Common Raven Density and Greater Sage-Grouse Nesting Success in Southern Wyoming: Potential Conservation and Management Implications

My research was focused on greater sage-grouse (Centrocercus urophasianus; hereafter sage-grouse ) nest-site selection, nest success, and hen survival in relation to avian predators. The trade-off between using habitat and avoiding predators is a common decision for prey species including sage-grouse. In Chapter 2, I compared avian predator densities at sage-grouse nest and brood locations to random locations. Sage-grouse were located where densities of small, medium, and large avian predators were 65-68% less than random locations. The effects of anthropogenic and landscape features on habitat use of sage-grouse hens have not been evaluated relative to avian predator densities. In Chapter 3, I compared anthropogenic and landscape features and avian predator densities among sage-grouse locations (nest, early-brood, late-brood) and random locations. I found sage-grouse hens chose locations with lower avian predator densities compared to random locations, and selected locations farther away from anthropogenic and landscape features. Depredation of sage-grouse nests can be an influential factor limiting their productivity. Predator removal has been simultaneously proposed and criticized as a potential mitigation measure for low reproductive rates of sage-grouse. In Chapter 4, I hypothesized that sage-grouse nest success would be greater in areas where Wildlife Services lowered common raven (Corvus corax: hereafter raven ) density. I found that Wildlife Services decreased raven density by 61% during 2008-2011 but I did not detect a direct improvement to sage-grouse nest success. However, sage-grouse nest success was 22% when ravens were detected within 550 m of a sage-grouse nest and 41% when no raven was detected within 550 m. In Chapter 5, I assessed interactive effects of corvid densities relative to anthropogenic and landscape features on sage-grouse nest success. I found that sage-grouse nest success was positively correlated with rugged habitat. Survival of breeding-age birds is the most important demographic parameter driving sage-grouse abundance. In Chapter 6, I evaluated the effect of raptor densities, proximity to anthropogenic and landscape features, and hen behavior on survival of sage-grouse hens. I found that sage-grouse hen survival was negatively correlated with golden eagle (Aquila chrysaetos) density, proximity to anthropogenic and landscape features, and hen parental investment (nesting and brood-rearing)

Comparison of Conservation Policy Benefits for an Umbrella and Related Sagebrush-Obligate Species

Many conservation strategies promote the potential of multiple species benefitting from protection of large areas necessary for the continued viability of 1 species. One prominent strategy in western North America is Wyoming’s Sage-grouse Core Area Policy, which was designed to conserve greater sage-grouse (Centrocercus urophasianus; hereafter, sage-grouse) breeding habitat, but may also serve as an umbrella to conserve other sagebrush (Artemisia spp.)-obligate wildlife, including songbirds. Sagebrush-obligate songbirds and sage-grouse have undergone population declines throughout the western United States attributed to similar habitat issues. We compared trends of sagebrush-obligate songbirds from the Breeding Bird Survey and sage-grouse lek counts in 2 sage-grouse populations in Wyoming (Powder River Basin and Wyoming Basins), USA from 1996–2013. Our evaluation was focused on similarities among population performance of the umbrella species and the species under that umbrella. Sagebrush-obligate songbird and both sage-grouse populations occupied habitat within and outside of protected core areas. Trends of sagebrush-obligate songbirds were not parallel or consistently similar in trajectory to sage-grouse in either core or non-core areas. Our results indicated core areas were successful at maintaining higher sage-grouse trends compared to areas not protected under the core area policy. However, sagebrush-obligate songbird trends did not follow the same pattern. This suggests that protection of only the best sage-grouse habitat may not be a sufficient conservation strategy for other sagebrush-obligate birds

Do artificial nests simulate nest success of greater sage-grouse?

Artificial nests have been used to study factors affecting nest success because researchers can manipulate them more than natural bird nests. Many researchers have questioned the validity of generalizing the results from artificial nests onto naturally occurring nests. Other studies have assessed the validity of artificial nest studies by simultaneously comparing overall depredation or daily survival rates, depredation timing, predator species, or habitat characteristics of artificial and natural nests. To evaluate how well artificial nests simulated nest success of greater sage-grouse (Centrocercus urophasianus; hereafter, sagegrouse), we used the unique approach of monitoring artificial nests (n = 69) placed in the natural nest bowls of sage-grouse in southern Wyoming, USA, during 2010 to 2011. Brown chicken eggs were placed in natural sage-grouse nests 7 to 14 days after the hatch or depredation of natural sage-grouse nests to compare artificial nest fate to the fate of natural sage-grouse nests. As secondary objectives, we placed cameras next to a subset of artificial nests to identify which predator species were depredating nests, and we assessed the effects of corvid (black-billed magpie [Pica hudsonia] and common raven [Corvus corax]) density, nest-site characteristics (i.e., anthropogenic development, landscape variables, and microhabitat) date of depredation, and presence of a camera near nest bowls on the depredation rate of all artificial nests. We found that depredation of artificial nests paralleled the fate of natural sagegrouse nests. Depredations were more likely to occur earlier in the summer (June to early July rather than late July to early August). Depredation of artificial nests was negligible as time progressed past the typical sage-grouse nesting season, supporting the hypothesis of predators using a search image to detect eggs. We also found that shorter perennial grass height and greater magpie densities were positively associated with the depredation rates of artificial nests. Camera-recorded depredation events verified that 4 badgers (Taxidea taxus), 2 magpies, and 1 domestic cow depredated artificial nests. Artificial nests may give managers insight into the expected nest success rates of sage-grouse in areas of conservation interest. However, care must be taken regarding placement and timing of artificial nests for reliable conclusions to be drawn from artificial nest studies. Furthermore, identifying predators based on artificial nests likely leads to inaccurate assessment of local species composition of nest depredators

Increased Abundance of the Common Raven Within the Ranges of Greater and Gunnison Sage-grouse: Influence of Anthropogenic Subsidies and Fire

The common raven (Corvus corax; raven) is native to North America and has increased in abundance, especially throughout western North America, during the last century. Human subsidies have facilitated raven dispersal into less suitable habitats and enabled these populations to maintain higher annual survival and reproduction. Concomitantly, overabundant raven populations are impacting other native at-risk species such as the greater sage-grouse (Centrocercus urophasianus) and potentially the Gunnison sage-grouse (C. minimus). Using Breeding Bird Survey data from 1995–2014, we evaluated raven count data to quantitatively describe changes in abundance and expansion into sagebrush (Artemisia spp.) ecosystems, specifically sage-grouse habitat. We focused our analyses on the 7 sage-grouse management zones (MZs) delineated across 11 western U.S. states and 2 Canadian provinces. We assessed the effects of land cover and anthropogenic disturbance on instantaneous growth rate (r) or carrying capacity (K) of ravens. Abundance of ravens in western and southeastern MZs was greater than northeastern MZs within the greater sage-grouse range. While raven abundance was lower in MZ I and II (Alberta, Canada; Dakotas, Montana, and northwestern Colorado, USA; Saskatchewan, Canada; and Wyoming, USA), raven expansion and percent increase were equivalent or greater than all other MZs. High abundance in MZ VII indicated Gunnison sage-grouse have been exposed to increased raven populations for several decades. Areas with greater electric power transmission line density had higher r; higher K was positively related to proportion of urban land cover within 25 km and burned area within 3 km and negatively related to greater distance from landfills and proportion of forest land cover within 15 km. Ravens have capitalized on human subsidies to increase abundance and expand into sagebrush ecosystems that did not historically support high raven populations. As such, managers are now faced with a new dilemma of reducing populations of a native species to benefit other native sagebrush obligate species

Greater Sage-Grouse (Centrocercus Urophasianus) Select Nest Sites and Brood Sites Away From Avian Predators

Greater Sage-Grouse (Centrocercus urophasianus) have declined in distribution and abundance in western North America over the past century. Depredation of nests and predation of chicks can be two of the most influential factors limiting their productivity. Prey species utilize antipredation behaviors, such as predator avoidance, to reduce the risk of predation. Birds in general balance the dual necessity of selecting cover to hide from visual and olfactory predators to enhance prospects of survival and reproductive success, which may also be achieved by selecting habitat with relatively fewer predators. We compared avian predator densities at Greater Sage-Grouse nests and brood locations with those at random locations within available sage-grouse habitat in Wyoming. This comparison allowed us to assess the species\u27 ability to avoid avian predators during nesting and early brood rearing. During 2008–2010, we conducted 10-min point-count surveys at 218 nests, 249 brood locations from 83 broods, and 496 random locations. We found that random locations had higher densities of avian predators compared with nest and brood locations. Greater Sage-Grouse nested in areas where there were lower densities of Common Ravens (Corvus corax), Black-billed Magpies (Pica hudsonia), Golden Eagles (Aquila chrysaetos), and hawks (Buteo spp.) compared with random locations. Additionally, they selected brood-rearing locations with lower densities of those same avian predators and of American Kestrels (Falco sparverius), compared with random locations. By selecting nest and brood-rearing locations with lower avian predator densities, Greater Sage-Grouse may reduce the risk of nest depredation and predation on eggs, chicks, and hens

Microhabitat Selection for Nesting and Brood-Rearing by the Greater Sage-Grouse in Xeric Big Sagebrush

Understanding selection of breeding habitat is critical to conserving and restoring habitats for the Greater Sage-Grouse (Centrocercus urophasianus), particularly in xeric landscapes (≤25 cm annual precipitation). We monitored radio-marked female sage-grouse in south-central Wyoming in 2008 and 2009 to assess microhabitat use during nesting and brood rearing. For each model we grouped variables into three hypothesis sets on the basis of the weight of support from previous research (a priori information). We used binary logistic regression to compare habitat used by grouse to that at random locations and used an information-theoretic approach to identify the best-supported models. Selection of microhabitat for nests was more positively correlated with mountain big sagebrush (Artemisia tridentata vaseyana) than with Wyoming big sagebrush (A. t. wyomingensis) and negatively correlated with cheatgrass. Nesting hens also selected microhabitats with greater litter cover. Microhabitat for brood-rearing had more perennial grass and sagebrush cover than did random locations. Microhabitat variables most supported in the literature, such as forb cover and perennial grass cover, accounted for only 8% and 16% of the pure variation in our models for early and late brood rearing, respectively. Our findings suggest sage-grouse inhabiting xeric sagebrush habitats rely on sagebrush cover and grass structure for nesting as well as brood-rearing and that at the microhabitat scale these structural characteristics may be more important than forb availability. Therefore, in xeric sagebrush, practices designed to increase forb production by markedly reducing sagebrush cover, as a means to increase sage-grouse productivity, may not be justified

A Novel Technique to Improve Capture Success of Common Ravens

Traditional trapping techniques for common ravens (Corvus corax; raven) require significant effort, often produce low capture rates, and cannot be used in some situations. We designed a 3-m noose pole to secure ravens from nocturnal roost locations while using a strobe spotlight to temporarily disorient them. We collected measures of trapping efficiency and contrasted them with padded leghold traps also used in the study. We effectively implemented our noose pole method in July and August of 2018, 2019, and 2020 in the Baker and Cow Lakes sage-grouse (Centrocercus urophasianus) Priority Areas of Conservation in eastern Oregon, USA, which yielded trapping efficiency of 0.48 trap-hours/raven (37 total captured ravens). Our trapping efficiency using leghold traps during the same summer months was 76.42 trap-hours/raven (3 total captured ravens). Our new trapping method constitutes an inexpensive and simple way to safely trap ravens at accessible communal roosts and merits further refinement to increase utility and capitalize on the vulnerability of ravens to capture at night

Greater Sage-Grouse (Centrocercus Urophasianus) Select Habitat Based on Avian Predators, Landscape Composition, and Anthropogenic Features

Prey species minimize the risk of predation directly by avoiding predators and indirectly by avoiding risky habitat. Habitat loss and fragmentation have been prevalent in Greater Sage-Grouse (Centrocercus urophasianus; hereafter “sage-grouse”) habitat, which has necessitated a better understanding of mechanisms driving habitat use. Using multinomial logistic regression, we compared landscape attributes and anthropogenic features (indirect mechanisms) and densities of avian predators (direct mechanisms) among 792 sage-grouse locations (340 nests, 331 early brood, and 121 late brood) and 660 random locations in Wyoming, USA, in 2008–2011. Anthropogenic features included oil and gas structures, communication towers, power lines, roads, and rural houses; and landscape attributes included a normalized difference vegetation index (NDVI), topographic ruggedness, the proportion of big sagebrush (Artemisia spp.), and proximity and proportion variables for forested and riparian habitats. Sage-grouse locations were best described with models that included multiple habitat variables and densities of small, medium, and large avian predators. Thus, both indirect and direct mechanisms of predator avoidance were employed by sage-grouse to select habitat and presumably lower their exposure to predation and nest predation. At all reproductive stages, sage-grouse selected flatter locations with a greater proportion of big sagebrush, a higher NDVI, and lower densities of oil and gas structures. Nest locations had a lower density of major roads and were farther away from riparian habitat; early-brood locations had a lower density of power lines and were closer to rural houses; and late-brood locations were closer to riparian habitat. The magnitudes of direct and indirect avoidance by sage-grouse hens were dependent on a sage-grouse\u27s reproductive stage. Differential habitat use of female sage-grouse relative to predation risk and food availability was a means for sage-grouse hens to lower their risk of predation and nest predation, while using habitat to meet their energetic requirements and those of their chicks

Greater Sage-Grouse Select Nest Sites to Avoid Visual Predators but Not Olfactory Predators

Birds can hide from visual predators by locating nests where there is cover and from olfactory predators where habitat features create updrafts, high winds, and atmospheric turbulence, but sites optimal for hiding from visual and olfactory predators often differ. We examined how Greater Sage-Grouse (Centrocercus urophasianus) balance the dual needs of hiding from both visual and olfactory predators on Parker Mountain, Utah, where the Common Raven (Corvus corax) is the main visual predator and the striped skunk (Mephitis mephitis) and American badger (Taxidea taxus) are the main olfactory predators. By comparing nest sites to random sites during 2005 and 2006, we found that sage-grouse nest at sites where their nests were obscured from visual predators but were exposed to olfactory predators. To validate these findings, we replicated the study in southwest Wyoming during 2008. Again, we found that visual obscurity at nest sites was greater than at control sites but olfactory obscurity was less. Our results indicate that Greater Sage-Grouse select nest sites where they will be concealed from visual predators but at the cost of locating nests where they are exposed to olfactory predators. In southwest Wyoming, we found that olfactory predators (mammals) and visual predators (birds) depredated an equal number of nests. By selecting nest sites with visual obscurity, Greater Sage-Grouse have reduced the threat from visual predators to where it was similar to the threat posed by olfactory predators

Estimating Trends of Common Raven Populations in North America, 1966–2018

Over the last half century, common raven (Corvus corax; raven) populations have increased in abundance across much of North America. Ravens are generalist predators known to depredate the eggs and young of several sensitive species. Quantifying raven population increases at multiple spatial scales across North America will help wildlife resource managers identify areas where population increases present the greatest risk to species conservation. We used a hierarchical Bayesian modeling approach to analyze trends of standardized raven counts from 1966 to 2018 using Breeding Bird Survey data within each Level I and II ecoregion of the United States and Canada. We also compared raven abundance within and outside the distributions of 9 sensitive or endangered species. Although we found substantial evidence that raven populations have increased across North America, populations varied in growth rates and relative abundances among regions. We found 73% of Level I (11/15) and II (25/34) ecoregions demonstrated positive annual population growth rates ranging from 0.2–9.4%. We found higher raven abundance inside versus outside the distributions of 7 of the 9 sensitive species included in our analysis. Gunnison sage-grouse (Centrocercus minimus) had the highest discrepancy, with 293% more ravens within compared to outside of their range, followed by greater sandhill crane (Antigone canadensis tabida; 280%), and greater sage-grouse (C. urophasianus; 204%). Only 2 species, least tern (Sternula antillarum) and piping plover (Charadrius melodus), indicated lower raven abundance within relative to outside their distributions. Our findings will help wildlife resource managers identify regional trends in abundance of ravens and anticipate which sensitive species are at greatest risk from elevated raven populations. Future research directed at identifying the underlying regional drivers of these trends could help elucidate the most appropriate and responsive management actions and, thereby, guide the development of raven population management plans to mitigate impacts to sensitive species