Mark E. Seamans

Modeling species occurrence dynamics with multiple states and imperfect detection

Recent extensions of occupancy modeling have focused not only on the distribution of species over space, but also on additional state variables (e.g., reproducing or not, with or without disease organisms, relative abundance categories) that provide extra information about occupied sites. These biologist-driven extensions are characterized by ambiguity in both species presence and correct state classification, caused by imperfect detection. We first show the relationships between independently published approaches to the modeling of multistate occupancy. We then extend the pattern-based modeling to the case of sampling over multiple seasons or years in order to estimate state transition probabilities associated with system dynamics. The methodology and its potential for addressing relevant ecological questions are demonstrated using both maximum likelihood (occupancy and successful reproduction dynamics of California Spotted Owl) and Markov chain Monte Carlo estimation approaches (changes in relative abundance of green frogs in Maryland). Just as multistate capture-recapture modeling has revolutionized the study of individual marked animals, we believe that multistate occupancy modeling will dramatically increase our ability to address interesting questions about ecological processes underlying population-level dynamics.

OCCUPANCY ESTIMATION AND MODELING WITH MULTIPLE STATES AND STATE UNCERTAINTY

The distribution of a species over space is of central interest in ecology, but species occurrence does not provide all of the information needed to characterize either the well-being of a population or the suitability of occupied habitat. Recent methodological development has focused on drawing inferences about species occurrence in the face of imperfect detection. Here we extend those methods by characterizing occupied locations by some additional state variable (e.g., as producing young or not). Our modeling approach deals with both detection probabilities <1 and uncertainty in state classification. We then use the approach with occupancy and reproductive rate data from California Spotted Owls (Strix occidentalis occidentalis) collected in the central Sierra Nevada during the breeding season of 2004 to illustrate the utility of the modeling approach. Estimates of owl reproductive rate were larger than naïve estimates, indicating the importance of appropriately accounting for uncertainty in detection and state classification.

Climate change, uncertainty, and natural resource management†

ABSTRACT Climate change and its associated uncertainties are of concern to natural resource managers. Although aspects of climate change may be novel (e.g., system change and nonstationarity), natural resource managers have long dealt with uncertainties and have developed corresponding approaches to decision-making. Adaptive resource management is an application of structured decision-making for recurrent decision problems with uncertainty, focusing on management objectives, and the reduction of uncertainty over time. We identified 4 types of uncertainty that characterize problems in natural resource management. We examined ways in which climate change is expected to exacerbate these uncertainties, as well as potential approaches to dealing with them. As a case study, we examined North American waterfowl harvest management and considered problems anticipated to result from climate change and potential solutions. Despite challenges expected to accompany the use of adaptive resource management to address problems associated with climate change, we conclude that adaptive resource management approaches will be the methods of choice for managers trying to deal with the uncertainties of climate change.

Spotted owl demography in the Central Sierra Nevada

There has been a great deal of controversy over the status of spotted owl (Strix occidentalis) populations in western North America. Therefore, we estimated the population dynamics of a population of California spotted owls (S. o. occidentalis) in the central Sierra Nevada. We assessed reproductive status at 62 territories on 253 occasions over a 10-year period and recorded 714 captures of 210 individuals over a 14-year period. Reproduction varied temporally but did not exhibit any noticeable trends, whereas survival followed a quadratic pattern: owls experienced higher survival during the middle years of the study. Demographic parameter estimates indicate that the population declined 5.2% (SE = 2.6) per year from 1990-1999. Although conditions behind this decline may change in the future, these results suggest that management of spotted owl populations and maintaining their habitat are high conservation priorities in the central Sierra Nevada.

A sampling design framework for monitoring secretive marshbirds

Abstract. A framework for a sampling plan for monitoring marshbird populations in the contiguous 48 states is proposed here. The sampling universe is the breeding habitat (i.e. wetlands) potentially used by marshbirds. Selection protocols would be implemented within each of large geographical strata, such as Bird Conservation Regions. Site selection will be done using a two-stage cluster sample. Primary sampling units (PSUs) would be land areas, such as legal townships, and would be selected by a procedure such as systematic sampling. Secondary sampling units (SSUs) will be wetlands or portions of wetlands in the PSUs. SSUs will be selected by a randomized spatially balanced procedure. For analysis, the use of a variety of methods as a means of increasing confidence in conclusions that may be reached is encouraged. Additional effort will be required to work out details and implement the plan.

Investigating the population dynamics of California spotted owls without marked individuals

Understanding population dynamics is of great interest in many different contexts. Traditionally, population dynamics have often been considered in terms of individual-based demographic parameters (e.g., abundance, survival, and reproductive rates), estimation of which generally requires information from marked individuals. Alternatively, in some situations, it may be appropriate to consider population dynamics at a landscape level where the focus is shifted from numbers of individuals to the status of the population at places on the landscape. One consequence of doing so is that information from marked individuals is no longer required. Recently developed methods allow the estimation of landscape-level population vital rates in the realistic situation where the current status of the population might be misclassified via field methods (e.g., because of imperfect detection). Here, we consider the case of the California spotted owl (Strix occidentalis occidentalis) at the Eldorado study area in central Sierra Nevada, California, USA, where interest is in the occupancy rate of potential nesting territories, and in whether owls in an occupied territory successfully reproduced each year during 1997–2004. We analyzed the data using multistate occupancy models and found no evidence of annual variation in dynamic occupancy probabilities. There was strong evidence of annual variation in successful reproduction, with the pattern of variation being different depending on whether there was successful reproduction in the territory in the previous year. Of the three environmental variables considered, the Southern Oscillation Index appeared to be most important and explained some of the annual variation in reproduction probabilities.

MEXICAN SPOTTED OWL (STRIX OCCIDENTALIS) POPULATION DYNAMICS: INFLUENCE OF CLIMATIC VARIATION ON SURVIVAL AND REPRODUCTION

Abstract Understanding the mechanisms causing temporal variability in demographic parameters is essential to understanding fluctuations in populations. As part of a long-term demographic study, we evaluated influence of climate on Mexican Spotted Owl (Strix occidentalis lucida) annual survival and reproduction in two study areas, one in Arizona and one in New Mexico. Spotted Owl survival in New Mexico and reproductive output in both study areas were positively related to total amounts of precipitation from the previous year, previous winter, or monsoon season. For both study areas, temporal process variation in reproductive output (CV[R] = 51.2 and 75.2% for Arizona and New Mexico, respectively) was greater than that for survival (CV[ϕ] = 12.9 and 7.1% for Arizona and New Mexico, respectively). Precipitation from the previous year explained 73% of σ̂2temporal reproductive output for Arizona owls and precipitation from the previous monsoon explained 42% of σ̂2temporal in reproductive output for New Mexico owls. Precipitation from the previous monsoon season explained 53% of σ̂2temporal in Arizona owl survival and precipitation from the previous winter explained 56% of σ̂2temporal in New Mexico owl survival. The two populations of Spotted Owls we studied appeared to have the same life-history strategy hypothesized for a population of Northern Spotted Owls (S. o. caurina), although the Mexican subspecies apparently responded quite differently to climatic variation.

Population Dynamics of Spotted Owls in the Sierra Nevada, California

Abstract The California spotted owl (Strix occidentalis occidentalis) is the only spotted owl subspecies not listed as threatened or endangered under the United States Endangered Species Act despite petitions to list it as threatened. We conducted a meta-analysis of population data for 4 populations in the southern Cascades and Sierra Nevada, California, USA, from 1990 to 2005 to assist a listing evaluation by the United States Fish and Wildlife Service. Our study areas (from N to S) were on the Lassen National Forest (LAS), Eldorado National Forest (ELD), Sierra National Forest (SIE), and Sequoia and Kings Canyon National Parks (SKC). These study areas represented a broad spectrum of habitat and management conditions in these mountain ranges. We estimated apparent survival probability, reproductive output, and rate of population change for spotted owls on individual study areas and for all study areas combined (meta-analysis) using model selection or model-averaging based on maximum-likelihood estimation. We followed a formal protocol to conduct this analysis that was similar to other spotted owl meta-analyses. Consistency of field and analytical methods among our studies reduced confounding methodological effects when evaluating results. We used 991 marked spotted owls in the analysis of apparent survival. Apparent survival probability was higher for adult than for subadult owls. There was little difference in apparent survival between male and female owls. Model-averaged mean estimates of apparent survival probability of adult owls varied from 0.811 ± 0.021 for females at LAS to 0.890 ± 0.016 for males at SKC. Apparent survival increased over time for owls of all age classes at LAS and SIE, for adults at ELD, and for second-year subadults and adults at SKC. The meta-analysis of apparent survival, which included only adult owls, confirmed an increasing trend in survival over time. Survival rates were higher for owls on SKC than on the other study areas. We analyzed data from 1,865 observations of reproductive outcomes for female spotted owls. The proportion of subadult females among all territorial females of known age ranged from 0.00 to 0.25 among study areas and years. The proportion of subadults among female spotted owls was negatively related to reproductive output (no. of young fledged/territorial F owl) for ELD and SIE. Eldorado study area and LAS showed an alternate-year trend in reproductive output, with higher output in even-numbered years. Mean annual reproductive output was 0.988 ± 0.154 for ELD, 0.624 ± 0.140 for LAS, 0.478 ± 0.106 for SIE, and 0.555 ± 0.110 for SKC. Eldorado Study Area exhibited a declining trend and the greatest variation in reproductive output over time, whereas SIE and SKC, which had the lowest reproductive output, had the lowest temporal variation. Meta-analysis confirmed that reproductive output varied among study areas. Reproductive output was highest for adults, followed by second-year subadults, and then by first-year subadults. We used 842 marked subadult and adult owls to estimate population rate of change. Modeling indicated that λt (λt is the finite rate of population change estimated using the reparameterized Jolly–Seber estimator [Pradel 1996]) was either stationary (LAS and SIE) or increasing after an initial decrease (ELD and SKC). Mean estimated λt for the 4 study areas was 1.007 (95% CI  =  0.952–1.066) for ELD; 0.973 (95% CI  =  0.946–1.001) for LAS; 0.992 (95% CI  =  0.966–1.018) for SIE; and 1.006 (95% CI  =  0.947–1.068) for SKC. The best meta-analysis model of population trend indicated that λ varied across time but was similar in trend among the study areas. Our estimates of realized population change (Δt; Franklin et al. 2004), which we estimated as the product 1 × λˆ3 × λˆ4 × …× λˆk−1, were based on estimates of λt from individual study areas and did not require estimating annual population size for each study area. Trends represented the proportion of the population size in the first year that remained in each subsequent year. Similar to λˆt on which they were based, these Δˆt showed evidence of decline over the study period for LAS and SIE. The best model indicated recruitment of male and female adult and subadults varied from 0.10 to 0.31 new territorial individuals at time t/number of territorial individuals at time t − 1 and similarly among areas. We also conducted a population viability analysis (PVA) based on results of our meta-analysis. This PVA was of limited utility for ELD and SKC study areas because 95% confidence intervals on the probability of decline or increase spanned the interval [0, 1] within 5–10 years. When we restricted inferences to 7 years, estimated probability of a >10% decline for SIE was 0.41 (95% CI  =  0.09–0.78); for LAS the probability was 0.64 (95% CI  =  0.27–0.94). In contrast, estimated probability of a >10% increase in 7 years for SIE was 0.23 (95% CI  =  0.01–0.55) and for LAS was 0.10 (95% CI  =  0.00–0.34). For comparisons, we simulated a PVA for a hypothetical population with mean λ  =  1.0 and the same temporal variation as observed in our owl populations. Our PVA suggested that both the SIE and LAS populations had higher probabilities of declining in a 7-year period than increasing but that it would be difficult to determine if a population was in a slight gradual decline. Our analysis and the repository of information on our 4 study populations provide a data-rich template for managers to monitor impacts of future management actions on the owl. Specifically, our data can be used to evaluate the effect of management strategies on spotted owls that are being implemented by the United States Forest Service to reduce the risk of wildfire in the Sierra Nevada ecosystem. Our information also provides baseline information for evaluating the status of the owl for potential listing as a threatened species by the United States Fish and Wildlife Service.

Does the presence of barred owls suppress the calling behavior of spotted owls

Abstract ABSTRACT Barred Owls (Strix varia) have expanded their range throughout the ranges of Northern (Strix occidentalis caurina) and California Spotted Owls (S. o. occidentalis). Field observations have suggested that Barred Owls may be behaviorally dominant to Spotted Owls. Therefore, we conducted a test of behavioral dominance by assessing responsiveness of Spotted Owls to conspecific calls when they were in the simulated presence (i.e., imitation of Barred Owl vocalizations) of a Barred Owl. We hypothesized that Spotted Owls would be less likely to respond to conspecific calls in areas where Barred Owls were common. We used a binary 2 × 2 crossover experimental design to examine male Spotted Owl responses at 10 territories randomly selected within two study areas that differed in abundance of Barred Owls. We also conducted a quasi experiment at four study areas using response data from any Spotted Owl (male or female) detected following exposure to Barred Owl calls. We inferred from the crossover experiment that the simulated presence of a Barred Owl might negatively affect Spotted Owl responsiveness. Both subspecies of Spotted Owl responded less to Spotted Owl calls after exposure to Barred Owl calls, Northern Spotted Owls responded less frequently in areas having higher numbers of Barred Owls, and California Spotted Owls responded less frequently than Northern Spotted Owls overall.

Effects of forest management on California Spotted Owls: implications for reducing wildfire risk in fire-prone forests

Management of many North American forests is challenged by the need to balance the potentially competing objectives of reducing risks posed by high-severity wildfires and protecting threatened species. In the Sierra Nevada, California, concern about high-severity fires has increased in recent decades but uncertainty exists over the effects of fuel-reduction treatments on species associated with older forests, such as the California Spotted Owl (Strix occidentalis occidentalis). Here, we assessed the effects of forest conditions, fuel reductions, and wildfire on a declining population of Spotted Owls in the central Sierra Nevada using 20 years of demographic data collected at 74 Spotted Owl territories. Adult survival and territory colonization probabilities were relatively high, while territory extinction probability was relatively low, especially in territories that had relatively large amounts of high canopy cover (≥70%) forest. Reproduction was negatively associated with the area of medium-intensity timber harvests characteristic of proposed fuel treatments. Our results also suggested that the amount of edge between older forests and shrub/sapling vegetation and increased habitat heterogeneity may positively influence demographic rates of Spotted Owls. Finally, high-severity fire negatively influenced the probability of territory colonization. Despite correlations between owl demographic rates and several habitat variables, life stage simulation (sensitivity) analyses indicated that the amount of forest with high canopy cover was the primary driver of population growth and equilibrium occupancy at the scale of individual territories. Greater than 90% of medium-intensity harvests converted high-canopy-cover forests into lower-canopy-cover vegetation classes, suggesting that landscape-scale fuel treatments in such stands could have short-term negative impacts on populations of California Spotted Owls. Moreover, high-canopy-cover forests declined by an average of 7.4% across territories during our study, suggesting that habitat loss could have contributed to declines in abundance and territory occupancy. We recommend that managers consider the existing amount and spatial distribution of high-canopy forest before implementing fuel treatments within an owl territory, and that treatments be accompanied by a rigorous monitoring program.