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.     

An experimental analysis of self-thinning in juvenile steelhead trout

Mobile animal populations have been proposed to decline in density according to a slope based on the allometry of metabolic requirements or space requirements. In salmonid fishes, metabolic rate and food consumption scale to body mass by the exponent 0.87 and 0.73, respectively; whereas the territory size of steelhead trout scales to body mass by the exponent 0.86. Experimental cohorts of juvenile steelhead trout ( Oncorhynchus mykiss ) were used to test the hypothesis that mobile animal populations composed of individuals with indeterminate growth decline in density as a result of self-thinning. After controlling for experimentally manipulated levels of food abundance and stocking density, cohorts of steelhead trout declined in density with increasing body size according to a slope closest to the allometry of food consumption. Densities of steelhead trout were inversely related to average mass by the exponent −0.74. Despite the similarity to the food consumption slope, a relatively wide confidence interval also precluded distinguishing the slope either the metabolic rate or territory size slopes. Data from the literature were also examined to determine if there was general support for the idea of self-thinning in natural populations of stream-dwelling salmonid fish. Although not all data suggest that populations of salmonids in streams decline as a result of density-dependent intraspecific competition, at least some appear to fit the idea of self-thinning; especially when density is above a minimum level of habitat saturation.     

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. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

Effect of water temperature and density of juvenile salmonids on growth of young-of-the-year Atlantic salmon Salmo salar

A von Bertalanffy growth model for young-of the-year Atlantic salmon Salmo salar in a small French coastal stream was fitted using water temperatures and densities of juvenile salmonids (S. salar and brown trout Salmo trutta) as covariates influencing daily growth rate. The Bayesian framework was used as a template to integrate prior information from external data sets. The relative influence of the covariates on parr growth was quantified and results showed that growth of S. salar juveniles depended on both water temperatures and densities, but that most of the spatiotemporal variability of growth resulted from local spatiotemporal variations of 0+ age salmonid (S. salar and S. trutta) densities. Further analysis revealed that the fluctuations in young-of-the-year salmonid densities are likely to dominate the effects of potential future warming of water temperature due to climate change. It is concluded that factors that could affect salmonid densities might well have a greater effect on S. salar population dynamics than factors influencing water temperatures.     

Geomorphological features drive spatiotemporal dynamics of young-of-the-year brown trout populations in a large New Zealand river catchment

1. Determining the factors driving population dynamics of stream salmonids across broad landscapes is important for understanding stream ecosystem functioning and for the management and conservation of the populations of such fish. A few studies have described the determinants of the early life history of salmonids across catchments, but none of them have examined how this spatial variation develops temporally.
2. We hypothesised that: (1) spatiotemporal dynamics of young-of-the-year (YoY) trout populations vary within the catchment, with loss rates (emigration plus mortality) being higher in lower elevation, downstream reaches than in higher elevation, upstream ones; and (2) loss rate would be density dependent, leading to reduced intra- and inter-cohort competition and supporting establishment of resident populations.
3. We tested the first hypothesis by examining relationships between geomorphic characteristics of spawning streams and temporal dynamics of YoY brown trout density, loss rate and biological traits through an austral summer across the catchment of New Zealand's fourth-longest river, the Taieri River. To test the second hypothesis, we examined whether initial YoY density in spring and other biotic determinants affected loss rate and cohort structure dynamics, resulting in density-dependent self-thinning of YoY fish to avoid resource limitation.
4. Spring YoY density and subsequent loss rate was higher in low elevation sites, corresponding to stream accessibility to migratory spawners. These significant correlations of YoY density and related biotic variables with elevation disappeared later in the season, indicating that regardless of their origin, brown trout YoY tend to form stream resident populations with multicohort structure and densities low enough to limit intraspecific competition.
5. Our findings provide evidence that geomorphological features of the landscape affect temporal dynamics of YoY trout populations driven by density-dependent self-regulatory mechanisms that can control the abundance of growing fish.

     

Life-History and Spatial Determinants of Somatic Growth Dynamics in Komodo Dragon Populations

Somatic growth patterns represent a major component of organismal fitness and may vary among sexes and populations due to genetic and environmental processes leading to profound differences in life-history and demography. This study considered the ontogenic, sex-specific and spatial dynamics of somatic growth patterns in ten populations of the world’s largest lizard the Komodo dragon (Varanus komodoensis). The growth of 400 individual Komodo dragons was measured in a capture-mark-recapture study at ten sites on four islands in eastern Indonesia, from 2002 to 2010. Generalized Additive Mixed Models (GAMMs) and information-theoretic methods were used to examine how growth rates varied with size, age and sex, and across and within islands in relation to site-specific prey availability, lizard population density and inbreeding coefficients. Growth trajectories differed significantly with size and between sexes, indicating different energy allocation tactics and overall costs associated with reproduction. This leads to disparities in maximum body sizes and longevity. Spatial variation in growth was strongly supported by a curvilinear density-dependent growth model with highest growth rates occurring at intermediate population densities. Sex-specific trade-offs in growth underpin key differences in Komodo dragon life-history including evidence for high costs of reproduction in females. Further, inverse density-dependent growth may have profound effects on individual and population level processes that influence the demography of this species.     

Is the shape of the density-growth relationship for stream salmonids evidence for exploitative rather than interference competition?

1. Empirical studies show that average growth of stream-dwelling salmon and trout often declines with increasing density in a characteristic concave relationship. However, the mechanisms that generate negative density-growth relationships in populations in natural streams are not certain. 2. In a recent study, Imre, Grant & Cunjak (2005; Journal of Animal Ecology, 74, 508-516) argue that density-dependent growth due to exploitative competition for prey causes the negative density-growth relationships for stream salmonids. They argue that the concave shape of empirical density-growth relationships is consistent with a simple model of exploitative competition and not consistent with interference competition for space. 3. We use a simple model to show that competition for space can yield concave density-growth relationships consistent with the empirical pattern when individuals compete for foraging sites that vary spatially in quality and lower-quality sites predominate. Thus, the predictions of the exploitative competition and spatial competition models overlap. 4. The shape of the density-growth relationship does not differentiate between candidate mechanisms underlying density-dependent growth for stream salmonids. Our results highlight the general problem with determining the mechanism driving an ecological process from patterns in observational data within the context of linking population demographics to habitat structure and animal behaviour.     

Changes over time in the spatiotemporal dynamics of cyclic populations of field voles (Microtus agrestis L.)

We demonstrate changes over time in the spatial and temporal dynamics of an herbivorous small rodent by analyzing time series of population densities obtained at 21 locations on clear cuts within a coniferous forest in Britain from 1984 to 2004. Changes had taken place in the amplitude, periodicity, and synchrony of cycles and density-dependent feedback on population growth rates. Evidence for the presence of a unidirectional traveling wave in rodent abundance was strong near the beginning of the study but had disappeared near the end. This study provides empirical support for the hypothesis that the temporal (such as delayed density dependence structure) and spatial (such as traveling waves) dynamics of cyclic populations are closely linked. The changes in dynamics were markedly season specific, and changes in overwintering dynamics were most pronounced. Climatic changes, resulting in a less seasonal environment with shorter winters near the end of the study, are likely to have caused the changes in vole dynamics. Similar changes in rodent dynamics and the climate as reported from Fennoscandia indicate the involvement of large-scale climatic variables.     

Sea lice as a density-dependent constraint to salmonid farming

Fisheries catches worldwide have shown no increase over the last two decades, while aquaculture has been booming. To cover the demand for fish in the growing human population, continued high growth rates in aquaculture are needed. A potential constraint to such growth is infectious diseases, as disease transmission rates are expected to increase with increasing densities of farmed fish. Using an extensive dataset from all farms growing salmonids along the Norwegian coast, we document that densities of farmed salmonids surrounding individual farms have a strong effect on farm levels of parasitic sea lice and efforts to control sea lice infections. Furthermore, increased intervention efforts have been unsuccessful in controlling elevated infection levels in high salmonid density areas in 2009-2010. Our results emphasize host density effects of farmed salmonids on the population dynamics of sea lice and suggest that parasitic sea lice represent a potent negative feedback mechanism that may limit sustainable spatial densities of farmed salmonids.     

Density, aggregation, and body size of northern pikeminnow preying on juvenile salmonids in a large river

Predation by northern pikeminnow Ptychocheilus oregonensis on juvenile salmonids Oncorhynchus spp. occurred probably during brief feeding bouts since diets were either dominated by salmonids (>80% by weight), or contained other prey types and few salmonids (<5%). In samples where salmonids had been consumed, large rather than small predators were more likely to have captured salmonids. Transects with higher catch-per-unit of effort of predators also had higher incidences of salmonids in predator guts. Predators in two of three reservoir areas were distributed more contagiously if they had preyed recently on salmonids. Spatial and temporal patchiness of salmonid prey may be generating differences in local density, aggregation, and body size of their predators in this large river.     

The effect of food abundance on territory size and population density of juvenile steelhead trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>)

Optimal territory size models predict a decrease in territory size with increasing food abundance. However, most of these models may not be applicable to juvenile salmonids in streams, because they defend contiguous territories at high densities. The optimal size of a contiguous territory is predicted to (1) be independent of food abundance when food is rare and (2) decrease only when food abundance is high enough to induce a reduction in territory size below the contiguous optimum. To test these predictions, we raised equal densities of juvenile steelhead trout in outdoor stream channels over a 32-fold range of food abundance in the absence of emigration for 25 days. Increasing competition for scarce food resulted in increasing mortality, higher willingness to emigrate, higher variance in body mass, lower growth, lower population density and lower biomass. The size of territories decreased with increasing local population density, and increased with increasing body size. However, territory size did not change with food abundance, a result consistent with the prediction of a contiguous territory size model. On average, total salmonid biomass increased 5.7 times in response to the 32-fold increase in food abundance. Our data provide strong support for an earlier quantitative relationship between the abundance of stream salmonids and their food.     

Translocation of stream-dwelling salmonids in headwaters: insights from a 15-year reintroduction experience

Translocation programs are a common strategy to increase the number of viable populations of threatened freshwater fishes. Yet, only in a minority of cases the success or failure of translocations has been assessed through a quantitative analysis of demographic traits, compensatory responses, life-histories and population dynamics of the threatened species. A paradigmatic case a translocation program combining both management- and research-oriented activities is represented by the Marble Trout Conservation Program, which started in 1993 in the upper reaches of the Soca, Idirjca and Baca river basins (Slovenia) for the conservation of stream-dwelling marble trout Salmo marmoratus. In order to enhance the viability of the species, two new populations were created in 1996 by stocking 500 marble trout aged 1+ in previously fishless streams (Gorska and Zakojska) within the core habitat of the species. The new populations have been systematically monitored for 15 years by individually tagging and sampling marble trout. Our analyses show that deterministic extinction of marble trout populations are unlikely and that high-magnitude environmental stochasticity (i.e., severe floods) is the only main cause of local population extinction, despite the high resilience to flood-induced massive mortalities exhibited by marble trout through compensatory mechanisms (e.g., relaxation of density-dependent body growth and survival at low densities). Fishless headwaters, probably characterized by a history of recurrent severe floods, should not be considered as candidate sites for the creation of new populations. Fewer individuals than originally reintroduced (i.e., 500 fish aged 1+ in each stream) might be sufficient to establish viable populations, since compensatory mechanisms are likely to regulate population size around stream carrying capacity in a few years. Besides enhancing the species viability, translocation programs can provide an excellent framework for the estimation of ecological traits (e.g., life-histories, demography, population dynamics etc.), identify potential vulnerabilities and thus guide well-formed management actions for the threatened species.     

Prey size,prey abundance,and temperature as correlates of growth in stream populations of cutthroat trout

Growth and maximum size of stream fishes can be highly variable across populations. For salmonid fishes in streams, individuals from populations confined to headwater streams often exhibit small size at maturity in comparison to populations with access to main-stem rivers. Differences in prey size, prey availability, and metabolic constraints based on temperature may explain patterns of maximum size and growth. In this study, cutthroat trout from headwater stream populations that were isolated above a waterfall were compared to individuals from populations in similar sized streams without a movement barrier and from large main-stem rivers. Cutthroat trout from smaller streams with or without a movement barrier were significantly smaller at a given age than fish from main-stem rivers, where individuals were able to achieve a much larger maximum size. Comparisons of invertebrate drift abundance and size in the three types of streams revealed that drift size did not differ between stream categories, but was highest per volume of water in large main-stem rivers. Across all stream types, prey abundance declined from summer to fall. Temperature declined over the course of the season in a similar manner across all stream types, but remained relatively high later in the season in main-stem river habitats. Prey availability and temperature conditions in main-stem rivers may provide more optimal growing conditions for fish as individuals increase in size and become constrained by prey availability and temperature conditions in small streams. Maintaining connectivity between small spawning and rearing tributary streams and main-stem river habitats may be critical in maintaining large-bodied populations of stream salmonids.     

Distribution and growth of brown trout in pristine headwaters of Central Europe

The majority of stream-dwelling salmonid populations in Europe are affected by artificial stocking and the fragmentation of riverine ecosystems. The present study was performed in the unique pristine headwaters of the Otava River in the Elbe catchment area of the Czech Republic. The aim was to investigate the spatial distribution and individual growth pattern of brown trout, Salmo trutta, populations. Twenty sites in two main streams and their tributaries were sampled twice a year (spring and autumn) during the period 2005–2011. The sampling sites were grouped into fourteen so-called synchronised population units within the boundaries of three populations, according to analyses of synchrony in population abundance. The individual growth of juveniles (age-0, age-1) varied between all three spatial units (sampling sites, synchronised population units and populations), while the individual growth of adults (age-2 and older) did not. The distinctiveness regarding individual growth and demographic independence among the synchronised population units and populations indicates their suitability for use as population units for management purposes.     

Density dependence and the control of helminth parasites

1. The transient dynamics and stability of a population are determined by the interplay between species density, its spatial distribution and the positive and negative density-dependent processes regulating population growth. 2. Using the human-helminth parasite system as an example, we propose that the life-stage upon which negative density dependence operates will influence the rate of host reinfection following anthelmintic chemotherapy, and the likely success of control programmes. 3. Simple deterministic models are developed which highlight how a parasite species whose population size is down-regulated by density-dependent establishment will reinfect a host population at a faster rate than a species with density-dependent parasite fecundity. 4. Different forms of density dependence can produce the same equilibrium behaviour but different transient dynamics. Under-representing the nature and magnitude of density-dependent mechanisms, and in particular those operating upon establishing life-stages, may cause the resilience of the parasite population to a control perturbation to be underestimated.     

Shrinking body size and climate warming: Many freshwater salmonids do not follow the rule

Declining body size is believed to be a universal response to climate warming and has been documented in numerous studies of marine and anadromous fishes. The Salmonidae are a family of coldwater fishes considered to be among the most sensitive species to climate warming; however, whether the shrinking body size response holds true for freshwater salmonids has yet to be examined at a broad spatial scale. We compiled observations of individual fish lengths from long-term surveys across the Northern Hemisphere for 12 species of freshwater salmonids and used linear mixed models to test for spatial and temporal trends in body size (fish length) spanning recent decades. Contrary to expectations, we found a significant increase in length overall but with high variability in trends among populations and species. More than two-thirds of the populations we examined increased in length over time. Secondary regressions revealed larger-bodied populations are experiencing greater increases in length than smaller-bodied populations. Mean water temperature was weakly predictive of changes in body length but overall minimal influences of environmental variables suggest that it is difficult to predict an organism's response to changing temperatures by solely looking at climatic factors. Our results suggest that declining body size is not universal, and the response of fishes to climate change may be largely influenced by local factors. It is important to know that we cannot assume the effects of climate change are predictable and negative at a large spatial scale.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Aquaculture, non-native salmonid invasions and associated declines of native fishes in Northern Patagonian lakes

1. Even though intensive aquaculture production of salmonids in lakes occurs in many locations around the world published studies on the survival and reproductive success of escaped cultured salmonids in freshwater ecosystems are not common. A recent expansion of aquaculture in Chile has led it to become the world's second largest producer of cultured salmonids.
2. We document the recent history of escaped and self-sustaining salmonid populations over a wide spatial scale and a long temporal scale in Chilean Patagonian lakes. Our hypotheses are that salmonid density in lakes will be higher where there is intensive aquaculture, due to greater numbers of potential escapees. Secondly, if non-native salmonids have adverse impacts on native fishes, increases in the abundance of non-native species should be associated with decreases in relative abundance of native species. Finally, if the first two hypotheses are correct we anticipate that diets of salmonids may show evidence of predation on native fishes, diet overlap with native species, and evidence of the influence of feed from aquaculture operations in the diets of salmonids and native fishes.
3. We sampled six lakes with gill nets from 1992 to 2001. Our results show that the relative abundance of free-living salmonids is closely related to the level of fish farming production. Salmonids are the top predators and in lakes with fish farming the main prey item is native fishes. The relative abundance of native fishes has decreased, most likely due to predation by salmonids.
4. Our study contributes to the understanding of the effects of non-native salmonids in oligotrophic lakes, and it provides a starting point to judge the establishment of new fish farming sites in lakes around the world.     

Linking individual phenotype to density-dependent population growth: the influence of body size on the population dynamics of malaria vectors

Understanding the endogenous factors that drive the population dynamics of malaria mosquitoes will facilitate more accurate predictions about vector control effectiveness and our ability to destabilize the growth of either low- or high-density insect populations. We assessed whether variation in phenotypic traits predict the dynamics of Anopheles gambiae sensu lato mosquitoes, the most important vectors of human malaria. Anopheles gambiae dynamics were monitored over a six-month period of seasonal growth and decline. The population exhibited density-dependent feedback, with the carrying capacity being modified by rainfall (97% wAIC(c) support). The individual phenotypic expression of the maternal (p = 0.0001) and current (p = 0.040) body size positively influenced population growth. Our field-based evidence uniquely demonstrates that individual fitness can have population-level impacts and, furthermore, can mitigate the impact of exogenous drivers (e.g. rainfall) in species whose reproduction depends upon it. Once frontline interventions have suppressed mosquito densities, attempts to eliminate malaria with supplementary vector control tools may be attenuated by increased population growth and individual fitness.