Seasonal reproductive costs contribute to reduced survival of female greater sage‐grouse

Tradeoffs among demographic traits are a central component of life history theory. We investigated tradeoffs between reproductive effort and survival in female greater sage‐grouse breeding in the American Great Basin, while also considering reproductive heterogeneity by examining covariance among current and future reproductive success. We analyzed survival and reproductive histories from 328 individual female greater sage‐grouse captured between 2003 and 2011, and examined the effect of reproductive effort on survival and future reproduction. Monthly survival of females was variable within years, and this within‐year variation was associated with distinct biological seasons. Monthly survival was greatest during the winter (November–March; Φ_W= 0.99 ± 0.001 SE), and summer (June–July; Φ_S= 0.98 ± 0.01 SE), and lower during nesting (April–May; Φ_N= 0.93 ± 0.02 SE) and fall (August–October; Φ_F= 0.92 ± 0.02 SE). Successful reproduction was associated with reduced monthly survival during summer and fall, and this effect was greatest during fall. Females that successfully fledged chicks had lower annual survival (0.47 ± 0.05 SE) than females who were not successful (0.64 ± 0.04 SE). Annual survival did not vary across years, consistent with a slow‐paced life history strategy in greater sage‐grouse. In contrast, reproductive success varied widely, and was positively correlated with annual rainfall. We found evidence for heterogeneity among females with respect to reproductive success; compared with unsuccessful females, females that raised a brood successfully in year t were more than twice as likely to be successful in year t+ 1. Female greater sage‐grouse incur costs to survival associated with reproduction, however, variation in quality among females may override costs to subsequent reproductive output.     

Survival of Gunnison sage‐grouse Centrocercus minimus in Colorado,USA

Gunnison sage‐grouse Centrocercus minimus has declined from their historic range and recent monitoring has provided evidence that some populations are continuing to decline. The evaluation of long‐term, population‐specific survival rates is important to assess population stability, and is necessary for conservation of this species of concern. We evaluated adult and yearling survival in two dynamically different populations of Gunnison sage‐grouse (a relatively large, more stable population and a small, declining population). Our goal was to examine the relationship between annual survival and population, and test hypotheses with regards to temporal effects (across years and within year) and individual effects (sex and age). We also evaluated the effects of snow depth on sage‐grouse survival. We tracked 214 radiomarked birds in the large population from 2005–2010 and 25 birds in the small population from 2007–2010. We found no evidence for a difference in survival between yearlings and adults nor did we find an influence of snow depth on survival. Males had the lowest survival during the lekking season (March–April); females had lower survival during the nesting and chick rearing season (May–July) and late‐summer and fall (August–October). The annual survival rate was 0.61 (SE 0.06) for females and 0.39 (SE 0.08) for males. Survival was constant across years and between the populations suggesting observed population changes during this time period are not a result of changes in adult survival.     

Equal nonbreeding period survival in adults and juveniles of a long‐distant migrant bird

In migrant birds, survival estimates for the different life‐history stages between fledging and first breeding are scarce. First‐year survival is shown to be strongly reduced compared with annual survival of adult birds. However, it remains unclear whether the main bottleneck in juvenile long‐distant migrants occurs in the postfledging period within the breeding ranges or en route. Quantifying survival rates during different life‐history stages and during different periods of the migration cycle is crucial to understand forces driving the evolution of optimal life histories in migrant birds. Here, we estimate survival rates of adult and juvenile barn swallows (Hirundo rustica L.) in the breeding and nonbreeding areas using a population model integrating survival estimates in the breeding ranges based on a large radio‐telemetry data set and published estimates of demographic parameters from large‐scale population‐monitoring projects across Switzerland. Input parameters included the country‐wide population trend, annual productivity estimates of the double‐brooded species, and year‐to‐year survival corrected for breeding dispersal. Juvenile survival in the 3‐week postfledging period was low (S = 0.32; SE = 0.05), whereas in the rest of the annual cycle survival estimates of adults and juveniles were similarly high (S > 0.957). Thus, the postfledging period was the main survival bottleneck, revealing the striking result that nonbreeding period mortality (including migration) is not higher for juveniles than for adult birds. Therefore, focusing future research on sources of variation in postfledging mortality can provide new insights into determinants of population dynamics and life‐history evolution of migrant birds.     

Spatially explicit models of seasonal habitat for greater sage‐grouse at broad spatial scales: Informing areas for management in Nevada and northeastern California

Defining boundaries of species' habitat across broad spatial scales is often necessary for management decisions, and yet challenging for species that demonstrate differential variation in seasonal habitat use. Spatially explicit indices that incorporate temporal shifts in selection can help overcome such challenges, especially for species of high conservation concern. Greater sage‐grouse Centrocercus urophasianus (hereafter, sage‐grouse), a sagebrush obligate species inhabiting the American West, represents an important case study because sage‐grouse exhibit seasonal habitat patterns, populations are declining in most portions of their range and are central to contemporary national land use policies. Here, we modeled spatiotemporal selection patterns for telemetered sage‐grouse across multiple study sites (1,084 sage‐grouse; 30,690 locations) in the Great Basin. We developed broad‐scale spatially explicit habitat indices that elucidated space use patterns (spring, summer/fall, and winter) and accounted for regional climatic variation using previously published hydrographic boundaries. We then evaluated differences in selection/avoidance of each habitat characteristic between seasons and hydrographic regions. Most notably, sage‐grouse consistently selected areas dominated by sagebrush with few or no conifers but varied in type of sagebrush selected by season and region. Spatiotemporal variation was most apparent based on availability of water resources and herbaceous cover, where sage‐grouse strongly selected upland natural springs in xeric regions but selected larger wet meadows in mesic regions. Additionally, during the breeding period in spring, herbaceous cover was selected strongly in the mesic regions. Lastly, we expanded upon an existing joint–index framework by combining seasonal habitat indices with a probabilistic index of sage‐grouse abundance and space use to produce habitat maps useful for sage‐grouse management. These products can serve as conservation planning tools that help predict expected benefits of restoration activities, while highlighting areas most critical to sustaining sage‐grouse populations. Our joint–index framework can be applied to other species that exhibit seasonal shifts in habitat requirements to help better guide conservation actions.     

Range‐wide patterns of greater sage‐grouse persistence

Aim Greater sage‐grouse (Centrocercus urophasianus), a shrub‐steppe obligate species of western North America, currently occupies only half its historical range. Here we examine how broad‐scale, long‐term trends in landscape condition have affected range contraction. Location Sagebrush biome of the western USA. Methods Logistic regression was used to assess persistence and extirpation of greater sage‐grouse range based on landscape conditions measured by human population (density and population change), vegetation (percentage of sagebrush habitat), roads (density of and distance to roads), agriculture (cropland, farmland and cattle density), climate (number of severe and extreme droughts) and range periphery. Model predictions were used to identify areas where future extirpations can be expected, while also explaining possible causes of past extirpations. Results Greater sage‐grouse persistence and extirpation were significantly related to sagebrush habitat, cultivated cropland, human population density in 1950, prevalence of severe droughts and historical range periphery. Extirpation of sage‐grouse was most likely in areas having at least four persons per square kilometre in 1950, 25% cultivated cropland in 2002 or the presence of three or more severe droughts per decade. In contrast, persistence of sage‐grouse was expected when at least 30 km from historical range edge and in habitats containing at least 25% sagebrush cover within 30 km. Extirpation was most often explained (35%) by the combined effects of peripherality (within 30 km of range edge) and lack of sagebrush cover (less than 25% within 30 km). Based on patterns of prior extirpation and model predictions, we predict that 29% of remaining range may be at risk. Main Conclusions Spatial patterns in greater sage‐grouse range contraction can be explained by widely available landscape variables that describe patterns of remaining sagebrush habitat and loss due to cultivation, climatic trends, human population growth and peripherality of populations. However, future range loss may relate less to historical mechanisms and more to recent changes in land use and habitat condition, including energy developments and invasions by non‐native species such as cheatgrass (Bromus tectorum) and West Nile virus. In conjunction with local measures of population performance, landscape‐scale predictions of future range loss may be useful for prioritizing management and protection. Our results suggest that initial conservation efforts should focus on maintaining large expanses of sagebrush habitat, enhancing quality of existing habitats, and increasing habitat connectivity.     

Phenology largely explains taller grass at successful nests in greater sage‐grouse

Much interest lies in the identification of manageable habitat variables that affect key vital rates for species of concern. For ground‐nesting birds, vegetation surrounding the nest may play an important role in mediating nest success by providing concealment from predators. Height of grasses surrounding the nest is thought to be a driver of nest survival in greater sage‐grouse (Centrocercus urophasianus; sage‐grouse), a species that has experienced widespread population declines throughout their range. However, a growing body of the literature has found that widely used field methods can produce misleading inference on the relationship between grass height and nest success. Specifically, it has been demonstrated that measuring concealment following nest fate (failure or hatch) introduces a temporal bias whereby successful nests are measured later in the season, on average, than failed nests. This sampling bias can produce inference suggesting a positive effect of grass height on nest survival, though the relationship arises due to the confounding effect of plant phenology, not an effect on predation risk. To test the generality of this finding for sage‐grouse, we reanalyzed existing datasets comprising >800 sage‐grouse nests from three independent studies across the range where there was a positive relationship found between grass height and nest survival, including two using methods now known to be biased. Correcting for phenology produced equivocal relationships between grass height and sage‐grouse nest survival. Viewed in total, evidence for a ubiquitous biological effect of grass height on sage‐grouse nest success across time and space is lacking. In light of these findings, a reevaluation of land management guidelines emphasizing specific grass height targets to promote nest success may be merited.     

Ectoparasitism and the role of green nesting material in the European starling

Summary The use of green nesting material is widespred among birds. Recent evidence suggests that birds use secondary chemicals contained in green plants to control ectoparasites. We manipulated green nesting material and ectoparasites of European starlings (Sturnus vulgaris) to test two hypotheses: (1) ectoparasites adversely affect prefledging survival and morphometrics or postfledging survival, and (2) green nesting material ameliorates the effects of ectoparasites. We recorded fat score, numbers of scabs, tarsal length, body mass, and hematocrit level on each nestling 17 days after hatching. We also fitted each nestling with unique patagial tags and resighted the starlings for 6–8 weeks after fledging to estimate survival and sighting rates. Nests devoid of green nesting material and dusted with the insecticide, carbaryl, had fewer high ectoparasite infestations, and nestlings had significantly lower scab scores, and significantly higher body masses than nestlings in undusted boxes. However, there was no difference in postfledging survival between birds from carbaryl-treated and undusted nests. There also was no difference in prefledging survival and morphometrics or postfledging survival between nestlings from boxes with and without green nesting material. These results do not support the hypothesis that starlings use green nesting material to control nest ectoparasites. We suggest an alternative hypothesis; green nesting material is used for mate selection or pairbonding in the starling.     

Modeling ecological minimum requirements for distribution of greater sage‐grouse leks: implications for population connectivity across their western range,U.S.A

Greater sage‐grouse Centrocercus urophasianus (Bonaparte) currently occupy approximately half of their historical distribution across western North America. Sage‐grouse are a candidate for endangered species listing due to habitat and population fragmentation coupled with inadequate regulation to control development in critical areas. Conservation planning would benefit from accurate maps delineating required habitats and movement corridors. However, developing a species distribution model that incorporates the diversity of habitats used by sage‐grouse across their widespread distribution has statistical and logistical challenges. We first identified the ecological minimums limiting sage‐grouse, mapped similarity to the multivariate set of minimums, and delineated connectivity across a 920,000 km² region. We partitioned a Mahalanobis D² model of habitat use into k separate additive components each representing independent combinations of species–habitat relationships to identify the ecological minimums required by sage‐grouse. We constructed the model from abiotic, land cover, and anthropogenic variables measured at leks (breeding) and surrounding areas within 5 km. We evaluated model partitions using a random subset of leks and historic locations and selected D² (k = 10) for mapping a habitat similarity index (HSI). Finally, we delineated connectivity by converting the mapped HSI to a resistance surface. Sage‐grouse required sagebrush‐dominated landscapes containing minimal levels of human land use. Sage‐grouse used relatively arid regions characterized by shallow slopes, even terrain, and low amounts of forest, grassland, and agriculture in the surrounding landscape. Most populations were interconnected although several outlying populations were isolated because of distance or lack of habitat corridors for exchange. Land management agencies currently are revising land‐use plans and designating critical habitat to conserve sage‐grouse and avoid endangered species listing. Our results identifying attributes important for delineating habitats or modeling connectivity will facilitate conservation and management of landscapes important for supporting current and future sage‐grouse populations.     

Prefledging Growth and Recruitment of Female Lesser Scaup

Annual variation in juvenile recruitment is an important component of duck population dynamics, yet little is known about the factors affecting the probability of surviving and breeding in the first year of life. Two hypothesized mechanisms to explain annual variability are indirect carry-over effects (COEs) from conditions experienced during the prefledging period and direct effects from climatic conditions during the postfledging period. We used Cormack-Jolly-Seber models to estimate apparent survival and detection rates of 643 juvenile female lesser scaup (Aythya affinis) marked just prior to fledging at Red Rock Lakes National Wildlife Refuge in southwestern Montana, USA, 2010–2018. We evaluated COEs from hatch date, a hatch date × spring phenology interaction, and conspecific duckling density in addition to a direct climatic effect of winter conditions (indexed by the El Niño Southern Oscillation [ENSO]) and spring habitat conditions on the study area. We used growth data from a subset (n = 190) of known-aged ducklings to estimate the influence of hatch date and conspecific density on prefledging growth to help identify mechanisms underlying COEs. Prefledging growth and juvenile apparent survival were negatively related to measures of conspecific duckling density. We found evidence that detection probability varied annually for juvenile (but not adult) scaup, possibly representing decisions to delay breeding and not return to or remain at the study site in their first year of life. Like with apparent survival, there was suggestive evidence that detection probability decreased with increasing duckling density in the previous year. Hatching date was weakly negatively related to detection probability, but unrelated to apparent survival, whereas neither vital rate was related to winter ENSO index. Our results are consistent with a process where density-dependent growth rates in the prefledging period carry over to influence fitness in subsequent life-cycle stages. If this pattern generalizes to other systems, this density COE may have important implications for our understanding of duck population dynamics and reaffirms the importance of maintaining abundant brood-rearing habitats in conservation and management of ducks. © 2021 The Wildlife Society.     

Precipitation and winter temperature predict long‐term range‐scale abundance changes in Western North American birds

Predicting biodiversity responses to climate change remains a difficult challenge, especially in climatically complex regions where precipitation is a limiting factor. Though statistical climatic envelope models are frequently used to project future scenarios for species distributions under climate change, these models are rarely tested using empirical data. We used long‐term data on bird distributions and abundance covering five states in the western US and in the Canadian province of British Columbia to test the capacity of statistical models to predict temporal changes in bird populations over a 32‐year period. Using boosted regression trees, we built presence‐absence and abundance models that related the presence and abundance of 132 bird species to spatial variation in climatic conditions. Presence/absence models built using 1970–1974 data forecast the distributions of the majority of species in the later time period, 1998–2002 (mean AUC = 0.79 ± 0.01). Hindcast models performed equivalently (mean AUC = 0.82 ± 0.01). Correlations between observed and predicted abundances were also statistically significant for most species (forecast mean Spearman′s ρ = 0.34 ± 0.02, hindcast = 0.39 ± 0.02). The most stringent test is to test predicted changes in geographic patterns through time. Observed changes in abundance patterns were significantly positively correlated with those predicted for 59% of species (mean Spearman′s ρ = 0.28 ± 0.02, across all species). Three precipitation variables (for the wettest month, breeding season, and driest month) and minimum temperature of the coldest month were the most important predictors of bird distributions and abundances in this region, and hence of abundance changes through time. Our results suggest that models describing associations between climatic variables and abundance patterns can predict changes through time for some species, and that changes in precipitation and winter temperature appear to have already driven shifts in the geographic patterns of abundance of bird populations in western North America.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.     

Variation in nestling body condition and wing development predict cause‐specific mortality in fledgling dickcissels

Phenotypic traits developed in one life‐history stage can carryover and affect survival in subsequent stages. For songbirds, carryover effects from the pre‐ to post‐fledging period may be crucial for survival but are poorly understood. We assessed whether juvenile body condition and wing development at fledging influenced survival during the post‐fledging period in the dickcissel Spiza americana. We found pre‐ to post‐fledging carryover effects on fledgling survival for both traits during the ‘early part’ – first four days – of the post‐fledging period. Survival benefits of each trait depended on cause‐specific sources of mortality; individuals in better body condition were less likely to die from exposure to adverse environmental conditions, whereas those with more advanced wing development were less likely to be preyed upon. Fledglings with more advanced wing development were comparatively more active and mobile earlier in the post‐fledging period, suggesting they were better able to avoid predators. Our results provide some of the first evidence linking development of juvenile phenotypic traits to survival against specific sources of post‐fledging mortality in songbirds. Further investigation into pre‐ to post‐fledging carryover effects may yield important insights into avian life‐history evolution.     

A recipe for postfledging survival in great tits Parus major: be large and be early (but not too much)

Survival of juveniles during the postfledging period can be markedly low, which may have major consequences on avian population dynamics. Knowing which factors operating during the nesting phase affect postfledging survival is crucial to understand avian breeding strategies. We aimed to obtain a robust set of predictors of postfledging local survival using the great tit (Parus major) as a model species. We used mark–recapture models to analyze the effect of hatching date, temperatures experienced during the nestling period, fledging size and body mass on first‐year postfledging survival probability of great tit juveniles. We used data from 5192 nestlings of first clutches ringed between 1993 and 2010. Mean first‐year postfledging survival probability was 15.2%, and it was lower for smaller individuals, as well as for those born in either very early or late broods. Our results stress the importance of choosing an optimum hatching period, and raising large chicks to increase first‐year local survival probability in the studied population.     

Seasonal variability in survivorship of a cooperatively breeding tropical passerine

Survival of tropical passerines is thought to be higher than those in northern temperate regions, but relatively few tropical studies have addressed this issue, particularly in tropical Asia. We examined factors that may have influenced the survival rate of a cooperatively breeding bird, the puff-throated bulbul (Alophoixus pallidus), in an evergreen forest in northeastern Thailand. These factors included year, season (breeding and non-breeding), sex, and presence of helper(s) in a family group. We present evidence of breeding season-dependent survival in a tropical passerine using an information theoretic approach based on both mark-recapture and resighting data collected during 6 years of study. Based on colour-banded adults the annual survival rate did not vary significantly among years (average = 0.85 ± 0.02 SE). The mean lifespan (MLS) for the population was 6.22 ± 4.38 SE years. Survivorship was lower during the breeding season (0.89 ± 0.02 SE) than during the non-breeding season (0.96 ± 0.02 SE). The MLS of males and females was 6.70 ± 7.73 SE and 5.87 ± 4.88 SE years, respectively. The annual survival rate we observed was high compared to the estimates of other tropical and temperate passerines, possibly due to the relatively stable climatic conditions in tropical latitudes and puff-throated bulbuls being generalists that exploit a wide range of food resources both in space and time.     

West Nile virus: pending crisis for greater sage-grouse

Scientists have feared that emerging infectious diseases could complicate efforts to conserve rare and endangered species, but quantifying impacts has proven difficult until now. We report unexpected impacts of West Nile virus (WNv) on radio‐marked greater sage‐grouse (Centrocercus urophasianus), a species that has declined 45–80% and is endangered in Canada and under current consideration for federal listing in the US. We show that WNv reduced late‐summer survival an average of 25% in four radio‐marked populations in the western US and Canada. Serum from 112 sage‐grouse collected after the outbreak show that none had antibodies, suggesting that they lack resistance. The spread of WNv represents a significant new stressor on sage‐grouse and probably other at‐risk species. While managing habitat might lessen its impact on sage‐grouse populations, WNv has left wildlife and public health officials scrambling to address surface water and vector control issues in western North America.     

Restoring Sage‐grouse nesting habitat through removal of early successional conifer

Conifer woodlands have expanded into sagebrush (Artemisia spp.) ecosystems and degrade habitat for sagebrush obligate species such as the Greater Sage‐grouse (Centrocercus urophasianus). Conifer management is increasing despite a lack of empirical evidence assessing outcomes to grouse and their habitat. Although assessments of vegetation recovery after conifer removal are common, comparisons of successional trends with habitat guidelines or actual data on habitat used by sage‐grouse is lacking. We assessed impacts of conifer encroachment on vegetation characteristics known to be important for sage‐grouse nesting. Using a controlled repeated measures design, we then evaluated vegetation changes for 3 years after conifer removal. We compared these results to data from 356 local sage‐grouse nests, rangewide nesting habitat estimates, and published habitat guidelines. We measured negative effects of conifer cover on many characteristics important for sage‐grouse nesting habitat including percent cover of forbs, grasses, and shrubs, and species richness of forbs and shrubs. In untreated habitat, herbaceous vegetation cover was slightly below the cover at local nest sites, while shrub cover and sagebrush cover were well below cover at the nest sites. Following conifer removal, we measured increases in herbaceous vegetation, primarily grasses, and sagebrush height. Our results indicate that conifer abundance can decrease habitat suitability for nesting sage‐grouse. Additionally, conifer removal can improve habitat suitability for nesting sage‐grouse within 3 years, and trajectories indicate that the habitat may continue to improve in the near future.     

The effects of acaricide treatment of sheep on red grouse Lagopus lagopus scotica tick burdens and productivity in a multi‐host system

Ixodes ricinus (Ixodida: Ixodidae) ticks are of economic and pathogenic importance across Europe. Within the uplands of the U.K., management to reduce ticks is undertaken to benefit red grouse Lagopus lagopus scotica (Galliformes: Phasianidae). Management strategies focus on the acaricide treatment of domestic sheep Ovis aries (Artiodactyla: Bovidae), but the effectiveness of this is less certain in the presence of wild hosts, particularly red deer Cervus elaphus (Artiodactyla: Cervidae) and mountain hare Lepus timidus (Lagomorpha: Leporidae). This study examines the effects of sheep management on grouse tick burdens and productivity using sites with a range of wild host densities. Sites at which applications of acaricide were more frequent had lower tick burdens; this relationship was similar on sites with a range of deer densities. However, no direct link was detected between acaricide treatment interval and grouse productivity. Sites with higher deer densities had higher grouse tick burdens and lower productivity [mean ± standard error (SE) young : adult ratio: 1.2 ± 0.2] compared with sites with lower deer densities (mean ± SE young : adult ratio: 1.8 ± 0.1). Sites with higher grouse brood sizes and higher proportions of hens with broods were also those with higher mountain hare abundance indices. This study highlights the importance of the frequent treatment of sheep with acaricide to reduce tick burdens on grouse, even in the presence of wild hosts.     

Sagebrush treatments influence annual population change for greater sage‐grouse

Vegetation management practices have been applied worldwide to enhance habitats for a variety of wildlife species. Big sagebrush (Artemisia tridentata spp.) communities, iconic to western North America, have been treated to restore herbaceous understories through chemical, mechanical, and prescribed burning practices thought to improve habitat conditions for greater sage‐grouse (Centrocercus urophasianus) and other species. Although the response of structural attributes of sagebrush communities to treatments is well understood, there is a need to identify how treatments influence wildlife population dynamics. We investigated the influence of vegetation treatments occurring in Wyoming, United States, from 1994 to 2012 on annual sage‐grouse population change using yearly male sage‐grouse lek counts. We investigated this response across 1, 3, 5, and 10‐year post‐treatment lags to evaluate how the amount of treated sagebrush communities and time since treatment influenced population change, while accounting for climate, wildfire, and anthropogenic factors. With the exception of chemical treatments exhibiting a positive association with sage‐grouse population change 11 years after implementation, population response to treatments was either neutral or negative for at least 11 years following treatments. Our work supports a growing body of research advocating against treating big sagebrush habitats for sage‐grouse, particularly in Wyoming big sagebrush (A. t. wyomingensis). Loss and fragmentation of sagebrush habitats has been identified as a significant threat for remaining sage‐grouse populations. Because sagebrush may take decades to recover following treatments, we recommend practitioners use caution when designing projects to alter remaining habitats, especially when focused on habitat requirements for one life stage and a single species.     

Age‐dependence and individual heterogeneity in reproductive success of greater sage‐grouse

Research on iteroparous species has shown that reproductive success may increase with age until the onset of senescence. However, from the population perspective, increased reproductive success with age could be a consequence of within‐individual variation (e.g. ageing, breeding experience, foraging ability hypotheses), between‐individual variation (e.g. individual heterogeneity, frailty, selection, delayed breeding hypotheses), or a combination thereof. We evaluated within‐ and between‐individual variation in reproductive success of greater sage‐grouse (Centrocercus urophasianus; sage‐grouse), a galliforme of conservation concern throughout western North America. We monitored female reproductive activity from 1998–2010 and used generalized linear mixed models incorporating within‐subject centering to evaluate and separate within‐ and between‐individual effects. We detected positive effects of within‐individual variation on nest initiation and success where ageing increased the likelihood of both parameters, which appears to support the breeding experience and/or foraging ability hypotheses. However, nest initiation was also affected by between‐individual variation whereby the likelihood of initiation was higher for individuals with higher mean age (i.e. survived longer), as is predicted by the frailty and selection hypotheses. Our results indicate both within‐ and between‐individual variation affect reproductive output of sage‐grouse, but the effects of each varied by measure of reproductive output. Our results corroborate previous findings that suggest population age parameters (i.e. cross‐sectional) should be interpreted with caution due to potential entanglement of within‐ and between‐individual processes. Moreover, the relative role and strength of within‐ and between‐individual processes appeared to vary by measure of reproductive output in our results, which further emphasizes the need for longitudinal analysis of age effects, even in relatively short‐lived iteroparous animals, to adequately interpret biological processes.     

Homoeostatic maintenance of nonstructural carbohydrates during the 2015–2016 El Niño drought across a tropical forest precipitation gradient

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.