Raymond J. Davis

The roles of competition and habitat in the dynamics of populations and species distributions.

The role of competition in structuring biotic communities at fine spatial scales is well known from detailed process-based studies. Our understanding of competitions importance at broader scales is less resolved and mainly based on static species distribution maps. Here, we bridge this gap by examining the joint occupancy dynamics of an invading species (Barred Owl, Strix varia) and a resident species (Northern Spotted Owl, Strix occidentalis caurina) in a 1000-km study area over a 22-year period. Past studies of these competitors have focused on the dynamics of one species at a time, hindering efforts to parse out the roles of habitat and competition and to forecast the future of the resident species. In addition, while these studies accounted for the imperfect detection of the focal species, no multi-season analysis of these species has accounted for the imperfect detection of the secondary species, potentially biasing inference. We analyzed survey data using models that combine the general multistate-multi-season occupancy modeling framework with autologistic modeling, allowing us to account for important aspects of our study system. We found that local extinction probability increases for each species when the other is present; however, the effect of the invader on the resident is greater. Although the species prefer different habitats, these habitats are highly correlated at the patch scale, and the impacts of invader on the resident are greatest in patches that would otherwise be optimal. As a consequence, competition leads to a weaker relationship between habitat and Northern Spotted Owl occupancy. Colonization and extinction rates of the invader are closely related to neighborhood occupancy, and over the first half of the study the availability of colonists limited the rate of population growth. Competition is likely to exclude the resident species, both through its immediate effects on local extinction and by indirectly lowering colonization rates as Northern Spotted Owl occupancy declines. Our analysis suggests that dispersal limitation affects both the invasion dynamics and the scale at which the effects of competition are observed. We also provide predictions regarding the potential costs and benefits of managing Barred Owl populations at different target levels.

Neighborhood and habitat effects on vital rates: expansion of the Barred Owl in the Oregon Coast Ranges

In this paper, we modify dynamic occupancy models developed for detection-nondetection data to allow for the dependence of local vital rates on neighborhood occupancy, where neighborhood is defined very flexibly. Such dependence of occupancy dynamics on the status of a relevant neighborhood is pervasive, yet frequently ignored. Our framework permits joint inference about the importance of neighborhood effects and habitat covariates in determining colonization and extinction rates. Our specific motivation is the recent expansion of the Barred Owl (Strix varia) in western Oregon, USA, over the period 1990-2010. Because the focal period was one of dramatic range expansion and local population increase, the use of models that incorporate regional occupancy (sources of colonists) as determinants of dynamic rate parameters is especially appropriate. We began our analysis of 21 years of Barred Owl presence/nondetection data in the Tyee Density Study Area (TDSA) by testing a suite of six models that varied only in the covariates included in the modeling of detection probability. We then tested whether models that used regional occupancy as a covariate for colonization and extinction outperformed models with constant or year-specific colonization or extinction rates. Finally we tested whether habitat covariates improved the AIC of our models, focusing on which habitat covariates performed best, and whether the signs of habitat effects are consistent with a priori hypotheses. We conclude that all covariates used to model detection probability lead to improved AIC, that regional occupancy influences colonization and extinction rates, and that habitat plays an important role in determining extinction and colonization rates. As occupancy increases from low levels toward equilibrium, colonization increases and extinction decreases, presumably because there are more and more dispersing juveniles. While both rates are affected, colonization increases more than extinction decreases. Colonization is higher and extinction is lower in survey polygons with more riparian forest. The effects of riparian forest on extinction rates are greater than on colonization rates. Model results have implications for management of the invading Barred Owl, both through habitat alteration and removal.

Integrating count and detection–nondetection data to model population dynamics

There is increasing need for methods that integrate multiple data types into a single analytical framework as the spatial and temporal scale of ecological research expands. Current work on this topic primarily focuses on combining capture-recapture data from marked individuals with other data types into integrated population models. Yet, studies of species distributions and trends often rely on data from unmarked individuals across broad scales where local abundance and environmental variables may vary. We present a modeling framework for integrating detection-nondetection and count data into a single analysis to estimate population dynamics, abundance, and individual detection probabilities during sampling. Our dynamic population model assumes that site-specific abundance can change over time according to survival of individuals and gains through reproduction and immigration. The observation process for each data type is modeled by assuming that every individual present at a site has an equal probability of being detected during sampling processes. We examine our modeling approach through a series of simulations illustrating the relative value of count vs. detection-nondetection data under a variety of parameter values and survey configurations. We also provide an empirical example of the model by combining long-term detection-nondetection data (1995-2014) with newly collected count data (2015-2016) from a growing population of Barred Owl (Strix varia) in the Pacific Northwest to examine the factors influencing population abundance over time. Our model provides a foundation for incorporating unmarked data within a single framework, even in cases where sampling processes yield different detection probabilities. This approach will be useful for survey design and to researchers interested in incorporating historical or citizen science data into analyses focused on understanding how demographic rates drive population abundance.

Patterns of Change across the Forested Landscape

The scope and extent of past natural disturbances and human-derived changes to the forest landscape often provide the historical context for management but are often insufficiently accounted for in forest planning. In particular, static components of many management plans are not easily adapted to unforeseen system dynamics. For example, when the Northwest Forest Plan was designed in 1993, the inherently dynamic nature of the forest ecosystem and landscape was acknowledged, but there was a general lack of scientific information about the ecological processes that would shape forests of the future. The expectation was that both management and natural disturbances would influence change in the forested landscape, but how management would then adapt to these altered conditions was not clear. At the time, climate change was not well understood and was just beginning to be discussed in relation to forests.

Estimating density of a territorial species in a dynamic landscape

ContextConservation planning for at-risk species requires understanding of where species are likely to occur, how many individuals are likely to be supported on a given landscape, and the ability to monitor those changes through time.ObjectivesWe developed a distribution model for northern spotted owls that incorporates both habitat suitability and probability of territory occupancy while accounting for interspecies competition.MethodsWe developed range-wide habitat suitability maps for two time periods (1993 and 2012) for northern spotted owls that accounted for regional differences in habitat use and home range size. We used these maps for a long-term demographic monitoring study area to assess habitat change and estimate the number of potential territories based on available habitat for both time periods. We adjusted the number of potential territories using known occupancy rates to estimate owl densities for both time periods. We evaluated our range-wide habitat suitability model using independent survey data.ResultsOur range-wide habitat maps predicted areas suitable for territorial spotted owl presence well. On the demographic study area, the amount of habitat declined 19.7% between 1993 and 2012, while our estimate of the habitat-based carrying capacity declined from 150 to 146 territories. Estimated number of occupied territories declined from 94 to 57.ConclusionsConservation and recovery of at-risk species depends on understanding how habitat changes over time in response to factors such as wildfire, climate change, biological invasions, and interspecies competition, and how these changes influence species distribution. We demonstrate a model-based approach that provides an effective planning tool.