Cascading life-history interactions: alternative density-dependent pathways drive recruitment dynamics in a freshwater fish

Although density-dependent mechanisms in early life-history are important regulators of recruitment in many taxa, consequences of such mechanisms on other life-history stages are poorly understood. To examine interacting and cascading effects of mechanisms acting on different life-history stages, we stocked experimental ponds with fathead minnow (Pimephales promelas) at two different densities. We quantified growth and survival of the stocked fish, the eggs they produced, and the resulting offspring during their first season of life. Per-capita production and survival of eggs were inversely related to density of stocked fish; significant egg cannibalism by stocked minnows resulted in initial young-of-the-year (YOY) densities that were inversely related to adult densities. Subsequent growth and survival of YOY were then inversely related to these initial YOY densities, and survival of YOY was selective for larger fish. Because of these compensatory processes in the egg and YOY stages, treatments did not differ in YOY abundance and mean size at the end of the growing season. Because of differences in the intensity of size-selective mortality, however, variation in end-of season sizes of YOY was strongly (and inversely) related to densities of stocked fish. When mortality was severe in the egg stage (high densities of stocked fish), final YOY size distributions were more variable than when the dominant mortality was size-selective in the YOY stage (low stocked fish densities). These differences in size variation could have subsequent recruitment consequences, as overwinter survival is typically selective for YOY fish larger than a critical threshold size. Density-dependent effects on a given life stage are not independent, but will be influenced by earlier stages; alternative recruitment pathways can result when processes at earlier stages differ in magnitude or selectivity. Appreciation of these cascading effects should enhance our overall understanding of the dynamics of stage-structured populations.     

Seasonal temperature and precipitation regulate brook trout young‐of‐the‐year abundance and population dynamics

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The contributions of age and sex to variation in common tern population growth rate

1. The decomposition of population growth rate into contributions from different demographic rates has many applications, ranging from evolutionary biology to conservation and management. Demographic rates with low variance may be pivotal for population persistence, but variable rates can have a dramatic influence on population growth rate. 2. In this study, the mean and variance in population growth rate (lambda) is decomposed into contributions from different ages and demographic rates using prospective and retrospective matrix analyses for male and female components of an increasing common tern (Sterna hirundo) population. 3. Three main results emerged: (1) subadult return was highly influential in prospective and retrospective analyses; (2) different age-classes made different contributions to variation in lambda: older age classes consistently produced offspring whereas young adults performed well only in high quality years; and (3) demographic rate covariation explained a significant proportion of variation in both sexes. A large contribution to lambda did not imply a large contribution to its variation. 4. This decomposition strengthens the argument that the relationship between variation in demographic rates and variation in lambda is complex. Understanding this relationship and its consequences for population persistence and evolutionary change demands closer examination of the lives, and deaths, of the individuals within populations within species.     

Survival,site fidelity,and the population dynamics of Piping Plovers in Saskatchewan

ABSTRACT To conserve threatened species, managers require predictions about the effects of natural and anthropogenic factors on population growth that in turn require accurate estimates of survival, birth, and dispersal rates, and their correlation with natural and anthropogenic factors. For Piping Plovers (Charadrius melodus), fledging rate is often more amenable to management than adult survival, and population models can be used to estimate the productivity (young produced per breeding female) necessary to maintain or increase populations for given levels of survival. We estimated true survival and site fidelity of adult and subadult (from fledging to second year) Piping Plovers breeding in Saskatchewan using mark‐resight data from 2002 to 2009. By estimating true survival rather than apparent survival (which is confounded with permanent emigration), we were able to provide more accurate projections of population trends. Average adult and subadult survival rates during our study were 0.80 and 0.57, respectively. Adult survival declined over time, possibly due in part to the loss of one breeding site to flooding. Average adult and subadult site fidelity were 0.86 and 0.46, respectively. Adult site fidelity declined during our study at two study sites, most strongly at the flooded site. Male and female Piping Plovers had similar survival rates, but males had greater site fidelity than females in some years. Based on our survival estimates, productivity needed for a stationary population was 0.75, a benchmark used for plover management on the Atlantic Coast, but not previously estimated for Prairie Canada. In stochastic simulations incorporating literature‐based variation in survival rates, productivity needed for a stationary population increased to 0.86, still lower than that previously estimated for western populations. Mean productivity for our study sites ranged from 0.87 to 0.96 fledged young per pair. Our results suggest that fledging rates of Piping Plovers in Saskatchewan were sufficient to ensure a stationary or increasing population during our study period. However, large‐scale habitat changes such as drought or anthropogenic flooding may lead to dispersal of breeding adults and possibly mortality that will increase the fledging rate needed for a stationary population.     

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

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

Spatio-temporal abundance of flushing leaves shapes host selection in the willow leaf beetle, Plagiodera versicolora

In the willow leaf beetle, Plagiodera versicolora (Laicharting) (Coleoptera: Chrysomelidae), food resources available for adults are severely restricted by leaf toughness, which increases with age. Nevertheless, females require their own food almost all their life in order to produce eggs. In this paper, we have focused our attention on the spatio-temporal abundance of flushing leaves and have examined its effect on host-plant selection by adults among four co-occurring willow species ( Salix chaenomeloides , Salix eriocarpa , Salix integra , and Salix serissaefolia ) (Salicaceae) by field observations and experiments at two spatial scales. Among the various factors associated with this, the amount of new leaf production contributed maximally to variation in adult abundance. By conducting two experiments, we confirmed that the adults preferentially flew towards willow trees with abundant flushing leaves. Furthermore, we detected substantial seasonal changes in new leaf abundance and realized fecundity in the field, and a strong positive correlation was observed between them. Availability of adult food resources limited the reproductive performance of adults, particularly in mid-summer when only S. serissaefolia produced a few new leaves. These results supported the substantial effect of new leaf abundance on adult abundance in the field. Thus, we concluded that adult feeding is a critical factor that shapes the host-plant selection of P. versicolora and determines its seasonal occurrence through the dispersal and settlement of adults.     

The overwintering of Antarctic krill, Euphausia superba, from an ecophysiological perspective

A major aim of this review is to determine which physiological functions are adopted by adults and larvae to survive the winter season with low food supply and their relative importance. A second aim is to clarify the extent to which seasonal variation in larval and adult krill physiology is mediated by environmental factors with a strong seasonality, such as food supply or day light. Experimental studies on adult krill have demonstrated that specific physiological adaptations during autumn and winter, such as reduced metabolic rates and feeding activity, are not caused simply by the scarcity of food, as was previously assumed. These adaptations appear to be influenced by the local light regime. The physiological functions that larval krill adopt during winter (reduced metabolism, delayed development, lipid utilisation, and variable growth rates) are, in contrast to the adults, under direct control by the available food supply. During winter, the adults often seem to have little association with sea ice (at least until early spring). The larvae, however, feed within sea ice but mainly on the grazers of the ice algal community rather than on the algae themselves. In this respect, a miss-match in timing of the occurrence of the last phytoplankton blooms in autumn and the start of the sea ice formation, as has been increasingly observed in the west Antarctic Peninsula (WAP) region, will impact larval krill development during winter in terms of food supply and consequently the krill stock in this region.     

Demography of alpine red squirrel populations in relation to fluctuations in seed crop size

Vertebrate population dynamics, social organisation and space use often are closely associated with the distribution of critical resources, such as food. Tree squirrels are ideal models to study these relationships, since both key demographic parameters (reproduction, survival and dispersal) and spatio-temporal variation in food supplies (measured as seed-crop size) can be reliably estimated. In this paper we test the following two predictions underlying the association between annual food abundance and demography in six alpine red squirrel populations, both with and without time-lag effects: 1) between-season and between-year fluctuations in survival rate, population density and increase parallel those in food availability; and 2) individuals follow a resource tracking strategy and increase in density mainly the year after a rich seed-crop. Red squirrels occurred at higher densities in Scots pine forest, characterised by stable seed-crops, than in Norway spruce with more abundant but more variable seed crops. Fluctuations in numbers were positively correlated with food availability, measured as annual conifer seed-crop sizes. Overall, adult survival rates were higher than those of subadults, and survival substantially fluctuated between seasons and years. Autumn densities and rates of population increase (summer-autumn) were strongly correlated with the same year's autumn seed-crop, while correlations with the previous year's seed-crop (time-lag models) were either weak (population density) or absent (population increase). Results of this paper show that fluctuations in red squirrel densities in habitats with strong temporal variation in seed production are more closely linked with food availability than in more stable habitats. In addition, in the Alpine conifer forests squirrel population sizes, in autumn, increase in synchrony with food resources, eliminating the population lag normally present when resources are produced in pulses.     

Spatial patterns in age- and colony-specific survival in a long-lived seabird across 14 contrasting colonies

Demographic rates such as recruitment and survival probability can vary considerably among populations of the same species due to variation in underlying environmental processes. If environmental processes are spatially correlated, nearby populations are expected to have more similar demographic rates than those further apart. Breeding populations and foraging ranges are spatially segregated in colonial seabirds, making them ideal for studying spatial patterns in demographic rates and their effects on local population dynamics. Here we explored variation in age-dependent survival probabilities across 14 colonies of Herring Gulls Larus argentatus breeding along the Dutch North Sea coast. We used long-term mark–recapture data of marked fledglings to estimate survival, and estimated spatial autocorrelation of survival probabilities. We assessed whether survival until recruitment age or until 10 years old (close to their expected lifespan) explained variation in population trajectories of each colony. Juvenile and adult survival showed a strong, but different, north-to-south gradient in survival probability, with lower juvenile but higher adult survival in northern colonies than southern colonies, whereas the spatial pattern of immature survival was less distinct. Neither recruitment nor the proportion of 10-year-old adults alive predicted whether a colony collapsed, declined, remained stable or increased. The distinct spatial pattern in survival suggests variation in regional food availability, which do not seem to drive local population dynamics. The absence of a link between survival and colony trajectories implies that connectivity between populations plays an important role affecting population dynamics.     

Contrasting patterns of survival and dispersal in multiple habitats reveal an ecological trap in a food-caching bird

A comprehensive understanding of how natural and anthropogenic variation in habitat influences populations requires long-term information on how such variation affects survival and dispersal throughout the annual cycle. Gray jays Perisoreus canadensis are widespread boreal resident passerines that use cached food to survive over the winter and to begin breeding during the late winter. Using multistate capture-recapture analysis, we examined apparent survival and dispersal in relation to habitat quality in a gray jay population over 34 years (1977–2010). Prior evidence suggests that natural variation in habitat quality is driven by the proportion of conifers on territories because of their superior ability to preserve cached food. Although neither adults (>1 year) nor juveniles (<1 year) had higher survival rates on high-conifer territories, both age classes were less likely to leave high-conifer territories and, when they did move, were more likely to disperse to high-conifer territories. In contrast, survival rates were lower on territories that were adjacent to a major highway compared to territories that did not border the highway but there was no evidence for directional dispersal towards or away from highway territories. Our results support the notion that natural variation in habitat quality is driven by the proportion of coniferous trees on territories and provide the first evidence that high-mortality highway habitats can act as an equal-preference ecological trap for birds. Reproductive success, as shown in a previous study, but not survival, is sensitive to natural variation in habitat quality, suggesting that gray jays, despite living in harsh winter conditions, likely favor the allocation of limited resources towards self-maintenance over reproduction.     

Stochastic density effects on adult fish survival and implications for population fluctuations

The degree to which population fluctuations arise from variable adult survival relative to variable recruitment has been debated widely for marine organisms. Disentangling these effects remains challenging because data generally are not sufficient to evaluate if and how adult survival rates are regulated by stochasticity and/or population density. Using unique time series for a largely unexploited reef fish, we found both population density and stochastic food supply impacted adult survival. The estimated effect of variable survival on adult abundance (both mean and variability) rivalled that of variable recruitment. Moreover, we show density‐dependent adult survival can dampen impacts of stochastic recruitment. Thus, food variability may alter population fluctuations by simultaneously regulating recruitment and compensatory adult survival. These results provide an additional mechanism for why intensified density‐independent mortality (via harvest or other means) amplifies population fluctuations and emphasises need for research evaluating the causes and consequences of variability in adult survival.     

Facultative prioritization of wing growth in the Welcome Swallow Hirundo neoxena

Bird species adapted to variable environments tend to have slow lean tissue growth rates and high fat deposition rates, allowing survival during food shortages. This emphasis on fat deposition may be a fixed physiological trait. Alternatively, tissue allocation may be adjusted facultatively according to the proximate food supply. We consider two models of facultative adjustment that could account for the emphasis on fat deposition: (1) the fat-priority model, in which no lean growth occurs when food is scarce, and (2) the lean-priority model, in which a minimal level of lean growth always occurs but nutrients are otherwise allocated to fat deposition. We tested these two models using Welcome Swallows Hirundo neoxena , a species we show to have a variable food supply that is influenced by weather. We reduced food supply to chicks experimentally, by enlarging broods or excluding parents from chicks, and tested for reduction in wing growth (an indicator of lean growth) and mass growth (an indicator of fat deposition). Mass growth was retarded by both manipulations, but not wing growth, corroborating the lean-priority model. This growth strategy may function not to cope with violent variation in food supply, but to maintain development and symmetry of wings and feathers in the face of moderate variation in food supply. Our results contrast with those of a similar experiment on the Black Noddy Anous minutus , a species with more severe variation in food supply.     

Seasonal survival of young and adult black grouse females in boreal forests

Direct demographical information from different populations of species in concern may reveal processes to target by management. We estimated seasonal survival rates using data from 253 individual females in a continuous population of black grouse Tetrao tetrix in Central Finland in 2005–2007. The information theoretic model selection included the effects of age class, ecological season, year, and their relevant interactions. We showed a clear difference in the survival of young and older females during late winter, a decrease in female survival during spring, and variation between the study years. The most frequent proximate mortality cause was avian predation, and the avian predation rate was highest in spring.     

Effects of Wind-induced Spatial Variation in Water Temperature and Zooplankton Concentration on the Growth of Young-of-the-year Smallmouth Bass, Micropterus dolomieu

Synopsis Young-of-the-year (YOY) smallmouth bass, Micropterus dolomieu, spend their first summer in littoral areas near their nests. Evidence indicates that nests of smallmouth bass in Lake Opeongo, Ontario, Canada are more abundant in downwind locations than in upwind areas. We hypothesize that wind-induced lower water temperatures and food availability in upwind nesting areas lead to lower growth rates of YOY bass in upwind than in downwind nesting areas. We show that water temperatures were 0.6–1.3 °C higher in downwind than upwind littoral areas during the period from mid-June to mid-July, when the YOY bass were on or near their nests. Although quite variable, zooplankton concentrations were also higher at downwind sites. In addition, bioenergetic simulations based on time series of field-measured temperatures predicted higher growth rates of YOY bass in the downwind sites. Growth rates based on sequential sampling of bass fry from their nests did not, however, differ statistically between upwind and downwind sites, although fry consistently weighed more downwind than upwind in the basin with the longer fetch possibly due to earlier spring warming. Our hypothesis is thus only partially supported and we call for further research on effects of spatial gradients on smallmouth bass life history.     

Density‐dependent survival and recruitment in a long‐distance Palaearctic migrant,the Sand Martin Riparia riparia

Long‐term studies can provide powerful insights into the relative importance of different demographic and environmental factors determining avian population dynamics. Here we use 23 years of capture–mark–recapture data (1981–2003) to estimate recruitment and survival rates for a Sand Martin Riparia riparia population in Cheshire, NW England. Inter‐annual variation in recruitment and adult survival was positively related to rainfall in the sub‐Saharan wintering grounds, but unrelated to weather conditions on the breeding grounds. After allowing for the effects of African rainfall, both demographic rates were negatively density‐dependent: adult survival was related to the size of the western European Sand Martin population (probably reflecting competition for resources in the shared wintering grounds) while recruitment was related to the size of the local study population in Cheshire (potentially reflecting competition for nesting sites or food). Local population size was more sensitive to variation in adult survival than to variation in recruitment, and an increase in population size after 1995 was driven mainly by the impact of more favourable conditions in the African wintering grounds on survival rates of adults. Overwinter survival in this long‐distance Palaearctic migrant is determined partly by the amount of suitable wetland foraging habitat in the sub‐Saharan wintering grounds (which is limited by the extent of summer rainfall) and partly by the number of birds exploiting that habitat.     

Reproductive phenology of a food-hoarding mast-seed consumer: resource- and density-dependent benefits of early breeding in red squirrels

The production of offspring by vertebrates is often timed to coincide with the annual peak in resource availability. However, capital breeders can extend the energetic benefits of a resource pulse by storing food or fat, thus relaxing the need for synchrony between energy supply and demand. Food-hoarding red squirrels (Tamiasciurus hudsonicus) breeding in the boreal forest are reliant on cones from a masting conifer for their nutrition, yet lactation is typically completed before the annual crop of cones is available for consumption such that peaks in energy supply and demand are not synchronized. We investigated the phenological response of red squirrels to annual variation in environmental conditions over a 20-year span and examined how intra- and inter-annual variation in the timing of reproduction affected offspring recruitment. Reproductive phenology was strongly affected by past resource availability with offspring born earlier in years following large cone crops, presumably because this affected the amount of capital available for reproduction. Early breeders had higher offspring survival and were more likely to renest following early litter loss when population density was high, perhaps because late-born offspring are less competitive in obtaining a territory when vacancies are limited. Early breeders were also more likely to renest after successfully weaning their first litter, but renesting predominantly occurred during mast years. Because of their increased propensity to renest and the higher survival rates of their offspring, early breeders contribute more recruits to the population but the advantage of early breeding depends on population density and resource availability.     

Census data aggregation decisions can affect population‐level inference in heterogeneous populations

1. Conservation and population management decisions often rely on population models parameterized using census data. However, the sampling regime, precision, sample size, and methods used to collect census data are usually heterogeneous in time and space. Decisions about how to derive population‐wide estimates from this patchwork of data are complicated and may bias estimated population dynamics, with important implications for subsequent management decisions.
2. Here, we explore the impact of site selection and data aggregation decisions on pup survival estimates, and downstream estimates derived from parameterized matrix population models (MPMs), using a long‐term dataset on grey seal (Halichoerus grypus) pup survival from southwestern Wales. The spatiotemporal and methodological heterogeneity of the data are fairly typical for ecological census data and it is, therefore, a good model to address this topic.
3. Data were collected from 46 sampling locations (sites) over 25 years, and we explore the impact of data handling decisions by varying how years and sampling locations are combined to parameterize pup survival in population‐level MPMs. We focus on pup survival because abundant high‐quality data are available on this developmental stage.
4. We found that survival probability was highly variable with most variation being at the site level, and poorly correlated among sampling sites. This variation could generate marked differences in predicted population dynamics depending on sampling strategy. The sample size required for a confident survival estimate also varied markedly geographically.
5. We conclude that for populations with highly variable vital rates among sub‐populations, site selection and data aggregation methods are important. In particular, including peripheral or less frequently used areas can introduce substantial variation into population estimates. This is likely to be context‐dependent, but these choices, including the use of appropriate weights when summarizing across sampling areas, should be explored to ensure that management actions are successful.

     

The spatial scale of competition from recruits on an older cohort in Atlantic salmon

Competitive effects of younger cohorts on older ones are frequently assumed to be negligible in species where older, larger individuals dominate in pairwise behavioural interactions. Here, we provide field estimates of such competition by recruits on an older age class in Atlantic salmon (Salmo salar), a species where observational studies have documented strong body size advantages which should favour older individuals in direct interactions. By creating realistic levels of spatial variation in the density of underyearling (YOY) recruits over a 1-km stretch of a stream, and obtaining accurate measurements of individual growth rates of overyearlings (parr) from capture–mark–recapture data on a fine spatial scale, we demonstrate that high YOY density can substantially decrease parr growth. Models integrating multiple spatial scales indicated that parr were influenced by YOY density within 16 m. The preferred model suggested parr daily mass increase to be reduced by 39% when increasing YOY density from 0.0 to 1.0 m⁻², which is well within the range of naturally occurring densities. Reduced juvenile growth rates will in general be expected to reduce juvenile survival (via increased length of exposure to freshwater mortality) and increase generation times (via increased age at seaward migrations). Thus, increased recruitment can significantly affect the performance of older cohorts, with important implications for population dynamics. Our results highlight that, even for the wide range of organisms that rely on defendable resources, the direction of competition among age classes cannot be assumed a priori or be inferred from behavioural observations alone.     

Decrease in young-of-the-year yellow perch growth rates following <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> invasion

Bythotrephes longimanus, an invasive zooplankter from Eurasia, has caused severe declines in native zooplankton communities in Rainy and Kabetogama lakes in northern Minnesota. Both lakes have experienced a 40–60% decrease in peak summer zooplankton biomass following B. longimanus establishment around 2006–2007. In these lakes, yellow perch (Perca flavescens) are a key fishery species, and young-of-the-year (YOY) yellow perch are mainly planktivorous during their first summer. This led to concern that their growth could be detrimentally affected by the depletion of zooplankton forage. We used seining data to compare growth rates of YOY yellow perch before (2001–2005) and after (2008–2012) B. longimanus establishment in Rainy and Kabetogama lakes. Nearby Lake Vermilion, assumed to have been unaffected by B. longimanus during this time period, was used as a reference for natural variation in YOY growth in the region. YOY yellow perch length was modeled as a linear function of cumulative growing degree days (GDD) throughout the summer, and the slope of the relationship was compared between pre- and post-B. longimanus time periods for the three study lakes. The two lakes with B. longimanus showed similar decreases in YOY yellow perch growth rate relative to GDD, whereas Lake Vermilion showed no evidence of a decline in growth rates during this period. The reduction in growth rates resulted in an approximate 10% decrease in mean length of YOY yellow perch at the end of the summer after B. longimanus establishment, which could lead to further effects of this invasive zooplankter at higher trophic levels.     

Adaptive variation in offspring size in the terrestrial isopod,Armadillidium vulgare

Summary Variation in the birth size of offspring of the terrestrial isopod, Armadillidium vulgare, was observed in laboratory experiments and in field populations. In the laboratory, larger offspring were produced when the mother's food supply was reduced. In field populations, larger offspring were produced during the summer, a period of reduced food availability. Smaller offspring are produced in the spring, when food is readily available. Females may be making larger young to increase survival during the more severe conditions of the summer breeding period.