Estimating vital rates of avian species is important to understand population dynamics and develop potential conservation strategies that target rates for management. Avian species have reduced potential for high annual fecundity in alpine ecosystems due to a short breeding window and harsh weather conditions. We located nests from Southern White-tailed Ptarmigan (Lagopus leucura altipetens) across six study sites in the Southern Rocky Mountains of Colorado to estimate daily nest survival from 2013–2017. We used a known-fate hierarchical nest survival model and fit several covariates, including environmental conditions representing daily weather events and shrub cover, to describe variation in daily survival and derive estimates of nest success. We located and monitored 198 nests from 128 radio-marked ptarmigan hens. The mean nest success estimated as a derived parameter from daily nest survival was 45.6% (95% credible interval [CI]: 31.2–59.6%) and ranged from 40.3% to 50.3% across sites. Variation in daily nest survival was poorly described by the covariates we fit (95% CI of most slope coefficients overlapped 0), although there was some support for a negative effect of relative elevation (nests at lower elevations within a site survived at higher rates) and a positive effect of nest age (older nests survived at higher rates). We examined how variation in nest success was likely to influence the finite rate of population growth using a simple simulation with an age-transition matrix parameterized with previously reported fecundity and survival estimates. We found that the finite growth rate was predicted to increase 18.7% when evaluated from the lower to upper 95% CI estimated values of nest success, conditional on the other vital rates used in our simulation. We discuss the broader implications of these findings in the context of managing for nest survival of Southern White-tailed Ptarmigan.

Band-tailed pigeons (Patagioenas fasciata) congregate at and use mineral sites (mineralized water or soil) primarily within the breeding range of the Pacific Coast band-tailed pigeon (P. f. monilis), but the Interior band-tailed pigeon (P. f. fasciata) generally does not exhibit this behavior. Differences in supplemental minerals used between subspecies could be explained by differences in the diet (grit and foods) of these birds between their regional ranges. We necropsied 1,169 band-tailed pigeons and sampled 35 primary foods of band-tailed pigeons in the United States range of the Interior (AZ, CO, NM, and UT) and Pacific Coast (CA, OR, and WA) band-tailed pigeons to evaluate regional differences in mineral and moisture in gastroliths and food items, and the association of grit with food items consumed. Grit and food types consumed by pigeons were similar between subspecies, but grit amount varied with foods consumed and food items varied with season and region in association with plant distribution and phenology. Gastroliths contained small amounts (<= 3.3% each) of calcium, sodium, and potassium and were smooth and polished, indicating relatively long-term retention and use primarily in food abrasion. Foods universally contained low amounts of sodium (0-328 ppm), moderate calcium (100-7,250 ppm), and high potassium (4,400-22,600 ppm), but varied somewhat with food type (e.g., acorns, berries, cultivated grains, pine seeds). Moisture and mineral content of grit consumed and foods used by Interior and Pacific Coast band-tailed pigeons did not differ between their regional ranges in explaining differential use of supplemental mineral sites. Band-tailed pigeon's use of supplemental sodium may not be necessary for survival and reproduction because Interior pigeons rarely use supplemental mineral sites unlike Pacific Coast pigeons despite similarities in moisture and mineral content of grit and foods consumed. Band-tailed pigeons are able to concentrate sodium and calcium in crop milk to meet the nutritional needs of hatchlings despite a diet low in sodium and calcium.

Numerous studies provide estimates of nesting propensity rates (proportion of females attempting to nest at least once in a given year) for greater sage-grouse Centrocercus urophasianus. However, females may initiate nests without being detected during the course of normal research, leading to negatively biased estimates. We evaluated nesting propensity rates (rate of females laying >= 1 egg/y) by examining ovaries from 941 female sage-grouse collected at hunter-check stations in North Park, Colorado, during 1975-1984. Mean rate estimates of nesting propensity were lower for yearlings (0.926, 95% CI = 0.895-0.948) than adults (0.964, 95% CI = 0.945-0.978). We did not attempt to estimate laying rates (number of eggs laid per year) because they were likely unreliable. Nesting success-estimated as the probability of females producing a successful clutch in a given year based on primary feather replacement from hunter-harvested wings-was lower for yearlings (0.398, 95% CI = 0370-0.427) than adults (0.571, 95% CI = 0.546-0.596). There were more chicks per female produced when nesting propensity rates were high, indicating nesting propensity rates correlate with the number of juveniles in the autumn population. Both nesting propensity rates and nesting success were positively related to precipitation during the lekking and brood-rearing seasons, respectively. Nesting propensity rates were positively related to spring abundance (as measured from annual lek counts), but nesting success was unrelated to spring abundance. A range-wide estimate of an unadjusted, apparent nesting propensity rate available from a previous study was approximately 7% lower than the North Park population. Postovulatory follicles provide a direct source of information on nesting propensity rates estimated from hunter-harvested sage-grouse. These estimated rates may prove useful to gain insights into annual variation of hunted populations' reproductive efforts.

Patterns in body mass and molt provide useful information about how birds interact with their environment and act as potential explanatory variables in behavioral and demographic differences among sex and age classes. We collected population data for Greater Sage-Grouse (Centrocercus urophasianus) in Jackson and Moffat counties, Colorado, from 1973 to 1993 prior to consideration of federal listing of the species under the Endangered Species Act. Using spotlights and long-handled nets, we located and captured Greater Sage-Crouse while they roosted (primarily at night). Each captured bird was banded and released near the site of capture. We recorded the molt status of the primary flight feathers (1-10) and measured body mass. Most birds were captured prior to and during breeding (March-May) and during the brood-rearing period (July-September). Replacement of primary (P) flight feathers was initiated in May, starting with the first primary (P1), and was completed at the last primary (P10) by late September for adult (ASY) and subadult (SY) males, and by October for hatch-year males (young of the year, HY). Adult and subadult females did not replace primary flight feathers until after nesting was completed, starting in June. Primaries 1-3 were replaced within a week of the hen leaving the nest site, with or without chicks. Primary replacement then was similar to that of adult and subadult males and was completed before early October. A few late-nesting hens retained PIO and P9 into early October. Young of the year rarely replaced juvenal P9 and PIO in the year of hatching. Body mass of males peaked ((x) over bar = 3000 g for adults, and 2200 g for yearlings) in March-April when birds were primarily feeding on sagebrush and then decreased to late May and reached seasonal lows ((x) over bar = 2200 g) in July-August. Mass of females peaked in April ((x) over bar = 1640 g) and decreased to 1250 gin August.

ABSTRACT Species distributions are influenced by climate and topography in alpine ecosystems, yet resource selection studies of alpine species are uncommon. Basic characteristics of habitats used by alpine‐endemic white‐tailed ptarmigan ( Lagopus leucura ) have been described to explain foraging behavior, morphology, and survival in many alpine regions; however, there is a lack of information about fine‐scale habitat selection for nesting and brood‐rearing, particularly in the southern extent of the species’ range. Few studies have tested whether nest and brood‐site selection by white‐tailed ptarmigan are influenced by fine‐scale components such as vegetation and arthropod communities. We assessed these fine‐scale habitat characteristics analyzing paired use‐available resource selection for nest ( n = 61) and brood ( n = 54) sites. We used conditional logistic regression for data collected in 2 alpine areas along the Front Range of Colorado, USA, during 2014 and 2015. We evaluated resource selection at larger (patch) and finer (nest site) scales. Nest‐site selection at the patch scale was best predicted by cover (%) of forage forbs, rock and gravel, and shrubs. Forage forb cover explained more variation in our top nest model at the patch scale when compared to models with specific vegetation species. Females placed their nests along elevational gradients but more so at lower elevations and selected for less graminoid cover at the nest‐site scale. Brood habitat selection at the patch level was influenced by cover (%) of rock and gravel and proximity to shrubs (m). Analysis of a subset of our brood data ( n = 34) revealed females selected brood habitat that contained high arthropod abundance (e.g., Cicadellidae) over high vegetation cover, likely as a response to meet dietary requirements of chicks. Our results demonstrate how and where white‐tailed ptarmigan are currently selecting these different breeding sites in Colorado's alpine, giving us insight into consequences this alpine‐endemic bird may face if their breeding habitat is altered. © 2019 The Wildlife Society.

Phenological mismatches-defined here as the difference in reproductive timing of an individual relative to the availability of its food resources-occur in many avian species. Mistiming breeding activities in environments with constrained breeding windows may have severe fitness costs due to reduced opportunities for repeated breeding attempts. Therefore, species occurring in alpine environments may be particularly vulnerable. We studied fitness consequences of timing of breeding in an alpine-endemic species, the white-tailed ptarmigan (Lagopus leucura), to investigate its influence on chick survival. We estimated phenological mismatch by measuring plant and arthropods used by ptarmigan in relation to their timing of breeding. We monitored 120 nests and 67 broods over a three-year period (2013-2015) at three alpine study sites in the Rocky Mountains of Colorado. During this same period, we actively monitored food resource abundance in brood-use areas to develop year and site-specific resource phenology curves. We developed several mismatch indices from these curves that were then fit as covariates in mark-recapture chick survival models. A correlation analysis between seasonal changes in arthropod and food plant abundance indicated that a normalized difference vegetation index (NDVI) was likely the best predictor for food available to hens and chicks. A survival model that included an interaction between NDVI mismatch and chick age received strong support and indicated young chicks were more susceptible to mismatch than older chicks. We provide evidence that individual females of a resident alpine species can be negatively affected by phenological mismatch. Our study focused on individual females and did not examine if phenological mismatch was present at the population level. Future work in animal populations occurring in mountain systems focusing on a combination of both individual- and population-level metrics of mismatch will be beneficial.

Knowledge of population fluctuations of Aleutian Islands Rock Ptarmigan (Lagopus muta) is limited because of isolation and access. We reviewed the available but limited data on ptarmigan counts on islands in North America and evaluated the use of point counts to estimate changes in apparent numbers of Rock Ptarmigan on three islands (Adak, Amchitka, and Attu) in the Western Aleutian Islands in Alaska. We developed a standardized protocol to count numbers of Rock Ptarmigan (males and females) seen and/or heard on 5-minute point counts at 0.8 km intervals along marked global positioning system routes on Adak (2015–2017), Amchitka (2015), and Attu (2015) islands. Apparent densities based on Rock Ptarmigan seen and/or heard at 98 stops on 10 routes varied and were highest (1.9 birds per stop in 2015, 1.4 in 2016, and 1.0 in 2017) on Adak, lower (0.4 birds per stop) on Amchitka, and lowest (0.0 birds per stop) on Attu in late May–early June 2015. These island populations represent three subspecies and unique conservation units. Continuation of point-count surveys of these three subspecies in future years will provide baseline data over time and lead to a better understanding of any fluctuations in and synchrony among Rock Ptarmigan populations on these islands. This information is necessary for both theoretical (how are ptarmigan breeding populations regulated on islands) and practical reasons (identifying the optimal period for possible translocation to islands where ptarmigan were extirpated by introduced Arctic Fox [Vulpes lagopus]).

The delineation of intraspecific units that are evolutionarily and demographically distinct is an important step in the development of species-specific management plans. Neutral genetic variation has served as the primary data source for delineating “evolutionarily significant units,” but with recent advances in genomic technology, we now have an unprecedented ability to utilize information about neutral and adaptive variation across the entire genome. Here, we use traditional genetic markers (microsatellites) and a newer reduced-representation genomic approach (single nucleotide polymorphisms) to delineate distinct groups of white-tailed ptarmigan (Lagopus leucura), an alpine-obligate species that is distributed in naturally fragmented habitats from Alaska to New Mexico. Five subspecies of white-tailed ptarmigan are currently recognized but their distinctiveness has not been verified with molecular data. Based on analyses of 436 samples at 12 microsatellite loci and 95 samples at 14,866 single nucleotide polymorphism loci, we provide strong support for treating two subspecies as distinct intraspecific units—L. l. altipetens, found in Colorado and neighboring states; and L. l. saxatilis, found on British Columbia’s Vancouver Island—but our findings reveal more moderate patterns of divergence within the remainder of the species’ range. Results based on genetic and genomic datasets generally agreed with one another, indicating that in many cases microsatellite loci may be sufficient for describing major patterns of genetic structure across species’ ranges. This work will inform future conservation and management decisions for the white-tailed ptarmigan, a species that may be vulnerable to future changes in climate.

We analyzed banding (3259) and recovery (six years) data from a hunted population of greater sage-grouse Centrocercus urophasianus in northwestern Moffat County, Colorado to examine vulnerability to hunter harvests and annual survival of adult and hatch-year (juvenile) birds. Additionally, we combined the recovery data with hunter-harvested wings and applied the Lincoln estimator to provide unbiased estimates of tertiary sex-ratio. Our results yielded the following findings: juveniles were harvested at twice the rate of adults, but harvest vulnerability was similar between adult males and females. Annual survival of juveniles was highly variable but similar between sexes. Sex ratios of adults and juveniles largely conformed to previously assumed proportions despite having adjusted those estimates with harvest rates. We suggest there is potential to effectively model populations of game birds using a combination of band recovery and hunter-harvested samples of species that can be readily captured, marked and have reasonable harvest rates.

‘‘Before there was Google, the solution was to ask Ron Ryder,’’ a colleague recently remarked. How true: He was a fountain of information about all things avian, as well as related topics. Dr. Ronald Arch Ryder, a longtime professor of wildlife biology at Colorado State University (CSU), passed away on August 2, 2016, in Fort Collins, Colorado. He was born in Kansas City, Kansas, on February 3, 1928, to Florence and Wendell Ryder, and attended high school in Kansas City, Missouri. He spent much of his younger life, during breaks from school, helping on farms of relatives near Colby, Kansas, and his interest in birds started early (in grade school, ca. 1934). He joined the Burroughs Nature Club in Kansas City, Missouri, and participated in Christmas Bird Counts in the early 1940s. We remember him describing some of his birding trips to Swope Park in Kansas City, Missouri. World War II was under way when Ron graduated from high school at 17 and enlisted in the U.S. Army, which sent him to the University of Illinois and the University of Wyoming to study engineering. When the war ended, Ron worked for the U.S. Forest Service for a summer in the Medicine Bow Mountains in southern Wyoming. At this time he started keeping a journal, which he continued well into retirement. He then enrolled at Colorado State College of Agriculture and Mechanic Arts (now CSU) and completed a B.S. in game management in 1949. He immediately launched into an M.S. program, also at CSU, in the Cooperative Wildlife Research Unit under Dr. Lee Yeager, studying waterfowl production in the San Luis Valley, Colorado. He completed his M.S. and was employed by the Colorado Game and Fish Department as a research biologist. The war in Korea was on the horizon, and he enlisted in the U.S. Navy, becoming a self-described ‘‘120-day wonder’’ after a brief period in Officer Candidate School. Thus, he was sent to the Arctic for 32 months to serve as an ensign on an icebreaker and also on an attack cargo ship. He was thoroughly impressed by the seabirds as well as by other marine animals there. After active duty (1951–1954), Ron continued in the Naval Reserve until retirement as a lieutenant commander in 1974. Ron returned to Colorado after military service and met Audrey Teele, who later became his wife, on a Colorado Mountain Club snowshoe trip. He used the G.I. Bill to earn a Ph.D. at the Utah State University Ronald A. Ryder in 1994. Photo credit: D. A. Leatherman

Abstract. We found 13 specimens (11 museum skins, 2 skeletons) of White-tailed Ptarmigan (Lagopus leucura) collected in Wyoming. All were collected in the 1911–1967 period: 12 from the Snowy Range (Albany County) and one from near Encampment (Carbon County). The last verifiable observations of White-tailed Ptarmigan in the Snowy Range were 2 separate reports in 1974, both of 4 individuals in the same location. A more recent report of 2 birds north of the Snowy Range on 15 October 2005 has also been verified. None of the verifiable observations or collection events was of more than 6 individuals. Most of the published observation records from areas other than in the Snowy Range were of single individuals and were likely Dusky Grouse (Dendragapus obscurus). Field investigations in the Beartooth Plateau, Bighorn Mountains, and Wind River Range outside of the Snowy Range revealed no ptarmigan or their sign. However, all unoccupied areas examined could potentially support White-tailed Ptarmigan populations, with the Wind River Range being most suitable, followed by the Bighorn Mountains and Beartooth Plateau. The lack of White-tailed Ptarmigan in alpine areas in Wyoming outside of the Snowy Range is most likely related to geologic events in the late Pleistocene period and not in the Holocene.

Data to inform population assessment of the Interior subspecies of band-tailed pigeon, Patagioenas fasciata fasciata (breeding range from Colorado and Utah south into Sierra Madre Occidental of Mexico), have been lacking despite substantial past banding efforts. We used a data set of more than 26,000 bandings from Colorado, with 3,500 live recaptures and 780 recoveries from the harvest of banded individuals to estimate annual survival, fidelity, and harvest rates. Most birds were harvested in Colorado (62%) followed by Mexico (18%); New Mexico (16%); Arizona (3%); and 1% or less each in California, Washington, and Utah. On average, each year 15% (range 0-30%) of surviving band-tailed pigeons did not return to Colorado. From 1969 to 1981 mean annual survival was 0.633 (standard error [SE] = 0.031) for hatch-year and 0.719 (SE = 0.016) for after-hatch-year birds, with a mean annual recovery rate of 0.015 (SE = 0.002) for hatch-year and 0.011 (SE = 0.001) for after-hatch-year birds. From 1970 to 1974, mean annual abundance of band-tailed pigeons in Colorado on 1 September was 59,911-88,290. These data provide a baseline for additional data collection for band-tailed pigeons in the range of the Interior subspecies.

Animal populations occurring at high elevations are often assumed to be in peril of extinctions or local extirpations due to elevational-dispersal limitations and thermoregulatory constraints as habitats change and warm. However, long-term monitoring of high-elevation populations is uncommon relative to those occurring at lower elevations, and evidence supporting this assumption is limited. We analyzed 45 years of reproductive data for two Colorado populations of white-tailed ptarmigan (Lagopus leucura), an alpine-endemic species with restricted distribution in western North America. Seasonal temperatures measured by the number of growing degree days warmed significantly at our study sites for pre-nesting, nesting, and brood-rearing seasonal periods (mean advance of 8 growing degree days per decade), and both populations advanced their reproductive phenology over the study period based on median hatch dates (median advance of 3.7 and 1.9 days per decade for the northern and southern sites, respectively). Reproductive performance measured by the number of chicks per hen declined significantly at one study site but not the other, and differences between sites may have been due to habitat degradation at one study area. Annual variability in chicks per hen was large at both sites but only weakly related to seasonal weather. An index of precipitation and temperature during the brood-rearing period was the best predictor for reproductive success with warm and dry conditions relating positively to number of chicks per hen. Our results provide evidence for two alpine ptarmigan populations that are remarkably invariant to fluctuations in seasonal weather with respect to reproductive success as measured by number of chicks per hen in the breeding population. These results are surprising given the general perception of alpine animal populations as being highly sensitive to warming temperatures.

Each year, the Wilson Ornithological Society recognizes a member who has provided extensive service to the Society with the William and Nancy Klamm Service Award. The award honors the memory of extensive service and commitment to the Wilson Ornithological Society shown by Bill and Nancy Klamm, who generously supported the Society with both their time and a substantial financial bequest. This year’s award is being given to a member whose involvement with the Society overlapped Bill’s work on the Audit Committee. In 1988, Edward H. Burtt, Jr., known to all as Jed, began his regular attendance at WOS meetings. In 1989 he was elected to a two-year term as an elective councilor. The following year, he co-chaired the scientific program committee for the Wilson meeting in Norton, Massachusetts. In 1993, he was elected as Second Vice President of the Society and after serving as Second Vice President and then First Vice President, he served as President from 1997 to 1999. During this time, he organized an interactive workshop on undergraduate teaching in ornithology and a workshop in the management of ornithological collections at teaching institutions and nature centers. Jed wanted to provide resources for teaching faculty, so after the former workshop, Jed shepherded the process of converting those exercises to a laboratory manual that was posted online free of charge. As First Vice President of the Wilson Society, Jed initiated the Margaret Morse Nice medal and plenary lecture in 1997. The award recognizes a lifetime of ornithological research and work as a mentor. Jed’s introductions to these lectures have been quite inspiring by providing information about Margaret Morse Nice’s life and her influential career as a scientist. As he introduced the recipients, he was able to find direct connections between their career and Nice’s. The detailed and individual information in the introductions of the recipients were typical of Jed’s work in the Society, integrating a commitment to high quality ornithological research with personal touches and a family feel. Although he completed his work on the executive committee in 1999, Jed continued to be active both on the Council and in the Society. In fact, Jed hosted the 2003 Wilson meeting at Ohio Wesleyan University, in joint session with the Association of Field Ornithologists and the Clark Ornithology Symposium. Jed’s interest in creating a special feel for the students at the meeting included scheduling the poster session during an extended lunch period, with lunch in the poster venue included in the registration. His interest in providing additional opportunities for students and professionals to interact included having the Ohio Wesleyan Jazz Band perform at the end of the banquet. Most years, Jed willingly served on the committee that judged student presentations and he served for many years on the nominating committee, helping to identify ornithologists who would be able to bring fresh perspectives to Council and many who could benefit from mentoring relationships with other Council members. Perhaps Jed’s most influential involvement with the Wilson Society is his attendance at meetings with students. Most meetings included several students from Ohio Wesleyan. His mentorship of

ABSTRACT We redefine and clarify procedures to classify sex and age (juveniles, yearlings, adults, and breeding‐age) of greater ( Centrocercus urophasianus ) and Gunnison sage‐grouse ( C. minimus ) from wings. Existing keys for greater sage‐grouse age and sex classification do not incorporate more recent information on timing and sequence of molt or regional variation. We evaluated keys with the aid of gonadally inspected, hunter‐harvested sage‐grouse in Colorado (1973–1990) and with birds captured and measured in Washington (1992–1997) and Oregon (2008–2012). The technique is accurate and transferable among biologists who have basic training in reading a key and examining wings (primaries, secondaries, tertials, and coverts). Accurate information on sex and age of grouse, particularly during harvest, is a fundamental component of our understanding of population dynamics, which ultimately enables improved management. © 2014 The Wildlife Society.

We studied the population structure of sage-grouse (Centrocercus spp.) based on collection and analysis of 67,679 wings from hunter-harvested birds in 10 areas in Colorado and 12 areas in Oregon during 1973-1998 and 1993-2013, respectively. The harvest age structure for greater sage-grouse (C. urophasianus) varied from 42 to 63% juveniles in Colorado and 27 to 58% in Oregon. Approximately 59% of the Gunnison sage-grouse (C. minimus) harvest was juvenile. The overall adult male:female sex ratio was 28:72 for greater sage-grouse in Colorado, 41:59 (this includes an unknown proportion of yearlings) for greater sage-grouse in Oregon, and 34:66 for Gunnison sage- grouse in Colorado. Proportions of females increased in all fall populations from juvenile to yearling to adult age classes. Estimated breeding success was similar for greater sage- grouse in Colorado (47%) and Oregon (49%), but Gunnison sage-grouse appeared to have higher (60%) breeding success. The average number of juveniles in the harvest per breeding-age female varied from 1.2 to 2.4. There was high annual variation within and among areas. Composite estimated annual survival varied from 46 to 48% for adult males and 56 to 59% for adult females.

We reviewed the literature and observations of the occurrence and status of the White-tailed Ptarmigan (Lagopus leucura) in New Mexico. Historical reports were infrequent, likely because of an inadequate system for recording observations from the public, although by 1928 biologists had a good understanding of the distribution and status of the species in the state. By 1980, ptarmigan persisted in small numbers in the northern portion of the New Mexico range but were uncommon or absent in the southern portion of the range, prompting a transplant of White-tailed Ptarmigan from Colorado into the southern area in 1981. Following that successful transplant, observations initially increased and subsequently continued at a relatively low level with most reports coming from the southern portion but including others from throughout the historical range. White-tailed Ptarmigan are localized in suitable habitats, but their abundance in New Mexico may be affected by the decreasing size of alpine snowfields in summer, grazing in areas dominated by willow (Salix spp.), and the shift to a warmer and drier climate.