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Management of Amphibians, Reptiles, and Small Mammals in North America

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Management of Amphibians, Reptiles, and Small Mammals in North America Abstract.-Pine seed, primarily available in the fall, Foraging Patterns of Tassel- and hypogeous fungi, potentially available in all Eared Squirrels in Selected seasons, were major food items whose consumption was associated with an increase in biweekly body Ponderosa Pine Stands1 weights of marked squirrels. Use of alternative foods such as twigs (inner bark) and apical buds occurred when these food items were unavailable. Consump- Jack S. state^,^ William S. Gaud,3W. tion of inner bark and buds was highest in winter Sylvester Allred,' and William J. Austin5 (93%) and spring (86%). Although feed tree prefer- ence was noted, widespread feeding occurred in more than half of the trees in both study sites. The resulting variability in physical evidence of foraging suggests caution in its use for indexing squirrel popu- lation densities. The tassel-eared tree squirrel (Sciurus on the ecology of Abert squirrels (S. bark in the absence of supplemental aberti) and its several subspecies, has a. aberti), Keith (1965) attributed short foods could potentially threaten a unique and apparent obligatory as- term fluctuations to changes in quan- squirrel survival during adverse sociation with Southwestern ponder- tity and quality of major food items weather conditions (Patton 1974). osa pine Pinus ponderosa). The die- assumed to be provided by pine. The obvious selection of certain trees tary dependence of these squirrels on However, high mortality in some by squirrels for inner bark consump- pine, including inner bark and buds years apparently resulted from some tion supports the assumption that of terminal twigs and both staminate factors other than food. Hall (19811, there are differences in quality of and ovulate cones, identifies the in a study of Kaibab squirrels.6. a. trees in the same stand. In a food squirrel as an herbivore having a po- knibabensis) also observed population preference study using captive Abert tentially major influence on the fluctuations. He suggested that sea- squirrels, Farentinos et al. (1981) growth and reproduction of ponder- sonal differences in food resources were able to show a significant rela- ' osa pine. Conversely, extensive har- and snowfall were potential causes of tionship between selective consump vest of pine for wood products has declines and recovery. Availability tion of inner bark and low oleoresin resulted in deterioration of the squir- and use of various food items have content. However, Pederson and rel's habitat since the turn of the cen- not been adequately studied. Welch (1987) noted a strong feeding tury (Keith 1965). Stephenson (1974) in a study of preference for trees with inner bark A number of studies have at- Abert diets discovered that fungi that was easily peeled with no appar- tempted to explain the "boom and were a major part of the total food ent relationship between inner bark bust" population fluctuations that consumed. The fungi in Stephenson's oleoresin content and "feed tree" se- seem to be characteristic of tassel- samples were identified as belonging lec tion. eared squirrels. In his observations to a subterranean group of mush- Studies on the impacts of squirrel rooms popularly called truffles (J. herbivory on ponderosa pine have States, unpublished data), which are lead to mixed conclusions. Hall 'Paper presented at symposium, Man- agement of Amphibians, Reptiles, and known to form mycorrhizal associa- (1981) and Ffolliott and Patton (1978) Small Mammals in North America. [Flag- tions with pine roots. Some of these found that heavy utilization of pine staff, AZ, July 19-21, 1988.) fungi were found to be new records twigs had negligible effect on stand 2Jack S. States is Professor of Biology, for the Southwest (States 1983,1984) productivity, although Hall demon- Department of Bidogical Sciences, North- and they were found to be primarily strated significant growth decreases ern Ar~onaUnivetsiiy, Flagstaff,AZ 860 1 1. associated with blackjack age-class of individual feed trees. Soderquist 3WilliamS. Gaud is Associate Professor of Biology, Department of Biological Sciences, ponderosa pine stands with high can- (1987) reported twig clipping to de- Northern Arizona University. Flagstaff.AZ opy densities (States 1985). crease tree growth in ecotonal stands 8601 1. A telltale sign of the activity of the of ponderosa pine. Pearson (1950) 4W.Sylvester Allred is a doctoral candi- tassel-eared squirrel is the presence and Larson and Schubert (1970) date in the Department of Biological Sci- of clipped twigs on the ground under noted extensive, but seasonally vari- ences. Northern Arizona University,Flagstaff, AZm11. a tree after the squirrel has removed able, damage to cone crops. They 5WilliamJ. Austin is a doctoral candi- the terminal shoot from a branch. were unable to determine the causes date in the Department of Biological Sci- The nutritional value of the inner of the highly variable pattern of her- ences. Northern Arizona University, Flagstaff. bark consumed by the squirrel is bivory during several years of obser- AZ8601 I. low. A diet comprised solely of inner vation. The tassel-eared squirrel's variety of diet and use of the forest has lead to differences of opinion regarding the best management plan for both squirrel and forest. Patton et al. (1985) considered tree density, size, and patchy distribution to be major factors constituting habitat quality since squirrels use pine for cover and nesting as well as for food. The purpose of this study was to determine the seasonal patterns of food resource utilization by Abert squirrels in selected ponderosa pine stands and to relate the results to squirrel population levels within the stands. The use of fungi and inner bark as major food items is discussed as it pertains to stand characteristics and the potential impact of herbivory on ponderosa pine. Study Areas and Methods Two sites in clumped, uneven-aged ponderosa pine stands were studied in areas that had not been disturbed by fire and timber harvest for the past 35 years. A 9.3 ha site was lo- cated on the property of the Lowell Observatory and adjacent to the Co- summer fall conino National Forest. The other site of 2.5 ha was located in the Mount BA -branch innerbark MI- mistletoe SC -staminate cones Elden Environmental Study area of the Coconino National Forest. The BU -terminal buds MU-mushrooms TR -truffles elevation of both sites was 2150 m, TW-twig innerbark and they were within 10 km of the CO-pine cones OP -open pine cones BA Flagstaff airport where weather data 1 used in the study was collected (NOAA 1987). Squirrels were captured, marked, and released at the observatory site. At this location there were 90 plots, 625 m2 each, in a nested trapping grid similar to the one described by Patton et a1.(1985). The grid con- tained 42 systematically spaced Tomahawk live-traps baited and set for eight daylight hours once each week. Trapping was conducted from September 1985 through June 1987 ' 3 winter spring and squirrel body weight was re- corded. Fecal pellets deposited in Figure 1 .-Percentage of feeding time by Abert squirrels for each diet item in each season traps were collected and analyzed to (from eighteen two-hour periods per season, Coconino National Forest, AZ). determine the ratio of dietary fungus (spring); Season 111, July-August entire 2.5 ha Mount Elden site was to plant matter and to identify the (summer); Season IV, September- sampled for seasonal foraging pat- fungi through spore characteristics. November (fall). terns. Permanently marked pines and Observational data on foraging Resource availability and physical oaks in 20 plots were censused yearly behavior was collected using focal signs of foraging activity were re- in September for acorn and cone pro- animal sampling (Altmann 1974). We corded over a 20-month period (June duction. Cones and acorns were observed four individually marked 1986 through January 1988). Cone counted on a quarter of the tree and males from July 1986 through No- consumption, twig clipping, and dig- multiplied by four to obtain a pro- vember 1987 as they foraged in the ging activity were documented on 26 duction estimate. Truffle production study site. Data were collected in 18 contiguous 625 m2 plots (1.6 ha) on estimates were made according to two-hour observation periods in each the observatory site. During each of States (1985). Relative mushroom of four "ecological" seasons. These the biweekly censuses, all twig abundance was determined by seasons were established by combin- remnants, cone cores, and digs counting numbers of mushrooms ing months with similar temperature (truffle excavations) were recorded present within 10 randomly placed 50 and precipitation means. The seasons with a notation of the nearest tree. m2quadrats sampled in the fall. correspond to periods of truffle pro- Trees were characterized by age-class duction: Season I, December-March (blackjack, or yellow pine) and di- (winter); Season 11, April-June ameter at breast height (DBH). The Results Resources available as food for tas- sel-eared squirrels showed consider- able annual variability (table 1). The four food items were all relatively abundant in 1983, but availability subsequently dropped. Production of cones and mushrooms was relatively high again in 1986, but there were considerably fewer truffles and acorns present than in 1983. In gen- eral, the quantity of truffle produc- tion was more consistent that it was JJASONDJFMAYJJASONDJ for the other foods. Seasonal foraging behavior of the squirrels (fig. 1) reflected changes in availability of food items. The ani- mals heavily utilized a large cone crop in 1986 before the seeds were mature, and continued to utilize it into November when the remaining seeds were released. Cone and truffle use dropped abruptly from a fall high of 80% feeding time when the squirrels switched to intensive feed- ing on buds and inner bark of twigs.
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