Michael A. Gregg

Ecology and management of sage-grouse and sage-grouse habitat

Abstract Sage-grouse (Centrocercus urophasianus and C. minimus) historically inhabited much of the sagebrush-dominated habitat of North America. Today, sage-grouse populations are declining throughout most of their range. Population dynamics of sage-grouse are marked by strong cyclic behavior. Adult survival is high, but is offset by low juvenile survival, resulting in low productivity. Habitat for sage-grouse varies strongly by life-history stage. Critical habitat components include adequate canopy cover of tall grasses (≥ 18 cm) and medium height shrubs (40–80 cm) for nesting, abundant forbs and insects for brood rearing, and availability of herbaceous riparian species for late-growing season foraging. Fire ecology of sage-grouse habitat changed dramatically with European settlement. In high elevation sagebrush habitat, fire return intervals have increased (from 12–24 to > 50 years) resulting in invasion of conifers and a consequent loss of understory herbaceous and shrub canopy cover. In lower elevation sagebrush habitat, fire return intervals have decreased dramatically (from 50–100 to < 10 years) due to invasion by annual grasses, causing loss of perennial bunchgrasses and shrubs. Livestock grazing can have negative or positive impacts on sage-grouse habitat depending on the timing and intensity of grazing, and which habitat element is being considered. Early season light to moderate grazing can promote forb abundance/availability in both upland and riparian habitats. Heavier levels of utilization decrease herbaceous cover, and may promote invasion by undesirable species. At rates intended to produce high sagebrush kill, herbicide-based control of big sagebrush may result in decreased habitat quality for sage-grouse. Light applications of tebuthiuron (N-[5-(1,1-dimethylethyl)-1,3,4-thiadiazol-2-yl]-N,N′-dimethylurea) can decrease canopy cover of sagebrush and increase grass and forb production which may be locally important to nesting and foraging activities. The ability of resource managers to address sage-grouse habitat concerns at large scales is aided greatly by geomatics technology and advances in landscape ecology. These tools allow unprecedented linkage of habitat and population dynamics data over space and time and can be used to retroactively assess such relationships using archived imagery. The present sage-grouse decline is a complex issue that is likely associated with multiple causative factors. Solving management issues associated with the decline will require unprecedented cooperation among wildlife biology, range science, and other professional disciplines.

Vegetational Cover and Predation of Sage Grouse Nests in Oregon

Because of long-term declines in sage grouse (Centrocercus urophasianus) abundance and productivity in Oregon, we investigated the relationship between vegetational cover and nesting by sage grouse in 2 study areas. Medium height (40-80 cm) shrub cover was greater (P 18 cm), residual grass cover was greater (P < 0.001) at nonpredated nests (x = 18%) than in areas surrounding nonpredated nests (x = 6%) or random locations (x = 3%)

Predictive modeling and mapping sage grouse (Centrocercus urophasianus) nesting habitat using Maximum Entropy and a long-term dataset from Southern Oregon

Predictive modeling and mapping based on the quantitative relationships between a species and the biophysical features (predictor variables) of the ecosystem in which it occurs can provide fundamental information for developing sustainable resource management policies for species and ecosystems. To create management strategies with the goal of sustaining a species such as sage grouse (Centrocercus urophasianus), whose distribution throughout North America has declined by approximately 50%, land management agencies need to know what attributes of the range they now inhabit will keep populations sustainable and which attributes attract disproportionate levels of use within a home range. The objectives of this study were to 1) quantify the relationships between sage grouse nest-site locations and a set of associated biophysical attributes using Maximum Entropy, 2) find the best subset of predictor variables that explain the data adequately, 3) create quantitative sage grouse distribution maps representing the relative likelihood of nest-site habitat based on those relationships, and 3) evaluate the implications of the results for future management of sage grouse. Nest-site location data from 1995 to 2003 were collected as part of a long-term research program on sage grouse reproductive ecology at Hart Mountain National Antelope Refuge. Two types of models were created: 1) with a set of predictor variables derived from digital elevation models, a field-validated vegetation classification, and UTM coordinates and 2) with the same predictors and UTM coordinates excluded. East UTM emerged as the most important predictor variable in the first type of model followed by the vegetation classification which was the most important predictor in the second type of model. The average training gain from ten modeling runs using all presence records and randomized background points was used to select the best subset of predictors. A predictive map of sage grouse nest-site habitat created from the application of the model to the study area showed strong overlap between model predictions and nest-site locations.

Survival of Greater Sage-Grouse Chicks and Broods in the Northern Great Basin

Abstract Reduced annual recruitment because of poor habitat quality has been implicated as one of the causative factors in the range-wide decline of sage-grouse (Centrocercus urophasianus) populations since the 1950s. Because chick and brood survival are directly linked to annual recruitment and may be the primary factors that limit sage-grouse population growth, we estimated 28-day survival rates of radiomarked chicks and broods from 2000 to 2003. We examined relationships between survival and several habitat variables measured at brood sites, including food availability (insects and forbs); horizontal cover of sagebrush, grasses, and forbs; and vertical cover of sagebrush and grass. We monitored 506 radiomarked chicks from 94 broods; chick survival was 0.392 (SE  =  0.024). We found evidence that both food and cover variables were positively associated with chick survival, including Lepidoptera availability, slender phlox (Phlox gracilis) frequency, total forb cover, and grass cover. The effect of total grass cover on chick survival was dependent on the proportion of short grass. The hazard of an individual chicks death decreased 8.6% (95% CI  =  −1.0 to 18.3) for each percentage point increase in total grass cover when the proportion of short grass was >70%. Survival of 83 radiomarked broods was 0.673 (SE  =  0.055). Lepidoptera availability and slender phlox frequency were the only habitat variables related to brood survival. Risk of total brood loss decreased by 11.8% (95% CI  =  1.2–22.5) for each additional Lepidoptera individual and 2.7% (95% CI  =  −0.4 to 5.8) for each percentage point increase in the frequency of slender phlox found at brood sites. Model selection results revealed that temporal differences in brood survival were associated with variation in the availability of Lepidoptera and slender phlox. Years with high brood survival corresponded with years of high Lepidoptera availability and high slender phlox frequency. These foods likely provided high-quality nutrition for chicks during early growth and development and enhanced survival. Habitat management that promotes Lepidoptera and slender phlox abundance during May and June (i.e., early brood rearing) should have a positive effect on chick and brood survival in the short term and potentially increase annual recruitment.

Total Plasma Protein and Renesting by Greater Sage-Grouse

Abstract Greater sage-grouse (Centrocercus urophasianus) population declines have been attributed to reduced productivity. Although renesting by sage-grouse may contribute significantly to annual productivity during some years, little information is available on this aspect of sage-grouse reproductive ecology. We investigated the relationship between total plasma protein, age of hen, time of first nest initiation, and time of first nest loss on occurrence of renesting. We captured, assigned age, extracted blood, and radiomarked prelaying, female sage-grouse on 4 study areas during 1999–2004. We monitored radiomarked females from mid-April through June to identify period of nest initiation (early, mid, or late), nest loss (early or late), and renesting activity. We only considered hens that were available to renest (n = 143) for analysis, and we censored those that nested successfully or died during their first nest attempt. Depredation and abandonment accounted for 85% (122/143) and 15% (21/143) of the unsuccessful first nests, respectively. The proportion of hens renesting was 34% (48/143) across all study areas and years. Akaikes Information Criterion model selection indicated that occurrence of renesting varied by age, nest initiation period, nest loss period, and total plasma protein. The best model had low predictive power for any given hen (r 2 = 0.296), but validation of the best model indicated that our predictor variables were important for distinguishing renesting status and likely explained substantial temporal and spatial variation in renesting rates. A greater proportion of adults than yearlings renested, and hens that nested early in the nesting season and lost nests early during incubation were the most likely to renest. Hens that renested had greater total plasma protein levels than non-renesting hens independent of age, nest initiation period, and nest loss period. Because sage-grouse depend on exogenous sources of protein for reproduction, land management practices that promote high-quality, prelaying hen habitat could increase dietary protein intake and sage-grouse renesting rates.

Use of Implanted Radiotransmitters to Estimate Survival of Greater Sage-Grouse Chicks

Abstract Reduced chick survival has been implicated in declines of greater sage-grouse (Centrocercus urophasianus) populations. Because monitoring survival of unmarked sage-grouse chicks is difficult, radiotelemetry may be an effective technique to estimate survival rates, identify causes of mortality, and collect ecological data. Previous studies have used subcutaneous implants to attach radiotransmitters to hatchlings of several species of birds with precocial young. Previous researchers who used subcutaneous implants in free-ranging populations removed chicks from the capture location and implanted transmitters at an alternate site. Because logistics precluded removing newly hatched greater sage-grouse chicks from the field, we evaluated a method for implanting transmitters at capture locations. We captured 288 chicks from 52 broods and monitored 286 radiomarked chicks daily for 28 days following capture during May and June 2001–2002. Two (<1%) chicks died during surgery and we did not radiomark them. At the end of the monitoring period, 26 chicks were alive and 212 were dead. Most (98%, 207/212) radiomarked chick mortality occurred ≤21 days posthatch and predation (82%, 174/212) was the primary cause of death. Necropsies of 22 radiomarked chicks did not indicate inflammation or infection from implants, and they were not implicated in the death of any chicks. Fate of 48 chicks was unknown because of transmitter loss (n = 16), radio failure (n = 29), and brood mixing (n = 3). Overall, the 28-day chick survival rate was 0.220 (SE = 0.028). We found that mortalities related to the implant procedure and transmitter loss were similar to rates reported by previous researchers who removed chicks from capture sites and implanted transmitters at an alternate location. Subcutaneous implants may be a useful method for attaching transmitters to newly hatched sage-grouse chicks to estimate survival rates, identify causes of mortality, and collect ecological data.

Temporal Variation in Diet and Nutrition of Preincubating Greater Sage-Grouse

Abstract Greater sage-grouse (Centrocercus urophasianus) habitat management involves vegetation manipulations to increase or decrease specific habitat components. For sage-grouse habitat management to be most effective, an understanding of the functional response of sage-grouse to changes in resource availability is critical. We investigated temporal variation in diet composition and nutrient content (crude protein, calcium, and phosphorus) of foods consumed by preincubating female sage-grouse relative to food supply and age of hen. We collected 86 preincubating female greater sage-grouse at foraging areas during early (18–31 March) and late (1–12 April) preincubation periods during 2002–2003. Females consumed 22 food types including low sagebrush (Artemisia arbuscula Nutt.), big sagebrush (Artemisia tridentata Nutt.), 15 forb species, 2 insect taxa, sagebrush galls, moss, and a trace amount of unidentified grasses. Low sagebrush was the most common food item, but forbs were found in 89% of the crops and composed 30.1% aggregate dry mass (ADM) of the diet. ADM and species composition of female diets were highly variable between collection periods and years, and coincided with temporal variation in forb availability. Adult females consumed more forbs and less low sagebrush compared to yearling females. Because of higher levels of crude protein, calcium, and phosphorus, forbs were important diet components in comparison with low sagebrush, which had the lowest nutrient content of all foods consumed. Our results indicate that increased forb abundance in areas used by female sage-grouse prior to nesting would increase their forb consumption and nutritional status for reproduction. We recommend that managers should emphasize delineation of habitats used by preincubating sage-grouse and evaluate the need for enhancing forb abundance and diversity.

Burrowing Owls, Pulex irritans , and Plague

Western Burrowing Owls (Athene cunicularia hypugaea) are small, ground-dwelling owls of western North America that frequent prairie dog (Cynomys spp.) towns and other grasslands. Because they rely on rodent prey and occupy burrows once or concurrently inhabited by fossorial mammals, the owls often harbor fleas. We examined the potential role of fleas found on burrowing owls in plague dynamics by evaluating prevalence of Yersinia pestis in fleas collected from burrowing owls and in owl blood. During 2012-2013, fleas and blood were collected from burrowing owls in portions of five states with endemic plague-Idaho, Oregon, Washington, Colorado, and South Dakota. Fleas were enumerated, taxonomically identified, pooled by nest, and assayed for Y. pestis using culturing and molecular (PCR) approaches. Owl blood underwent serological analysis for plague antibodies and nested PCR for detection of Y. pestis. Of more than 4750 fleas collected from owls, Pulex irritans, a known plague vector in portions of its range, comprised more than 99.4%. However, diagnostic tests for Y. pestis of flea pools (culturing and PCR) and owl blood (PCR and serology) were negative. Thus, even though fleas were prevalent on burrowing owls and the potential for a relationship with burrowing owls as a phoretic host of infected fleas exists, we found no evidence of Y. pestis in sampled fleas or in owls that harbored them. We suggest that studies similar to those reported here during plague epizootics will be especially useful for confirming these results.