D. Ryan Norris

Carry‐over effects as drivers of fitness differences in animals

1. Carry-over effects occur when processes in one season influence the success of an individual in the following season. This phenomenon has the potential to explain a large amount of variation in individual fitness, but so far has only been described in a limited number of species. This is largely due to difficulties associated with tracking individuals between periods of the annual cycle, but also because of a lack of research specifically designed to examine hypotheses related to carry-over effects. 2. We review the known mechanisms that drive carry-over effects, most notably macronutrient supply, and highlight the types of life histories and ecological situations where we would expect them to most often occur. We also identify a number of other potential mechanisms that require investigation, including micronutrients such as antioxidants. 3. We propose a series of experiments designed to estimate the relative contributions of extrinsic and intrinsic quality effects in the pre-breeding season, which in turn will allow an accurate estimation of the magnitude of carry-over effects. To date this has proven immensely difficult, and we hope that the experimental frameworks described here will stimulate new avenues of research vital to advancing our understanding of how carry-over effects can shape animal life histories. 4. We also explore the potential of state-dependent modelling as a tool for investigating carry-over effects, most notably for its ability to calculate optimal rates of acquisition of a multitude of resources over the course of the annual cycle, and also because it allows us to vary the strength of density-dependent relationships which can alter the magnitude of carry-over effects in either a synergistic or agonistic fashion. 5. In conclusion carry-over effects are likely to be far more widespread than currently indicated, and they are likely to be driven by a multitude of factors including both macro- and micronutrients. For this reason they could feasibly be responsible for a large amount of the observed variation in performance among individuals, and consequently warrant a wealth of new research designed specifically to decompose components of variation in fitness attributes related to processes across and within seasons.

Seasonal interactions, habitat quality, and population dynamics in migratory birds

Abstract Abstract. Historically, studies of habitat selection have focused on quantifying how current patterns of habitat occupancy influence condition and survival within a season. This approach, however, is overly simplistic, especially for migratory birds that spend different periods of the year in geographically distinct places. Habitat occupancy and the resulting condition of individual birds is likely to be affected by events in the previous season, and the consequences of habitat occupancy will influence individuals and populations in subsequent seasons. Thus, for migratory birds, variation in habitat quality (and quantity) needs to be understood in the context of how events interact throughout periods of the annual cycle. Seasonal interactions can occur at the individual level or population level. Individual-level interactions occur when events in one season produce nonlethal, residual effects that carry over to influence individuals the following season. Population-level interactions occur when a change in population size in one season influences per capita rates the following season. We review various methods for estimating seasonal interactions and highlight a number of examples in the literature. Using a variety of techniques, including intrinsic and extrinsic markers, the vast majority of studies to date have measured seasonal interactions at the individual level. Obtaining estimates of density and changes in per capita rates across multiple seasons to determine population-level interactions has been more challenging. Both types of seasonal interactions can influence population dynamics, but predicting their effects requires detailed knowledge of how populations are geographically connected (i.e., migratory connectivity). We recommend that researchers studying habitat occupancy and habitat selection consider how events in previous seasons influence events within a season.

Cross-hemisphere migration of a 25 g songbird

The northern wheatear (Oenanthe oenanthe) is a small (approx. 25 g), insectivorous migrant with one of the largest ranges of any songbird in the world, breeding from the eastern Canadian Arctic across Greenland, Eurasia and into Alaska (AK). However, there is no evidence that breeding populations in the New World have established overwintering sites in the Western Hemisphere. Using light-level geolocators, we demonstrate that individuals from these New World regions overwinter in northern sub-Sahara Africa, with Alaskan birds travelling approximately 14 500 km each way and an eastern Canadian Arctic bird crossing a wide stretch of the North Atlantic (approx. 3500 km). These remarkable journeys, particularly for a bird of this size, last between one to three months depending on breeding location and season (autumn/spring) and result in mean overall migration speeds of up to 290 km d−1. Stable-hydrogen isotope analysis of winter-grown feathers sampled from breeding birds generally support the notion that Alaskan birds overwinter primarily in eastern Africa and eastern Canadian Arctic birds overwinter mainly in western Africa. Our results provide the first evidence of a migratory songbird capable of linking African ecosystems of the Old World with Arctic regions of the New World.

Carry‐over effects in a Pacific seabird: stable isotope evidence that pre‐breeding diet quality influences reproductive success

1. Understanding the interactions between different periods of the annual cycle in migratory animals has been constrained by our inability to track individuals across seasons. In seabirds, virtually nothing is known about how diet quality during the non-breeding period, away from the breeding grounds, might influence subsequent reproductive success. 2. We used stable nitrogen (delta(15)N) and carbon (delta(13)C) isotopes to evaluate the effects of non-breeding diet quality on the timing of breeding and egg size in a population of Cassins auklets (Ptychoramphus aleuticus) breeding on Triangle Island, British Columbia. Adult feathers are grown during two different periods of the annual cycle, which allowed us to estimate diet quality from the previous fall (October-November) and pre-breeding (February-March) period. 3. We found that the estimated proportion of energetically superior copepods (Neocalanus spp.) in the pre-breeding diet tended to be higher in females that bred earlier and laid larger eggs, whereas energetically poor juvenile rockfish (Sebastes spp.) were dominant in the pre-breeding diets of females that bred later and laid smaller eggs. We detected no effect of fall diet quality on breeding date or egg size, and no effect of pre-breeding diet quality on breeding date in males. 4. Pre-breeding diet quality was not related to body condition measured 1-2 days after laying, which suggests that females may need to attain a threshold condition before they initiate breeding and successfully rear young. 5. Our results suggest that changes in climatic conditions during the pre-breeding period may have severe consequences for reproductive success by influencing breeding date and egg size. Our work emphasizes the importance of determining how events are linked throughout the annual cycle for understanding the fitness and population dynamics of migratory animals.

IMPROVED ESTIMATES OF CERTAINTY IN STABLE-ISOTOPE-BASED METHODS FOR TRACKING MIGRATORY ANIMALS

The use of stable-hydrogen isotopes (deltaD) has become a common tool for estimating geographic patterns of movement in migratory animals. This method relies on broad and relatively predictable geographic patterning in deltaD values of precipitation, but these patterns are not estimated without error. In addition, deltaD measurements are relatively imprecise, particularly for organic tissue. Most models for estimating geographic locations have ignored these sources of error. Common modeling approaches include regression, range-matching, and likelihood-based assignment tests (including discriminant analysis). Here, we show the benefits of a simple stochastic extension to likelihood-based assignment tests that incorporates two estimable sources of error and describe the resulting influence on the certainty of assigning breeding origins for wintering American Redstarts (Setophaga ruticilla), a small Nearctic-Neotropical migratory bird. Through simulation, we incorporated both spatial interpolation error associated with models of deltaD in precipitation and analytical error associated with the measurement of deltaD in tissue samples. In general, assignments that did not include these sources of error fell within the ranges of the stochastic results, but the difference in proportion of birds assigned to any one breeding region varied by as much as 54%. To explore how the distribution of assignments generated from error models influenced the application of these results, we developed a simple model of winter habitat loss. We removed the proportion of Redstarts wintering at a particular site from the global population and then used the isotope-based assignments to predict the resulting population declines for each breeding region. This gave distributions of change in population sizes, some of which included no change or even a population increase. The sources of error we modeled may challenge the degree of certainty in the use of stable-isotope-based data on connectivity to predict population dynamics of migratory animals. We suggest that stronger inference will result from incorporating these sources of error into future studies that use deltaD or other stable isotopes to infer the geographic origin of individuals.

Population dynamics in migratory networks

Migratory animals are comprised of a complex series of interconnected breeding and nonbreeding populations. Because individuals in any given population can arrive from a variety of sites the previous season, predicting how different populations will respond to environmental change can be challenging. In this study, we develop a population model composed of a network of breeding and wintering sites to show how habitat loss affects patterns of connectivity and species abundance. When the costs of migration are evenly distributed, habitat loss at a single site can increase the degree of connectivity (mixing) within the entire network, which then acts to buffer global populations from declines. However, the degree to which populations are buffered depends on where habitat loss occurs within the network: a site that has the potential to receive individuals from multiple populations in the opposite season will lead to smaller declines than a site that is more isolated. In other cases when there are equal costs of migration to two or more sites in the opposite season, habitat loss can result in some populations becoming segregated (disconnected) from the rest of the network. The geographic structure of the network can have a significant influence on relative population sizes of sites in the same season and can also affect the overall degree of mixing in the network, even when sites are of equal intrinsic quality. When a migratory network is widely spaced and migration costs are high, an equivalent habitat loss will lead to a larger decline in global population size than will occur in a network where the overall costs of migration are low. Our model provides an important foundation to test predictions related to habitat loss in real-world migratory networks and demonstrates that migratory networks will likely produce different dynamics from traditional metapopulations. Our results provide strong evidence that estimating population connectivity is a prerequisite for successfully predicting changes in migratory populations.

Storms drive altitudinal migration in a tropical bird

Although migration is a widespread and taxonomically diverse behaviour, the ecological factors shaping migratory behaviour are poorly understood. Like other montane taxa, many birds migrate along elevational gradients in the tropics. Forty years ago, Alexander Skutch postulated that severe storms could drive birds to migrate downhill. Here, we articulate a novel mechanism that could link storms to mortality risks via reductions in foraging time and provide, to our knowledge, the first tests of this hypothesis in the White-ruffed Manakin (Corapipo altera), a small partially migratory frugivore breeding on the Atlantic slope of Costa Rica. As predicted, variation in rainfall was associated with plasma corticosterone levels, fat stores, plasma metabolites and haematocrit. By collecting data at high and low elevation sites simultaneously, we also found that high-elevation residents were more adversely affected by storms than low elevation migrants. These results, together with striking temporal capture patterns of altitudinal migrants relative to storms, provide, to our knowledge, the first evidence that weather-related risks incurred by species requiring high food intake rates can explain altitudinal migrations of tropical animals. These findings resolve conflicting evidence for and against food limitation being important in the evolution of this behaviour, and highlight how endogenous and exogenous processes influence life-history trade-offs made by individuals in the wild. Because seasonal storms are a defining characteristic of most tropical ecosystems and rainfall patterns will probably change in ensuing decades, these results have important implications for understanding the ecology, evolution and conservation of tropical animals.

Animal Migration: A Context for Using New Techniques and Approaches

Publisher Summary The movement of organisms in space and time defines their interaction with their environments and, therefore, comprises a fundamental aspect of their ecology and evolutionary history. The degree to which organisms move also characterizes the range of resources they encounter, the array of hazards they experience from predators to hurricanes, and the degree to which they interact with other life forms. For animals, movement is very much a question of geospatial scale. While some species occupy a landscape of a few square meters for their entire lives or wander nomadically, others travel across thousands of kilometers in regular movements that constitute some of the most spectacular natural phenomena on the planet. The lack of tools available to scientists to infer or determine large- and small-scale animal movements has been a challenge. This chapter explores the recent developments in stable isotope methods that can be used to contribute tremendously to the field of understanding animal movement in terrestrial ecosystems. It discusses the migratory populations, connectivity, and conservation, the scientific tools used to study migration, and the intrinsic markers of animals.

Predicting conditions for migration: effects of density dependence and habitat quality

Migration is widespread among animals, but the factors that influence the decision to migrate are poorly understood. Within a single species, populations may be completely migratory, completely sedentary or partially migratory. We use a population model to derive conditions for migration and demonstrate how migratory survival, habitat quality and density dependence on both the breeding and non-breeding grounds influence conditions for migration and the proportion of migrants within a population. Density dependence during the season in which migratory and sedentary individuals use separate sites is necessary for partial migration. High levels of density dependence at the non-shared sites widen the range of survival values within which we predict partial migration, whereas increasing the strength of density dependence at the shared sites narrows the range of survival values within which we predict partial migration. Our results have important implications for predicting how contemporary populations with variable migration strategies may respond to changes in the quality or quantity of habitat.

The relationship between personality and plasticity in tree swallow aggression and the consequences for reproductive success

Both personality and plasticity can influence fitness, but few studies have investigated these two sources of individual variation simultaneously for the same behaviour. The individual quality hypothesis proposes that individuals with high personality scores will also be better able to respond to the changes in the environment (i.e. have a high degree of plasticity). Alternatively, the ‘compensatory’ hypothesis proposes that personality and plasticity are negatively correlated, because only individuals with low personality scores need to be able to rapidly adjust to environmental conditions. To examine these two hypotheses, we investigated the overall level of aggressiveness (personality) in nest defence, the ability of individuals to adjust this behaviour to respond to changes in temperature (plasticity), and their consequences for reproductive success in male and female tree swallows, Tachycineta bicolor. Using linear mixed effect models with individual as a random effect, we found that consistent differences between individuals explained approximately 55% of variation in aggressiveness and that more aggressive individuals were better able to adjust to variation in temperature, providing support for the individual quality hypothesis. However, although more aggressive males tended to fledge a higher number of young, the degree of plasticity only conferred a reproductive advantage for nonaggressive males, providing support for the compensatory hypothesis. For females, neither personality, nor plasticity was a good predictor of reproductive success. Our results suggest that personality and plasticity in aggressiveness are important components of individual variation but that the fitness advantages of each are context dependent.