Ranking Alaska Moose Nutrition: Signals to Begin Liberal Antlerless Harvests

Abstract: We focused on describing low nutritional status in an increasing moose (Alces alces gigas) population with reduced predation in Game Management Unit (GMU) 20A near Fairbanks, Alaska, USA. A skeptical public disallowed liberal antlerless harvests of this moose population until we provided convincing data on low nutritional status. We ranked nutritional status in 15 Alaska moose populations (in boreal forests and coastal tundra) based on multiyear twinning rates. Data on age-of-first-reproduction and parturition rates provided a ranking consistent with twinning rates in the 6 areas where comparative data were available. Also, short-yearling mass provided a ranking consistent with twinning rates in 5 of the 6 areas where data were available. Data from 5 areas implied an inverse relationship between twinning rate and browse removal rate. Only in GMU 20A did nutritional indices reach low levels where justification for halting population growth was apparent, which supports prior findings that nutrition is a minor factor limiting most Alaska moose populations compared to predation. With predator reductions, the GMU 20A moose population increased from 1976 until liberal antlerless harvests in 2004. During 1997-2005, GMU 20A moose exhibited the lowest nutritional status reported to date for wild, noninsular, North American populations, including 1) delayed reproduction until moose reached 36 months of age and the lowest parturition rate among 36-month-old moose (29%, n = 147); 2) the lowest average multiyear twinning rates from late-May aerial surveys (x = 7%, SE = 0.9%, n = 9 yr, range = 3-10%) and delayed twinning until moose reached 60 months of age; 3) the lowest average mass of female short-yearlings in Alaska (x̄ = 155 ± 1.6 [SE] kg in the Tanana Flats subpopulation, up to 58 kg below average masses found elsewhere); and 4) high removal (42%) of current annual browse biomass compared to 9-26% elsewhere in boreal forests. When average multiyear twinning rates in GMU 20A (sampled during 1960-2005) declined to <10% in the mid- to late 1990s, we began encouraging liberal antlerless harvests, but only conservative annual harvests of 61-76 antlerless moose were achieved during 1996-2001. Using data in the context of our broader ranking system, we convinced skeptical citizen advisory committees to allow liberal antlerless harvests of 600-690 moose in 2004 and 2005, with the objective of halting population growth of the 16,000-17,000 moose; total harvests were 7-8% of total prehunt numbers. The resulting liberal antlerless harvests served to protect the moose population's health and habitat and to fulfill a mandate for elevated yield. Liberal antlerless harvests appear justified to halt population growth when multiyear twinning rates average ≤10% and ≥1 of the following signals substantiate low nutritional status: <50% of 36-month-old moose are parturient, average multiyear short-yearling mass is <175 kg, or >35% of annual browse biomass is removed by moose.     

Activity Budgets Derived From Time-Depth Recorders in a Diving Mammal

Abstract: We describe a method to convert continuously collected time-depth data from archival time-depth recorders (TDRs) into activity budgets for a benthic-foraging marine mammal. We used data from 14 TDRs to estimate activity-specific time budgets in sea otters (Enhydra lutris) residing near Cross Sound, southeast Alaska, USA. From the TDRs we constructed a continuous record of behavior for each individual over 39-46 days during summer of 1999. Behaviors were classified as foraging (diving to the bottom), other diving (traveling, grooming, interacting), and nondiving (assumed resting). The overall average activity budget (proportion of 24-hr/d) was 0.37 foraging (8.9 hr/d), 0.11 in other diving (2.6 hr/d), and 0.52 nondiving time (12.5 hr/d). We detected significant differences in activity budgets among individuals and between groups within our sample. Historically, the sea otter population in our study area had been expanding and sequentially reoccupying vacant habitat since their reintroduction to the area in the 1960s, and our study animals resided in 2 adjacent yet distinct locations. Males (n = 5) and individuals residing in recently occupied habitat (n = 4) spent 0.28-0.30 of their time foraging (6.7-7.2 hr/d), 0.17-0.18 of their time in other diving behaviors (4.1-4.3 hr/d), and 0.53-0.54 of their time resting (12.7-13.0 hr/d). In contrast, females (n = 9) and individuals residing in longer occupied habitat (n = 10) spent 0.40 of their time foraging (9.6 hr/d), 0.08-0.09 of their time in other diving behaviors (1.9-2.2 hr/d), and 0.51-0.52 of their time resting (12.2-12.5 hr/d). Consistent with these differences, sea otters residing in more recently occupied habitat captured more and larger clams (Saxidomus spp., Protothaca spp., Macoma spp., Mya spp., Clinocardium spp.) and other prey, and intertidal clams were more abundant and larger in this area. We found that TDRs provided data useful for measuring activity time budgets and behavior patterns in a diving mammal over long and continuous time periods. Fortuitous contrasts in time budgets between areas where our study animals resided suggest that activity time budgets estimated from TDRs may be a sensitive indicator of population status, particularly in relation to prey availability.     

Landward and eastward shift of Alaskan polar bear denning associated with recent sea ice changes

Polar bears (Ursus maritimus) in the northern Alaska region den in coastal areas and on offshore drifting ice. We evaluated changes in the distribution of polar bear maternal dens between 1985 and 2005, using satellite telemetry. We determined the distribution of maternal dens occupied by 89 satellite collared female polar bears between 137°W and 167°W longitude. The proportion of dens on pack ice declined from 62% in 1985–1994 to 37% in 1998–2004 (P = 0.044) and among pack ice dens fewer occurred in the western Beaufort Sea after 1998. We evaluated whether hunting, attraction to bowhead whale remains, or changes in sea ice could explain changes in den distribution. We concluded that denning distribution changed in response to reductions in stable old ice, increases in unconsolidated ice, and lengthening of the melt season. In consort, these changes have likely reduced the availability and quality of pack ice denning habitat. Further declines in sea ice availability are predicted. Therefore, we expect the proportion of polar bears denning in coastal areas will continue to increase, until such time as the autumn ice retreats far enough from shore that it precludes offshore pregnant females from reaching the Alaska coast in advance of denning.     

Using dive behavior and active acoustics to assess prey use and partitioning by fin and humpback whales near Kodiak Island,Alaska

Near the Kodiak Archipelago, fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales frequently overlap spatially and temporally. The Gulf Apex Predator‐prey study (GAP) investigated the prey use and potential prey partitioning between these sympatric species by combining concurrent analysis of vertical whale distribution with acoustic assessment of pelagic prey. Acoustic backscatter was classified as consistent with either fish or zooplankton. Whale dive depths were determined through suction cup tags. Tagged humpback whales (n = 10) were most often associated with distribution of fish, except when zooplankton density was very high. Associations between the dive depths of tagged fin whales (n = 4) and the vertical distribution of either prey type were less conclusive. However, prey assessment methods did not adequately describe the distribution of copepods, a potentially significant resource for fin whales. Mean dive parameters showed no significant difference between species when compared across all surveys. However, fin whales spent a greater proportion of dive time in the foraging phase than humpbacks, suggesting a possible difference in foraging efficiency between the two. These results suggest that humpback and fin whales may target different prey, with the greatest potential for diet overlap occurring when the density of zooplankton is very high.     

Effects of the landscape on boreal toad gene flow: does the pattern–process relationship hold true across distinct landscapes at the northern range margin?

Understanding the impact of natural and anthropogenic landscape features on population connectivity is a major goal in evolutionary ecology and conservation. Discovery of dispersal barriers is important for predicting population responses to landscape and environmental changes, particularly for populations at geographic range margins. We used a landscape genetics approach to quantify the effects of landscape features on gene flow and connectivity of boreal toad (Bufo boreas) populations from two distinct landscapes in south-east Alaska (Admiralty Island, ANM, and the Chilkat River Valley, CRV). We used two common methodologies for calculating resistance distances in landscape genetics studies (resistance based on least-cost paths and circuit theory). We found a strong effect of saltwater on genetic distance of CRV populations, but no landscape effects were found for the ANM populations. Our discordant results show the importance of examining multiple landscapes that differ in the variability of their features, to maximize detectability of underlying processes and allow results to be broadly applicable across regions. Saltwater serves as a physiological barrier to boreal toad gene flow and affects populations on a small geographic scale, yet there appear to be few other barriers to toad dispersal in this intact northern region.     

Egg production and early development of Thysanoessa inermis and Euphausia pacifica (Crustacea: Euphausiacea) in the northern Gulf of Alaska

Early life history patterns were studied in the dominant euphausiids from the northern Gulf of Alaska (GOA) in 2001-2004. Gravid females of Thysanoessa inermis were observed in April and May. Brood size varied from 10 to 1021 eggs with an average of 138 ± 19 (95% CI) eggs female^− 1. Most gravid females started to release eggs within the first 2 days of incubation. The average number of eggs released per female was similar in incubation Day 1 and 2, but significantly smaller on Day 3 and 4. About 25% of the females were continuously releasing eggs over 3 days rather than producing a single distinctive brood. In contrast, gravid females of Euphausia pacifica were observed from early July through October. Most gravid females released eggs on the first day of observation, while only 2% of females produced eggs repeatedly. Brood size varied from 20 to 246 eggs with an average of 102 ± 12 (95% CI) eggs female^− 1. The relationship between E. pacifica brood size and ambient chlorophyll-a concentration was sigmoidal (r² = 0.73), with food saturated brood size of 144 ± 14(SE, P < 0.001) eggs, and half-saturation occurring at 0.46 ± 0.02(SE, P < 0.001) mg chlorophyll-a m^− 3. The average interbrood interval of E. pacifica reared at 12 °C and satiated food conditions in the laboratory was ∼ 8 days, suggesting their potential individual fecundity in the GOA was 1148-1530 eggs per spawning season. Hatching and early development (from egg to furcilia stage) was studied under 5 °C, 8 °C and 12 °C. Hatching was nearly synchronous and lasted 3-6 h, depending on incubation temperature. Development times from egg to the first furcilia stage ranged between 20 and 33 days for T. inermis, and 15 and 45 days for E. pacifica at 12 °C and 5 °C, respectively.     

Human Impacts on the Fire Regime of Interior Alaska: Interactions among Fuels, Ignition Sources, and Fire Suppression

Wildfire is the major natural agent of disturbance in interior Alaska. We examined the magnitude of human impact on fire by comparing fire regime between individual 1-km² grid cells designated for fire suppression with lands where fires are allowed to burn naturally. Two-thirds of interior Alaska has an essentially natural fire regime, with few human ignitions, negligible suppression activity, and many large lightning-caused fires. In the 17% of land that is designated for fire suppression due to its proximity to communities and roads, there was a 50% reduction in the proportion of area burned from 1992–2001, relative to areas without suppression. The remaining 16% of land serves as a buffer, receives some suppression, and has an intermediate fire regime. Even though there were 50 times more fires and the fire season began two months earlier in lands designated for suppression, most of these fires were lit by people and remained small because fires tended to occur at times and places less favorable for fire spread and were more accessible to fire fighters compared to lands not designated for suppression. Even in the absence of fire suppression, human-caused fires were less likely to exceed 400 ha compared to lightning-caused fires. Fire suppression reduced area burned in all fuel types but was somewhat more effective in less flammable (non-forest) vegetation. Alaska’s fire policy of focusing suppression efforts on a small proportion of the fire-prone region maximizes the ecological and social benefits associated with fire-dependent ecosystem services, while minimizing the social and ecological costs of suppression. Application of this policy to other areas would require well-informed managers and stakeholders to make difficult decisions about the relative costs and benefits of fire across ecologically and culturally variable landscapes.     

Correlates of Mortality in an Exploited Wolf Population

Abstract We investigated the influence of habitat use on risk of death from hunting and trapping of 55 radiocollared gray wolves (Canis lupus) from an exploited insular population in Southeast Alaska, USA. We compared mortality rates for resident and nonresident wolves and used Cox proportional hazards regression to relate habitat composition within 100-m circular buffers around radiolocations to risk of death of resident and nonresident wolves. In addition, we included covariates representing distances to roads, logged stands, and lakes and streams in those analyses. We also compiled harvest data from 31 harvest units within the study area to compare densities of roads and distances from human settlements with rates of harvest. During our study 39 wolves died, of which 18 were harvested legally, 16 were killed illegally, and 5 died from natural causes. Legal and illegal harvest accounted for >87% of the mortality of radiocollared resident and nonresident wolves. Mean annual survival was 0.54 (SE = 0.17) for all wolves. Annual survival was 0.65 (SE = 0.17) for resident wolves and 0.34 (SE = 0.17) for nonresidents. Very few (19%) nonresident wolves survived to colonize vacant territories or join existing wolf packs. Roads, muskegs, and distances from lakes and streams were covariates positively associated with death of resident wolves. Clear-cuts were positively associated with risk of death of nonresident wolves. Rate of harvest increased with density of roads; however, road densities >0.9 km/km² had little additional effect on harvest rates. Harvest rates decreased with ocean distances from nearest towns or settlements. Roads clearly increased risk of death for wolves from hunting and trapping and contributed to unsustainable rates of harvest. Wildlife managers should consider effects of roads and other habitat features on harvest of wolves when developing harvest recommendations. They should expect substantial illegal harvest where wolf habitat is accessible to humans. Moreover, high rates of mortality of nonresident wolves exposed to legal and illegal harvest may reduce or delay successful dispersal, potentially affecting linkages between small disjunct wolf populations or population segments. We conclude that a combination of conservative harvest regulations and large roadless reserves likely are the most effective measures for conserving wolves where risks from human-caused mortality are high.     

Factors Influencing Nesting Success of King Eiders on Northern Alaska's Coastal Plain

Abstract: King eider (Somateria spectabilis) populations have declined markedly in recent decades for unknown reasons. Nest survival is one component of recruitment, and a female's chance of reproductive success increases with her ability to choose an appropriate nesting strategy. We estimated variation in daily nest survival of king eiders at 2 sites, Teshekpuk and Kuparuk, Alaska, USA, 2002–2005. We evaluated both a priori and exploratory competing models of nest survival that considered importance of nest concealment, seclusion, and incubation constancy as strategies to avoid 2 primary egg predators, avian (Larus spp., Stercorarius spp., and Corvus corax) and fox (Alopex lagopus). We used generalized nonlinear techniques to examine factors affecting nest survival rates and information-theoretic approaches to select among competing models. Estimated nest survival, accounting for a nest visitation effect, varied considerably across sites and years (0.21–0.57); however, given our small sample size, much of this variation may be attributable to sampling variation (s̀²_process = 0.007, 95% CI: 0.003–0.070). Nest survival was higher at Kuparuk than Teshekpuk in all years; however, due to the correlative nature of our data, we cannot determine the underlying causes with any certainty. We found mixed support for the concealed breeding strategy; females derived no benefit from nesting in areas with more willow (Salix spp.; measure of concealment) except that the observer effect diminished as willow cover increased. We suggest these patterns are due to conflicting predation pressures. Nest survival was not higher on islands (measure of seclusion) or with increased incubation constancy but was higher post-fox removal, indicating that predator control on breeding grounds could be a viable management option. Nest survival was negatively affected by our nest visitations, most likely by exposing the nest to avian scavengers. We recommend precautions be taken to limit the effects of nest visits in future studies and to consider them as a possible negative bias in estimated nest survival. Future models of the impacts of development within the breeding grounds of king eider should consider the influence of humans in the vicinity of nests.     

Seasonal Survival of Radiomarked Emperor Geese in Western Alaska

Abstract The population of emperor geese (Chen canagica) in western Alaska, USA, declined by >50% from the 1960s to the mid-1980s and has increased only slightly since. Rates of population increase among arctic geese are especially sensitive to changes in adult survival. Improving adult survival in seasons or geographic areas where survival is low may be the best means of increasing the emperor goose population. We monitored fates of 133 adult female emperor geese that were radiomarked with surgically implanted very high frequency or satellite radiotransmitters from 1999 to 2004 to assess whether monthly survival varied among years, seasons, or geographic areas. Because of uncertainties in determining whether a bird had died based on the radio signal, we analyzed 2 versions of the data. One version used conservative criteria to identify which birds had died based on radio signals and the other used more liberal criteria. In the conservative version of the data we detected 12 mortalities of emperor geese, whereas in the liberal interpretation there were 18 mortalities. In both versions, the models with greatest support indicated that monthly survival varied seasonally and that compared to most seasons estimated monthly survival was lower (φ = 0.95–0.98) in May and August when emperor geese were mainly on the Yukon-Kuskokwim Delta. From 44% to 47% of annual mortality occurred in those months. Estimated monthly survival was higher (φ = 0.98–1.0) from September through March when emperor geese were at autumn staging or wintering areas and in June and July when birds were nesting, rearing broods, or molting. Estimated annual survival was 0.85 (95% CI = 0.77–0.92) in the best-supported model when we used conservative criteria to identify mortalities and 0.79 (95% CI = 0.74–0.85) under the best model using liberal mortality criteria. Lower survival in August and May corresponded to periods when subsistence harvest of emperor geese was likely highest. Managers may be able to most effectively influence population growth rate of emperor geese by reducing subsistence harvest on the Yukon-Kuskokwim Delta in May and August.