Fluctuating optimum and temporally variable selection on breeding date in birds and mammals.

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection.

Hydrology influences breeding time in the white-throated dipper.

BACKGROUND: Earlier breeding is one of the strongest responses to global change in birds and is a key factor determining reproductive success. In most studies of climate effects, the focus has been on large-scale environmental indices or temperature averaged over large geographical areas, neglecting that animals are affected by the local conditions in their home ranges. In riverine ecosystems, climate change is altering the flow regime, in addition to changes resulting from the increasing demand for renewable and clean hydropower. Together with increasing temperatures, this can lead to shifts in the time window available for successful breeding of birds associated with the riverine habitat. Here, we investigated specifically how the environmental conditions at the territory level influence timing of breeding in a passerine bird with an aquatic lifestyle, the white-throated dipper Cinclus cinclus. We relate daily river discharge and other important hydrological parameters, to a long-term dataset of breeding phenology (1978-2015) in a natural river system. RESULTS: Dippers bred earlier when winter river discharge and groundwater levels in the weeks prior to breeding were high, and when there was little snow in the catchment area. Breeding was also earlier at lower altitudes, although the effect dramatically declined over the period. This suggests that territories at higher altitudes had more open water in winter later in the study period, which permitted early breeding also here. Unexpectedly, the largest effect inducing earlier breeding time was territory river discharge during the winter months and not immediately prior to breeding. The territory river discharge also increased during the study period. CONCLUSIONS: The observed earlier breeding can thus be interpreted as a response to climate change. Measuring environmental variation at the scale of the territory thus provides detailed information about the interactions between organisms and the abiotic environment.

Long-term annual and spatial variation of polygyny in the white-throated dipper (Cinclus cinclus).

Mating strategies are key components in the fitness of organisms, and notably in birds the occurrence of monogamy versus polygyny has attracted wide interest. We address this by a very comprehensive dataset (2899 breeding events spanning the years 1978-2019) of the white-throated dipper Cinclus cinclus. Though the mating system of this species has been regarded as generally monogamous, we find that 7% of all breeding events were performed by polygynous males (approximately 15% of all pairs). The fraction of polygyny has been stable over the entire study period irrespective of population size. The assumption that polygyny is most common at low population density was not supported. Surprisingly, there was no difference between polygynous and monogamous males with regard to the quality of the territories they inhabited, ranked according to their overall frequency of use. The most common age group, first-year breeders, dominated among monogamous males, while among polygynous males second-year breeders were most common, followed by third and first-year breeders. The primary females were in general older than females mated to monogamous males, also when controlled for their general frequency in the population. The majority of the two females mated to a polygynous male, bred in the vicinity of each other. The probability for a male to be involved in polygyny more than once, was significantly higher than by chance, suggesting phenotypic quality differences among males.

Climate effects on population fluctuations of the white-throated dipper Cinclus cinclus.

1. Climate change may have profound consequences for many organisms. We have studied fluctuations in a population of the white-throated dipper Cinclus cinclus during 31 years (1978-2008) in a river system in southern Norway in relation to both large-scale and local weather conditions occurring during the non-breeding season. 2. Multiple regression and partial least squares regression were used to model the growth rate of the population, accounting for population size in the previous year. 3. Population growth was influenced by North Atlantic Oscillation (NAO), mean winter temperature, precipitation and timing of ice formation on the main lake in the river system in autumn. These variables explained 84% of the variation in population growth over the 31-year study period. 4. Local winter conditions played a prominent role in explaining the population fluctuations, which is plausible because the dipper depends on open water for foraging. In the study area, winters can be harsh and rivers and lakes may freeze and severely affect the subsequent population size of the dipper in spring. 5. The breeding population of the dipper does not seem yet to have reached a level where all possible territories in the area have been occupied, even after mild winters, and the estimated carrying capacity is also decidedly lower (66 breeding pairs) than the number of available territories. If the trend of milder winters continues, the population might increase in the future. However, strong climate variation is expected to continue in the future, and hence periods of rapid growth of the dipper population will probably be followed by severe declines.

To make the most of what we have: extracting phenological data from nestling measurements.

To study the ecological and evolutionary effects of climate change on timing of annual events, scientists need access to data that have been collected over long time periods. High-quality long-term phenology data are rare and costly to obtain and there is therefore a need to extract this information from other available data sets. Many long-term studies on breeding birds include detailed information on individually marked parents and offspring, but do not include information on timing of breeding. Here, we demonstrate how a study of repeated standard measurements of white-throated dipper Cinclus cinclus nestlings in our study system in southernmost Norway can be used for modeling nestling growth, and how this statistical model can be used to estimate timing of breeding for birds with sparser data. We also evaluate how the accuracies of nestling growth models based on different morphological traits (mass and feather length) differ depending on the nestling age, present user guidelines and demonstrate how they can be applied to an independent data set. In conclusion, the approach presented is likely to be useful for a wide variety of species, even if the preferred measurement may differ between species.

Environmental drivers of varying selective optima in a small passerine: A multivariate, multiepisodic approach.

In changing environments, phenotypic traits are shaped by numerous agents of selection. The optimal phenotypic value maximizing the fitness of an individual thus varies through time and space with various environmental covariates. Selection may differ between different life-cycle stages and act on correlated traits inducing changes in the distribution of several traits simultaneously. Despite increasing interests in environmental sensitivity of phenotypic selection, estimating varying selective optima on various traits throughout the life cycle, while considering (a)biotic factors as potential selective agents has remained challenging. Here, we provide a statistical model to measure varying selective optima from longitudinal data. We apply our approach to analyze environmental sensitivity of phenotypic selection on egg-laying date and clutch size throughout the life cycle of a white-throated dipper population. We show the presence of a joint optimal phenotype that varies over the 35-year period, being dependent on altitude and temperature. We also find that optimal laying date is density-dependent, with high population density favoring earlier laying dates. By providing a flexible approach, widely applicable to free-ranging populations for which long-term data on individual phenotypes, fitness, and environmental factors are available, our study improves the understanding of phenotypic selection in varying environments.

The potential influence of Atlantic salmon Salmo salar and brown trout Salmo trutta on density and breeding of the white-throated dipper Cinclus cinclus.

Interactions between birds and fish are often overlooked in aquatic ecosystems. We studied the influence of Atlantic salmon and brown trout on the breeding population size and reproductive output of the white-throated dipper in a Norwegian river. Acidic precipitation led to the extinction of salmon, but salmon recolonized after liming was initiated in 1991. We compared the dipper population size and reproductive output before (1978-1992) and after (1993-2014) salmon recolonization. Despite a rapid and substantial increase in juvenile salmon, the breeding dipper population size and reproductive output were not influenced by juvenile salmon, trout, or total salmonid density. This might be due to different feeding strategies in salmonids and dippers, where salmonids are mainly feeding on drift, while the dipper is a benthic feeder. The correlation between the size of the dipper population upstream and downstream of a salmonid migratory barrier was similar before and after recolonization, indicating that the downstream territories were not less attractive after the recolonization of salmon. Upstream dipper breeding success rates declined before the recolonization event and increased after, indicating improved water quality due to liming, and increasing invertebrate prey abundances and biodiversity. Surprisingly, upstream the migratory barrier, juvenile trout had a weak positive effect on the dipper population size, indicating that dippers may prey upon small trout. It is possible that wider downstream reaches might have higher abundances of alternative food, rending juvenile trout unimportant as prey. Abiotic factors such as winter temperatures and acidic precipitation with subsequent liming, potentially mediated by prey abundance, seem to play the most important role in the life history of the dipper.

Interactions between demography and environmental effects are important determinants of population dynamics.

Climate change will affect the population dynamics of many species, yet the consequences for the long-term persistence of populations are poorly understood. A major reason for this is that density-dependent feedback effects caused by fluctuations in population size are considered independent of stochastic variation in the environment. We show that an interplay between winter temperature and population density can influence the persistence of a small passerine population under global warming. Although warmer winters favor an increased mean population size, density-dependent feedback can cause the local population to be less buffered against occasional poor environmental conditions (cold winters). This shows that it is essential to go beyond the population size and explore climate effects on the full dynamics to elaborate targeted management actions.

Demographic routes to variability and regulation in bird populations.

There is large interspecific variation in the magnitude of population fluctuations, even among closely related species. The factors generating this variation are not well understood, primarily because of the challenges of separating the relative impact of variation in population size from fluctuations in the environment. Here, we show using demographic data from 13 bird populations that magnitudes of fluctuations in population size are mainly driven by stochastic fluctuations in the environment. Regulation towards an equilibrium population size occurs through density-dependent mortality. At small population sizes, population dynamics are primarily driven by environment-driven variation in recruitment, whereas close to the carrying capacity K, variation in population growth is more strongly influenced by density-dependent mortality of both juveniles and adults. Our results provide evidence for the hypothesis proposed by Lack that population fluctuations in birds arise from temporal variation in the difference between density-independent recruitment and density-dependent mortality during the non-breeding season.

Climate warming decreases the survival of the little auk (Alle alle), a high Arctic avian predator.

Delayed maturity, low fecundity, and high adult survival are traits typical for species with a long-life expectancy. For such species, even a small change in adult survival can strongly affect the population dynamics and viability. We examined the effects of both regional and local climatic variability on adult survival of the little auk, a long-lived and numerous Arctic seabird species. We conducted a mark-resighting study for a period of 8 years (2006-2013) simultaneously at three little auk breeding sites that are influenced by the West Spitsbergen Current, which is the main carrier of warm, Atlantic water into the Arctic. We found that the survival of adult little auks was negatively correlated with both the North Atlantic Oscillation (NAO) index and local summer sea surface temperature (SST), with a time lag of 2 and 1 year, respectively. The effects of NAO and SST were likely mediated through a change in food quality and/or availability: (1) reproduction, growth, and development of Arctic Calanus copepods, the main prey of little auks, are negatively influenced by a reduction in sea ice, reduced ice algal production, and an earlier but shorter lasting spring bloom, all of which result from an increased NAO; (2) a high sea surface temperature shortens the reproductive period of Arctic Calanus, decreasing the number of eggs produced. A synchronous variation in survival rates at the different colonies indicates that climatic forcing was similar throughout the study area. Our findings suggest that a predicted warmer climate in the Arctic will negatively affect the population dynamics of the little auk, a high Arctic avian predator.