Wildfire is the most impactful disturbance regime in the North American boreal region, driving the structure and composition of forests across the region. Recent climate models predict that increasing fire intensity and frequency will result in a shift from a largely coniferous forest to one with a greater dominance by deciduous species. We investigated how an iconic predator of the boreal system, the Canada lynx ( Lynx canadensis ), moves through a range of burn scars (4–73 years old). Using GPS collars at 4‐h fix rates, we fitted integrated step selection models to lynx movements across an 80‐year post‐fire chronosequence to assess habitat selection in both deciduous and coniferous forests. We predicted that lynx would primarily select intermediately aged spruce and young deciduous stands, mirroring previous research on the habitat selection of their main prey, snowshoe hares ( Lepus americanus ). We found, however, that lynx habitat selection peaked at intermediately aged stands in both forest types, with selection for younger deciduous stands in the winter months. There was no seasonal change in coniferous stands as they experience little change in cover across seasons. We hypothesize that lynx select for habitats that maximize capture probability as opposed to simply habitats with the highest hare density. Together, these results show that lynx can be resilient to short‐term shifts toward intermediate‐aged stands. However, these benefits will likely diminish in the longer term as the decrease in fire return interval may reduce the prevalence of intermediate‐aged stands.
Connectivity is determined by the degree to which inhospitable habitat matrices are permeable to gene flow. Successful gene flow requires that dispersing individuals travel to new populations, survive there, and finally, reproduce. Consequently, examinations of connectivity must consider both the permeability of the landscape and individual survival patterns, as surviving individuals are the only ones that have the potential to reproduce. We investigated functional connectivity of Canada lynx (Lynx canadensis) populations in the northwestern boreal forest using an unprecedented dataset of 207,957 relocations from 142 individual lynx. As a specialist predator of snowshoe hare (Lepus americanus), lynx populations follow a 10- to 12-year population cycle, which lags a few years behind the population cycles of hares. We used integrated step selection analyses to quantify habitat selection, and Bayesian counting process implementation of wrapped Cauchy survival models to evaluate mortality risk during dispersal. We compared movement patterns and survival across behavioral states and population phases, following the a population decline. Our results showed few complete physical barriers to dispersal in the study region, with many lynx dispersing greater than 3000 km. Step-selection analysis of lynx movement revealed a higher tolerance for poor quality habitats in dispersing lynx as compared to resident lynx, primarily in selection for rugged terrain and closer proximity to roads. These differences resulted in a higher estimate of landscape permeability for dispersing lynx compared to residents. When survival patterns were considered, there were increasingly stark differences between resident and dispersing lynx during population declines. Dispersing lynx faced markedly lower survival rates than resident lynx. Overall, this paints a picture of a more connected landscape during population highs, which becomes increasingly fragmented during population lows. Canada lynx populations appeared to have few impermeable barriers across the landscape, but they face periods of low functional connectivity driven by lower survival during population declines. This emphasizes the significance of maintaining high-quality refugia habitats from which population rebounds can occur following cyclic lows.
Cyclical population dynamics are a common phenomenon in populations worldwide, yet the spatial organization of these cycles remains poorly understood. In this study, we investigated the spatial form and timing of a population collapse from 2018 to 2022 in Canada lynx (Lynx canadensis) across the northwest boreal forest. We analyzed survival, reproduction, and dispersal data from 143 individual global positioning system (GPS) collared lynx from populations across five study sites spanning interior Alaska to determine whether lynx displayed characteristics of a population wave following a concurrent wave in snowshoe hare (Lepus americanus) abundance. Reproductive rates declined across the study sites; however, site-level reproduction declined first in our easternmost study sites, supporting the idea of a population wave. Despite a clear increase in percent of dispersing lynx, there was no evidence of directional bias in dispersal following a hare population wave. Analysis did show increasingly poor survival for lynx dispersing to the east compared to combined resident and westward dispersal. This pattern is consistent with a survival-mediated population wave in lynx as the driver of the theorized population wave. The combination of these factors supports the idea of a hierarchical response to snowshoe hare population declines with a drop in lynx reproduction followed by increased dispersal, and finally reduced survival. All of this evidence is consistent with the expected characteristics of a population undergoing a traveling wave and supports the hypothesis that lynx presence may facilitate and mirror the underlying wave patterns in snowshoe hare.
A pilot study for mapping the Arctic wetlands was conducted in the Yukon Flats National Wildlife Refuge (Refuge), Alaska. It included commissioning the HySpex VNIR-1800 and the HySpex SWIR-384 imaging spectrometers in a single-engine Found Bush Hawk aircraft, planning the flight times, direction, and speed to minimize the strong bidirectional reflectance distribution function (BRDF) effects present at high latitudes and establishing improved data processing workflows for the high-latitude environments. Hyperspectral images were acquired on two clear-sky days in early September, 2018, over three pilot study areas that together represented a wide variety of vegetation and wetland environments. Steps to further minimize BRDF effects and achieve a higher geometric accuracy were added to adapt and improve the Hyspex data processing workflow, developed by the German Aerospace Center (DLR), for high-latitude environments. One-meter spatial resolution hyperspectral images, that included a subset of only 120 selected spectral bands, were used for wetland mapping. A six-category legend was established based on previous U.S. Geological Survey (USGS) and U.S. Fish and Wildlife Service (USFWS) information and maps, and three different classification methods—hybrid classification, spectral angle mapper, and maximum likelihood—were used at two selected sites. The best classification performance occurred when using the maximum likelihood classifier with an averaged Kappa index of 0.95; followed by the spectral angle mapper (SAM) classifier with a Kappa index of 0.62; and, lastly, by the hybrid classifier showing lower performance with a Kappa index of 0.51. Recommendations for improvements of future work include the concurrent acquisition of LiDAR or RGB photo-derived digital surface models as well as detailed spectra collection for Alaska wetland cover to improve classification efforts.
ABSTRACT Understanding how species respond to environmental conditions can assist with conservation strategies and harvest management, especially in arctic and boreal regions that are experiencing rapid climate change. Although climatic influences on species distributions have been studied, broad‐scale effects of climate on survival are less well known. We examined the interactive effects of meteorological and remotely sensed environmental variables on survival of Dall's sheep ( Ovis dalli dalli ) lambs and adults by synthesizing radio‐telemetry data across their range. We used data from 9 studies of adult sheep and 2 studies of lambs that were conducted between 1997 and 2012 at sites spanning the species' range in Alaska, USA, and northwestern Canada. We obtained environmental variables throughout the range of Dall's sheep, including the normalized difference vegetation index (NDVI) from optical remote sensing, freeze‐thaw frequency (FTF) from passive microwave remote sensing, and gridded climate variables such as snow water equivalent, temperature, and precipitation. We used Cox proportional hazard regression to investigate the effects of environmental variables recorded during summer, winter, and the previous winter on annual survival rates of Dall's sheep lambs and adults. Summer NDVI was the most influential environmental factor affecting lamb survival, with improved lamb survival occurring in years with a high maximum NDVI. Also, lamb predation by coyotes ( Canis latrans ) and golden eagles ( Aquila chrysaetos ) decreased substantially with increasing NDVI. The previous winter FTF had the strongest effect on adult survival, with decreased survival occurring after winters with high FTF. In addition, these remotely sensed environmental factors interacted with meteorological factors to affect survival, such that effects of winter temperature depended on summer NDVI and winter FTF. Warm winters increased lamb survival only when preceded by summers with high NDVI, and warm winters increased adult survival only when winter FTF was low. Thus, potential benefits of climate warming may be counteracted if wintertime freeze‐thaw events markedly increase. Correlations among environmental variables across sites were low, and regional climate cycles such as the Pacific Decadal Oscillation (PDO) had weak effects, indicating substantial local variability in climatic conditions experienced by Dall's sheep across their range. These findings can help managers anticipate how Dall's sheep populations will respond to changes in local environmental conditions. Our results also highlight the utility of multiple remotely sensed environmental conditions for ungulate management, especially passive microwave products that provide valuable information on winter icing events. © 2020 The Wildlife Society.
Abstract AirSWOT, an experimental airborne Ka‐band interferometric synthetic aperture radar, was developed for hydrologic research and validation of the forthcoming Surface Water and Ocean Topography (SWOT) satellite mission (to be launched in 2021). AirSWOT and SWOT aim to improve understanding of surface water processes by mapping water surface elevation (WSE) and water surface slope (WSS) in rivers, lakes, and wetlands. However, the utility of AirSWOT for these purposes remains largely unexamined. We present the first investigation of AirSWOT WSE and WSS surveys over complex, low‐relief, wetland‐river hydrologic environments, including (1) a field‐validated assessment of AirSWOT WSE and WSS precisions for lakes and rivers in the Yukon Flats Basin, an Arctic‐Boreal wetland complex in eastern interior Alaska; (2) improved scientific understanding of surface water flow gradients and the influence of subsurface permafrost; and (3) recommendations for improving AirSWOT precisions in future scientific and SWOT validation campaigns. AirSWOT quantifies WSE with an RMSE of 8 and 15 cm in 1 and 0.0625 km2 river reaches, respectively, and 21 cm in lakes. This indicates good utility for studying hydrologic flux, WSS, geomorphic processes, and coupled surface/subsurface hydrology in permafrost environments. This also suggests that AirSWOT supplies sufficient precision for validating SWOT WSE and WSS over rivers, but not lakes. However, improvements in sensor calibration and flight experiment design may improve precisions in future deployments as may modifications to data processing. We conclude that AirSWOT is a useful tool for bridging the gap between field observations and forthcoming global SWOT satellite products.
Shrinking lakes were recently observed for several Arctic and Subarctic regions due to increased evaporation and permafrost degradation. Along with lake drawdown, these processes often boost aquatic chemical concentrations, potentially impacting trophic dynamics. In particular, elevated chemical levels may impact primary productivity, which may in turn influence populations of primary and secondary consumers. We examined trophic dynamics of 18 shrinking lakes of the Yukon Flats, Alaska, that had experienced pronounced increases in nutrient (>200 % total nitrogen, >100 % total phosphorus) and ion concentrations (>100 % for four major ions combined) from 1985–1989 to 2010–2012, versus 37 stable lakes with relatively little chemical change over the same period. We found that phytoplankton stocks, as indexed by chlorophyll concentrations, remained unchanged in both shrinking and stable lakes from the 1980s to 2010s. Moving up the trophic ladder, we found significant changes in invertebrate abundance across decades, including decreased abundance of five of six groups examined. However, these decadal losses in invertebrate abundance were not limited to shrinking lakes, occurring in lakes with stable surface areas as well. At the top of the food web, we observed that probabilities of lake occupancy for ten waterbird species, including adults and chicks, remained unchanged from the period 1985–1989 to 2010–2012. Overall, our study lakes displayed a high degree of resilience to multi-trophic cascades caused by rising chemical concentrations. This resilience was likely due to their naturally high fertility, such that further nutrient inputs had little impact on waters already near peak production.
ABSTRACT Identification of ecological factors that drive animal distributions allows us to understand why distributions vary temporally and spatially, and to develop models to predict future changes to populations–vital tools for effective wildlife management and conservation. For waterbird broods in the boreal forest, distributions are likely driven by factors affecting quality of nesting and brood‐rearing habitats, and the influence of these factors may extend beyond singles species, affecting the entire waterbird community. We used occupancy models to assess factors influencing species richness of waterbird broods on 72 boreal lakes, along with brood distributions of 3 species of conservation concern: lesser scaup ( Aythya affinis ), white‐winged scoters ( Melanitta fusca ), and horned grebe ( Podiceps auritus ). Factors examined included abundance of invertebrate foods (Amphipoda, Diptera, Gastropoda, Hemiptera, Odonata), physical lake attributes (lake area, emergent vegetation), water chemistry (nitrogen, phosphorus, chlorophyll a concentrations), and nesting habitats (water edge, non‐forest cover). Of the 5 invertebrates, only amphipod density was related to richness and occupancy, consistently having a large and positive relationship. Despite this importance to waterbirds, amphipods were the most patchily distributed invertebrate, with 17% of the study lakes containing 70% of collected amphipods. Lake area was the only other covariate that strongly and positively influenced species richness and occupancy of scaup, scoters, and grebes. All 3 water chemistry covariates, which provided alternative measures of lake productivity, were positively related to species richness but had little effect on scaup, scoter, and grebe occupancy. Conversely, emergent vegetation was negatively related to richness, reflecting avoidance of overgrown lakes by broods. Finally, nesting habitats had no influence on richness and occupancy, indicating that, at a broad spatial scale, brood distributions are largely driven by the presence of quality brood‐rearing lakes, not nesting habitats. Our findings are relevant to generating conservation plans or management goals; specifically, boreal lakes with abundant amphipods and surface areas >25 ha are important habitat for waterbird broods and merit conservation, especially given the patchy distribution of amphipods. Moreover, these high quality brood‐rearing lakes are much rarer, and thus more constraining, than are quality nesting habitats, which are likely abundant in the boreal. © 2015 The Wildlife Society.
1 Hyperspectral Imaging Laboratory (HyLab), Geophysical Institute (GI), University of Alaska Fairbanks (UAF), 903 Koyukuk Dr., Fairbanks, AK 99775-7320, USA; 2 Alaska Geobotany Center (AGC), Institute fo Arctic Biology (IAB), University of Alaska Fairbanks (UAF), 902 N. Koyukuk Dr., Fairbanks, AK 99775-7000, USA; 3 Spectroscopy Laboratory, U.S. Geological Survey, Denver Federal Center, Denver, CO 80225, USA; 4 Mineral Resources Section, Alaska Division of Geological & Geophysical Surveys, Fairbanks, AK 99709, USA; 5 Yukon Flats National Wildlife Refuge, U.S. Fish & Wildlife Service, Fairbanks, AK 99701, USA.
We investigated the winter density and territory size of wolves ( Canis lupus ) on the Yukon Flats, Alaska, where moose ( Alces alces ) was the sole ungulate prey, occurring at a low density and representing a biomass of ungulate food lower than previously studied in North America. Using locations (GPS coordinates) from collars deployed on seven wolves, we estimated territory sizes with adaptive kernel and minimum convex polygon methods. We then estimated wolf density from a population area defined by these territory sizes and counts of wolves in five marked packs. From November 2009 to April 2010, we obtained 6263 GPS locations. Pack size ranged from two to 10 wolves, with average size of 5.0 in November 2009 and 4.8 in March 2010. Average winter territory size for five packs was 1433 km2 with the 95% adaptive kernel method and 1608 km2 with the minimum convex polygon method. Density (wolves/1000 km2) was 3.6 in November and 3.4 in March with the 95% adaptive kernel method and 3.4 in both November and March with the minimum convex polygon method. Territories were large and estimates produced by the two methods differed by 11%. Densities were low, and the two analysis methods yielded densities that differed from each other by 3% to 6%. Low wolf density corresponded with low biomass of ungulate food, suggesting that moose availability on the Yukon Flats likely limited wolf density.
Fires are the major natural disturbance in the boreal forest, and their frequency and intensity will likely increase as the climate warms. Terrestrial nutrients released by fires may be transported to boreal lakes, stimulating increased primary productivity, which may radiate through multiple trophic levels. Using a before-after-control-impact (BACI) design, with pre- and postfire data from burned and unburned areas, we examined effects of a natural fire across several trophic levels of boreal lakes, from nutrient and chlorophyll levels, to macroinvertebrates, to waterbirds. Concentrations of total nitrogen and phosphorus were not affected by the fire. Chlorophyll a levels were also unaffected, likely reflecting the stable nutrient concentrations. For aquatic invertebrates, we found that densities of three functional feeding groups did not respond to the fire (filterers, gatherers, scrapers), while two groups increased (shredders, predators). Amphipods accounted for 98% of shredder numbers, and we hypothesize that fire-mediated habitat changes may have favored their generalist feeding and habitat ecology. This increase in amphipods may, in turn, have driven increased predator densities, as amphipods were the most numerous invertebrate in our lakes and are commonly taken as prey. Finally, abundance of waterbird young, which feed primarily on aquatic invertebrates, was not affected by the fire. Overall, ecosystems of our study lakes were largely resilient to forest fires, likely due to their high initial nutrient concentrations and small catchment sizes. Moreover, this resilience spanned multiple trophic levels, a significant result for ecologically similar boreal regions, especially given the high potential for increased fires with future climate change.
Losses in lake area have been observed for several Arctic and Subarctic regions in recent decades, with unknown consequences for lake ecosystems. These reductions are primarily attributed to two climate‐sensitive mechanisms, both of which may also cause changes in water chemistry: (i) increased imbalance of evaporation relative to inflow, whereby increased evaporation and decreased inflow act to concentrate solutes into smaller volumes; and (ii) accelerated permafrost degradation, which enhances sublacustrine drainage while simultaneously leaching previously frozen solutes into lakes. We documented changes in nutrients [total nitrogen ( TN ), total phosphorus ( TP )] and ions (calcium, chloride, magnesium, sodium) over a 25 year interval in shrinking, stable, and expanding Subarctic lakes of the Yukon Flats, Alaska. Concentrations of all six solutes increased in shrinking lakes from 1985–1989 to 2010–2012, while simultaneously undergoing little change in stable or expanding lakes. This created a present‐day pattern, much weaker or absent in the 1980s, in which shrinking lakes had higher solute concentrations than their stable or expanding counterparts. An imbalanced evaporation‐to‐inflow ratio (E/I) was the most likely mechanism behind such changes; all four ions, which behave semiconservatively and are prone to evapoconcentration, increased in shrinking lakes and, along with TN and TP , were positively related to isotopically derived E/I estimates. Moreover, the most conservative ion, chloride, increased >500% in shrinking lakes. Conversely, only TP concentration was related to probability of permafrost presence, being highest at intermediate probabilities. Overall, the substantial increases of nutrients ( TN >200%, TP >100%) and ions (>100%) may shift shrinking lakes towards overly eutrophic or saline states, with potentially severe consequences for ecosystems of northern lakes.
Wolf (Canis lupus) kill rates are fundamental to understanding predation, but are not well known at low moose (Alces alces) densities. We investigated kill rates of 6 wolf packs (2-10 wolves/pack) during 2 winters on the Yukon Flats, a region of eastern Interior Alaska where moose were the sole ungulate prey of wolves occurring at densities <0.2moose/km(2). Our objectives were to compare kill rates with those from areas of greater moose densities, and to determine potential trends in kill rates across the winter. We located moose killed by wolves in February-March 2009, and November 2009-March 2010 using aerial tracking techniques and global positioning system (GPS) location clusters. Wolves killed more moose in early than late winter ((MONTH)=-0.02 moose/pack/day, 95% CI=-0.01 to -0.04), and kill rate estimates (mean, 95% CI) were greatest in November (0.033moose/wolf/day, 0.011-0.055) and least in February (0.011, 0.002-0.02). Kill rates were similar between February and March 2009 (0.019moose/wolf/day, 0.01-0.03) and 2010 (0.018, 0.01-0.03). Prey composition was primarily adult females (39%) and young-of-the-year (35%). We attribute an elevated kill rate in early winter to predation on more vulnerable young-of-the-year. Kill rates in our study were similar to those from other studies where moose occurred at greater densities. We suggest that very few, if any, wolf-moose systems in Alaska and the Yukon experience a density-dependent phase in the functional response, and instead wolves respond numerically to changes in moose density or availability in the absence of alternative prey. Through a numerical response, wolf predation rates may approximate the annual growth potential of the moose population, contributing to persistent low densities of moose and wolves on the Yukon Flats. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.
The movement and transmission of avian influenza viral strains via wild migratory birds may vary by host species as a result of migratory tendency and sympatry with other infected individuals. To examine the roles of host migratory tendency and species sympatry on the movement of Eurasian low‐pathogenic avian influenza (LPAI) genes into North America, we characterized migratory patterns and LPAI viral genomic variation in mallards ( Anas platyrhynchos ) of Alaska in comparison with LPAI diversity of northern pintails ( Anas acuta ). A 50‐year band‐recovery data set suggests that unlike northern pintails, mallards rarely make trans‐hemispheric migrations between Alaska and Eurasia. Concordantly, fewer (14.5%) of 62 LPAI isolates from mallards contained Eurasian gene segments compared to those from 97 northern pintails (35%), a species with greater inter‐continental migratory tendency. Aerial survey and banding data suggest that mallards and northern pintails are largely sympatric throughout Alaska during the breeding season, promoting opportunities for interspecific transmission. Comparisons of full‐genome isolates confirmed near‐complete genetic homology (>99.5%) of seven viruses between mallards and northern pintails. This study found viral segments of Eurasian lineage at a higher frequency in mallards than previous studies, suggesting transmission from other avian species migrating inter‐hemispherically or the common occurrence of endemic Alaskan viruses containing segments of Eurasian origin. We conclude that mallards are unlikely to transfer Asian‐origin viruses directly to North America via Alaska but that they are likely infected with Asian‐origin viruses via interspecific transfer from species with regular migrations to the Eastern Hemisphere.
A study to estimate the kill rate of wolves on moose was initiated in eastern interior Alaska. This study is the first to examine kill rates in a system with such a low prey density (0.08 moose/km) and the presence of only a single prey species. Wolves were radio tracked daily during early February and March to locate kills. The estimated kill rate was 0.019 moose/wolf/day (95% CI: 0.011-0.028). This estimate was intermediary relative to previous work. The management implications of this result are discussed.
Information on the ecology of waterfowl breeding in the boreal forest is lacking, despite the boreal region's importance to continental waterfowl populations and to duck species that are currently declining, such as lesser scaup (Aythya affinis). We estimated breeding probability and breeding season survival of female lesser scaup on the Yukon Flats National Wildlife Refuge, Alaska, USA, in 2005 and 2006. We captured and marked 93 lesser scaup with radiotransmitters during prelaying and nesting periods. Although all marked lesser scaup females were paired throughout prelaying and incubation periods, we estimated breeding probability over both years as 0.12 (SE = 0.05, n = 67) using telemetry. Proportion of lesser scaup females undergoing rapid follicle growth at capture in 2006 was 0.46 (SE = 0.11, n = 37), based on concentration of yolk precursors in blood plasma. By combining methods based on telemetry, yolk precursors, and postovulatory follicles, we estimated maximum breeding probability as 0.68 (SE = 0.08, n = 37) in 2006. Notably, breeding probability was positively related to female body mass. Survival of female lesser scaup during the nesting and brood-rearing periods was 0.92 (SE = 0.05) in 2005 and 0.86 (SE = 0.08) in 2006. Our results suggest that breeding probability is lower than expected for lesser scaup. In addition, the implicit assumption of continental duck-monitoring programs that all paired females attempt to breed should be reevaluated. Recruitment estimates based on annual breeding-pair surveys may overestimate productivity of scaup pairs in the boreal forest. (JOURNAL OF WILDLIFE MANAGEMENT 73(6): 914-923; 2009)
Over the last 20 years scaup numbers have declined, and these declines have been greatest in the northern boreal forests of Canada and Alaska where most lesser scaup (Aythya affinis) nest. We studied nest success and duckling survival of lesser scaup over 3 field seasons, 2001-2003, on the Yukon Flats National Wildlife Refuge in northeastern Alaska, USA. Daily survival rate (DSR) of nests on our study area across all 3 years was 0.943 (n = 177 nests, 95% CI: 0.930-0.954), corresponding to a nest success of only 12.3%, considerably lower than published estimates of an average nest success as high as 57% for lesser scaup in the northern boreal forest. With Mayfield logistic regression, we investigated effects on nest survival of year, clutch initiation date, and nesting habitat type (large wetlands > 10 ha, small wetlands < 10 ha, and wooded creeks). Neither year nor clutch initiation date influenced nest survival; however, the odds of nest success on large wetlands was 49% lower than on wooded creeks (odds ratio = 0.512, 95% CI = 0.286, 0.918). Based on the model that used only habitat type for estimation, DSR on large wetlands was 0.931 (corresponding nest success = 7.6%), DSR on small wetlands was 0.941 (nest success = 11.1%), and DSR on wooded creeks was 0.963 (nest success = 26.2%). To estimate duckling survival, we monitored 10 broods (n = 75 ducklings) over 3 field seasons by radiotagging hens at nest hatch. Most duckling mortality (94%) occurred in the first 10 days after hatch. Average duckling survival during 1-10 days was 0.321 (95% CI: 0.122-0.772), during 11-20 days was 0.996 (95% CI: 0.891-1.040), and during 21-30 days was 0.923 (95% CI: 0.769-1.041). Three of 10 hens moved all or part of their broods overland between nesting and brood-rearing wetlands for distances of 0.3-1.6 km. Our estimates of lesser scaup nest success and duckling survival on the Yukon Flats were among the lowest ever reported for ducks nesting at northern latitudes, even though the study site was in pristine boreal forest. Estimating and comparing scaup demographic rates from different geographic areas can contribute to improved conservation. Given the scarcity of information on scaup nesting in the boreal forest, basic nesting parameters are important to those trying to model scaup population dynamics.