Range-wide sources of variation in reproductive rates of northern spotted owls.

We conducted a range-wide investigation of the dynamics of site-level reproductive rate of northern spotted owls using survey data from 11 study areas across the subspecies geographic range collected during 1993-2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did or did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Invader removal triggers competitive release in a threatened avian predator.

Changes in the distribution and abundance of invasive species can have far-reaching ecological consequences. Programs to control invaders are common but gauging the effectiveness of such programs using carefully controlled, large-scale field experiments is rare, especially at higher trophic levels. Experimental manipulations coupled with long-term demographic monitoring can reveal the mechanistic underpinnings of interspecific competition among apex predators and suggest mitigation options for invasive species. We used a large-scale before-after control-impact removal experiment to investigate the effects of an invasive competitor, the barred owl (Strix varia), on the population dynamics of an iconic old-forest native species, the northern spotted owl (Strix occidentalis caurina). Removal of barred owls had a strong, positive effect on survival of sympatric spotted owls and a weaker but positive effect on spotted owl dispersal and recruitment. After removals, the estimated mean annual rate of population change for spotted owls stabilized in areas with removals (0.2% decline per year), but continued to decline sharply in areas without removals (12.1% decline per year). The results demonstrated that the most substantial changes in population dynamics of northern spotted owls over the past two decades were associated with the invasion, population expansion, and subsequent removal of barred owls. Our study provides experimental evidence of the demographic consequences of competitive release, where a threatened avian predator was freed from restrictions imposed on its population dynamics with the removal of a competitively dominant invasive species.

Polar bear population dynamics in the southern Beaufort Sea during a period of sea ice decline.

In the southern Beaufort Sea of the United States and Canada, prior investigations have linked declines in summer sea ice to reduced physical condition, growth, and survival of polar bears (Ursus maritimus). Combined with projections of population decline due to continued climate warming and the ensuing loss of sea ice habitat, those findings contributed to the 2008 decision to list the species as threatened under the U.S. Endangered Species Act. Here, we used mark-recapture models to investigate the population dynamics of polar bears in the southern Beaufort Sea from 2001 to 2010, years during which the spatial and temporal extent of summer sea ice generally declined. Low survival from 2004 through 2006 led to a 25-50% decline in abundance. We hypothesize that low survival during this period resulted from (1) unfavorable ice conditions that limited access to prey during multiple seasons; and possibly, (2) low prey abundance. For reasons that are not clear, survival of adults and cubs began to improve in 2007 and abundance was comparatively stable from 2008 to 2010, with ~900 bears in 2010 (90% CI 606-1212). However, survival of subadult bears declined throughout the entire period. Reduced spatial and temporal availability of sea ice is expected to increasingly force population dynamics of polar bears as the climate continues to warm. However, in the short term, our findings suggest that factors other than sea ice can influence survival. A refined understanding of the ecological mechanisms underlying polar bear population dynamics is necessary to improve projections of their future status and facilitate development of management strategies.

Effects of airgun sounds on bowhead whale calling rates: evidence for two behavioral thresholds.

In proximity to seismic operations, bowhead whales (Balaena mysticetus) decrease their calling rates. Here, we investigate the transition from normal calling behavior to decreased calling and identify two threshold levels of received sound from airgun pulses at which calling behavior changes. Data were collected in August-October 2007-2010, during the westward autumn migration in the Alaskan Beaufort Sea. Up to 40 directional acoustic recorders (DASARs) were deployed at five sites offshore of the Alaskan North Slope. Using triangulation, whale calls localized within 2 km of each DASAR were identified and tallied every 10 minutes each season, so that the detected call rate could be interpreted as the actual call production rate. Moreover, airgun pulses were identified on each DASAR, analyzed, and a cumulative sound exposure level was computed for each 10-min period each season (CSEL10-min). A Poisson regression model was used to examine the relationship between the received CSEL10-min from airguns and the number of detected bowhead calls. Calling rates increased as soon as airgun pulses were detectable, compared to calling rates in the absence of airgun pulses. After the initial increase, calling rates leveled off at a received CSEL10-min of ~94 dB re 1 µPa2-s (the lower threshold). In contrast, once CSEL10-min exceeded ~127 dB re 1 µPa2-s (the upper threshold), whale calling rates began decreasing, and when CSEL10-min values were above ~160 dB re 1 µPa2-s, the whales were virtually silent.

Pacific walrus (Odobenus rosmarus divergens) resource selection in the Northern Bering Sea.

The Pacific walrus is a large benthivore with an annual range extending across the continental shelves of the Bering and Chukchi Seas. We used a discrete choice model to estimate site selection by adult radio-tagged walruses relative to the availability of the caloric biomass of benthic infauna and sea ice concentration in a prominent walrus wintering area in the northern Bering Sea (St. Lawrence Island polynya) in 2006, 2008, and 2009. At least 60% of the total caloric biomass of dominant macroinfauna in the study area was composed of members of the bivalve families Nuculidae, Tellinidae, and Nuculanidae. Model estimates indicated walrus site selection was related most strongly to tellinid bivalve caloric biomass distribution and that walruses selected lower ice concentrations from the mostly high ice concentrations that were available to them (quartiles: 76%, 93%, and 99%). Areas with high average predicted walrus site selection generally coincided with areas of high organic carbon input identified in other studies. Projected decreases in sea ice in the St. Lawrence Island polynya and the potential for a concomitant decline of bivalves in the region could result in a northward shift in the wintering grounds of walruses in the northern Bering Sea.

The point process use-availability or presence-only likelihood and comments on analysis.

1. Use-availability and presence-only analyses are synonyms. Both require two samples (one containing known locations, one containing potential locations), both estimate the same parameters, and both use the same fundamental likelihood. 2. Use-availability and presence-only designs compare characteristics of points where an organism was located to those where the organism could have been located. These designs can be generalized to estimate the relative probability that any event occurred at a set of locations. 3. This article generalizes the use-availability likelihood given in Johnson et al. (Resource selection functions based on use-availability data: theoretical motivation and evaluation methods, Journal of Wildlife Management, 2006) to point locations. This derivation arrives at the same likelihood as Fithian & Hastie (Statistical Models for Presence-Only Data: Finite-Sample Equivalence and Addressing Observer Bias, 2012) but uses a different technique and allows a more general link function. Fithian & Hastie (2012) use a case-control argument and Bayes theorem to derive the likelihood. This article uses Lagrangian multipliers to maximize the two-sample likelihood. 4. Resource selection functions (RSF) defined here are ratios of density functions. RSFs must be positive and unbounded. Proper link functions must provide proportionality over their entire range. Given these conditions, the exponential link is the most logical and appropriate link function for RSFs. These conditions exclude the logistic link. 5. This article affirms that estimation of a RSF does not involve 'running logistic regression'. By assigning 0 and 1 (pseudo-)responses to vectors of covariates associated with locations in the used and available sample, it is possible to 'trick' logistic regression software into maximizing the use-availability likelihood. Representing the analysis as 'logistic regression' is misleading because that implies use of the logistic link, which is inappropriate for RSF's. It is more accurate to state that the 'use-availability likelihood was maximized'. 6. RSFs are more general, intuitive and useful than resource selection probability functions (RSPF). RSPFs depend heavily on sampling mechanisms and the number of used and available locations selected. Consequently, the objective of estimation in use-availability studies should be the RSF, not the RSPF. 7. Two simple examples and R code in the Supporting Information illustrate computations. These examples maximize the general log likelihood without the aid of logistic regression software.

Automated detection and localization of bowhead whale sounds in the presence of seismic airgun surveys.

An automated procedure has been developed for detecting and localizing frequency-modulated bowhead whale sounds in the presence of seismic airgun surveys. The procedure was applied to four years of data, collected from over 30 directional autonomous recording packages deployed over a 280 km span of continental shelf in the Alaskan Beaufort Sea. The procedure has six sequential stages that begin by extracting 25-element feature vectors from spectrograms of potential call candidates. Two cascaded neural networks then classify some feature vectors as bowhead calls, and the procedure then matches calls between recorders to triangulate locations. To train the networks, manual analysts flagged 219 471 bowhead call examples from 2008 and 2009. Manual analyses were also used to identify 1.17 million transient signals that were not whale calls. The network output thresholds were adjusted to reject 20% of whale calls in the training data. Validation runs using 2007 and 2010 data found that the procedure missed 30%-40% of manually detected calls. Furthermore, 20%-40% of the sounds flagged as calls are not present in the manual analyses; however, these extra detections incorporate legitimate whale calls overlooked by human analysts. Both manual and automated methods produce similar spatial and temporal call distributions.

Polar bear population status in the northern Beaufort Sea, Canada, 1971-2006.

Polar bears (Ursus maritimus) of the northern Beaufort Sea (NB) population occur on the perimeter of the polar basin adjacent to the northwestern islands of the Canadian Arctic Archipelago. Sea ice converges on the islands through most of the year. We used open-population capture-recapture models to estimate population size and vital rates of polar bears between 1971 and 2006 to: (1) assess relationships between survival, sex and age, and time period; (2) evaluate the long-term importance of sea ice quality and availability in relation to climate warming; and (3) note future management and conservation concerns. The highest-ranking models suggested that survival of polar bears varied by age class and with changes in the sea ice habitat. Model-averaged estimates of survival (which include harvest mortality) for senescent adults ranged from 0.37 to 0.62, from 0.22 to 0.68 for cubs of the year (COY) and yearlings, and from 0.77 to 0.92 for 2-4 year-olds and adults. Horvtiz-Thompson (HT) estimates of population size were not significantly different among the decades of our study. The population size estimated for the 2000s was 980 +/- 155 (mean and 95% CI). These estimates apply primarily to that segment of the NB population residing west and south of Banks Island. The NB polar bear population appears to have been stable or possibly increasing slightly during the period of our study. This suggests that ice conditions have remained suitable and similar for feeding in summer and fall during most years and that the traditional and legal Inuvialuit harvest has not exceeded sustainable levels. However, the amount of ice remaining in the study area at the end of summer, and the proportion that continues to lie over the biologically productive continental shelf (< 300 m water depth) has declined over the 35-year period of this study. If the climate continues to warm as predicted, we predict that the polar bear population in the northern Beaufort Sea will eventually decline. Management and conservation practices for polar bears in relation to both aboriginal harvesting and offshore industrial activity will need to adapt.

Estimating habitat selection when GPS fix success is less than 100%.

Inferences about habitat selection by animals derived from sequences of relocations obtained with global positioning system (GPS) collars can be influenced by GPS fix success. Environmental factors such as dense canopy cover or rugged terrain can reduce GPS fix success, making subsequent modeling problematic if fix success depends on the selected habitat. Ignoring failed fix attempts may affect estimates of model coefficients and lead to incorrect conclusions about habitat selection. Here, we present a habitat selection model that accounts for missing locations due to habitat-induced data losses, called a resource selection function (RSF) for GPS fix success. The model's formulation is similar to adjusting estimates of probability of occupancy when detection is less than 100% in patch occupancy sampling. We demonstrate use of the model with GPS data collected from an adult female mule deer (Odocoileus hemionus) and discuss how to analyze data from multiple animals. In the simulations presented, our habitat selection model was generally unbiased for GPS data sets missing up to 50% of the locations.

Abundance, behavior, and movement patterns of western gray whales in relation to a 3-D seismic survey, Northeast Sakhalin Island, Russia.

A geophysical seismic survey was conducted in the summer of 2001 off the northeastern coast of Sakhalin Island, Russia. The area of seismic exploration was immediately adjacent to the Piltun feeding grounds of the endangered western gray whale (Eschrichtius robustus). This study investigates relative abundance, behavior, and movement patterns of gray whales in relation to occurrence and proximity to the seismic survey by employing scan sampling, focal follow, and theodolite tracking methodologies. These data were analyzed in relation to temporal, environmental, and seismic related variables to evaluate potential disturbance reactions of gray whales to the seismic survey. The relative numbers of whales and pods recorded from five shore-based stations were not significantly different during periods when seismic surveys were occurring compared to periods when no seismic surveys were occurring and to the post-seismic period. Univariate analyses indicated no significant statistical correlation between seismic survey variables and any of the eleven movement and behavior variables. Multiple regression analyses indicated that, after accounting for temporal and environmental variables, 6 of 11 movement and behavior variables (linearity, acceleration, mean direction, blows per surfacing, and surface-dive blow rate) were not significantly associated with seismic survey variables, and 5 of 11 variables (leg speed, reorientation rate, distance-from-shore, blow interval, and dive time) were significantly associated with seismic survey variables. In summary, after accounting for environmental variables, no correlation was found between seismic survey variables and the linearity of whale movements, changes in whale swimming speed between theodolite fixes, mean direction of whale movement, mean number of whale exhalations per minute at the surface, mean time at the surface, and mean number of exhalations per minute during a whales surface-to-dive cycle. In contrast, at higher received sound energy exposure levels, whales traveled faster, changed directions of movement less, were recorded further from shore, and stayed under water longer between respirations.

Directional frequency and recording (DIFAR) sensors in seafloor recorders to locate calling bowhead whales during their fall migration.

Bowhead whales, Balaena mysticetus, migrate west during fall approximately 10-75 km off the north coast of Alaska, passing the petroleum developments around Prudhoe Bay. Oil production operations on an artificial island 5 km offshore create sounds heard by some whales. As part of an effort to assess whether migrating whales deflect farther offshore at times with high industrial noise, an acoustical approach was selected for localizing calling whales. The technique incorporated DIFAR (directional frequency and recording) sonobuoy techniques. An array of 11 DASARs (directional autonomous seafloor acoustic recorders) was built and installed with unit-to-unit separation of 5 km. When two or more DASARs detected the same call, the whale location was determined from the bearing intersections. This article describes the acoustic methods used to determine the locations of the calling bowhead whales and shows the types and precision of the data acquired. Calibration transmissions at GPS-measured times and locations provided measures of the individual DASAR clock drift and directional orientation. The standard error of the bearing measurements at distances of 3-4 km was approximately 1.35 degrees after corrections for gain imbalance in the two directional sensors. During 23 days in 2002, 10,587 bowhead calls were detected and 8383 were localized.

Review of environmental monitoring methods: survey designs.

During the past decade and a half, environmental monitoring programs have increased in number and importance. Large scale environmental monitoring programs often present design difficulties because they tend to measure many (sometimes hundreds) of parameters through space and time. This paper reviewed and summarized one important component of environmental monitoring programs, the statistical survey design. Survey designs used for long-term monitoring programs lasting multiple (> or = 3) occasions were reviewed, paying special attention to those published after 1985. During this review, two key components of the overall survey design were identified. The first key component was the membership design. Groups of population units sampled the same occasion were called panels here, and the membership design specified which units were members of which panels. The second component was the revisit design that specified when panels were to be revisited. Membership designs varied, but some form of simple random or systematic design was popular. Among revisit designs, four basic patterns were found in the literature and their relative strengths and weaknesses were summarized. To efficiently discuss revisit designs, a new unified short-hand notation was proposed and adopted.