Christopher K. Williams

Phylogenetic Measures of Biodiversity

We developed a theoretical framework based on phylogenetic comparative methods to integrate phylogeny into three measures of biodiversity: species variability, richness, and evenness. These metrics can be used in conjunction with permutation procedures to test for phylogenetic community structure. As an illustration, we analyzed data on the composition of 58 lake fish communities in Wisconsin. The fish communities showed phylogenetic underdispersion, with communities more likely to contain closely related species. Using information about differences in environmental characteristics among lakes, we demonstrated that phylogenetic underdispersion in fish communities was associated with environmental factors. For example, lakes with low pH were more likely to contain species in the same clade of acid‐tolerant species. Our metrics differ from existing metrics used to calculate phylogenetic community structure, such as net relatedness index and Faith’s phylogenetic diversity. Our metrics have the advantage of providing an integrated and easy‐to‐understand package of phylogenetic measures of species variability, richness, and evenness with well‐defined statistical properties. Furthermore, they allow the easy evaluation of contributions of individual species to different aspects of the phylogenetic organization of communities. Therefore, these metrics should aid with the incorporation of phylogenetic information into strategies for understanding biodiversity and its conservation.

The northern bobwhite decline: scaling our management for the twenty-first century

Abstract Northern bobwhites (Colinus virginianus) are one of the most broadly researched and intensively managed species in North America. However, we argue that a disadvantage of this status is that traditional management principles currently are incompatible with the spatial scale necessary to address the nationwide decline in bobwhite abundance. We maintain that halting or reversing this decline will entail 2 principal changes in the scale of management. Primarily we suggest that habitat oversight must switch from historical fine-scale management (promotion of edge habitat, weedy fencelines, disked strips, living hedges, and food plots) to regional management of usable space. Secondly, within these regional management areas, we should apply harvest management that employs risk-sensitive strategies that conservatively avoid undermining the primary goal. This entails narrowing the scale of harvest management from statewide to regional levels. If these ideological changes cannot be made and historical policies remain in force, we risk failing to stabilize, let alone increase, bobwhite populations.

Demographic Sensitivity of Population Change in Northern Bobwhite

Abstract The northern bobwhite (Colinus virginianus) is an economically important gamebird that is currently undergoing widespread population declines. Despite considerable research on the population ecology of bobwhites, there have been few attempts to model population dynamics of bobwhites to determine the contributions of different demographic parameters to variance of the finite rate of population change (λ). We conducted a literature review and compiled 405 estimates of 9 demographic parameters from 49 field studies of bobwhites. To identify demographic parameters that might be important for management, we used life-stage simulation analyses (LSA) to examine sensitivity of λ to simulated variation in 9 demographic parameters for female bobwhites. In a baseline LSA based on uniform distributions bounded by the range of estimates for each demographic parameter, bobwhite populations were predicted to decline (λ = 0.56) and winter survival of adults made the greatest contribution to variance of λ (r2 = 0.453), followed by summer survival of adults (r2 = 0.163), and survival of chicks (r2 = 0.120). Population change was not sensitive to total clutch laid, nest survival, egg hatchability, or 3 parameters associated with the number of nesting attempts (r2 <0.06). Our conclusions were robust to alternative simulation scenarios, and parameter rankings changed only if we adjusted the lower bounds of winter survival upwards. Bobwhite populations were not viable with survival rates reported from most field studies. Survival rates may be depressed below sustainable levels by environmental conditions or possibly by impacts of capture and telemetry methods. Overall, our simulation results indicate that management practices that improve seasonal survival rates will have the greatest potential benefit for recovery of declining populations of bobwhites.

Life on the Edge: Northern Bobwhite Ecology at the Northern Periphery of Their Range

ABSTRACT Over the past 40 years, Northern bobwhite (Colinus virginianus) populations have declined range-wide. The Mid-Atlantic once held the highest densities in the country and now shows some of the worst declines. Although population parameters have been quantified throughout most of the bobwhite range, Mid-Atlantic populations have been largely unstudied. To better quantify the dynamics of this declining system, we sought to not only gather annual data on home range, movement, and habitat selection, but also examine how some of these metrics might impact survival. We captured and radio-tracked 154 bobwhites between May 2006 and April 2008 on a 125 km2 area of Cumberland County, New Jersey, USA. Seasonal daily movement ranged from 146 m to 158 m but several extreme movements were notable. Across seasons, grassland habitat was used in greater proportion to its availability, shrub-scrub and agriculture habitats were used equally with their availability and forests and other habitats were used less than their availability. Differences in second-order selection occurred between seasons with lower use of shrub-scrub and forest habitats and higher use of other habitat in breeding seasons. Pooled breeding season survival was 0.343, nonbreeding season survival was 0.183, and annual survival was 0.063. Although mortality was dominated by avian predators, house cat mortalities were noteworthy. Cox proportional hazard analysis revealed that risk of breeding season mortality risk was increased by longer daily movement, lower grassland use, and higher forest and other use. During the nonbreeding season, risk of mortality increased with shorter daily movement and proximity to occupied buildings and barns. This information could inform management decisions in the greater Mid-Atlantic as well as other areas of their range where they exist at very low abundances.

An improved multi-scale approach to modeling habitat occupancy of northern bobwhite†

ABSTRACT Predicting species presence requires knowledge of detection of individuals, scale of model variables, model selection uncertainty, and spatial autocorrelation. Our objective was to incorporate recent modeling advances to predict potential habitat occupancy of northern bobwhite (Colinus virginianus). From 15 May—15 August 2008 and 2009, we conducted repeat-visit surveys at 360 sites within Delaware to sample presence of bobwhite. We randomly selected half the data to model scale-dependent relationships of bobwhite presence with metrics of landscape- and site-scale habitat composition and configuration. The final averaged habitat-occupancy model fit the remainder testing dataset with an area under the receiver operating characteristic curve value of 0.62. At the site scale, bobwhite presence was negatively related to interspersion and juxtaposition of early successional habitat (ESH; grassland and shrubland), ESH to forest edge density, and agriculture to forest edge density, though relative effect sizes were weak to moderate after accounting for model selection uncertainty. At the landscape scale, bobwhite presence was negatively related to patch cohesion of human development within 2.5 km and positively related to patch cohesion of ESH within 2.0 km, with both variables exerting strong effects. Bobwhite presence was also weakly and positively related to percentage of shrubland habitat within 1.0 km of the sampling point. We applied our habitat occupancy model to map the predicted presence of breeding bobwhite within the Delmarva Peninsula, USA. The modeling results and distribution map will provide guidance to State and Federal private land management programs in the Mid-Atlantic to identify where habitat management efforts will be most effective. Our methodology can also serve as a basis for future habitat modeling of bobwhite and other grassland—shrubland species across their range.

Winter fidelity and apparent survival of lesser snow goose populations in the Pacific flyway

Abstract The Beringia region of the Arctic contains 2 colonies of lesser snow geese (Chen caerulescens caerulescens) breeding on Wrangel Island, Russia, and Banks Island, Canada, and wintering in North America. The Wrangel Island population is composed of 2 subpopulations from a sympatric breeding colony but separate wintering areas, whereas the Banks Island population shares a sympatric wintering area in California, USA, with one of the Wrangel Island subpopulations. The Wrangel Island colony represents the last major snow goose population in Russia and has fluctuated considerably since 1970, whereas the Banks Island population has more than doubled. The reasons for these changes are unclear, but hypotheses include independent population demographics (survival and recruitment) and immigration and emigration among breeding or wintering populations. These demographic and movement patterns have important ecological and management implications for understanding goose population structure, harvest of admixed populations, and gene flow among populations with separate breeding or wintering areas. From 1993 to 1996, we neckbanded molting birds at their breeding colonies and resighted birds on the wintering grounds. We used multistate mark–recapture models to evaluate apparent survival rates, resighting rates, winter fidelity, and potential exchange among these populations. We also compared the utility of face stain in Wrangel Island breeding geese as a predictor of their wintering area. Our results showed similar apparent survival rates between subpopulations of Wrangel Island snow geese and lower apparent survival, but higher emigration, for the Banks Island birds. Males had lower apparent survival than females, most likely due to differences in neckband loss. Transition between wintering areas was low (<3%), with equal movement between northern and southern wintering areas for Wrangel Island birds and little evidence of exchange between the Banks and northern Wrangel Island populations. Face staining was an unreliable indicator of wintering area. Our findings suggest that northern and southern Wrangel Island subpopulations should be considered a metapopulation in better understanding and managing Pacific Flyway lesser snow geese. Yet the absence of a strong population connection between Banks Island and Wrangel Island geese suggests that these breeding colonies can be managed as separate but overlapping populations. Additionally, winter population fidelity may be more important in lesser snow geese than in other species, and both breeding and wintering areas are important components of population management for sympatric wintering populations.

Assessing Uncertainty in Coastal Marsh Core Sampling for Waterfowl Foods

Abstract Quantifying foraging resources available to waterfowl in different habitat types is important for estimating energetic carrying capacity. To accomplish this, most studies collect soil-core samples from the marsh substrate, sieve and sort food items, and extrapolate energy values to wetland or landscape scales. This is a costly and time-intensive process; furthermore, extrapolation methods yield energy estimates with large variances relative to the mean. From both research and management perspectives, it is important to understand sources of this variation and estimate the number of soil cores needed to reduce the variance to desired levels. Using 2,341 cores collected from freshwater and salt marsh habitats at four sites along the Atlantic Coast, we examined sampling variation and biological variation among sites and habitats. When we removed extreme outliers in the data caused by large animal food items found in a small core sample, estimates of energy density decreased by an order of magnitude ...

Time Energy Budgets and Food use of Atlantic Brant Across Their Wintering Range

ABSTRACT We conducted extensive behavioral and food sampling of Atlantic brant (Branta bernicla hrota) across their winter range and used time-activity budgets for brant to determine daily energy expenditure (DEE). Sampling occurred 1 December—31 May 2006–2008 in 11,225-km2 sites between Rhode Island and Virginia containing important estuarine and upland habitat. To calculate DEE we used instantaneous scan sampling to estimate time-activity budgets. We also determined foods eaten by brant and energy density of food plants. Last, we quantified body condition of brant, which differed among years, months, regions, and ages, and sexes. Overall DEE for brant was 1,530 ± 64 kJ/day. There was considerable variation in time— activity budgets among years, months, regions, habitat, tide, temperature, and time-of-day, but we detected no significant difference in DEE of brant between years or among regions. However, DEE in January (2,018 ± 173 kJ/day) was nearly double the DEE of brant in May (1,048 ± 137 kJ/day). Brant spent their time feeding (32.3%), swimming (26.2%), resting (16.2%), and flying (14.5%). The percent of brant foreguts sampled contained macroalgae (53%) eelgrass (Zostera marina; 18%), salt marsh cordgrass (Spartina alterniflora; 17%), and terrestrial grass (Poa. sp.) and clover (Trifollium sp.; 9%). Energy density differed by vegetation type: macroalgae (12.6 ± 0.1 kJ/g), eelgrass (14.1 ± 0.1 kJ/g), new-growth salt marsh cordgrass (16.9 ± 0.2 kJ/g), and terrestrial grass and clover (17.7 ± 0.1 kJ/g). Atlantic brant exhibited behavioral plasticity thereby allowing modification of daily activity budgets to meet seasonally varying energetic requirements associated with wintering and spring staging. Recognizing a variable DEE can be used along with eventual estimates of food biomass and total metabolizable energy on the landscape to calculate carrying capacity (goose use days) on state, region, or range-wide scales.

Spatially-explicit land use effects on nesting of Atlantic Flyway resident Canada geese in New Jersey

Atlantic Flyway resident population (AFRP) Canada geese Branta canadensis in New Jersey, USA, have grown dramatically during the last thirty years and are considered as overabundant in many areas. Development of corporate parks and urban areas with manicured lawns and artificial ponds offer ideal nesting habitat for AFRP geese, with limited pressure from hunting or natural predators. As a result, spatial heterogeneity in reproduction must be taken into account in managing the population. We identified the site and landscape spatial scale extents at which land use features influenced nest site selection and nest success. Nest searches were conducted throughout the State during 2009—2010, and 309 nests were monitored through hatch to determine their fates. We ran a spatial correlation analysis of land use composition to identify spatial scale extents at which geese most considerably respond to their environment for nest site selection and nest success. All significant spatial scale extents were at or below 2.25 km for the five classified land use types. We emphasize that habitat-goose associations in densely urban areas were strongest at extents < 1 km, while rural and natural areas were strongest at extents > 1 km. Geese responded to human-dominated land uses at a smaller spatial scale extent than land uses with low human density. The strength of all nest-land use univariate relationships was low; however, our primary objective was to identify the scales extent at which geese associate with land use, rather than the intensity. We encourage managers to consider these scale-dependent associations in identifying important habitat variables in multivariate models; and if population control of AFRP Canada geese is of primary interest, then focusing on local habitat management will most likely have the largest influence in managing this population.

Assuring the future of prairie grouse: dogmas, demagogues, and getting outside the box

Abstract We discuss the necessity of a paradigm shift among managers toward dealing with the recovery and management of prairie grouse (Tympanuchus spp). To assure the future of these species, we will need to test dogmatic assumptions about grouse and their management and challenge the demagogues who insist on perpetuating untested “principles.” Tolerance for descriptive and qualitative studies is needed. Additionally, managers will need to remove themselves from the box and embrace landowners, theoretical biologists, economists, human-dimensions researchers, marketing and advertising specialists, and many other professionals outside the normal sphere of wildlife management. There is crucial need for a willingness to devise, test, and apply innovative ideas that are not normally considered in the management of grouse species, especially applying management to large areas within ecosystems. A comprehensive plan is needed to guide rangewide prairie grouse management. Planned management systems are needed to provide operational guidance in implementing species plans.