Erik K. Fritzell

Habitat use by prairie raccoons during the waterfowl breeding season

Mobility and habitat use of raccoons (Procyon lotor) in an intensively farmed area of the prairie pothole region were studied during the waterfowl breeding seasons (April-July) of 1973-75. Over 5700 locations of 30 raccoons were analyzed. Movement patterns varied with sex, age, and reproductive status. Adult males moved regularly throughout slightly overlapping ranges that averaged 2560 ha. Yearling males dispersed during May-June but their movements before and after dispersal were similar. Parous or pregnant females (mostly adults) had ranges averaging 806 ha but their movements were confined to smaller areas near the litter site after parturition. Nulliparous yearling females did not disperse and their ranges averaged 656 ha. Building sites, wooded areas, and wetlands were the only habitats preferentially used both at night and during the day. Eighty-one percent of all nocturnal locations and 94 percent of all diurnal locations were in these 3 habitats which comprised only 10 percent of the study area. Use of building sites decreased concomitantly with increased use of wetlands. Upland habitats were seldom used. J. WILDL. MANAGE. 42(l):118-127 Raccoons have increased dramatically in North Dakota in the past 50 years. Prior to 1920 they occurred in low densities, restricted to the wooded valleys (Bailey 1926). Today they are common throughout the state in all habitat types. Waterfowl nesting success in the prairie pothole region has declined markedly during the past 35 years (Miller 1971) and raccoon predation has been considered a major influence in that decline (Duebbert and Kantrud 1974, Nelson and Duebbert 1974, Trauger and Stoudt 1974). There have been numerous studies of raccoon mobility and habitat use (Ellis 1964, Geis 1966, Schnell 1969-70, Johnson 1970, Urban 1970, Schneider et al. 1971, VanDruff 1971, Cowan 1973), but none have been conducted in intensively farmed prairie habitat. The objective of this study was to describe the movements, home ranges, and habitat use characteristics of raccoons in the prairie pothole region during the waterfowl breeding season. A. B. Sargeant provided encouragement, guidance, and assistance in this study from its inception through its editorial phase. R. J. Greenwood freely exchanged data collected by R. D. Eliason and himself. D. A. Davenport, D. B. Siniff, D. H. Johnson, and E. Multer provided valuable advice and assistance. Northern Prairie Wildlife Research Center financed the study.

Habitat use by migrant sandhill cranes in Nebraska

The principal spring staging areas of the midcontinent population of sandhill cranes (Grus can- adensis) are along the Platte and North Platte rivers in south-central Nebraska. Most of these lands are privately owned and managed for corn and cattle production. Diurnal habitat use by radio-tagged cranes was primarily in cropland (55%), native grassland (28%), and tame hayland (15%). Ninety-nine percent of the cropland use was in cornfields; 55% as grazed stubble, 36% as disced, cultivated, and plowed stubble, 7% as ungrazed stubble, and 1% unclassified. Grazed pastures accounted for 93% of the grassland locations and mowed alfalfa fields 77% of the tame hayland locations. Other habitats were seldom used. Time budget analyses indicated that cranes, while in croplands, grasslands, and haylands, spent 35, 36, and 50% of the time foraging, respectively Cranes roosted in the shallows and on nearby sandbars of about 111 km of river channel. Cranes usuallv roosted where the channel was at least 150 m wide and avoided stretches narrower than 50 m. Height of woody vegetation along shorelines and on islands influenced where cranes roosted when unobstructed channel width was less than 150 m; bridges or roads adjacent to the channel also reduced use by about half. Management recommendations are made for maintaining suitable habitat for sandhill cranes on their staging areas in Nebraska. J. WILDL. MANAGE. 48(2):407-4 17 Approximately one-half million sand- hill cranes (four-fifths of the continental population) gather annually along the Platte and North Platte rivers in Nebraska during March and early April while en route to their breeding grounds in central and arctic Canada, Alaska, and Siberia (US. Fish and Wildl. Serv., unpubl. data). Water developments in the upper Platte River Basin during this century have dras- tically reduced flows, causing major changes in channel width in the Big Bend reach (Williams 1978) where most of the cranes gather. Extensive encroachment by woody vegetation has accompanied chan- nel shrinkage (Currier 1982), and pro- posed developments would cause addi- tional habitat degradation and loss (Krapu et al. 1982), raising concern for the well-

Ruffed grouse winter roost site preference and influence on energy demands.

We located winter roosts of ruffed grouse (Bonasa umbellus) in Missouri with radiotelemetry and determined roost type preference. Thermostatic energy demands in 4 roost types were measured with a heated taxidermic mount calibrated with metabolic rates of captive grouse. Ruffed grouse preferred to roost in the canopies of eastern red cedar (Juniperus virginiana) and avoided roosting in deciduous cover. Roost sites had higher woody stem densities (9 = 5,494 stems/ha) than random plots (9 = 4,236 stems/ha). We predicted ruffed grouse metabolic rates (? 0.75%), at ambient temperatures (Ta) of -20 to 0 C, from power (P,,,) used by the heated taxidermic mount. Standard operative temperature (T,) was elevated 7.3, 2.6, 2.9, and -0.3 C in snow roosts, cedar tree roosts, cedar ground roosts, and deciduous roosts, respectively, above that at an open site for T, of -20 to 0 C. When wind speed was 3 m/second in the open, T, was elevated a mean of 12.9, 7.0, 6.7, and 2.5 C, in snow, cedar tree, cedar ground, and deciduous roosts, respectively. These increases in Tes resulted in a 33, 19, 18, and 6% reduction in metabolic rate in snow roosts, cedar tree roosts, cedar ground roosts, and deciduous roosts, respectively, from that in the open at -20 to 0 C and 3 m/second wind speed. About 40% of the elevation in T, resulted from reduced convection inside roost sites and 60% from a more favorable radiation balance in roosts. Low coniferous vegetation provided thermal benefits that may be important because snow roosts were rarely available. J. WILDL. MANAGE. 52(3):454-460 Birds show physiological and behavioral adaptations to cold stress. Physiological adaptations include increasing metabolic rate, accumulating fat reserves, acclimatizing, decreasing body temperature, and developing winter plumages. Behavioral adaptations include migrating, selecting micro-climates, changing activity patterns, adjusting posture, and responding as a group (Calder and King 1974). Ruffed grouse exhibit several of these adaptations. Grouse increase their metabolic rate in response to cold (Thompson and Fritzell 1988). Increased energy demands in winter may be met in part by fat reserves accrued during summer and fall (Norman and Kirkpatrick 1984) or by regular feeding (Thomas et al. 1975). Heat loss and energy demands are reduced by a lower than average lower-critical temperature and nocturnal depression in body temperature (Thompson and Fritzell 1988). Grouse reduce heat loss behaviorally through micro-habitat selection (Thomas et al. 1975). Snow roosting is documented in ruffed grouse (Bump et al. 1947, Grange 1948) and other tetraoninae can maintain a thermoneutral microenvironment in snow burrows (Hoglund 1980, Marjakangas et al. 1984). During cold weather, ruffed grouse also use conifer cover (Bump et al. 1947) where radiant temperature is higher (Brander 1965). Optimal winter cover for ruffed grouse in northern forests is provided by young aspen stands with 14,000-20,000 stems/ha, where grouse are relatively safe from predation, deep snow provides roosting cover with thermal benefits, and nearby mature aspens (Populus spp.) provide a winter food resource (Gullion 1977). In the southern portion of their range, grouse are often exposed to cold temperatures, but snow cover is insufficient for snow roosting and aspen is bsent. Grouse must use other foods and rely on vegetative cover to reduce thermostatic demands, increase their food consumption, or use accrued energy reserves. We determined microhabitat preferences of ruffed grouse for nocturnal wirter roost sites in Missouri, near the southern terminus of their range. Thermostatic energy demands and heat loss of grouse roosting in different microhabitats were compared. J. E. Roberts provided valuable assistance with the design and construction of the heated taxidermic mount. D. E. Figert, D. Hoffman, and D. G. Kusmec provided field assistance. M. R. Ryan, T. V. Dailey, and F. L. Thompson reviewed the manuscript. Financial support for this study was provided by the Ruffed Grouse Society, the U.S. Forest Service, and the Missouri Department of Conservation. This is Contribution 10381 of the Missouri Agricultural Experiment Station Project (J. Pap. 189) and the

Drumming, nesting, and brood habitats of ruffed grouse in an oak-hickory forest

We described drumming, nesting, and brood habitats selected by ruffed grouse (Bonasa umbellus) in central Missouri oak (Quercus spp.)-hickory (Carya spp.) forest with no recent timber harvest. Habitat at drumming sites differed from that at randomly located non-drumming logs (P < 0.001). Drumming logs had higher coniferous and deciduous shrub stem densities, coniferous canopy closure, understory foliage density, total woody stem densities, and slope position than random logs. Deciduous tree density and deciduous canopy closure were lower at drumming logs. Habitat at brood locations differed from that at random sites (P < 0.001). Brood locations had higher percent ground cover, deciduous shrub stem density, and total woody stem densities; a lower number of coniferous trees; and a lower slope position. Habitat at nest sites was not different from that at randomly selected sites (P = 0.843). Drumming grouse and broods selected habitats with higher stem densities than the average available to them but lower than selected by grouse in other portions of their range. Significant differences existed among habitats found at drumming, nesting, brood, and random sites (P < 0.001). Drumming and brood habitat had the highest number of total woody stems/ ha. Drumming habitat had the highest number of coniferous shrub stems/ha and highest slope position. Brood sites had the lowest number of coniferous shrub stems/ha and the lowest slope position. Nest and random locations had the lowest number of total woody stems/ha and, on average, occurred mid-slope. J. WILDL. MANAGE. 51(3):568-575 There are few published studies of habitat use by ruffed grouse along the southern periphery of their range where aspen (Populus spp.) is not dominant. Ruffed grouse used areas of brushland or forest stands with a dense understory of shrubs for drumming sites in oak-hickory forests in Ohio (Stoll et al. 1979), Georgia (Hale et al. 1982), and Indiana (Backs 1984). In oak-hickory forests of Tennessee ruffed grouse used habitat similarly during fall and winter (White and Dimmick 1978). Habitat use by drumming males, nesting hens, and broods was described in Minnesota (Gullion 1977) and in New York (Bump et al. 1947). Stauffer and Peterson (1985) compared habitat use between drumming males and broods, and delineated seasonal use by all sex-age groups in Idaho. Those studies emphasized the importance of juxtaposition of several habitat types because ruffed grouse change patterns of habitat use throughout the year. No studies have compared drumming, nesting, and brood habitat in oak-hickory forests of the central United States. Such information should enhance forest management for ruffed grouse in this portion of their range and should enable us to evaluate suitable habitats for reintroductions. We investigated habitat characteristics of drumming sites, nesting sites, and brood locations in an unmanaged oak-hickory forest in central Missouri to characterize and differentiate between those aspects of breeding habitat. Habitat use described in this paper was obtained during a pretreatment (no timber harvest) study of a long-term project investigating the impact of clearcutting. Financial support for this study was provided by The Ruffed Grouse Soc., the U.S. For. Serv., an E. K. Love Fellowship, and the Mo. Dep. Conserv. This is a contribution of Mo. Agric. Exp. Stn. Proj. 189, J. Pap. 10056 and the Mo. Coop. Fish and Wildl. Res. Unit (U.S. Fish and Wildl. Serv., Mo. Dep. Conserv., Univ. Missouri-Columbia, and Wildl. Manage. Inst., cooperating). We thank T. F. Glueck, K. J. Haroldson, B. W. Hunyadi, D. G. Kusmec, K. P. McDowell, and G. E. Schreckengast for assistance in various aspects of this study. G. W. Gullion, M. R. Ryan, and J. M. Sweeney provided constructive comments on the manu-

Effects of river recreationists on green-backed heron behavior

10% to 15% annually (Marnell et al. 1978). The major form of recreational activity on the ONSR is canoe float trips. Estimated use increased from 40,000 canoe floater days in 1968 to 243,000 in 1977; over 500,000 canoe floater days are projected for 1985 (Marnell et al. 1978). The objective of this study was to determine the behavior of green-backed herons in relation to the level of recreationist activity on the ONSR.

Bird densities and diversity in clearcut and mature oak-hickory forest.

Describes nongame bird densities and diversity in a central Missouri oak-hickory forest 1 year before and 3 years after portions were clearcut. Discusses changes in species density and diversity and their management implications.

Movements and habitat use of Franklin's ground squirrels in duck-nesting habitat

We used radio telemetry to determine the movements and habitat use patterns of adult Franklins ground squirrels (Spermophilus franklinii) on a 152-ha Waterfowl Production Area (WPA) in central North Dakota. The squirrels were diurnal. Mean total distance traveled daily was 213 and 153 m for males and females, respectively. Movements of females decreased during gestation and lactation; those of both sexes decreased prior to immergence. Annual home ranges averaged 24.6 and 8.7 ha for males and females, respectively; biweekly home-range sizes exhibited seasonal patterns. Ground squirrels restricted their activities almost exclusively to the WPA. Use of blocks of herbaceous cover on the WPA could not be explained by vegetation height, density, or litter depth. Ground squirrel movements were influenced by locations of burrow systems and patchily distributed food resources. Periodic cultivation or removal of herbaceous vegetation may inhibit use of upland duck-nesting habitat by ground squirrels. J. WILDL. MANAGE. 53(2):324-331 Planting dense nesting cover (DNC) to provide upland-nesting ducks with increased security from nest predators is a common practice on areas managed for duck production in North Americas prairie pothole region (Duebbert and Kantrud 1974, Duebbert and Lokemoen 1976, Higgins and Barker 1982). Although DNC is attractive to some duck species, its importance to the activities of predators of duck eggs is undetermined. The Franklins ground squirrel inhabits dense cover throughout much of the prairie pothole region (Hall 1981). It is a predator of duck eggs (Sargeant et al. 1987) and has been implicated in severe predation on duck nests in fields of DNC in east-central North Dakota (Greenwood 1986). Past studies of Franklins ground squirrels have emphasized natural history (Sowles 1948, Haggerty 1968, Haberman and Fleharty 1971, Iverson and Turner 1972, Murie 1973), but little information is available on their movements and use of habitat. Our objective was to describe the movements and use of habitat by Franklins ground squirrels on a federally owned WPA where upland habitat was primarily DNC fields of different ages. We thank D. H. Johnson and J. N. Burroughs for help with data processing, R. J. Greenwood, M. A. Sovada, and D. J. Norris for field assistance and helpful suggestions throughout the study, and H. F. Duebbert and D. H. Johnson for comments on manuscript drafts. The study was funded by the U.S. Fish and Wildlife Service, Northern Prairie Wildlife Research Center; the Missouri Cooperative Fish and Wildlife Research Unit; and the Edward K. Love Foundation. The U.S. Fish and Wildlife Service, Missouri Department of Conservation, University of Missouri-Columbia, and the Wildlife Management Institute contribute to the Missouri Cooperative Fish and Wildlife Research Unit. This is Journal Series 10008, Missouri Agricultural Experiment Station Project 189.

Ruffed Grouse Metabolic Rate and Temperature Cycles

We measured the effects of temperature and wind on ruffed grouse (Bonasa umbellus) metabolic rates in an open-circuit respiration system with a closed-circuit wind tunnel. Metabolic heat production was linearly related to operative temperature (Te) below a lower critical temperature of 1.5 ? 0.99 (SE) C. Standard metabolic rate was 3.2 + 0.11 W (66.1 kcal/day) for a grouse of mean weight (607 g). Metabolic rate was related linearly to wind speed 05. The regression predicting net metabolic heat production (metabolic heat production evaporative cooling [M E]) was 3.1589 0.1176T, + 0.4141(wind speed)s5 (r2 = 0.98, P mean low nocturnal cloacal temperature (38.3 + 0.54 C) (P = 0.006). J. WILDL. MANAGE. 52(3):450-453 Ruffed grouse are distributed throughout north temperate and subarctic North America (Aldrich 1963) where winter thermostatic energy demands are potentially high. Grouse may feed frequently (Thomas et al. 1975) or use fat reserves (Norman and Kirkpatrick 1984) to meet winter metabolic demands. Rasmussen and Brander (1973) reported effects of cold temperatures on ruffed grouse metabolic rates under free convection (no wind). Effect of forced convection on heat loss by birds has been estimated from predictions based on heat transfer theory of cylinders and flat plates (Calder and King 1974, Robinson et al. 1976). Only a few studies have directly measured effects of forced convection on birds (Gessaman 1972, Robinson et al. 1976, Bakken et al. 1981), and none have been on galliforms. Most birds exhibit daily cyclical changes of 1-3 C body temperature between active and inactive periods (Calder and King 1974). This variation may increase with food deprivation or cold exposure (Calder and King 1974, Chaplin 1976, Bucher and Worthington 1982). Body temperature of ruffed grouse has been reported (Rasmussen and Brander 1973), but cyclic fluctuations or response to cold temperatures have not been measured. We report on effects of forced convection and perature on the metabolic heat production of ruffed grouse. We also measured daily fluctuations in body temperature and changes due to cold exposure. H. C. Gerhardt, J. Maruniak, and the Division of Biological Sciences, University of MissouriColumbia provided equipment necessary for this study. We thank D. E. Figert, D. E. Hoffman, B. J. Wilson, and C. S. James for assistance with various aspects of this study. M. R. Ryan and T. V. Dailey reviewed this manuscript. Financial support for this study was provided by the Ruffed Grouse Society and the Missouri Department of Conservation. This is Contribution 10382 of Missouri Agricultural Experiment Station ProjThis content downloaded from 157.55.39.45 on Fri, 02 Sep 2016 05:31:36 UTC All use subject to http://about.jstor.org/terms J. Wildl. Manage. 52(3):1988 GROUSE METABOLIC RATES * Thompson and Fritzell 451 ect 189 and the Missouri Cooperative Fish and Wildlife Research Unit (U.S. Fish and Wildl. Serv., Mo. Dep. Conserv., Univ. Missouri-Columbia, and Wildl. Manage. Inst., cooperating).