Little is known about the population status of many marsh-dependent birds in North America but recent efforts have focused on collecting more reliable information and estimates of population trends. ...As part of that effort, a standardized survey protocol was developed in 1999 that provided guidance for conducting marsh bird surveys throughout North America such that data would be consistent among locations. The original survey protocol has been revised to provide greater clarification on many issues as the number of individuals using the protocol has grown. The Standardized North American Marsh Bird Monitoring Protocol instructs surveyors to conduct an initial 5-minute passive point-count survey followed by a series of 1-minute segments during which marsh bird calls are broadcast into the marsh following a standardized approach. Surveyors are instructed to record each individual bird from the suite of 26 focal species that are present in their local area on separate lines of a datasheet and estimate the distance to each bird. Also, surveyors are required to record whether each individual bird was detected within each 1-minute subsegment of the survey. These data allow analysts to use several different approaches for estimating detection probability. The Standardized North American Marsh Bird Monitoring Protocol provides detailed instructions that explain the field methods used to monitor marsh birds in North America.
Natal dispersal is a key demographic trait that affects population dynamics, and intraspecific variation in dispersal affects gene flow among populations and source-sink dynamics. However, relatively ...little is known about the selective pressures and trade-offs that animals face when departing their natal area due to the logistical difficulties associated with monitoring animals during this critical life stage. We used a randomized block design to examine the selective pressure that influence dispersal timing in juvenile burrowing owls (Athene cunicularia) by experimentally altering both food and ectoparasites at 135 nests. We also examined the effects of local food abundance, ectoparasite loads, and parental departure on natal dispersal timing. Juvenile burrowing owls varied widely in natal dispersal timing, and phenotypic plasticity in dispersal timing was evident in juvenile owls' response to our experimental treatments, local conditions, and their parents' departure from the natal area. Moreover, juveniles responded differently than their parents to experimental manipulation of food and ectoparasite loads. Juveniles typically dispersed shortly after their parents departed the natal area, but delayed dispersing more than 2 weeks after parental departure if they did not receive experimental food supplements during a low-food year. In contrast, the experimental food supplements did not affect the migratory departure decisions of adult owls in either year. Juveniles at nests treated for ectoparasites initiated dispersal at a younger age (and prior to adults in the high-food year) compared to juveniles at control nests. In contrast, parents at nests treated for ectoparasites departed later than parents at control nests. Our results suggest that unfavorable conditions (low food or high ectoparasite loads) caused juveniles to delay dispersal, but prompted adults to depart sooner. Our results highlight the extent of intraspecific variation in natal dispersal timing, and demonstrate that ecological conditions affect dispersal decisions of parents and offspring differently, which can create important trade-offs that likely affect life history strategies and responses to climatic changes.
Avian reproductive strategies vary widely, and many studies of life-history variation have focused on the incubation and hatching stages of nesting. Birds make proximate decisions regarding ...reproductive investment during the laying stage, and these decisions likely constrain and tradeoff with other traits and subsequent behaviors. However, we know relatively little about egg-laying stage behaviors given the difficulty of locating and monitoring nest sites from the onset of laying. We used non-invasive continuous video recording to quantify variation in the egg-laying behaviors of burrowing owls (
Athene cunicularia)
along a 1400-km latitudinal gradient in western North America. Burrowing owls laid eggs disproportionately in the morning hours, and that tendency was strongest among first eggs in a clutch. However, selection appeared to act more strongly on laying intervals (the time between laying of consecutive eggs) than on the diel time of laying, and laying intervals varied widely among and within clutches. Laying intervals declined seasonally and with increasing clutch size but increased with increasing burrow temperature and as a function of laying stage nest attentiveness, which may be a strategy to preserve egg viability. Laying interval was positively correlated with the duration of hatching intervals, suggesting that laying interval duration is one mechanism (along with timing of incubation onset) that generates variation in hatching asynchrony. Our results lend support to two general hypotheses to explain laying schedules; selection favors laying eggs in the morning, but other selective pressures may override that pattern. These conclusions indicate that allocation decisions during laying are an important part of avian life-history strategies which are subject to energetic constraints and tradeoffs with other traits.
ABSTRACT
One enduring priority for ecologists has been to understand the cause(s) of variation in reproductive effort among species and localities. Avian clutch size generally increases with ...increasing latitude, both within and across species, but the mechanism(s) driving that pattern continue to generate hypotheses and debate. In 1961, a Ph.D. student at Oxford University, N. Philip Ashmole, proposed the influential hypothesis that clutch size varies in direct proportion to the seasonality of resources available to a population. Ashmole's hypothesis has been widely cited and discussed in the ecological literature. However, misinterpretation and confusion has been common regarding the mechanism that underlies Ashmole's hypothesis and the testable predictions it generates. We review the development of well‐known hypotheses to explain clutch size variation with an emphasis on Ashmole's hypothesis. We then discuss and clarify sources of confusion about Ashmole's hypothesis in the literature, summarise existing evidence in support and refutation of the hypothesis, and suggest some under‐utilised and novel approaches to test Ashmole's hypothesis and gain an improved understanding of the mechanisms responsible for variation in avian clutch size and fecundity, and life‐history evolution in general.
Variation in life-history strategies is central to our understanding of population dynamics and how organisms adapt to their environments. Yet we lack consensus regarding the ecological processes ...that drive variation in traits related to reproduction and survival. For example, we still do not understand the cause of two widespread inter- and intraspecific patterns: (1) the ubiquitous positive association between avian clutch size and latitude; and (2) variation in the extent of asynchronous hatching of eggs within a single clutch. Well-known hypotheses to explain each pattern have largely focused on biotic processes related to food availability and predation risk. However, local adaptation to maintain egg viability could explain both patterns with a single abiotic mechanism. The egg viability hypothesis was initially proposed to explain the cause of asynchronous hatching and suggests that asynchronous hatching results from early incubation onset in response to unfavorable nest microclimatic conditions, which otherwise reduce egg viability. However, allocation of resources to early incubation, prior to clutch completion, may energetically constrain clutch size and help explain the positive association between clutch size and latitude. We measured intraspecific variation in five functionally linked life-history traits of burrowing owls at five study sites spanning a 1,400-km latitudinal transect in western North America: clutch size, the timing of incubation onset, the degree of hatching asynchrony, the probability of hatching failure, and nestling survival. We found that most traits varied clinally with latitude, but all the traits were more strongly associated with individual nest microclimates than with latitude, and all varied with nest microclimate in the directions predicted by the egg viability hypothesis. Furthermore, incubation began earlier, hatching asynchrony increased, and clutch size declined across the breeding season. These results suggest that nest microclimate drives an important life-history trade-off and that thermal gradients are often sufficient to account for observed biogeographic and seasonal patterns in life-history strategies. Furthermore, our results reveal a potentially important indirect mechanism by which reproductive success and recruitment could be affected by climate change.
Aim
Predicting distributions is fundamental to ecology, yet hindered by spatially restricted sampling, scale‐dependent relationships and detection error associated with field surveys. Predictive ...species distribution models (SDMs) are nonetheless vital for conservation of many species. We developed a framework for building predictive SDMs with multi‐scale data and used it to develop range‐wide breeding‐season SDMs for 14 marsh bird species of concern.
Location
USA.
Methods
We built SDMs using data from range‐wide surveys conducted over 14 years, and habitat and disturbance covariates measured at multiple spatial scales. We built hierarchical occupancy models that included heterogeneity in detectability during sampling, and used Bayesian model selection to regulate model complexity (covariates and scales) based explicitly on spatial predictive abilities. We thus integrated model selection for optimizing out‐of‐sample prediction, range‐wide sampling over broad conditions, multi‐scale analyses and scale optimization, and species‐specific detectability for a suite of wide‐ranging species.
Results
Distributions of marsh birds were affected by local wetland conditions, but also by agricultural, urban and hydrologic disturbances operating from local scales (100–500 m) to the watershed level. Variables measuring human disturbances improved prediction for most species, and every species was affected by attributes at >1 scale. Five species showed evidence for continental‐scale range contraction during the study.
Main conclusions
We demonstrate how hierarchical occupancy models can be optimized for prediction across a species' range at the extent of a continent while also accounting for imperfect detection, and thus describe a generalizable approach that can be used for any species. We provide the first data‐driven, empirical SDMs built at the range‐wide extent for most of our 14 study species and demonstrate that previous studies focused on local distributions and the effects of fine‐scale wetland vegetation missed important broadscale drivers of occupancy for marsh birds.
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.
This study is the first to quantify variation in >10 life‐history traits along a single continuous elevational gradient. The data provide new insights into the ecological processes that promote diversification, the capacity for birds to respond to environmental change via phenotypic plasticity and the causes of altitudinal migration.
Setting land aside has long been a primary approach for protecting biodiversity; however, the efficacy of this approach has been questioned. We examined whether protecting lands positively influences ...bird species in the U.S., and thus overall biodiversity. We used the North American Breeding Bird Survey and Protected Areas Database of the U.S. to assess effects of protected and multiple-use lands on the prevalence and long-term population trends of imperiled and non-imperiled bird species. We evaluated whether both presence and proportional area of protected and multiple-use lands surrounding survey routes affected prevalence and population trends for imperiled and non-imperiled species. Regarding presence of these lands surrounding these survey routes, our results suggest that imperiled and non-imperiled species are using the combination of protected and multiple-use lands more than undesignated lands. We found no difference between protected and multiple-use lands. Mean population trends were negative for imperiled species in all land categories and did not differ between the land categories. Regarding proportion of protected lands surrounding the survey routes, we found that neither the prevalence nor population trends of imperiled or non-imperiled species was positively associated with any land category. We conclude that, although many species (in both groups) tend to be using these protected and multiple-use lands more frequently than undesignated lands, this protection does not appear to improve population trends. Our results may be influenced by external pressures (e.g., habitat fragmentation), the size of protected lands, the high mobility of birds that allows them to use a combination of all land categories, and management strategies that result in similar habitat between protected and multiple-use lands, or our approach to detect limited relationships. Overall, our results suggest that the combination of protected and multiple-use lands is insufficient, alone, to prevent declines in avian biodiversity at a national scale.
The risk that an animal’s offspring are eaten by predators is thought to strongly influence an animal’s decisions regarding reproductive effort. We found that birds breeding in locations with a high ...risk of nest predation laid fewer eggs than their conspecifics nesting in areas with a lower risk of nest predation. Montane birds nesting at higher elevations lay fewer eggs than at lower elevations because of the higher risk of nest predation at higher elevations.
Abstract
Questions about the ecological drivers of, and mechanistic constraints on, productivity have driven research on life-history evolution for decades. Resource availability and offspring mortality are considered among the 2 most important influences on the number of offspring per reproductive attempt. We used a factorial experimental design to manipulate food abundance and perceived offspring predation risk in a wild avian population (red-faced warblers; Cardellina rubrifrons) to identify the mechanistic cause of variation in avian clutch size. Additionally, we tested whether female quality helped explain the extant variation in clutch size. We found no support for the Food Limitation or Female Quality Hypotheses, but we did find support for both predictions of the Nest Predation Risk Hypothesis. Females that experienced an experimentally heightened perception of offspring predation risk responded by laying a smaller clutch than females in the control group. Additionally, predation rates at artificial nests were highest where red-faced warbler clutch size was smallest (at high elevations). Life-history theory predicts that an individual should invest less in reproduction when high nest predation risk reduces the likely benefit from that nesting attempt and, indeed, we found that birds exhibit phenotypic plasticity in clutch size by laying fewer eggs in response to increasing nest predation risk.
The strategies by which animals allocate reproductive effort across their lifetimes vary, and the causes of variation in those strategies are actively debated. In birds, most research has focused ...heavily on variation in clutch size and fecundity, but incubation behaviour and other functionally related traits have received less attention. Variation in incubation period duration is notable because time‐dependent sources of clutch mortality should impose strong directional selection to minimize the incubation period. However, life‐history theory predicts multiple mechanisms by which inter‐ and intraspecific variation in incubation behaviours may be adaptive.
We conducted one of the first studies of intraspecific latitudinal variation in avian incubation behaviours across a large portion of a single species’ range. We placed motion‐activated nest cameras inside burrowing owl nests at five study sites to quantify variation in daily nest attentiveness, cumulative nest attendance and incubation period duration. We tested predictions of two alterative hypotheses that have been proposed to explain variation in incubation periods: the parental risk tolerance hypothesis and the neonate quality hypothesis.
Daily nest attentiveness, cumulative nest attendance and incubation period duration in burrowing owls were all positively correlated with latitude. Burrowing owls reduced their daily nest attentiveness at low latitudes and on days when the average nest temperature was within the range that is optimal for embryo development. Further, longer incubation periods were most strongly associated with greater cumulative nest attendance instead of reduced daily nest attentiveness.
These results support predictions of the neonate quality hypothesis: longer incubation periods result from stronger selection on neonate quality rather than selection to reduce reproductive effort in response to low extrinsic mortality risk. However, some owls facultatively reduced their daily nest attentiveness, and this result supports the general hypothesis that incubation decisions reflect a trade‐off between reproduction and self‐maintenance, and that the optimal solution to that trade‐off varies systematically in response to latitudinal gradients in adult mortality.
A free Plain Language Summary can be found within the Supporting Information of this article.
A free Plain Language Summary can be found within the Supporting Information of this article.