Using complementary approaches to estimate survival of juvenile and adult Eastern Kingbirds

ABSTRACT Survival rates of young birds during the period between nest departure and their first breeding season is an important but difficult statistic to measure because of low natal site fidelity, especially for long‐distance migrants. From 2002 to 2008, we conducted a capture‐mark‐resight study of Eastern Kingbirds (Tyrannus tyrannus), a Nearctic‐Neotropical migrant, at Malheur National Wildlife Refuge, Oregon, to estimate juvenile (S_J) and adult (S_A) survival. The return rate of juvenile kingbirds was high (0.224) and not significantly different from program MARK's estimate of S_J (0.291). On average, and for both sexes, program MARK's estimate of S_A for birds banded as nestlings (0.64) was similar to that for birds first banded as adults (0.65). Enumeration methods and MARK yielded similar estimates of S_A, especially for males banded as adults. We attribute the similarity of resighting rate of birds banded as nestlings to S_J and the similarity of S_A estimated using program MARK and by enumeration methods to the high site fidelity of most juvenile and adult kingbirds at our ecologically isolated study site. An independent estimate of S_J calculated using local estimates of population growth and average annual production of young per year suggested that true S_J and S_A were probably slightly higher than program MARK's estimates because of undocumented permanent emigration, especially of birds first banded as adults. Demographic balancing suggests that true S_A and S_J were roughly 0.70 and 0.30, respectively. In general, our multiple estimates of S_J were similar. However, program MARK estimates of S_J tended to be lower than those produced by demographic balancing. Because of the difficulty in differentiating permanent emigration and mortality, and the affect it has on empirical estimates of survival, we urge researchers to use multiple methods of survival estimation, when possible, to validate the precision of their estimates.     

Estimates of direct and indirect effects for early juvenile survival in captive populations maintained for conservation purposes: the case of Cuvier's gazelle

Together with the avoidance of any negative impact of inbreeding, preservation of genetic variability for life‐history traits that could undergo future selective pressure is a major issue in endangered species management programmes. However, most of these programmes ignore that, apart from the direct action of genes on such traits, parents, as contributors of offspring environment, can influence offspring performance through indirect parental effects (when parental genotype and phenotype exerts environmental influences on offspring phenotype independently of additive genetic effects). Using quantitative genetic models, we estimated the additive genetic variance for juvenile survival in a population of the endangered Cuvier's gazelle kept in captivity since 1975. The dataset analyzed included performance recording for 700 calves and a total pedigree of 740 individuals. Results indicated that in this population juvenile survival harbors significant additive genetic variance. The estimates of heritability obtained were in general moderate (0.115–0.457) and not affected by the inclusion of inbreeding in the models. Maternal genetic contribution to juvenile survival seems to be of major importance in this gazelle's population as well. Indirect genetic and indirect environmental effects assigned to mothers (i.e., maternal genetic and maternal permanent environmental effects) roughly explain a quarter of the total variance estimated for the trait analyzed. These findings have major evolutionary consequences for the species as show that offspring phenotypes can evolve strictly through changes in the environment provided by mothers. They are also relevant for the captive breeding programme of the species. To take into account, the contribution that mothers have on offspring phenotype through indirect genetic effects when designing pairing strategies might serve to identify those females with better ability to recruit, and, additionally, to predict reliable responses to selection in the captive population.     

A single climate driver has direct and indirect effects on insect population dynamics

Weather drives population dynamics directly, through effects on vital rates, or indirectly, through effects on the population's competitors, predators or prey and thence on vital rates. Indirect effects may include non-additive interactions with density dependence. Detection of climate drivers is critical to predicting climate change effects, but identification of potential drivers may depend on knowing the underlying mechanisms. For the butterfly Speyeria mormonia, one climate driver, snow melt date, has multiple effects on population growth. Snow melt date in year t has density-dependent indirect effects. Through frost effects, early snow melt decreases floral resources, thence per-capita nectar availability, which determines fecundity in the lab. Snow melt date in year t?+?1 has density-independent direct effects. These effects explain 84% of the variation in population growth rate. One climate parameter thus has multiple effects on the dynamics of a species with non-overlapping generations, with one effect not detectable without understanding the underlying mechanism.     

Survival of Afro‐Palaearctic passerine migrants in western Europe and the impacts of seasonal weather variables

Populations of migratory songbirds in western Europe show considerable variation in population trends between species and regions. The demographic and environmental causes of these large‐scale patterns are poorly understood. Using data from Constant Effort mist‐netting studies, we investigated relationships between changes in abundance, adult survival and seasonal weather conditions among 35 western European populations of eight species of migratory warblers (Sylviidae). We used cross‐species and within‐species comparisons to assess whether annual variation in survival was correlated with weather conditions during passage or winter. We estimated survival using CJS mark‐recapture models accounting for variation in the proportion of transient individuals and recapture rates. Species wintering in the humid bioclimatic zone of western Africa had significantly higher annual survival probabilities than species wintering in the arid bioclimatic zone of Africa (the Sahel). Rainfall in the Sahel was positively correlated with survival in at least some populations of five species. We found substantially fewer significant relationships with indices of weather during the autumn and spring passage periods, which may be due to the use of broad‐scale indices. Annual population changes were correlated with adult survival in all of our study species, although species undergoing widespread declines showed the weakest relationships.     

The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long-lived seabird

1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.     

Contrasting effects of climate on juvenile body size in a Southern Hemisphere passerine bird

Despite extensive research on the topic, it has been difficult to reach general conclusions as to the effects of climate change on morphology in wild animals: in particular, the effects of warming temperatures have been associated with increases, decreases or stasis in body size in different populations. Here, we use a fine‐scale analysis of associations between weather and offspring body size in a long‐term study of a wild passerine bird, the cooperatively breeding superb fairy‐wren, in south‐eastern Australia to show that such variation in the direction of associations occurs even within a population. Over the past 26 years, our study population has experienced increased temperatures, increased frequency of heatwaves and reduced rainfall – but the mean body mass of chicks has not changed. Despite the apparent stasis, mass was associated with weather across the previous year, but in multiple counteracting ways. Firstly, (i) chick mass was negatively associated with extremely recent heatwaves, but there also positive associations with (ii) higher maximum temperatures and (iii) higher rainfall, both occurring in a period prior to and during the nesting period, and finally (iv) a longer‐term negative association with higher maximum temperatures following the previous breeding season. Our results illustrate how a morphological trait may be affected by both short‐ and long‐term effects of the same weather variable at multiple times of the year and that these effects may act in different directions. We also show that climate within the relevant time windows may not be changing in the same way, such that overall long‐term temporal trends in body size may be minimal. Such complexity means that analytical approaches that search for a single ‘best’ window for one particular weather variable may miss other relevant information, and is also likely to make analyses of phenotypic plasticity and prediction of longer‐term population dynamics difficult.     

Seasonal climate effects on the survival of a hibernating mammal

Global climate change and associated regional climate variability is impacting the phenology of many species, ultimately altering individual fitness and population dynamics. Yet, few studies have considered the effects of pertinent seasonal climate variability on phenology and fitness. Hibernators may be particularly susceptible to changes in seasonal climate since they have a relatively short active season in which to reproduce and gain enough mass to survive the following winter. To understand whether and how seasonal climate variability may be affecting hibernator fitness, we estimated survival from historical (1964–1968) and contemporary (2014–2017) mark–recapture data collected from the same population of Uinta ground squirrels (UGS, Urocitellus armatus), a hibernator endemic to the western United States. Despite a locally warming climate, the phenology of UGS did not change over time, yet season‐specific climate variables were important in regulating survival rates. Specifically, older age classes experienced lower survival when winters or the following springs were warm, while juveniles benefited from warmer winter temperatures. Although metabolic costs decrease with decreasing temperature in the hibernacula, arousal costs increase with decreasing temperature. Our results suggest that this trade‐off is experienced differently by immature and mature individuals. We also observed an increase in population density during that time period, suggesting resources are less limited today than they used to be. Cheatgrass is now dominating the study site and may provide a better food source to UGS than native plants did historically.     

Time‐lag effects of global vegetation responses to climate change

Climate conditions significantly affect vegetation growth in terrestrial ecosystems. Due to the spatial heterogeneity of ecosystems, the vegetation responses to climate vary considerably with the diverse spatial patterns and the time‐lag effects, which are the most important mechanism of climate–vegetation interactive effects. Extensive studies focused on large‐scale vegetation–climate interactions use the simultaneous meteorological and vegetation indicators to develop models; however, the time‐lag effects are less considered, which tends to increase uncertainty. In this study, we aim to quantitatively determine the time‐lag effects of global vegetation responses to different climatic factors using the GIMMS3g NDVI time series and the CRU temperature, precipitation, and solar radiation datasets. First, this study analyzed the time‐lag effects of global vegetation responses to different climatic factors. Then, a multiple linear regression model and partial correlation model were established to statistically analyze the roles of different climatic factors on vegetation responses, from which the primary climate‐driving factors for different vegetation types were determined. The results showed that (i) both the time‐lag effects of the vegetation responses and the major climate‐driving factors that significantly affect vegetation growth varied significantly at the global scale, which was related to the diverse vegetation and climate characteristics; (ii) regarding the time‐lag effects, the climatic factors explained 64% variation of the global vegetation growth, which was 11% relatively higher than the model ignoring the time‐lag effects; (iii) for the area with a significant change trend (for the period 1982–2008) in the global GIMMS3g NDVI (P < 0.05), the primary driving factor was temperature; and (iv) at the regional scale, the variation in vegetation growth was also related to human activities and natural disturbances. Considering the time‐lag effects is quite important for better predicting and evaluating the vegetation dynamics under the background of global climate change.     

Beyond a warming fingerprint: individualistic biogeographic responses to heterogeneous climate change in California

Understanding recent biogeographic responses to climate change is fundamental for improving our predictions of likely future responses and guiding conservation planning at both local and global scales. Studies of observed biogeographic responses to 20th century climate change have principally examined effects related to ubiquitous increases in temperature – collectively termed a warming fingerprint. Although the importance of changes in other aspects of climate – particularly precipitation and water availability – is widely acknowledged from a theoretical standpoint and supported by paleontological evidence, we lack a practical understanding of how these changes interact with temperature to drive biogeographic responses. Further complicating matters, differences in life history and ecological attributes may lead species to respond differently to the same changes in climate. Here, we examine whether recent biogeographic patterns across California are consistent with a warming fingerprint. We describe how various components of climate have changed regionally in California during the 20th century and review empirical evidence of biogeographic responses to these changes, particularly elevational range shifts. Many responses to climate change do not appear to be consistent with a warming fingerprint, with downslope shifts in elevation being as common as upslope shifts across a number of taxa and many demographic and community responses being inconsistent with upslope shifts. We identify a number of potential direct and indirect mechanisms for these responses, including the influence of aspects of climate change other than temperature (e.g., the shifting seasonal balance of energy and water availability), differences in each taxon's sensitivity to climate change, trophic interactions, and land‐use change. Finally, we highlight the need to move beyond a warming fingerprint in studies of biogeographic responses by considering a more multifaceted view of climate, emphasizing local‐scale effects, and including a priori knowledge of relevant natural history for the taxa and regions under study.     

Separating direct and indirect effects of global change: a population dynamic modeling approach using readily available field data

Two sources of complexity make predicting plant community response to global change particularly challenging. First, realistic global change scenarios involve multiple drivers of environmental change that can interact with one another to produce non‐additive effects. Second, in addition to these direct effects, global change drivers can indirectly affect plants by modifying species interactions. In order to tackle both of these challenges, we propose a novel population modeling approach, requiring only measurements of abundance and climate over time. To demonstrate the applicability of this approach, we model population dynamics of eight abundant plant species in a multifactorial global change experiment in alpine tundra where we manipulated nitrogen, precipitation, and temperature over 7 years. We test whether indirect and interactive effects are important to population dynamics and whether explicitly incorporating species interactions can change predictions when models are forecast under future climate change scenarios. For three of the eight species, population dynamics were best explained by direct effect models, for one species neither direct nor indirect effects were important, and for the other four species indirect effects mattered. Overall, global change had negative effects on species population growth, although species responded to different global change drivers, and single‐factor effects were slightly more common than interactive direct effects. When the fitted population dynamic models were extrapolated under changing climatic conditions to the end of the century, forecasts of community dynamics and diversity loss were largely similar using direct effect models that do not explicitly incorporate species interactions or best‐fit models; however, inclusion of species interactions was important in refining the predictions for two of the species. The modeling approach proposed here is a powerful way of analyzing readily available datasets which should be added to our toolbox to tease apart complex drivers of global change.     

Synergistic effects of climate and land‐use change influence broad‐scale avian population declines

Climate and land‐use changes are expected to be the primary drivers of future global biodiversity loss. Although theory suggests that these factors impact species synergistically, past studies have either focused on only one in isolation or have substituted space for time, which often results in confounding between drivers. Tests of synergistic effects require congruent time series on animal populations, climate change and land‐use change replicated across landscapes that span the gradient of correlations between the drivers of change. Using a unique time series of high‐resolution climate (measured as temperature and precipitation) and land‐use change (measured as forest change) data, we show that these drivers of global change act synergistically to influence forest bird population declines over 29 years in the Pacific Northwest of the United States. Nearly half of the species examined had declined over this time. Populations declined most in response to loss of early seral and mature forest, with responses to loss of early seral forest amplified in landscapes that had warmed over time. In addition, birds declined more in response to loss of mature forest in areas that had dried over time. Climate change did not appear to impact populations in landscapes with limited habitat loss, except when those landscapes were initially warmer than the average landscape. Our results provide some of the first empirical evidence of synergistic effects of climate and land‐use change on animal population dynamics, suggesting accelerated loss of biodiversity in areas under pressure from multiple global change drivers. Furthermore, our findings suggest strong spatial variability in the impacts of climate change and highlight the need for future studies to evaluate multiple drivers simultaneously to avoid potential misattribution of effects.     

Climate change,phenology, and habitat degradation: drivers of gosling body condition and juvenile survival in lesser snow geese

Nesting migratory geese are among the dominant herbivores in (sub) arctic environments, which have undergone unprecedented increases in temperatures and plant growing days over the last three decades. Within these regions, the Hudson Bay Lowlands are home to an overabundant breeding population of lesser snow geese that has dramatically damaged the ecosystem, with cascading effects at multiple trophic levels. In some areas the overabundance of geese has led to a drastic reduction in available forage. In addition, warming of this region has widened the gap between goose migration timing and plant green‐up, and this ‘mismatch’ between goose and plant phenologies could in turn affect gosling development. The dual effects of climate change and habitat quality on gosling body condition and juvenile survival are not known, but are critical for predicting population growth and related degradation of (sub) arctic ecosystems. To address these issues, we used information on female goslings marked and measured between 1978 and 2005 (4125 individuals). Goslings that developed within and near the traditional center of the breeding colony experienced the effects of long‐term habitat degradation: body condition and juvenile survival declined over time. In newly colonized areas, however, we observed the opposite pattern (increase in body condition and juvenile survival). In addition, warmer than average winters and summers resulted in lower gosling body condition and first‐year survival. Too few plant ‘growing days’ in the spring relative to hatch led to similar results. Our assessment indicates that geese are recovering from habitat degradation by moving to newly colonized locales. However, a warmer climate could negatively affect snow goose populations in the long‐run, but it will depend on which seasons warm the fastest. These antagonistic mechanisms will require further study to help predict snow goose population dynamics and manage the trophic cascade they induce.     

Dynamic size responses to climate change: prevailing effects of rising temperature drive long‐term body size increases in a semi‐arid passerine

Changes in animal body size have been widely reported as a correlate of contemporary climate change. Body size affects metabolism and fitness, so changing size has implications for resilience, yet the climatic factors that drive size variation remain poorly understood. We test the role of mean and extreme temperature, rainfall, and remotely sensed primary productivity (NDVI) as drivers of body size in a sedentary, semi‐arid Australian passerine, Ptilotula (Lichenostomus) penicillatus, over 23 years. To distinguish effects due to differential growth from changes in population composition, we analysed first‐year birds and adults separately and considered climatic variation at three temporal scales (current, previous, and preceding 5 years). The strongest effects related to temperature: in both age classes, larger size was associated with warmer mean temperatures in the previous year, contrary to Bergmann's Rule. Moreover, adults were larger in warmer breeding seasons, while first years was larger after heatwaves; these effects are more likely to be mediated through size‐dependent mortality, highlighting the role of body size in determining vulnerability to extinction. In addition to temperature, larger adult size was associated with lower primary productivity, which may reflect a trade‐off between vegetative growth and nectar production, on which adults rely. Finally, lower rainfall was associated with decreasing size in first year and adults, most likely related to decreased food availability. Overall, body size increased over 23 years, strongly in first‐year birds (2.7%) compared with adults (1%), with size outcomes a balance between competing drivers. As rainfall declined over time and productivity remained fairly stable, the temporal increase in body size appears largely driven by rising mean temperature and temperature extremes. Body size responses to environmental change are thus complex and dynamic, driven by effects on growth as well as mortality.     

Direct and indirect effects of winter harshness on the survival of Mallards Anas platyrhynchos in northwest Europe

To understand population dynamics it is necessary to understand vital rates, which may be affected by a wide range of factors including environmental variables such as weather. Weather conditions can affect birds’ vital rates directly through increased mortality due to impaired conditions, or indirectly via changes in body condition and/or behaviour. Most understanding of direct and indirect effects of weather comes from studies of breeding birds, whereas the situation in non‐breeding periods is less clear. Here, we analysed annual survival of non‐breeding Mallard Anas platyrhynchos, the most hunted waterfowl species in Europe, and assessed whether survival is related directly to winter harshness and/or indirectly via changes in winter recovery distributions. Recovery data on Mallards, initially marked in southeast Sweden, were analysed with an information‐theoretic approach using program mark . Over 10 000 Mallards were marked in two time periods, 1964–1982 and 2002–2008, of which 13.3 and 4.7%, respectively, were later recovered. Mallards had lower annual survival in the early trapping period (0.58–0.63) than in the later period (0.69–0.71), with no clear effects of sex, age or year. Within each study period, winter harshness did not directly correlate with survival. However, milder winters may have contributed indirectly to higher survival in the second period, as winter harshness data were correlated with the distances to recovery positions for females, and also because winter recovery areas have shifted northeast during the past decades, possibly indicating a shortened migratory distance. Migration is costly, and there is therefore a likely linkage between migration behaviour and survival of dabbling ducks, in which direct as well as indirect effects of winter harshness may play a role. Other factors, such as hunting pressure, are also likely to have changed in the past decades, and may also have contributed to improved survival of Mallards in northwest Europe.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

MHC‐dependent survival in a wild population: evidence for hidden genetic benefits gained through extra‐pair fertilizations

Females should prefer to be fertilized by males that increase the genetic quality of their offspring. In vertebrates, genes of the major histocompatibility complex (MHC) play a key role in the acquired immune response and have been shown to affect mating preferences. They are therefore important candidates for the link between mate choice and indirect genetic benefits. Higher MHC diversity may be advantageous because this allows a wider range of pathogens to be detected and combated. Furthermore, individuals harbouring rare MHC alleles might better resist pathogen variants that have evolved to evade common MHC alleles. In the Seychelles warbler, females paired with low MHC‐diversity males elevate the MHC diversity of their offspring to levels comparable to the population mean by gaining extra‐pair fertilizations. Here, we investigate whether increased MHC diversity results in higher life expectancy and whether there are any additional benefits of extra‐pair fertilizations. Our 10‐year study found a positive association between MHC diversity and juvenile survival, but no additional survival advantage of extra‐pair fertilizations. In addition, offspring with a specific allele (Ase‐ua4) had a fivefold longer life expectancy than offspring without this allele. Consequently, the interacting effects of sexual selection and pathogen‐mediated viability selection appear to be important for maintaining MHC variation in the Seychelles warbler. Our study supports the prediction that MHC‐dependent extra‐pair fertilizations result in genetic benefits for offspring in natural populations. However, such genetic benefits might be hidden and not necessarily apparent in the widely used fitness comparison of extra‐ and within‐pair offspring.     

Evolutionary importance of parental care performance,food resources,and direct and indirect genetic effects in a burying beetle

Indirect genetic effects (IGE) of parental care performance and the direct–indirect covariance contribute substantially to total heritability in domesticated and laboratory mammals. For animals from natural populations empirical estimates of IGE are sparse. Thus, despite recent models relating IGE to evolution, evolutionary interpretations of IGE are limited. To address this deficit, we used a reciprocal cross‐fostering breeding design to estimate environmental influences, direct and indirect genetic effects, and direct–indirect genetic covariances in the burying beetle Nicrophorus pustulatus to determine the evolutionary importance of IGE arising from variation in parental care performance. Carrion size positively affected adult mass and time on carrion, but had no effect on total development time. Males were slightly larger than females. For both mass and development, independent of these environmental influences, direct and indirect genetic effects were of moderate magnitude. Total genetic effects explained 36–50% of the phenotypic variance in mass and size and 27–37% of phenotypic variance in development time. Direct–indirect genetic covariances were zero or close to zero. Thus, for both mass and development time, the response to natural selection arising from environmental variation may be accelerated by the presence of IGE in N. pustulatus. The generality of this pattern and the evolutionary significance of IGE arising from parental care awaits further study of natural populations.