawning in the semelparous Barents Sea capelin (Mallotus villosus)

Capelin (Mallotus villosus) is a forage fish and a key species in the Barents Sea (BS). The BS capelin are semelparous and hence only spawn once along the north coasts of Norway and Russia before they die. The age at spawning ranges from 2 to 5 years and the spawning season peaks in March/April but starts in February and lasts until June, and the causes of the variability in timing of spawning are not well understood. Here, we aimed to find out whether early growth is associated with the timing of spawning in BS capelin, both on the individual and population level, and if there is an association between early life growth and the spatial distribution at the nursery areas and feeding grounds. For the analysis, we used an extensive dataset comprising >150000 otolith growth zone measurements carried out during surveys from 1976 to 2019 both from the spawning and feeding areas. The data from the feeding area showed that capelin with good first-year growth were found in the productive north-west part of the Barents Sea at both age 1 and 2, while capelin with relatively poor first-year growth were typically found in the south-east Barents Sea. The data from the spawning area showed on the individual level that capelin with good first-year growth tend to spawn both at a younger age and earlier in the season. The capelin spawning late in the season were also generally smaller than early spawners. On the population level, a contradictory pattern was observed where the proportion of maturing capelin at age 2 and 3 was negatively correlated with first-year growth indicating that the great variability in year-class strength masks the general effect found at the level of individual fish. Furthermore, first-year growth was positively associated with the abundance of 1-year-old capelin indicating that rapid growth early in life enhances recruitment. On the other hand, first-year growth was strongly negatively correlated with third-year growth suggesting an increased effect of density-dependent growth with age and/or compensatory growth, or reduced growth linked to earlier maturation. In sum, our results show that the first-year growth affects growth, maturation processes, and timing of spawning later in life, thus potentially strongly influencing capelin population dynamics.

No evidence for reduced growth in resident fish species in the era of de-eutrophication in a coastal area in NW Europe

Coastal areas in north-western Europe have been influenced by elevated nutrient levels starting in the 1960s. Due to efficient measures, both nitrate and phosphate levels decreased since the mid-1980s. The co-occurring declines in nutrient loadings and fish productivity are often presumed to be causally linked. We investigated whether four resident fish species (twaite shad, bull-rout, thick-lipped grey mullet and eelpout), that spend the majority of their life in the vicinity of the coast, differed in growth between the historic eutrophication period compared to the recent lower nutrient-level period. Based on Von Bertalanffy growth models of length at age, and the analysis of annual otolith increments, we investigated the difference in sex-specific growth patterns and related these to temperature, eutrophication level (Chlorophyll a), growth window and fish density. In all four species, annual otolith growth rates during the early life stages differed between the two periods, mostly resulting in larger lengths at age in the recent period. All species showed significant correlations between increment size and temperature, explaining the observed period differences. The lack of an effect of total fish biomass provided no evidence for density dependent growth. A correlation with chlorophyll was found in bull-rout, but the relationship was negative, thus not supporting the idea of growth enhanced by high nutrient levels. In conclusion, we found no evidence for reduced growth related to de-eutrophication. Our results indicate that temperature rise due to climate change had a greater impact on growth than reduced food availability due to de-eutrophication. We discuss potential consequences of growth changes for length-based indicators used in management.