Anthony M. Pagano

Detection Probabilities for Ground-Based Breeding Waterfowl Surveys

Abstract Current methods for conducting ground-based surveys of breeding waterfowl pairs make the unlikely assumption that detection probabilities are constant and approach 100%. To test this assumption, we conducted independent double-observer pair surveys in North Dakota, USA, to evaluate sources of variation in detection probabilities for 8 common species of prairie-nesting ducks. An experienced observer had 0.911 detection probability averaged over all 8 species (range = 0.866–0.944) versus 0.790 (range = 0.537–0.890) for a novice observer. Detection probabilities also varied substantially among species, but patterns were not consistent between observers. Detection probabilities declined as number of ducks per wetland increased, presumably due to difficulty in identifying large numbers of flushing ducks. Other covariates affecting detection probabilities included size of social groups, precipitation, survey methodology (roadside vs. walk-up), cloud cover, time of day, and amount of wetland vegetation, but these covariates only affected detection probabilities by 2–5%. Our results demonstrated that the assumption of 100% detection probabilities for ground-based waterfowl counts was clearly false and surveys based on this erroneous assumption underestimated population size by 10–29%. We recommend that future investigators measure detection probabilities explicitly by using double-observer methodologies.

Estimating Detection Probabilities of Waterfowl Broods From Ground‐Based Surveys

Abstract Brood:pair ratios could provide an economical method for assessing spatial or temporal variation in waterfowl productivity, but such estimators are severely biased by incomplete detection of broods. We conducted 3 sequential counts of 1,357 waterfowl broods in northeastern North Dakota, USA, and used closed-population mark–recapture models to estimate total brood abundance while controlling for variation in detection probabilities (p). Blue-winged teal (Anas discors) broods had the lowest average detection probability (p = 0.305), whereas diving-duck broods had the highest average detectability (p = 0.571). Detection was generally highest in morning or evening, but temporal patterns varied among species and there was no survey window that maximized detection probabilities for all species. Detection probabilities averaged 0.108 (SD = 0.056) higher for an experienced observer versus an inexperienced observer. Detection probabilities were 0.044 higher for roadside versus walk-up surveys and increased with increasing brood size, total brood abundance, survey date, wind speed, temperature, cloud cover, and amount of time spent surveying each wetland. Detection probabilities declined with increasing wetland size and amount of tall peripheral vegetation. Our mark–recapture results indicated that a traditional unreplicated brood survey would have missed 67.5% of estimated broods, summed over all species. Use of closed-population mark–recapture techniques provided an effective method for reducing this bias and identifying and quantifying factors that reduce detection probabilities of waterfowl broods. We recommend that future brood surveys incorporate 2 or 3 temporally segregated replicate counts to allow for formal estimation of detection probabilities.

Effects of capturing and collaring on polar bears: findings from long-term research on the southern Beaufort Sea population

Abstract Context. The potential for research methods to affect wildlife is an increasing concern among both scientists and the public. This topic has a particular urgency for polar bears because additional research is needed to monitor and understand population responses to rapid loss of sea ice habitat. Aims. This study used data collected from polar bears sampled in the Alaska portion of the southern Beaufort Sea to investigate the potential for capture to adversely affect behaviour and vital rates. We evaluated the extent to which capture, collaring and handling may influence activity and movement days to weeks post-capture, and body mass, body condition, reproduction and survival over 6 months or more. Methods. We compared post-capture activity and movement rates, and relationships between prior capture history and body mass, body condition and reproductive success. We also summarised data on capture-related mortality. Key results. Individual-based estimates of activity and movement rates reached near-normal levels within 2–3 days and fully normal levels within 5 days post-capture. Models of activity and movement rates among all bears had poor fit, but suggested potential for prolonged, lower-level rate reductions. Repeated captures was not related to negative effects on body condition, reproduction or cub growth or survival. Capture-related mortality was substantially reduced after 1986, when immobilisation drugs were changed, with only 3 mortalities in 2517 captures from 1987–2013. Conclusions. Polar bears in the southern Beaufort Sea exhibited the greatest reductions in activity and movement rates 3.5 days post-capture. These shorter-term, post-capture effects do not appear to have translated into any long-term effects on body condition, reproduction, or cub survival. Additionally, collaring had no effect on polar bear recovery rates, body condition, reproduction or cub survival. Implications. This study provides empirical evidence that current capture-based research methods do not have long-term implications, and are not contributing to observed changes in body condition, reproduction or survival in the southern Beaufort Sea. Continued refinement of capture protocols, such as the use of low-impact dart rifles and reversible drug combinations, might improve polar bear response to capture and abate short-term reductions in activity and movement post-capture.

High-energy, high-fat lifestyle challenges an Arctic apex predator, the polar bear

A demanding lifestyle Polar bears appear to be well adapted to the extreme conditions of their Arctic habitat. Pagano et al., however, show that the energy balance in this harsh environment is narrower than we might expect (see the Perspective by Whiteman). They monitored the behavior and metabolic rates of nine free-ranging polar bears over 2 years. They found that high energy demands required consumption of high-fat prey, such as seals, which are easy to come by on sea ice but nearly unavailable in ice-free conditions. Thus, as sea ice becomes increasingly short-lived annually, polar bears are likely to experience increasingly stressful conditions and higher mortality rates. Science, this issue p. 568; see also p. 514 Polar bears’ high-energy lifestyle makes then reliant on sea ice–associated prey. Regional declines in polar bear (Ursus maritimus) populations have been attributed to changing sea ice conditions, but with limited information on the causative mechanisms. By simultaneously measuring field metabolic rates, daily activity patterns, body condition, and foraging success of polar bears moving on the spring sea ice, we found that high metabolic rates (1.6 times greater than previously assumed) coupled with low intake of fat-rich marine mammal prey resulted in an energy deficit for more than half of the bears examined. Activity and movement on the sea ice strongly influenced metabolic demands. Consequently, increases in mobility resulting from ongoing and forecasted declines in and fragmentation of sea ice are likely to increase energy demands and may be an important factor explaining observed declines in body condition and survival.

Social Indices of Breeding Productivity in Parkland Mallards

Abstract Social indices were developed to assess breeding productivity of waterfowl based on weekly roadside surveys of social groupings (i.e., pairs, lone M, flocked M). We calculated social indices for mallard (Anas platyrhynchos) populations breeding on 16 study sites in the Canadian parklands from 1993 to 1998 using 7 previously developed indices. We also calculated duckling:pair ratios from our roadside counts, and we obtained independent measures of nesting effort, nesting success, female success, and fledging rate for these same 16 sites from a concurrent telemetry study. Social indices were correlated (r2 = 0.28–0.67) with telemetry-based measures of breeding productivity in 5 of 7 cases, with the strongest relationships deriving from indices that emphasized renesting effort. The 2 ineffective social indices (r2 ≤ 0.13) both measured early onset of nesting activity. Duckling:pair ratios could be calculated more easily from the same survey data and also were correlated (r2 = 0.26–0.48) with measures of breeding productivity. Because surveys measuring late-nesting effort also can enumerate early hatched ducklings, we recommend that waterfowl researchers use duckling:pair ratios rather than social indices because ducking:pair ratios are more easily interpretable. Development of sightability-adjustment factors for pair and duckling surveys could further enhance the utility of duckling:pair ratios as indices of breeding productivity in mallards.

Energetic costs of locomotion in bears: is plantigrade locomotion energetically economical?

ABSTRACT Ursids are the largest mammals to retain a plantigrade posture. This primitive posture has been proposed to result in reduced locomotor speed and economy relative to digitigrade and unguligrade species, particularly at high speeds. Previous energetics research on polar bears (Ursus maritimus) found locomotor costs were more than double predictions for similarly sized quadrupedal mammals, which could be a result of their plantigrade posture or due to adaptations to their Arctic marine existence. To evaluate whether polar bears are representative of terrestrial ursids or distinctly uneconomical walkers, this study measured the mass-specific metabolism, overall dynamic body acceleration, and gait kinematics of polar bears and grizzly bears (Ursus arctos) trained to rest and walk on a treadmill. At routine walking speeds, we found polar bears and grizzly bears exhibited similar costs of locomotion and gait kinematics, but differing measures of overall dynamic body acceleration. Minimum cost of transport while walking in the two species (2.21 J kg−1 m−1) was comparable to predictions for similarly sized quadrupedal mammals, but these costs doubled (4.42 J kg−1 m−1) at speeds ≥5.4 km h−1. Similar to humans, another large plantigrade mammal, bears appear to exhibit a greater economy while moving at slow speeds. Editors Choice: Polar bears and grizzly bears exhibit a greater economy while moving at slow speeds as a consequence of their plantigrade posture.

Habitat degradation affects the summer activity of polar bears

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

Validation of mercury tip-switch and accelerometer activity sensors for identifying resting and active behavior in bears

Abstract Activity sensors are often included in wildlife transmitters and can provide information on the behavior and activity patterns of animals remotely. However, interpreting activity-sensor data relative to animal behavior can be difficult if animals cannot be continuously observed. In this study, we examined the performance of a mercury tip-switch and a tri-axial accelerometer housed in collars to determine whether sensor data can be accurately classified as resting and active behaviors and whether data are comparable for the 2 sensor types. Five captive bears (3 polar [Ursus maritimus] and 2 brown [U. arctos horribilis]) were fitted with a collar specially designed to internally house the sensors. The bears’ behaviors were recorded, classified, and then compared with sensor readings. A separate tri-axial accelerometer that sampled continuously at a higher frequency and provided raw acceleration values from 3 axes was also mounted on the collar to compare with the lower resolution sensors. Both accelerometers more accurately identified resting and active behaviors at time intervals ranging from 1 minute to 1 hour (≥91.1% accuracy) compared with the mercury tip-switch (range = 75.5–86.3%). However, mercury tip-switch accuracy improved when sampled at longer intervals (e.g., 30–60 min). Data from the lower resolution accelerometer, but not the mercury tip-switch, accurately predicted the percentage of time spent resting during an hour. Although the number of bears available for this study was small, our results suggest that these activity sensors can remotely identify resting versus active behaviors across most time intervals. We recommend that investigators consider both study objectives and the variation in accuracy of classifying resting and active behaviors reported here when determining sampling interval.

Convergence of marine megafauna movement patterns in coastal and open oceans

Significance Understanding the key drivers of animal movement is crucial to assist in mitigating adverse impacts of anthropogenic activities on marine megafauna. We found that movement patterns of marine megafauna are mostly independent of their evolutionary histories, differing significantly from patterns for terrestrial animals. We detected a remarkable convergence in the distribution of speed and turning angles across organisms ranging from whales to turtles (epitome for the slowest animals on land but not at sea). Marine megafauna show a prevalence of movement patterns dominated by search behavior in coastal habitats compared with more directed, ballistic movement patterns when the animals move across the open ocean. The habitats through which they move will therefore need to be considered for effective conservation. The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals’ movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content.

Evaluating methods to assess the body condition of female polar bears

Abstract:  An animals body condition provides insight into its health, foraging success, and overall fitness. Measures of body composition including proportional fat content are useful indicators of condition. Isotopic dilution is a reliable non-destructive method for estimating the body composition of live mammals, but can require prolonged handling times. Alternatively, bioelectrical impedance analysis (BIA) has promise as a quick method for estimating the body composition of live mammals, but measurements can potentially be affected by field conditions. Body condition indices (BCI) and energy density models can also be used to assess body condition based on morphological measurements, but may not reliably reflect an animals energy stores. Here we evaluate BIA, BCI, and an energy density model in measuring the energy stores of female polar bears (Ursus maritimus). We examine the relationship between total body fat (TBF) derived from isotopic dilution to these alternative methods for 9 female polar bears from 14 captures on the sea ice of the southern Beaufort Sea in April 2014–2016. An energy density model, BCI, and BIA-derived measures of TBF were poor predictors of TBF derived from isotopic dilution. We suggest energy density, BCI, and BIA may not be predictive of an animals body fat at fine scales (e.g., among individuals within the same sex, reproductive status, and season). In particular, BIA should provide similar measures of body composition as isotopic dilution, but it failed to reliably measure TBF of individual bears. These limitations in the precision of body condition measures should be considered when planning future studies.