Branta Bernicla Nigricans Research Articles

Predicting the response of a long-distance migrant to changing environmental conditions in winter.

Access to high-quality food is critical for long-distance migrants to provide energy for migration and arrival at breeding grounds in good condition. We studied effects of changing abundance and availability of a marine food, common eelgrass (Zostera marina L.), on an arctic-breeding, migratory goose, black brant (Brant bernicla nigricans Lawrence 1846), at a key non-breeding site, Bahía San Quintín, Mexico. Eelgrass, the primary food of brant, is consumed when exposed by the tide or within reach from the water's surface. Using an individual-based model, we predicted effects of observed changes (1991-2013) in parameters influencing food abundance and availability: eelgrass biomass (abundance), eelgrass shoot length (availability, as longer shoots more within reach), brant population size (availability, as competition greater with more birds), and sea level (availability, as less food within reach when sea level higher). The model predicted that the ability to gain enough energy to migrate was most strongly influenced by eelgrass biomass (threshold January biomass for migration = 60 g m-2 dry mass). Conversely, annual variation in population size (except for 1998), was relatively low, and variation in eelgrass shoot length and sea level were not strongly related to ability to migrate. We used observed data on brant body mass at Bahía San Quintín and annual survival to test for effects of eelgrass biomass in the real system. The lowest observed values of body mass and survival were in years when biomass was below 60 g m-2, although in some years of low biomass body mass and/or survival was higher. This suggests that the real birds may have some capacity to compensate to meet their energy demands when eelgrass biomass is low. We discuss consequences for brant population trends and conservation.

Use of amino acid isotope analysis to investigate capital versus income breeding strategies in migratory avian species

Abstract Income and capital breeding represent opposing ends of a continuum of reproductive strategies. Quantifying nutrient allocation to reproduction is challenging, but recent advances in compound‐specific stable isotope analysis hold promise for tracing the source of individual compounds allocated to reproduction. Here, we describe a novel approach of using measured carbon (δ13C) and nitrogen (δ15N) isotope values of individual amino acids (AAs) in pectoral muscle of egg‐laying females and egg yolk as a useful tool to quantify the reliance on income versus capital breeding in migrating species. We used white‐fronted (Anser albifrons frontalis), lesser snow (A. caerulescens caerulescens) and black brant (Branta bernicla nigricans) geese breeding in tundra ecosystems of northern Alaska as model organisms. All three species relied on mixed capital–income breeding strategies, but models based on AA isotope data estimated higher proportional contributions of endogenous resources to yolk synthesis compared to results based on bulk tissue isotope analyses. Tracing income versus capital nutrient allocation in migratory species at the compound level is a major advance from the current ‘elemental’ perspective obtained from bulk tissue stable isotope analyses. Our framework is applicable to all taxonomic groups, as long as there is a sufficient spatial or temporal isotopic gradient between resources obtained during the breeding and non‐breeding periods.

Comparison of Indices to Infer Population Dynamics of Black Brant

Abstract To aid managers in assessing status of Pacific black brant Branta bernicla nigricans (hereafter brant), I examined pre-existing long-term data series from summer, fall staging, and wintering areas to infer overall population processes and assessed the utility of the various data sources. Variation in demographic parameters measured in subarctic and Arctic locations suggests some form of metapopulation structure likely exists for brant. I used serial autocorrelation coefficients to assess the ability of various indices to track population processes. Based on this approach, the Lincoln–Petersen estimator and the fall aerial survey estimate partitioned using age ratios of staging brant at Izembek Lagoon, Alaska, appear to be the best indicators. However, these two indexes show different trends for the overall brant population. The Lincoln–Petersen estimates showed biologically implausible changes in size among sequential years, whereas the fall Izembek index did not. Annual estimates of survival and productivity fit the patterns of annual variation in the fall Izembek index better than the Lincoln–Petersen estimates. I conclude that the fall age–partitioned Izembek Lagoon index appears to be the best for tracking population processes in brant.

Do hunters target auxiliary markers? An example using black brant

ABSTRACTAuxiliary markers play an essential role in understanding migration, movement, demography, and behavior of migratory birds. Use of such markers relies on the assumption that the markers do not affect the traits of interest. Neck collars, among the most conspicuous of markers, substantially affect risk of harvest, and survival even in the absence of harvest. Effects of less‐conspicuous markers, such as colored plastic tarsal bands, are not well understood. We used 30 years (1986–2015) of banding, recovery, and recapture data from the Yukon‐Kuskokwim Delta in Alaska, USA, to assess differences in direct band recovery rates (DRRs) between black plastic and brightly colored plastic bands applied to black brant (Branta bernicla nigricans). We also assessed the effect of the color of plastic tarsal bands on annual survival, risks of natural mortality harvest, and fidelity to the breeding colony of adult female black brant. When assessing only DRRs we found that brightly colored bands were recovered at higher rates than black plastic bands in the early 2000s, but DRRs for black bands increased more rapidly through time, resulting in similar DRRs for the 2 band colors at the end of the study. Using a Burnham model structure, our results demonstrated that individuals fitted with colored bands had slightly lower hazards of dying from natural causes or hunting than individuals carrying less‐conspicuous black tarsal bands. Differences on annual probability scales were small and credible intervals broadly overlapped between band types, indicating minimal differences between individuals with different band types; however, we could not resolve all confounding in our study design and we suggest that specific studies directed at assessing marker effects are warranted. We encourage education of hunters about their roles as citizen scientists and the potentially detrimental effect of targeting birds with auxiliary markers.

Life-history attributes of Arctic-breeding birds drive uneven responses to environmental variability across different phases of the reproductive cycle.

Animals exhibit varied life‐history traits that reflect adaptive responses to their environments. For Arctic‐breeding birds, traits related to diet, egg nutrient allocation, clutch size, and chick growth are predicted to be under increasing selection pressure due to rapid climate change and increasing environmental variability across high‐latitude regions. We compared four migratory birds (black brant [Branta bernicla nigricans], lesser snow geese [Chen caerulescens caerulescens], semipalmated sandpipers [Calidris pusilla], and Lapland longspurs [Calcarius lapponicus]) with varied life histories at an Arctic site in Alaska, USA, to understand how life‐history traits help moderate environmental variability across different phases of the reproductive cycle. We monitored aspects of reproductive performance related to the timing of breeding, reproductive investment, and chick growth from 2011 to 2018. In response to early snowmelt and warm temperatures, semipalmated sandpipers advanced their site arrival and bred in higher numbers, while brant and snow geese increased clutch sizes; all four species advanced their nest initiation dates. During chick rearing, longspur nestlings were relatively resilient to environmental variation, whereas warmer temperatures increased the growth rates of sandpiper chicks but reduced growth rates of snow goose goslings. These responses generally aligned with traits along the capital‐income spectrum of nutrient acquisition and altricial–precocial modes of chick growth. Under a warming climate, the ability to mobilize endogenous reserves likely provides geese with relative flexibility to adjust the timing of breeding and the size of clutches. Higher temperatures, however, may negatively affect the quality of herbaceous foods and slow gosling growth. Species may possess traits that are beneficial during one phase of the reproductive cycle and others that may be detrimental at another phase, uneven responses that may be amplified with future climate warming. These results underscore the need to consider multiple phases of the reproductive cycle when assessing the effects of environmental variability on Arctic‐breeding birds.

Individual heterogeneity in fitness in a long-lived herbivore.

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.

Simulation modeling accounts for uncertainty while quantifying ecological effects of development alternatives

AbstractWildlife management often involves trade‐offs between protecting species and allowing human activities and development. Ideally, these decisions are guided by scientific studies that quantify the impacts of proposed actions on the environment. However, critical information to assess impacts of proposed activities may be lacking, such as certainty in where actions will take place, which may hinder a robust impact assessment. To address this issue, we present the Development Impacts Analysis (DIA), which employs Monte Carlo simulation modeling to quantify the environmental consequences of proposed development scenarios, while accounting for uncertainty in the exact location of future development. We applied the DIA to five proposed oil leasing management scenarios under a revised management plan for the National Petroleum Reserve—Alaska. For each management scenario with differing levels of proposed development (“alternatives”), oil production pads and roads were randomly simulated in proportion to estimated undiscovered oil and following alternative‐specific restrictions. We assessed habitat displacement for two caribou (Rangifer tarandus) herds, eight shorebird species, and black brant (Branta bernicla) based on reported responses to development, repeating the process 100 times for each alternative. Some habitat loss was reported for each proposed alternative, but the amount of impact varied by alternative and species. One caribou herd and most bird species indicated greatest effects in the alternative with the least restrictions on development and lesser impacts under more protective alternatives. Our results emphasized the importance of considering spatial variation in development effects and species‐specific differences when evaluating management proposals. The DIA quantified potential impacts on a suite of species under proposed management alternatives, while accounting for uncertainty in where development will occur and providing confidence intervals on estimated impacts. This illustrates that uncertainty need not preclude management decisions about establishment of broad land use restrictions prior to submission of project‐level proposals but can instead be explicitly incorporated into decision making. While no single management approach will likely benefit all species, use of tools such as the DIA allows managers to quantify trade‐offs among species and pursue approaches that balance the needs of various taxa and other management objectives.

Predicting impacts of food competition, climate, and disturbance on a long‐distance migratory herbivore

AbstractClimate change is driving worldwide shifts in the distribution of biodiversity, and fundamental changes to global avian migrations. Some arctic‐nesting species may shorten their migration distance as warmer temperatures allow them to winter closer to their high‐latitude breeding grounds. However, such decisions are not without risks, since this intensifies pressure on resources when they are used for greater periods of time. In this study, we used an individual‐based model to predict how future changes in food abundance, winter ice coverage, and human disturbance could impact an Arctic/sub‐Arctic breeding goose species, black brant (Branta bernicla nigricans, Lawrence 1846), and their primary food source, common eelgrass (Zostera marina L.), at the Izembek Lagoon complex in southwest Alaska. Brant use the site during fall and spring migrations, and increasingly, for the duration of winter. The model was validated by comparing predictions to empirical observations of proportion of geese surviving, proportion of geese emigrating, mean duration of stay, mean rate of mass gain/loss, percentage of time spent feeding, number of bird days, peak population numbers, and distribution across the complex. The model predicted that reductions >50% of the current decadal (2007–2015) mean of eelgrass biomass, which have been observed in some years, or increases in the number of brant, could lead to a reduction in the proportion of birds that successfully migrate to their breeding grounds from the site. The model also predicted that access to eelgrass in lagoons other than Izembek was critical for overwinter survival and spring migration of brant, if overall eelgrass biomass was 50% of the decadal mean biomass. Geese were typically predicted to be more vulnerable to environmental change during winter and spring, when eelgrass biomass is lower, and thermoregulatory costs for the geese are higher than in fall. We discuss the consequences of these predictions for goose population trends in the face of natural and human drivers of change.

Mate fidelity improves survival and breeding propensity of a long-lived bird.

Evolutionary and behavioural ecologists have long been interested in factors shaping the variation in mating behaviour observed in nature. Although much of the research on this topic has focused on the consequences of mate choice and mate change on annual reproductive success, studies of a potential positive link between mate fidelity and adult demographic rates have been comparatively rare. This is particularly true for long-lived birds with multi-year, socially monogamous pair bonds. We used a 26-year capture-mark-recapture dataset of 3,330 black brent Branta bernicla nigricans to test whether breeding with a familiar mate improved future breeding propensity and survival. We predicted that experienced breeders nesting with a new partner would have rates of survival similar to familiar pairs because long-lived species avoid jeopardizing survival since their lifetime fitness is sensitive to this vital rate. In contrast, we expected that any costs of breeding with a new partner would be paid through skipping the subsequent breeding attempt. We found that unfamiliar pairs had lower subsequent breeding propensity than faithful partners. However, contrary to our expectations, individuals breeding with a new mate also suffered reduced survival. These results add to a small number of studies indicating that a positive relationship between mate retention and adult demographic rates may exist in a diverse array of avian species. Given these results, researchers should consider costs of mate change that extend beyond within-season reproductive success to fully understand the potential adaptive basis for perennial social monogamy. We caution that if mate retention enhances survival prospects, improvements in annual reproductive success with pair-bond length could be a secondary factor favouring perennial social monogamy, particularly in species with slower life-history strategies. Furthermore, some cases where annual reproductive success does not improve with pair-bond duration, yet multi-year pair bonds are common, could be explained by benefits afforded by mate fidelity to adult vital rates.

Long‐term research and hierarchical models reveal consistent fitness costs of being the last egg in a clutch

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.

Constraint versus restraint in the body mass dynamics during moult of a species with precocial young

Body mass declines during wing moult in numerous, but not all, populations of Anatidae. We assessed two leading hypotheses for body mass dynamics during wing moult: (1) body mass dynamics are adapted to attain a target body mass at the end of wing moult (restraint hypothesis) vs. (2) body mass dynamics reflect environmental constraint on the nutrient–energy balance during wing moult (constraint hypothesis). We used regressions of mass of breeding female Black Brant Branta bernicla nigricans on ninth primary length (a measure of moult stage) for each of 16 years to assess mass dynamics during wing moult and used regression equations to predict mass at the beginning and end of wing moult each year. We also included gosling mass at 30 days (an indicator of forage availability) in models of adult mass to assess how mass dynamics varied as a function of foraging conditions. Predicted body mass (± 95% CI) at the start of wing moult (ninth primary = 0 mm) varied significantly among years from 1032 ± 52 to 1169 ± 27 g. Similarly, predicted mass in late wing moult (ninth primary = 142 mm) ranged from 1048 ± 25 to 1222 ± 28 g. The rate of mass gain was significantly related to gosling mass at 30 days: interaction between adult ninth primary length and gosling mass = 0.0031 ± 0.0020 (P = 0.003). Females initiated wing moult at lower body masses, gained mass more rapidly and ended with wing moult heaviest when goslings were heaviest. Body mass dynamics of female Black Brant during wing moult were consistent with the constraint hypothesis. The positive association between gosling mass and rate of body mass gain by adult females during wing moult was also consistent with the constraint hypothesis.

Dynamics of Dispersed-Nesting Black Brant on the Yukon-Kuskokwim Delta

Abstract The number of black brant Branta bernicla nigricans nests within major breeding colonies on the Yukon-Kuskowkim Delta has declined since 1992. It has been hypothesized that these declines are partially explained by increased numbers of black brant breeding outside of these colonies. To assess this hypothesis, we analyzed spatiotemporal patterns in numbers of black brant nests occurring outside major colonies. Nesting densities of black brant vary among three strata: 1) peripheral to major colonies, 2) other coastal habitats, and 3) inland habitats. We sampled some substrata within each stratum type only between 1986 and 1999 (historical strata), whereas we sampled others over the period 1986–2016 (long-term strata). We fit regression models with number of nests on a plot as a response variable, a log link, and year as the explanatory variable. We allowed each stratum (e.g., historical peripheral) to have its own intercept to account for variation in mean nest density but constrained linear and quadratic regression coefficients to be the same for strata in similar habitats (e.g., historical and long-term peripheral). We used a negative binomial distribution for nests to allow for substantial variation in nests per plot. We fit models using Markov Chain Monte Carlo methods in JAGS. Ninety-five percent credible intervals for both linear and quadratic coefficients for the peripheral and coastal strata, where most nests occurred, broadly overlapped zero, indicating modest trends in numbers of nests in these strata. We estimated there were 6,584 (95% credible interval: 4,221−11,269) dispersed nests in 1986, increasing to 11,051 (95% credible interval: 7,450−17,460) in 2016. Our results indicated that increases in dispersed nests were unable to replace declines in colony nests. Furthermore, quadratic trends indicated that potential earlier annual increases in dispersed nests have declined to zero. We conclude that total numbers of black brant nests on the Yukon-Kuskokwim Delta are likely declining, consistent with the trend in fall age ratios over the same period. Uncertainty about trends in areas not sampled since 1994 adds to the uncertainty about the precise magnitude of the decline. We recommend that the area sampled by the random plot program be expanded to include strata sampled only before 1995.

Early Goose Arrival Increases Soil Nitrogen Availability More Than an Advancing Spring in Coastal Western Alaska

An understudied aspect of climate change-induced phenological mismatch is its effect on ecosystem functioning, such as nitrogen (N) cycling. Migratory herbivore arrival time may alter N inputs and plant–herbivore feedbacks, whereas earlier springs are predicted to increase N cycling rates through warmer temperatures. However, the relative importance of these shifts in timing and how they interact to affect N cycling are largely unknown. We conducted a 3-year factorial experiment in coastal western Alaska that simulated different timings of Pacific black brant (Branta bernicla nigricans) arrival (3 weeks early, typical, 3 weeks late, or no-grazing) and the growing season (ca. 3 weeks advanced and ambient) on adsorbed and mobile inorganic (NH4+–N, NO3−–N) and mobile organic N (amino acid) pools. Early grazing increased NH4+–N, NO3−–N, and amino acids by 103%, 119%, and 7%, respectively, whereas late grazing reduced adsorbed NH4+–N and NO3−–N by 16% and 17%, respectively. In comparison, the advanced growing season increased mobile NH4+–N by 26%. The arrival time by geese and the start of the season did not interact to influence soil N availability. While the onset of spring in our system is advancing at twice the rate of migratory goose arrival, earlier goose migration is likely to be more significant than the advances in springs in influencing soil N, although both early goose arrival and advanced springs are likely to increase N availability in the future. This increase in soil N resources can have a lasting impact on plant community composition and productivity in this N-limited ecosystem.

Phenological mismatch between season advancement and migration timing alters Arctic plant traits

Abstract Climate change is creating phenological mismatches between herbivores and their plant resources throughout the Arctic. While advancing growing seasons and changing arrival times of migratory herbivores can have consequences for herbivores and forage quality, developing mismatches could also influence other traits of plants, such as above‐ and below‐ground biomass and the type of reproduction, that are often not investigated. In coastal western Alaska, we conducted a 3‐year factorial experiment that simulated scenarios of phenological mismatch by manipulating the start of the growing season (3 weeks early and ambient) and grazing times (3 weeks early, typical, 3 weeks late, or no‐grazing) of Pacific black brant (Branta bernicla nigricans), to examine how the timing of these events influence a primary goose forage species, Carex subspathacea. After 3 years, an advanced growing season compared to a typical growing season increased stem heights, standing dead biomass, and the number of inflorescences. Early season grazing compared to typical season grazing reduced above‐ and below‐ground biomass, stem height, and the number of tillers; while late season grazing increased the number of inflorescences and standing dead biomass. Therefore, an advanced growing season and late grazing had similar directional effects on most plant traits, but a 3‐week delay in grazing had an impact on traits 3–5 times greater than a similarly timed shift in the advancement of spring. In addition, changes in response to treatments for some variables, such as the number of inflorescences, were not measurable until the second year of the experiment, while other variables, such as root productivity and number of tillers, changed the direction of their responses to treatments over time. Synthesis. Factors affecting the timing of migration have a larger influence than earlier springs on an important forage species in the breeding and rearing habitats of Pacific black brant. The phenological mismatch prediction for this site of earlier springs and later goose arrival will likely increase above‐ and below‐ground biomass and sexual reproduction of the often‐clonally reproducing C. subspathacea. Finally, the implications of mismatch may be difficult to predict because some variables required successive years of mismatch to respond.

Changes in behavior are unable to disrupt a trophic cascade involving a specialist herbivore and its food plant.

Changes in ecological conditions can induce changes in behavior and demography of wild organisms, which in turn may influence population dynamics. Black brant (Branta bernicla nigricans) nesting in colonies on the Yukon–Kuskokwim Delta (YKD) in western Alaska have declined substantially (~50%) since the turn of the century. Black brant are herbivores that rely heavily on Carex subspathacea (Hoppner's sedge) during growth and development. The availability of C. subspathacea affects gosling growth rates, which subsequently affect pre‐ and postfledging survival, as well as size and breeding probability as an adult. We predicted that long‐term declines in C. subspathacea have affected gosling growth rates, despite the potential of behavior to buffer changes in food availability during brood rearing. We used Bayesian hierarchical mixed‐effects models to examine long‐term (1987–2015) shifts in brant behavior during brood rearing, forage availability, and gosling growth rates at the Tutakoke River colony. We showed that locomotion behaviors have increased (β = 0.05, 95% CRI: 0.032–0.068) while resting behaviors have decreased (β = −0.024, 95% CRI: −0.041 to −0.007), potentially in response to long‐term shifts in forage availability and brood density. Concurrently, gosling growth rates have decreased substantially (β = −0.100, 95% CRI: −0.191 to −0.016) despite shifts in behavior, mirroring long‐term declines in the abundance of C. subspathacea (β = −0.191, 95% CRI: −0.355 to −0.032). These results have important implications for individual fitness and population viability, where shifts in gosling behavior putatively fail to mitigate long‐term declines in forage availability.

Migratory goose arrival time plays a larger role in influencing forage quality than advancing springs in an Arctic coastal wetland.

With warmer springs, herbivores migrating to Arctic breeding grounds may experience phenological mismatches between their energy demands and the availability of high quality forage. Yet, how the timing of the start of the season and herbivore arrival influences forage quality is often unknown. In coastal western Alaska, approximately one million migratory geese arrive each spring to breed, where foliar %N and C:N ratios are linked to gosling survival and population growth. We conducted a three-year experiment where we manipulated the start of the growing season using warming chambers and grazing times using captive Pacific black brant (Branta bernicla nigricans) to examine how the timing of these events influences the quality of an important forage species. Our results suggest that grazing timing plays a much greater role than an advanced growing season in determining forage quality. All top models included grazing timing, and suggested that compared to typical grazing timing, early grazing significantly reduced foliar %C by 6% and C:N ratios by 16%, while late goose grazing significantly reduced foliar %N by 15% and increased foliar C:N ratios by 21%. While second-ranking top models included the effect of season, the advanced growing season effect was not significant and only reduced %N by 4%, increased %C by <1%, and increased C:N ratios by 5% compared to an ambient growing season. In summary, in years where geese arrive early, they will consume higher quality forage when they arrive and throughout the season, while in years that geese arrive late they will consume lower quality forage when they arrive and for the remainder of the season. When the growing season starts has only a minor influence on this pattern. Our findings suggest that cues determining migration and arrival times to breeding areas are important factors influencing forage quality for geese in western Alaska.

Brood Size Affects Future Reproduction in a Long-Lived Bird with Precocial Young.

Estimation of trade-offs between current reproduction and future survival and fecundity of long-lived vertebrates is essential to understanding factors that shape optimal reproductive investment. Black brant geese (Branta bernicla nigricans) fledge more goslings, on average, when their broods are experimentally enlarged to be greater than the most common clutch size of four eggs. Thus, we hypothesized that the lesser frequency of brant clutches exceeding four eggs results, at least partially, from a future reduction in survival, breeding probability, or clutch size for females tending larger broods. We used an 8-year mark-recapture data set (Barker robust design) with 5 years of clutch and brood manipulations to estimate long-term consequences of reproductive decisions in brant. We did not find evidence of a trade-off between reproductive effort and true survival or future initiation date and clutch size. Rather, future breeding probability was maximized ( [SE]) for manipulated females tending broods of four goslings and was lower for females tending smaller (one gosling; [SE]) or larger (seven goslings; [SE]) broods. Our results suggest that demographic trade-offs for female brant tending large broods may reduce the fitness value of clutches larger than four and, therefore, contribute to the paucity of larger clutches. The lack of a trade-off between reproductive effort and survival provides evidence that survival, to which fitness is most sensitive in long-lived animals, is buffered against temporal variation in brant.

The black brant population is declining based on mark recapture

ABSTRACTAnnual survival and recruitment in black brant (Branta bernicla nigricans) have declined since the 1990s, yet aerial surveys of the global population have been stable or even increasing over the past decade. We used a combination of a Lincoln estimator based on harvest information and band recoveries, and marked‐unmarked ratios in bag checks in 1 harvest area in Mexico to estimate the number of adults in the population during 1992–2015. We produced weighted means from the 2 kinds of estimates for years in which we had data for both, with weights equal to the inverse of the variance of the individual estimates. We treated the black brant population as consisting of 2 subpopulations. One population consisted of breeding black brant on the Yukon‐Kuskokwim Delta (YKD), Alaska, USA, and the other consisted of Arctic (northern Alaska, western Canada, and eastern Russia) breeders, and nonbreeders and failed breeders from the YKD that underwent molt migration to the Arctic. For the global population estimates, we assessed potential bias due to differential marking and harvest of the 2 subpopulations, which was approximately 1%, probably because band recovery rates were similar for the 2 subpopulations. Population estimates declined from 229,980 (average for 1999–2002) to 161,504 (average for 2012–2015). Population estimates based on estimated harvest were variable but more stable in the later years of the study, when larger numbers of brant hunters were included in the sample. We suggest that the combination of Lincoln estimates and bag check data provides a reasonable and cost effective approach to monitoring the population. © 2018 The Wildlife Society.

Spatial distribution of band recoveries of black brant

ABSTRACTOn average, band recovery rates of adult black brant (Branta bernicla nigricans) more than doubled between the 2000s and 2010s. However, the spatial distribution of band recoveries of black brant has not been reported. Our objective was to describe the spatial distribution of band recoveries of black brant since 1990. We found that Alaska, California, and Mexico accounted for ≥89% of band recoveries for black brant released from the Arctic and the Tutakoke River Colony in southwestern Alaska, in each decade studied. Although recovery rates increased in each major harvest region from the 1990s to 2010s, increases were not even. For brant released from the Arctic, the recovery rate in Alaska has gone up 7‐fold, whereas the recovery rate in Mexico has increased 1.3‐fold since the 1990s. For brant banded in the Arctic, the increased recovery rates in Alaska resulted largely from a rise in recoveries from the Izembek Lagoon area. For brant banded at the Tutakoke River Colony, the recovery rate in Alaska increased 4.8‐fold, whereas recovery rates in Mexico increased 1.6‐fold. Despite the reduced relative contribution of Mexico to band recoveries in the 2010s, Bahia San Quintin, Mexico, still contributes more recoveries than any other wintering estuary. Because 57–60% of black brant band recoveries occur at the localities of Izembek Lagoon, Alaska; Humboldt Bay, California; and Bahia San Quintin, it is likely that accurate annual estimates of the black brant population using Lincoln's method could be derived from hunter bag checks at these 3 areas. If population managers are interested in managing harvest rates of black brant, they could focus efforts on the main hunting areas we highlight in this paper. Finally, it is unclear if the recent increases in band recovery rates are the result of increased harvest, a declining population, or both; therefore, we suggest refinement of population monitoring programs and continued monitoring of band recovery rates. © 2018 The Wildlife Society.

Delayed herbivory by migratory geese increases summer‐long CO2 uptake in coastal western Alaska

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to "phenological mismatches" as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three-year experiment to examine the consequences for CO2 exchange of advanced spring green-up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green-up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO2 uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green-up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2µmolm-2 s-1 toward a greater CO2 sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO2 uptake. Importantly, while the direct effect of climate warming on phenology of green-up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.