Simone Vincenzi

The role of density-dependent individual growth in the persistence of freshwater salmonid populations

Theoretical and empirical models of populations dynamics have paid little attention to the implications of density-dependent individual growth on the persistence and regulation of small freshwater salmonid populations. We have therefore designed a study aimed at testing our hypothesis that density-dependent individual growth is a process that enhances population recovery and reduces extinction risk in salmonid populations in a variable environment subject to disturbance events. This hypothesis was tested in two newly introduced marble trout (Salmo marmoratus) populations living in Slovenian streams (Zakojska and Gorska) subject to severe autumn floods. We developed a discrete-time stochastic individual-based model of population dynamics for each population with demographic parameters and compensatory responses tightly calibrated on data from individually tagged marble trout. The occurrence of severe flood events causing population collapses was explicitly accounted for in the model. We used the model in a population viability analysis setting to estimate the quasi-extinction risk and demographic indexes of the two marble trout populations when individual growth was density-dependent. We ran a set of simulations in which the effect of floods on population abundance was explicitly accounted for and another set of simulations in which flood events were not included in the model. These simulation results were compared with those of scenarios in which individual growth was modelled with density-independent Von Bertalanffy growth curves. Our results show how density-dependent individual growth may confer remarkable resilience to marble trout populations in case of major flood events. The resilience to flood events shown by the simulation results can be explained by the increase in size-dependent fecundity as a consequence of the drop in population size after a severe flood, which allows the population to quickly recover to the pre-event conditions. Our results suggest that density-dependent individual growth plays a potentially powerful role in the persistence of freshwater salmonids living in streams subject to recurrent yet unpredictable flood events.

Determining Individual Variation in Growth and Its Implication for Life-History and Population Processes Using the Empirical Bayes Method

The differences in demographic and life-history processes between organisms living in the same population have important consequences for ecological and evolutionary dynamics. Modern statistical and computational methods allow the investigation of individual and shared (among homogeneous groups) determinants of the observed variation in growth. We use an Empirical Bayes approach to estimate individual and shared variation in somatic growth using a von Bertalanffy growth model with random effects. To illustrate the power and generality of the method, we consider two populations of marble trout Salmo marmoratus living in Slovenian streams, where individually tagged fish have been sampled for more than 15 years. We use year-of-birth cohort, population density during the first year of life, and individual random effects as potential predictors of the von Bertalanffy growth functions parameters k (rate of growth) and (asymptotic size). Our results showed that size ranks were largely maintained throughout marble trout lifetime in both populations. According to the Akaike Information Criterion (AIC), the best models showed different growth patterns for year-of-birth cohorts as well as the existence of substantial individual variation in growth trajectories after accounting for the cohort effect. For both populations, models including density during the first year of life showed that growth tended to decrease with increasing population density early in life. Model validation showed that predictions of individual growth trajectories using the random-effects model were more accurate than predictions based on mean size-at-age of fish.

Efficiency of attack strategies on complex model and real-world networks

We investigated the efficiency of attack strategies to network nodes when targeting several complex model and real-world networks. We tested 5 attack strategies, 3 of which were introduced in this work for the first time, to attack 3 model networks (Erdos and Renyi, Barabasi and Albert preferential attachment network, and scale-free network configuration models) and 3 real networks (Gnutella peer-to-peer network, email network of the University of Rovira i Virgili, and immunoglobulin interaction network). Nodes were removed sequentially according to the importance criterion defined by the attack strategy, and we used the size of the largest connected component (LCC) as a measure of network damage. We found that the efficiency of attack strategies (fraction of nodes to be deleted for a given reduction of LCC size) depends on the topology of the network, although attacks based on either the number of connections of a node or betweenness centrality were often the most efficient strategies. Sequential deletion of nodes in decreasing order of betweenness centrality was the most efficient attack strategy when targeting real-world networks. The relative efficiency of attack strategies often changed during the sequential removal of nodes, especially for networks with power-law degree distribution.

The Effect of Recurrent Floods on Genetic Composition of Marble Trout Populations

A changing global climate can threaten the diversity of species and ecosystems. We explore the consequences of catastrophic disturbances in determining the evolutionary and demographic histories of secluded marble trout populations in Slovenian streams subjected to weather extremes, in particular recurrent flash floods and debris flows causing massive mortalities. Using microsatellite data, a pattern of extreme genetic differentiation was found among populations (global F ST of 0.716), which exceeds the highest values reported in freshwater fish. All locations showed low levels of genetic diversity as evidenced by low heterozygosities and a mean of only 2 alleles per locus, with few or no rare alleles. Many loci showed a discontinuous allele distribution, with missing alleles across the allele size range, suggestive of a population contraction. Accordingly, bottleneck episodes were inferred for all samples with a reduction in population size of 3–4 orders of magnitude. The reduced level of genetic diversity observed in all populations implies a strong impact of genetic drift, and suggests that along with limited gene flow, genetic differentiation might have been exacerbated by recurrent mortalities likely caused by flash flood and debris flows. Due to its low evolutionary potential the species might fail to cope with an intensification and altered frequency of flash flood events predicted to occur with climate change.

Spatial and temporal scale of density-dependent body growth and its implications for recruitment, population dynamics and management of stream-dwelling salmonid populations

Density-dependent variations in body growth and size have important consequences for the population dynamics of stream-dwelling salmonid populations, since body size is related to a variety of ecologically relevant characteristics. These include survival and fecundity, competitive and predatory abilities, and foraging behavior. However, little work has been done to understand how density-dependent body growth varies across temporal and spatial scales and when this compensatory process is relevant for recruitment and population dynamics of stream-dwelling salmonids. Increased intra- or inter-cohort competition reduces growth rates of juveniles. Both within- and among-cohort differences at the juvenile stage are likely to be maintained through the lifetime. Limited movement or dispersal can lead to subdivision of a population into several local populations with independent dynamics. The spatial and temporal variation in movement and the patchy distribution of resources make fish likely to experience density-dependence across location, life-stage, and season. The relaxation of density-dependent suppression of body growth at low densities constitutes a potential mechanism for salmonids to persist in the face of environmental perturbation and may contribute to explaining the peculiar resilience to population collapses often showed by salmonids. The inclusion of density-dependent growth in population models may increase the usefulness of model predictions in management contexts. Models not accounting for density-dependent growth may underestimate the recovery potential of resident salmonid populations when they collapse to low densities.

Density-dependent individual growth of marble trout (Salmo marmoratus) in the Soca and Idrijca river basins, Slovenia

Although the main features of salmonid life cycles are currently well known, marble trout (Salmo marmoratus) populations have been scarcely studied due to the present limited geographical distribution of the species. In this work we tested the hypothesis of density-dependent individual growth of marble trout with data gathered from multi-year on-going monitoring started in 1996 in three streams (Zakojska, Gorska and Gatsnik) in the Soca and Idrijca river basins (Slovenia). As observed for other salmonid species, marble trout exhibits high plasticity in body size and growth in response to environmental conditions. Age-specific mean lengths are significantly different among the three streams. Despite this variability, the statistical analyses outlined some clear patterns. Mean lengths of marble trout cohorts aged ≥1+ are correlated with total population density in the first year of life of the cohorts; the age-specific relationship between mean length and total density is well explained by negative power curves. Length of marble trout at age 1+ is significantly correlated with length at subsequent years up to age 4+. ANCOVAs performed on the stream-specific datasets showed a significant effect of total density of marble trout on annual age-specific individual growth rates. Von Bertalanffy’s body growth models were calibrated on length–age data of Zakojska and Gorska marble trout. Our analysis shows the existence of density-dependent effects on growth of marble trout that might be crucial in regulating population dynamics.

Eco-evolutionary dynamics induced by massive mortality events

An eco-genetic model tuned on a population of marble trout Salmo marmoratus subject to periodic flood events was used to explore how the evolution of growth rates interacting with density-dependent processes can modify size at age and population structure and in turn influence the resilience of populations. Fish with greater growth potential were assumed to have higher mortality rates. The results of simulations were compared between two scenarios, one in which populations may evolve growth rates and the other one in which the distribution of growth rates within a population is kept fixed. Evolving populations had a greater proportion of age 1 year individuals in the population, greater median length at age 3 years (the typical age at sexual maturity for S. marmoratus) and lower population sizes. The slightly smaller population sizes did not affect realized extinction risk. Resilience, defined as the number of years necessary to rebound from flood-induced population collapse, was on average from 2 to 3 years in both scenarios, with no significant difference between them. Moderate heritability of growth, relaxation of density-dependent processes at low densities and rapid recovery to a safe population size combine to limit the capacity to evolve faster recovery after flood-induced population collapses via changing growth rates.

Density-dependent and inter-specific interactions affecting European eel settlement in freshwater habitats

Identifying the factors influencing the settlement of European eel (Anguilla anguilla) juveniles in continental habitats is crucial to designing effective management and conservation measures for this endangered species. A long-term data series (1993–2008) of European eel and European catfish (Silurus glanis) abundance in a freshwater canal of the Camargue water system (southern France), along with parallel data on water salinity and glass eel abundance in the adjacent Vaccarès lagoon, was analysed to identify the possible causes of decline in eel abundance observed in the canal during the last two decades. A model including glass eel recruitment and catfish abundance as covariates explained 78% of the observed variation in eel settlement success. Results suggest that (1) salinity does not play a significant role in determining the fraction of eels moving from the brackish lagoon to the canal; (2) density dependence affects settlement success, possibly through a reduction of juvenile survival in the adjacent lagoon; and (3) catfish abundance is negatively correlated with eel settlement. We discuss this latter point in terms of possible predation of catfish upon eels and/or inter-specific competition between the two species.

Food availability affects onset of reproduction in a long-lived seabird

Life-history theory predicts that suboptimal developmental conditions may lead to faster life histories (younger age at recruitment and higher reproductive investment), but experimental testing of this prediction is still scarce in long-lived species. We report the effects of an experimental manipulation of food availability during early development and at recruitment on the onset of reproduction and reproductive performance (productivity at first breeding) in a long-lived seabird, the black-legged kittiwake Rissa tridactyla, breeding on Middleton Island, Alaska. Birds were born and raised in nests with supplemented food (‘fed’) or unsupplemented control nests (‘unfed’), and later recruited into either fed or unfed nests. Fed chicks grew faster than unfed chicks, and males grew faster than females. Birds were more likely to reproduce at younger ages when recruiting into fed nests. Faster growth during development tended to increase age at recruitment in all individuals. Social rank of individuals also affected age at recruitment: B-chicks recruited earlier than A-chicks and singletons recruited later than A- and B-chicks. Productivity increased with the age at recruitment and growth rate as chick, but much of the variability remained unexplained. We conclude that results of this study at least partially support predictions of life-history theory: younger age at first breeding for kittiwakes that experienced suboptimal natal conditions, as well as greater productivity of early recruiting kittiwakes that grew in control nests compared with those that grew in food-supplemented nests.

Selective consequences of catastrophes for growth rates in a stream-dwelling salmonid

Optimal life histories in a fluctuating environment are likely to differ from those that are optimal in a constant environment, but we have little understanding of the consequences of bounded fluctuations versus episodic massive mortality events. Catastrophic disturbances, such as floods, droughts, landslides and fires, substantially alter the population dynamics of affected populations, but little has been done to investigate how catastrophes may act as a selective agent for life-history traits. We use an individual-based model of population dynamics of the stream-dwelling salmonid marble trout (Salmo marmoratus) to investigate how trade-offs between the growth and mortality of individuals and density-dependent body growth can lead to the maintenance of a wide or narrow range of individual variation in body growth rates in environments that are constant (i.e., only demographic stochasticity), variable (i.e., environmental stochasticity), or variable with catastrophic events that cause massive mortalities (e.g., flash floods). We find that occasional episodes of massive mortality can substantially reduce persistent variability in individual growth rates. Lowering the population density reduces density dependence and allows for higher fitness of more opportunistic strategies (rapid growth and early maturation) during the recovery period.