Glen A. Sargeant

Survival of Adult Female Elk in Yellowstone Following Wolf Restoration

Abstract Counts of northern Yellowstone elk (Cervus elaphus) in northwestern Wyoming and adjacent Montana, USA, have decreased at an average rate of 6–8% per year since wolves (Canis lupus) were reintroduced in 1995. Population growth rates of elk are typically sensitive to variations in adult female survival; populations that are stable or increasing exhibit high adult female survival. We used survival records for 85 radiocollared adult female elk 1–19 years old to estimate annual survival from March 2000 to February 2004. Weighted average annual survival rates were approximately 0.83 (95% CI = 0.77–0.89) for females 1–15 years old and 0.80 (95% CI = 0.73–0.86) for all females. Our estimates were much lower than the rate of 0.99 observed during 1969–1975 when fewer elk were harvested by hunters, wolves were not present, and other predators were less numerous. Of 33 documented deaths included in our analysis, we attributed 11 to hunter harvest, 14 to predation (10 wolf, 2 unknown, 1 cougar [Puma concolor], and 1 bear [Ursus sp.]), 6 to unknown causes, and 2 to winter-kill. Most deaths occurred from December through March. Estimates of cause-specific annual mortality rates were 0.09 (0.05–0.14) for all predators, 0.08 (0.04–0.13) for hunting, and 0.07 (0.03–0.11) for wolves specifically. Wolf-killed elk were typically older (median = 12 yr) than hunter-killed elk (median = 9 yr, P = 0.03). However, elk that winter outside the park where they were exposed to hunting were also younger (median = 7 yr) than elk that we did not observe outside the park (median = 9 yr, P < 0.01). Consequently, differences in ages of elk killed by wolves and hunters may reflect characteristics of elk exposed to various causes of mortality, as well as differences in susceptibility. Unless survival rates of adult females increase, elk numbers are likely to continue declining. Hunter harvest is the only cause of mortality that is amenable to management at the present time.

Sampling designs for carnivore scent-station surveys

Scent stations usually are deployed in clusters to expedite data collection and increase the number of stations that can be operated for a given cost. Presumed benefits of cluster sampling may not be realized, however, unless cluster sizes are chosen with respect to sampling variation within and among clusters. To encourage and facilitate the use of efficient designs and reporting standards, we used data collected in Minnesota, USA, during 1986-1991 to (1) compare the performance of survey designs with various numbers of stations/cluster; (2) estimate relations between required sample sizes and visitation rates, changes in visitation rates, and error rates; and (3) compare 2 measures of carnivore response: proportions of scent stations (station index) and proportions of clusters (line index) visited by red foxes (Vulpes vulpes) and striped skunks (Mephitis mephitis). Despite broad ecological differences between the species, results were similar for foxes and skunks. Foxes visited 2-21% of stations and 15-84% of fines. Skunks visited 1-16% of stations and 3-54% of lines. Station and line indices were closely related (r 2 > 0.86) and were similarly sensitive indicators of change in visitation rates. Low visitation rates greatly limited the potential usefulness of scent-station surveys because required minimum sample sizes increased exponentially as visitation rates decreased. For visitation rates below 5-10%, required minimum sample sizes were very large and difficult to anticipate. Relative to single-stage sampling, cluster sampling with 10 stations/cluster inflated sample variances, hence sample sizes required to achieve a fixed level of precision, by a factor of 1.6-2.2. Cluster sampling is advantageous only when cost savings permit increases in sample sizes that outweigh concomitant increases in sampling variability. Costs and sampling variation both should be considered when choosing survey designs, and designs should be evaluated and refined as data accumulate.

MARKOV CHAIN MONTE CARLO ESTIMATION OF SPECIES DISTRIBUTIONS: A CASE STUDY OF THE SWIFT FOX IN WESTERN KANSAS

Abstract Accurate maps of species distributions are essential tools for wildlife research and conservation. Unfortunately, biologists often are forced to rely on maps derived from observed occurrences recorded opportunistically during observation periods of variable length. Spurious inferences are likely to result because such maps are profoundly affected by the duration and intensity of observation and by methods used to delineate distributions, especially when detection is uncertain. We conducted a systematic survey of swift fox (Vulpes velox) distribution in western Kansas, USA, and used Markov chain Monte Carlo (MCMC) image restoration to rectify these problems. During 1997–1999, we searched 355 townships (ca. 93 km2) 1–3 times each for an average cost of

PATTERNS OF VARIATION IN CLUTCH SIZES IN A GUILD OF TEMPERATE-NESTING DABBLING DUCKS

7,315 per year and achieved a detection rate (probability of detecting swift foxes, if present, during a single search) of θ̂ = 0.69 (95% Bayesian confidence interval [BCI] = [0.60, 0.77]). Our analysis produced an estimate of the underlying distribution, rather than a map of observed occurrences, that reflected the uncertainty associated with estimates of model parameters. To evaluate our results, we analyzed simulated data with similar properties. Results of our simulations suggest negligible bias and god good precision when probabilities of detection on ≥1 survey occasions (cumulative probabilities of detection) exceed 0.65. Although the use of MCMC image restoration has been limited by theoretical and computational complexities, alternatives do not possess the same advantages. Image models accommodate uncertain detection, do not require spatially independent data or a census of map units, and can be used to estimate species distributions directly from observations without relying on habitat covariates or parameters that must be estimated subjectively. These features facilitate economical surveys of large regions, the detection of temporal trends in distribution, and assessments of landscape-level relations between species and habitats. Requirements for the use of MCMC image restoration include study areas that can be partitioned into regular grids of mapping units, spatially contagious species distributions, reliable methods for identifying target species, and cumulative probabilities of detection ≥0.65.

Dynamics of newly established elk populations

Abstract We investigated patterns and causes of variation in clutch sizes in a guild of five species of temperate-nesting dabbling ducks (Mallard [Anas platyrhynchos], Northern Pintail [“pintail,” A. acuta], Gadwall [A. strepera], Blue-winged Teal [“teal,” A. discors], and Northern Shoveler [“shoveler,” A. clypeata]) during 1993-1995 in the Prairie Pothole Region of midcontinental North America. Clutch sizes (mean ± SE) were largest for teal (10.80 ± 0.03), followed in descending order by those of shoveler (10.31 ± 0.05), Gadwall (9.92 ± 0.04), Mallard (8.91 ± 0.04), and pintail (7.66 ± 0.06). In Mallard, pintail, and shoveler, predicted clutch sizes at onset of nesting exhibited minimal variation. Clutch sizes of Gadwall and teal displayed statistically significant variation among years at onset of nesting; pintail clutch sizes showed significant variation late in the nesting season. Clutch sizes declined seasonally in all species. Declines in clutch sizes of teal and shoveler were approximately linear; whereas clutch sizes of Mallard, pintail, and Gadwall usually declined at progressively decreasing rates. Linear declines in teal and shoveler clutches suggest that those species experienced greater difficulty securing lipids for egg production late in the nesting season than did Mallard, pintail, and Gadwall. That disparity may result because egg-laying female teal and shoveler feed almost exclusively on animal foods, which are primarily protein; whereas female Mallard, pintail, and Gadwall consume more carbohydrate-rich plant foods. Our findings, when examined in context with existing information, suggest that interspecific variation in clutch sizes results from innate differences in several traits—including body size, diet, timing of lipid acquisition, and nesting—all of which can affect the amount of lipid available for egg production. Temperate-nesting dabbling ducks have evolved traits that facilitate laying of large clutches early in the nesting season, because risk of mortality is lower among early-hatched young. Annual differences in clutch sizes of all five species were not significant when effects of annual variation in nest-initiation dates were accounted for, reflecting the key role of environmental influences on intraspecific variation in clutch sizes among years.

Implications of chronic wasting disease, cougar predation, and reduced recruitment for elk management

Abstract The dynamics of newly established elk (Cervus elaphus) populations can provide insights about maximum sustainable rates of reproduction, survival, and increase. However, data used to estimate rates of increase typically have been limited to counts and rarely have included complementary estimates of vital rates. Complexities of population dynamics cannot be understood without considering population processes as well as population states. We estimated pregnancy rates, survival rates, age ratios, and sex ratios for reintroduced elk at Theodore Roosevelt National Park, North Dakota, USA; combined vital rates in a population projection model; and compared model projections with observed elk numbers and population ratios. Pregnancy rates in January (early in the second trimester of pregnancy) averaged 54.1% (SE = 5.4%) for subadults and 91.0% (SE = 1.7%) for adults, and 91.6% of pregnancies resulted in recruitment at 8 months. Annual survival rates of adult females averaged 0.96 (95% CI = 0.94–0.98) with hunting included and 0.99 (95% CI = 0.97–0.99) with hunting excluded from calculations. Our fitted model explained 99.8% of past variation in population estimates and represents a useful new tool for short-term management planning. Although we found no evidence of temporal variation in vital rates, variation in population composition caused substantial variation in projected rates of increase (λ = 1.20–1.36). Restoring documented hunter harvests and removals of elk by the National Park Service led to a potential rate of λ = 1.26. Greater rates of increase substantiated elsewhere were within the expected range of chance variation, given our model and estimates of vital rates. Rates of increase realized by small elk populations are too variable to support inferences about habitat quality or density dependence.

Determinants of Mallard and Gadwall Nesting on Constructed Islands in North Dakota

ABSTRACT Emerging diseases and expanding carnivore populations may have profound implications for ungulate harvest management and population regulation. To better understand effects of chronic wasting disease (CWD) and cougar (Puma concolor) predation, we studied mortality and recruitment of elk (Cervus elaphus) at Wind Cave National Park (WICA) during 2005–2009. We marked 202 elk (83 subadult M and 119 subadult and ad F) with Global Positioning System (GPS) collars, observed 28 deaths during 74,220 days of monitoring, and investigated 42 additional deaths of unmarked elk found dead. Survival rates were similar for males and females and averaged 0.863 (SE = 0.025) annually. Leading causes of mortality included hunting (0.065, SE = 0.019), CWD (0.034, SE = 0.012), and cougar predation (0.029, SE = 0.012). Marked elk killed by hunters and cougars typically were in good physical condition and not infected with CWD. Effects of mortality on population growth were exacerbated by low rates of pregnancy (subadults = 9.5%, SE = 6.6%; ad = 76.9%, SE = 4.2%) and perinatal survival (0.49, SE = 0.085 from 1 Feb to 1 Sep). Chronic wasting disease, increased predation, and reduced recruitment reduced the rate of increase for elk at WICA to approximately &lgr; = 1.00 (SE = 0.027) during the past decade. Lower rates of increase are mitigating effects of elk on park vegetation, other wildlife, and neighboring lands and will facilitate population control, but may reduce opportunities for elk hunting outside the park.

DOES INCREASING DAYLENGTH CONTROL SEASONAL CHANGES IN CLUTCH SIZES OF NORTHERN PINTAILS (ANAS ACUTA)

Abstract Constructed islands with adequate nesting cover provide secure nesting sites for ducks because islands restrict access by mammalian predators. These islands are costly to construct and should be placed in areas that ensure the greatest use by nesting ducks. We studied mallard (Anas platyrhynchos) and gadwall (A. strepera) nesting on constructed islands in North Dakota in 1996 (n = 20) and 1997 (n = 22) to evaluate factors—particularly amount of perennial grass cover in the surrounding landscape and density of breeding pairs—that possibly influence numbers of initiated nests. We also examined effects of island characteristics, such as island vegetation, on numbers of nests. Numbers of mallard and gadwall nests on islands were negatively related to amounts of perennial grass cover in the surrounding uplands. Numbers of mallard nests were positively related to percentages of tall dense cover on islands. We found no effects of breeding-pair density on numbers of nests initiated by either species, possibly because breeding pairs were abundant on all study sites. Percent shrub cover on islands was a better predictor of island use than was percent tall dense cover. Island use by these species increased with island age and distance from mainland shore. Amounts of perennial cover in landscapes should be primary considerations in determining where to build islands. Our data suggest that use of islands by nesting mallards and gadwalls is greatest in landscapes with little perennial grass cover (i.e., high amounts of cropland). Other researchers documented a positive relation between nest success in upland covers and amount of perennial grass cover in the landscape. Therefore, islands constructed in landscapes with little perennial cover should provide greater gains in duck recruitment rates than islands constructed in landscapes with greater amounts of perennial grass cover.

Demographic response of black bears at Cold Lake, Alberta, to the removal of adult males

Abstract We evaluated spatiotemporal variation in clutch sizes of Northern Pintails (pintails; Anas acuta) nesting in California (1985 to 1996), North Dakota (1982 to 1985), Saskatchewan (1982 to 1985) and Alaska (1991 to 1993) to determine whether seasonal declines in clutch size varied in ways that were consistent with a controlling influence of increasing day length. Pintails began nesting in mid-March in California, mid-April in North Dakota and Saskatchewan, and mid-May in Alaska. Observed durations of nesting were 70 ± 2.6 days (SE) in California, 60 ± 6.3 days in North Dakota, 66 ± 1.3 days in Saskatchewan, and 42 ± 0.7 days in Alaska. Annual differences were the principal source of variation in mean clutch sizes (σ̂Y2 = 0.15, SE = 0.049), which varied little among study locations (σ̂A2 = 0.002, SE = 0.013). Predicted rates of seasonal decline in clutch sizes increased with latitude early in the nesting season, but declined as the nesting season progressed, except in California. Rates of decline in clutch sizes thus were not directly related to rates of increase in day length. Predicted declines in numbers of eggs per clutch over the nesting season were similar for all four locations (range, 3.05–3.12) despite wide variation in durations of nesting. Evidence suggests that reduced nutrient availability during nesting contributes to a higher rate of decline in clutch sizes in Alaska than in temperate regions. Pintails that nest early lay large initial clutches, but thereafter clutch sizes decline rapidly and breeding terminates early. This reproductive strategy is adaptive because young that hatch earliest exhibit the highest survival rates; however, the conversion of grassland to cropland on the primary prairie breeding grounds has reduced hatching rates of clutches laid early in the nesting season. Under these conditions, the limited capacity to renest in late spring on their prairie breeding grounds probably has contributed to Pintail population declines.