Comparative studies, across and within taxa, have made important contributions to our understanding of the evolutionary processes that promote phenotypic diversity. Trait variation along geographic gradients provides a convenient heuristic for understanding what drives and maintains diversity. Intraspecific trait variation along latitudinal gradients is well‐known, but elevational variation in the same traits is rarely documented. Trait variation along continuous elevational gradients, however, provides compelling evidence that individuals within a breeding population may experience different selective pressures. Our objectives were to quantify variation in a suite of traits along a continuous elevational gradient, evaluate whether individuals in the population experience different selective pressures along that gradient and quantify variation in migratory tendency along that gradient. We examined variation in a suite of 14 life‐history, morphological and behavioural traits, including migratory tendency, of yellow‐eyed juncos along a continuous 1000‐m elevational gradient in the Santa Catalina Mountains of Arizona. Many traits we examined varied along the elevational gradient. Nest survival and nestling growth rates increased, while breeding season length, renesting propensity and adult survival declined, with increasing elevation. We documented the migratory phenotype of juncos (partial altitudinal migrants) and show that individual migratory tendency is higher among females than males and increases with breeding elevation. Our data support the paradigm that variation in breeding season length is a major selective pressure driving life‐history variation along elevational gradients and that individuals breeding at high elevation pursue strategies that favour offspring quality over offspring quantity. Furthermore, a negative association between adult survival and breeding elevation and a positive association between nest survival and breeding elevation help explain both the downslope and reciprocal upslope seasonal migratory movements that characterize altitudinal migration in many birds. Our results demonstrate how detailed studies of intraspecific variation in suites of traits along environmental gradients can lend new insights into the evolutionary processes that promote diversification and speciation, the causes of migratory behaviour, and how animal populations will likely respond to climate change.
The concept of a pace‐of‐life syndrome describes inter‐ and intraspecific variation in several life‐history traits along a slow‐to‐fast pace‐of‐life continuum, with long lifespans, low reproductive and metabolic rates, and elevated somatic defences at the slow end of the continuum and the opposite traits at the fast end. Pace‐of‐life can vary in relation to local environmental conditions (e.g. latitude, altitude), and here we propose that this variation may also occur along an anthropogenically modified environmental gradient. Based on a body of literature supporting the idea that city birds have longer lifespans, we predict that urban birds have a slower pace‐of‐life compared to rural birds and thus invest more in self maintenance and less in annual reproduction. Our statistical meta‐analysis of two key traits related to pace‐of‐life, survival and breeding investment (clutch size), indicated that urban birds generally have higher survival, but smaller clutch sizes. The latter finding (smaller clutches in urban habitats) seemed to be mainly a characteristic of smaller passerines. We also reviewed urbanization studies on other traits that can be associated with pace‐of‐life and are related to either reproductive investment or self‐maintenance. Though sample sizes were generally too small to conduct formal meta‐analyses, published literature suggests that urban birds tend to produce lower‐quality sexual signals and invest more in offspring care. The latter finding is in agreement with the adult survival hypothesis, proposing that higher adult survival prospects favour investment in fewer offspring per year. According to our hypothesis, differences in age structure should arise between urban and rural populations, providing a novel alternative explanation for physiological differences and earlier breeding. We encourage more research investigating how telomere dynamics, immune defences, antioxidants and oxidative damage in different tissues vary along the urbanization gradient, and suggest that applying pace‐of‐life framework to studies of variation in physiological traits along the urbanization gradient might be the next direction to improve our understanding of urbanization as an evolutionary process.
more » « less- Award ID(s):
- 1637590
- NSF-PAR ID:
- 10047066
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Global Change Biology
- Volume:
- 24
- Issue:
- 4
- ISSN:
- 1354-1013
- Page Range / eLocation ID:
- p. 1452-1469
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
more » « less
Abstract -
more » « less
Synopsis The allocation of limited resources among life history traits creates trade-offs that constrain the range of possible phenotypes of organisms. In animals, the cost of maintaining an effective immune response may reduce the ability to invest in reproduction, resulting in altered susceptibility to disease. However, not all members of a population face identical constraints because differences in an individual's environmental context or physiological state can influence the degree to which traits are negatively associated. Here, we evaluated how variation in timing of breeding, a correlate of fitness, may result in different patterns of trait associations between immunity and reproduction. We measured constitutive immunity in breeding female tree swallows (Tachycineta bicolor) using a bacteria killing assay with blood plasma to assess the relationships between bacteria killing ability (BKA), reproductive effort, and reproductive success. We found that timing of breeding can influence the association between BKA and reproductive effort, but its effects are not homogeneous among all traits. Late-breeding tree swallows with stronger BKA laid smaller clutches, a pattern that was not apparent in early breeders. Regardless of the timing of breeding, birds with stronger BKA fed their nestlings less. Despite a negative association with reproductive effort, we found no association between immunity and reproductive success. We provide evidence that individual tree swallows do not experience some trade-offs equally, and that timing of breeding likely plays a role in how costs of immunity are weighed. To understand how investment in immunity can limit life history traits, we must consider how a variation among individuals influences the relative costs of immunity.
-
more » « less
Abstract Predator fear effects influence reproductive outcomes in many species. In non‐urban systems, passerines often respond to predator cues by reducing parental investment, resulting in smaller and lighter nestlings. Since trophic interactions in urban areas are highly altered, it is unclear how passerines respond to fear effects in human‐altered landscapes. Nestlings of passerines in urban areas also tend to be smaller and lighter than their rural counterparts and are often exposed to high densities of potential predators yet experience lower per capita predation—the predation paradox. We suggest fear effects in urban habitats could be a significant mechanism influencing nestling condition in birds, despite lowered predation rates. We manipulated exposure of nesting birds to adult‐consuming predator risk in residential yards across a gradient of urbanization to determine the relative influence of urbanization and fear on nestling condition. We found nestlings had reduced mass in nests exposed to predator playbacks as well as in more urban areas. Despite lower per capita predation rates in urban areas, fear effects from increased predator densities may influence passerine fitness through reduced nestling condition. As urban development expands, biodiversity conservation hinges on a deeper mechanistic understanding of how urbanization affects reproductive outcomes.
-
more » « less
Abstract Heterogeneity in the intrinsic quality and nutritional condition of individuals affects reproductive success and consequently fitness. Black brant (
Branta bernicla nigricans ) are long‐lived, migratory, specialist herbivores. Long migratory pathways and short summer breeding seasons constrain the time and energy available for reproduction, thus magnifying life‐history trade‐offs. These constraints, combined with long lifespans and trade‐offs between current and future reproductive value, provide a model system to examine the role of individual heterogeneity in driving life‐history strategies and individual heterogeneity in fitness. We used hierarchical Bayesian models to examine reproductive trade‐offs, modeling the relationships between within‐year measures of reproductive energy allocation and among‐year demographic rates of individual females breeding on the Yukon‐Kuskokwim Delta, Alaska, using capture–recapture and reproductive data from 1988 to 2014. We generally found that annual survival tended to be buffered against variation in reproductive investment, while breeding probability varied considerably over the range of clutch size‐laying date combinations. We provide evidence for relationships between breeding probability and clutch size, breeding probability and nest initiation date, and an interaction between clutch size and initiation date. Average lifetime clutch size also had a weak positive relationship with apparent survival probability. Our results support the use of demographic buffering strategies for black brant. These results also indirectly suggest associations among environmental conditions during growth, fitness, and energy allocation, highlighting the effects of early growth conditions on individual heterogeneity, and subsequently, lifetime reproductive investment. -
more » « less
Abstract Maintenance of phenotypic heterogeneity in the face of strong selection is an important component of evolutionary ecology, as are the consequences of such heterogeneity. Organisms may experience diminishing returns of increased reproductive allocation as clutch or litter size increases, affecting current and residual reproductive success. Given existing uncertainty regarding trade‐offs between the quantity and quality of offspring, we sought to examine the potential for diminishing returns on increased reproductive allocation in a long‐lived species of goose, with a particular emphasis on the effect of position in the laying sequence on offspring quality.
To better understand the effects of maternal allocation on offspring survival and growth, we estimated the effects of egg size, timing of breeding, inter‐ and intra‐annual variation, and position in the laying sequence on gosling survival and growth rates of black brant
Branta bernicla nigricans breeding in western Alaska from 1987 to 2007.We found that gosling growth rates and survival decreased with position in the laying sequence, regardless of clutch size. Mean egg volume of the clutch a gosling originated from had a positive effect on gosling survival (
β = 0.095, 95% CRI: 0.024, 0.165) and gosling growth rates (β = 0.626, 95% CRI: 0.469, 0.738). Gosling survival (β = −0.146, 95% CRI: −0.214, −0.079) and growth rates (β = −1.286, 95% CRI: −1.435, −1.132) were negatively related to hatching date.These findings indicate substantial heterogeneity in offspring quality associated with their position in the laying sequence. They also potentially suggest a trade‐off mechanism for females whose total reproductive investment is governed by pre‐breeding state.
