Great Basin Naturalist
Volume 52 Number 4 Article 1
12-30-1992
Winter nutrient content and deer use of Gambel oak twigs in north central Utah
Rosemary L. Pendleton Shrub Sciences Laboratory, Intermountain Research Station, USDA Forest Service, Provo, Utah
Fred J. Wagstaff Shrub Sciences Laboratory, Intermountain Research Station, USDA Forest Service, Provo, Utah
Bruce L. Welch Shrub Sciences Laboratory, Intermountain Research Station, USDA Forest Service, Provo, Utah
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Recommended Citation Pendleton, Rosemary L.; Wagstaff, Fred J.; and Welch, Bruce L. (1992) "Winter nutrient content and deer use of Gambel oak twigs in north central Utah," Great Basin Naturalist: Vol. 52 : No. 4 , Article 1. Available at: https://scholarsarchive.byu.edu/gbn/vol52/iss4/1
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ISSN 001 7-3614
VOLUME 52 DECEMBER 1992 NO.4
Great Basin Naturalist 52(4), pp. 293-299
WINTER NUTRIENT CONTENT AND DEER USE OF GAMBEL OAK TWIGS IN NORTH CENTRAL UTAH
Rosemary L. Pendleton" Fred J. Wagstaff" and Bruce L. Welch'
ABSTMCT.-We examined winter nutritional quality of current-year bud and stem tissues from burned and unburned stands of Gambel oak (Quemus garnhelii Nutt.), Nutritional analyses were based on the amount of forage consumed by wintering mule deer. Deer use along the Utah Valley foothills averaged 6.25-10.7 em ofcurrent-year growth. Ofthe tissues examined, post-fire bud tissue had the highest nutrient content, with a mean of9.51% crude protein, 0.19% phosphorus, and 34.0% in vitro digestibility. Composite values (bud + stem) for unburned stands were slightly higher in crude protein and phosphorus and lower in digestibility than those reported in previous studies. Nutrient values from bumed stands were Significantly higher than those ofunbumed stands for all three measures. Tannin content ofthe bumed-area regrowth was also higher. Overall forage value of Gambel oak to wintering mule deer is relatively low.
Key wonk: Quercus gambelii, Odocoileus hemionus, nutrients,jomging behavior, utiliz.ation, browse, winter.
Gambel oak (Quercus gamhelii Nutt.) is a 17 browse species basedon time spentbrowsing valuable year-round source offood andcover for and plant weight consumed. many wildlife species, Including deer, elk, bIg Although important to wintering mule deer horn sheep, small mammals, and a variety of in terms of forage availability and palatability, birds (Reynolds et aJ. 1970, Harpel et aJ. 1985, oak ranks among the bottom in nutritional value Tirmenstein 1988). Because of its abundance (Smith 1957, Bunderson et aJ. 1986). Nutri and location, oak is an importantfood source for tional studies report winter oak browse as being wintering mule deer, providing up to 75% ofthe low in essential nutrients and digestibility available winter browse along the Wasatch (Smith 1957, Kufeld et al. 1981, Meneely and Front (Perry 1980). Winter use of oak varies Schemnitz 1981). Smith and Hubbard (1954) with location, but it has been reported high in described oak as being well liked but of low some areas along the Wasatch Front, declining forage quality. Currently, little information is in the presence of more palatable rosaceous available on the nutrient content of different shrubs (SmIth 1952, Julander 1955). Deer use portions ofthe plant stem or on the selection of in western North America ranges from moder plant parts by deer. ate to heavy throughout the year (Kufeld et al. The effect offire on the nutritional status of 1973 and references therein). In winter prefer oak browse is also ofsome interest to land man ence trials, Smidl (1950) and Smith and Hub agers. Fire may provide an effective manage bard (1954) ranked oak as 7th or higher out of ment tool for opening the canopy of the more
1Shrub Sciences Laboratory, Intermountain Rese 293 294 GREAT BASIN NATURALIST [Volume 52 primarily of Gambel oak and sagehrush (Arte misia triikntata ssp. vaseyana), with scattered patches ofcliffrose (Cowania stansburiana) and J' bitterbrush (Purshia trid£ntata). All three loca e" 1-, tions are heavily used by \vintering mule deer. "- InAugust 1987 awildfire bumed approximately co'" 1- 1270 acres on thesouthwest-facing slopes above ? Orem and Lindon, Utah. Oak present on the 0 ~ burn showed considerable regrowth two t months following the fire. Two study sites were +'" established at the Lindon location, one on the j ...'r bum itself, the other in the adjacent unburned ~ vegetation. Study sites were also established at the Pleasant Grove and Hobble Creek Canyon .;:,...'" .. <{ locations, for a total offour study sites. V> Deer utilization was determined by measur ..: ing the length of marked twigs before and after ~ brOWSing. In November of 1987, 679 twigs on the Lindon burn site and 660 twigs on the adja I I to kul. cent unburned site were marked with colored plastiC tape. Twigs were selected from around Fig. 1. Location of four oakbrush stud)· ~tes in Utah the peripheryofmultiple clones to represent all County. Utah. PC = Ple~ant Grove; LB = Lindon-burned; directional aspects and a variety of heights LV - Lindon-unbumed; He ... Hobble Creek Canyon. accesSible to deer. Twig lengths were measured from the tape to the end ofthe terminal bud. In dense oak thickets to allow greater herbaceous March 1988 the twigs were remeasured and the growth (Anonymous 1966, Dills 1970, Hallisey number ofcentimeters browsed determined for and Wood 1976, Harper et al. 1985). Deer use each twig. The ratio ofbud tissue and twig tissue of browse species has been found to increase consumed by deer was then calculated. The following fire in some (Horn 1938, Hallisey and procedure was repeated at the Pleasant Grove Wood 1976), but not all (Kufeld 1983), cases. and Hobble Creek sites the foIlO\ving yeal; The nutrient content of some oak species has where 186 twigs were marked and measured at been reported higher following burning each site. (Hallisey and wood 1976, Meneely and Twenty-two samples for nutritioual analysis Schemnitz 1981). were collected at mid-winter from 12 burned The intent of this study was to provide a and 10 unburned oak clones at the Lindon loca more accurate assessment of the nutrient con tion. Portions ofeach ofthe burned clones were tent ofoak forage consumed by wintering mule fenced in early November to ensure availability deer on the Wasatcb Front. Specific objectives of mid-winter collection material. In late Janu were (1) to determine what portion of Gambel ary 200-300 stems were removed from each oak twigs was used by wintering mule deer in clone, packed in snow, and transported to the Utah Valley, (2) to determine the percent crude laboratory. Twigs were collected from all sides protein, phosphorus, and in vitro digestibility of ofthe peliphery ofeach clone to eliminate pos terminal buds and stems ofGamhel oak, and (3) sible differences due to directional aspect. At to compare values obtained from adjacent the laboratory, stems from each clone sample burned and unburned stands. were divided into a I-em terminal bud portion and an adjacent lO-em stem. Theproportion of current-year growth sampled (11 cm) was MATERIALS ",'1D METHODS approximately equal to that removed by winter ing mule deer. \\1lere stem lengths measured Deerutilization was studied at locations near less than II em, total current-year grO\vth was Lindon, Utah, above Pleasant Grove, Utah, and used in the analysis. 'IWig diameters at 1 and 5 . . in the mouth ofHobble Creek Canyon (Fig. 1). em from the tip were also recorded. Vegetation at these foothill locations consists The ensuing 44 bud and stem tissue samples 1992J WINTER NUTRIENT CONTENT OF GAMBEL OAK 295 TABLE L Summary ofdeer utilization on marked twigs ofGambel oak at four ~tudy sites in Utah County, Utah. No. twi~S No. twigs Percent Mean marke bro'NSed browsed utilization (cm)U Lindon-burned 679 194 28.6 10.7 ± 0.44>1. Unclan-unburned 660 308 55.8 10.7" 0.24 Hobble Creek 186 1I2 60.2 6.3" 0.39 Pleasant Grove 186 157 83.9 7.7::t 0.33 ~Meau :!: stan<;lard error were ground using liquid nitrogen and stored at Percentage data were arcsine transformed -80 C. In vitro digestibility, crude protein, and and analyzed using the General Linear Models phosphorus were determined for both bud and (GLM) routine available on SAS, The model stem portions. These three measures were con used was a 2 X 2 factorial design, with bum sidered sufficient to determine overall nutri treatment (burned, uuburned) and tissue type tional quality of oak as they are the nutrients (bud, twig) as main effects, Clone was used as most commonly deficient in winter diets of the error term for the burn treatment main range animals (Welch et a1. 1986). In vitro effect. Tissue differences were also examined digestibility was assessed using Pearson's (1970) separately for burned and unburned areas modification ofthe TIlley andTerry (1963) tech because ofa significant burn treatment X tissue nique, This technique, while possibly overesti interaction. mating in vivo digestion of cell contents in tannin-containing forages (Robbins et al, 1987, RESULTS Nastis and Malechek 1988), remains the easiest and most accurate of the in vitro techniques Deer use at the Lindon sites averaged 10,7 (Nastis and Malechek 1988) and is commonly em for bothburnedand unburned clones (Table employed in nutritional studies ofrange forages, 1), Individual twig use varied widely, ranging from 1.5 to 33 cm. Although mean use atthetwo Inoculum for the digestion trial was obtained Lindon sites was the same, the burued area had from a slaughter-house steer. The CO -injected 2 a greater proportion of small bites than the inoculum was processed within 45 minutes of unburned area (Fig, 2), Over 24% of the bites removal from the rumen (Milchunas and Baker were in the 1.5-5 cm category at the burned site 1982). Studies have shown that inoculaobtained as compared to 5,7% in this category at the from domestic ruminants can successfully unburned site, Also, a smaller percentage of approximate digestibility of range forages to marked twigs was browsed in the burned area deer (Palmer and Cowan 1979, Welch et al, (Table 1), Mean use at the Pleasant Grove and 1983), Phosphorus and crude protein determi Hobble Creek sites during the milder 1988-89 nations were made at the Plant and Soil Analysis winter was somewhat less than at the Lindon Laboratoryat Brigham Young University, Crude sites, averaging 7,7 and 6,3 em, respectively protein was based on Kjeldahl nitrogen content. (Table 1), A Technicon Auto Analyzer (Technicon Instru Results from the nutrient analysis, of sam- ment Corp" Tarrytown, NY) was used to deter pled tissues are given in Table 2, Main effects mine phosphorus content. To simplify from the analysis of variance were aU highly comparisons with values reported in the litera significant. Post-burn sprouts contained more ture, composite values for the complete ll-cm crude protein and phosphorus and were more sample were calculated as follows: composite digestible than unburned samples, Bud tissue value ~ [lO(twig value) + bud value]/I1. Bulk exceeded stem tissue in all three measures, The samples made up ofone twig from each sampled interaction term was also highly significant for clone were tested for tannin content. Twigs crude protein and phosphorus (p < ,0001 and were kept frozen at -80 C until use, then p ~ ,0021, respectively), Runningseparate anal ground under liquid nitrogen, Tannin content yses for burued and unburned areas revealed for each bulk sample was determined at the that the difference between bud and stem Plant and Soil Analysis Laboratory using values was greatest for post-bum sprouts, creat Hagerman's (1987) radial diffusion method, ing the Significant interaction term, Bud and 296 GREAT BASIN NATURAUST [Volume 52 lindon burned lindon unburned .. " , o 2. 4 • B 10 1t 14 " ,. 20 12 >2) I 4 • • 10 11 '4 III II 20 22 > 2~ Pleasant Grove Hobble Creek " .. " , o 1 4 • • 10 II 14 ,. I' ZO 22 >u 1 4 • • 10 12 14 1. " 20 U >23 Twig utilization (cm) FJg. 2. Dismbution of stem utilization at fOUT oakbrush study sites in Utah County, Utah. TABLE 2. Attained ,,;gn;ficance values from aoalyses of tentof4.1 mgper 100 mgplanttissue compared V'dri.mce for nutrient content of Gambel oak. with 3.4 mg for unburned clones. The stem tissue samplederived from burned clones had a SOUR--e of Crude vaJiution protein Phosphorus Digestibility tannin content of1.6 mg per100 mg plant tissue compared with 0.7 mg For unburned clones. Bum treatment 0.0001 0.0002 0.0001 1lssue type 0.0001 0.0001 0.0001 Bum X tissue 0.0001 0.0021 0.3519 DISCUSSION Clone 0.0001 0.0228 0.0015 Previous reports on fall-winter nutrient con tent of Gambel oak twigs from mature stands twig values from burned clones differed Signif range from 4.6% to 5.7%for crudeprotein, from icantly for all three variables (Table 3). Bud and 0.09% to 0.10% for phosphorus (Smith 1957, twig values from unburned clones differed only Kufeld et a1. 1981, Meneely and Schemnitz 1981), and from 26.6% to 40.2% for in vitro in nitrogen content. Twigs from burned and digestibility (Kufeld et a1. 1981, Meneely and unburned clones differed in appearance, also Schemnitz 1981). Similar values have been burned twigs being more slender at 1 cm (1.8 obtained for other oak species (Meneely and mm vs. 2.6 mm;p = .0002) and at5 cm (2.1 mm Schemnitz 1981). Composite values from vs. 2.9 mm; p = .0001). unburned stands in our study (Table 2) are sim Burning also had a significant effect (p = ilar to previous results, t1lOUgh slightly higher in .0001) on tannin content. The budtissue sample crude protein and phosphorus and slightly derived from burned clones had a tannin con- lower in digestibility. The use ofdifferent twig - ',"'", ,,-. "". ''''''''. '"",, " ".' '_. 1992] WINTER NUTRIENT CONTENT OF GAMBEL OAK 297 TABLE 3. Means and standard errors for percent dry matter nurnent content of bud and hvig samples collected from burned and unburned oak stands growing near Lindon, Utah. Letters following means indicate significant differences (p = .0001) between bud and twig values within bum treatment. Composite values ". [lO(twig value) + bud value]!lL Crude protein Phosphorus Digestibility Burned stands Bud 9.5 + 0.36 a 0.19 ± 0.0068 a 34.0 + 0.59 a Twig 7.5 ± 0.22 b 0.13 ± 0.0092 b 29.8 ± 0.70 b CompOSite 7.7 ± 0.23 0.14 ± 0.0087 30.2 ± 0.66 Unburned stands Bud 6.5 :t 0.09a 0.12:!: 0.0056 a 26.2:!: 1.68 a Twig 5.7 ± 0.10 b O.I! ± 0.0086 a 23.5 :!: 1.15 a Composite 5.8 :t 0.10 0.11 ± 0.0081 23.7± 1.16 lengths for nutrient analysis did not affect the tissues sampled, had a crude protein content of overall results significantly. Composite values less than 10%. TI,e composite value from based on twig lengths of 7.7 or 6.3 cm (mean burned clones was less than 8%, which ranks deer utilization at Pleasant Grove and Hobble below that of sagebrusb, aspen, and rosaceous Creek) differed very little from those based on shrubs (Smith 1957, Kufeld et aL 1981, DeByle the ll-cm sample. In the absence of foliage, et a1. 1989). Actual amounts ofprotein digested buds provide the highest source of nutrients may be somewhat less than predicted by the in during late fall and winter. Values for leaves of vitro technique. Tannins presentin summer oak Gambel oak are substantially higher in summer, and other forages have been found to increase but comparable to bud tissue by mid-winter the fecal excretion ofprotein by domestic live (Urness et aI. 1975, Meneely and Schemnitz stock (Robbins et aL 1987). In mule deer and 1981, Welch et a1. 1983, Austin and Urness other browsers, nitrogen excretion may be 1985). reduced by tannin-binding proteins present in Burning had a Significant influence on the the saliva (Robbins et al 1987). Winter digest nutrient content ofoak forage, particularly bud ibility ofGambel oak is also lowwhen compared tissue. Values obtained from burned stands to otherforages. Bunderson et aI. (1986) ranked were relatively higher than those obtained from digestibility ofwinter oak forage 25th out of27 unburned stands. Post-bum bud tissue had by species tested. Our results are similar in digest far the highest nutritional value of any tissues ibility to that listed in the ranking, shOwing examined. Increased nutrient content offorage slightlyhigherdigestiblityforburned stands and follOwing fire has been reported elsewhere, lower digestibility for unburned stands. although indications are that such an increase is Deer use of oak depends on many factors temporary (Dills 1970, HalliseyandWood 1976, including cover, exposure, density ofoakbrush, Meneely and Schemnitz 1981, DeByle et aI. and availability of other forages (Smith 1952, .1989). The evidence to date suggests that Julander 1955, Kufeld 1983, Austin and Urness improved phosphorus content offorage due to 1985). Fire affects the structure of vegetation fire is fairly short-lived, usually lasting only one and cover as well as quantity and quality of year. Nitrogen benefits may last longer, depend forage produced (Horn 1938, Hallisey and ing on the species and season. The higher values Wood 1976, Meneely and Schemnitz 1981, reported for burned stands in this study likely Kufeld 1983). Whether or not deer use an area occurred because sampling took place less than more after burning appears to depend on struc one year follOwing the burn. ture of the oak community and type ofvegeta Despite the slightly higher values for crude tion present on adjacent areas, as well as protein and phosphorus reported here for intensity and size of burn. Where oak stands unburned stands and the markedly higher form impenetr'lble thickets, or where little values for recently burned stands, the overall understory is available, burning has resulted in winter nutritional value ofCambe! oak remains increased use by deer (Horn 1938, Hallisey and relatively low. Even post-burn bud tissue, whicb Wood 1976). In contrast, Kufeld (1983) found had the highest nutrient content of any of the increased use by elk but not deer following 298 GHEAT BASIN NATURALIST [Volume 52 burning. Vegetation at this Colorado location ACKNOWLEDGMENTS consisted of a mixture of mature oak stands, sagebrush, snowberry (Symphoricarpos albus), Thanks are extended to Len Carpenter, chokecherry (Pmnus oirginiana), and service Steve Monsen, and Art Tiedemann for sugges berry (Amelanchier alnifolia). Burning elimi tions on research design; to Warren Clary, Philip nated big sagebrush plants and decreased Urness, Robert Ferguson, and an anonymous production of several other important browse reviewer for comments on the manuscript; and species, partially as a result of abnormally dry to John Allen and the Pleasant Grove Ranger weather conditions. District for access to the Lindon burn site. We found no evidence for increased use on tbe Lindon burned site (Table 1). The mean LITEHATUHE CITED number of centimeters browsed at the burned site was identical to that of adjacent unburned ANONYMOUS. 1966. CoD\·er~ion oftlJicket covered areas to stands, even though twigs (sprouts) from the productive grazing lands. USDA, ForestService. lnter burned stands tended to be longer. Also, a lower mountain Region. Rllnge Improvement Notes 11: 1-4. percentage of marked twigs was browsed at the AUSTIN, D. D., AND P. J. URNESS. 1985. Values of four communities for mule deer on ranges with limited burned site. Theapparently lower use ofburned summer habitat. JmunaI of Range Management 38: twigs by deer despite higher nutrient content 167-l7J.. may be due to several factors. Oak stands in the BUNDERSON, E. D., B. L. WELCH, AND D. J. WEBER. 1986. In vttro digestibility ofJunipena on:rosperma (Torr.) area fonn discrete clones rather than large Little from 17 Utah sites. Forest Science 32: 834-840. impenetrable thickets. Important browse spe DeBYLE. N. V., P. J. URNE.SS, AND D. L. BLANK. 1989. cies such as sagebrush and bitterbrush present Forage quality in burned and unbumedaspenoommu. nilies. USD..o\., ForestService, Intennountain Research on unburned areas were lost as a result of the Stafion, Ogden, Utah, Research Paper INT-404. 8 pp. fire. Also, a lack of cover and increased tannin DILLS, G. G. .1970. Effects of prescribed. burning on deer content of forage on the burn may have had browse. Journal ofWildl.ife Management 34: 540-545. HAGERMAN. A, E. 1987. Radial diffusion method for deter· some effect on deer preference. mining tannin in plant extracts. Journal of Chemical The oakbrush ?.one is critical to wintering Ecology 13, 437-449. deer populations along the Wasatch Front. HALLISEY, D. M., AND G. W. WOOD. 1976. prescribed Oro in scrub oak hahitat in central Pennsylvania. Journal of Although not the most preferredwinter food, its Wl1dlife Management 40: 507--516. protective cover and sheer abundance make it HARPER, K. T.. F. J. WACSTAFf,AND L. M. KUNZLlw.1985. one ofthe most widely used (Smith 1949, Smith Biology and management oftheGambel oa.kvegetatJve type: a literature review. USDA, Forest ScIVice, Inter and Hubbard 1954, Julander 1955). Current mountain Fore.!li: and Range Experiment Station. emphasis in the Intermountain region is to Ogden, Utah, General Technic--u RepOlt INT~ 179.31 pp. manage the oakbrush zone primarily for wildlife HORN, E. E. 1938. Some wildlife··forest relationships, TranSl'lCtioos of the North American Wildlife Confer (Winward 1985). Several management tools ence 3: 376--380. have been suggested, including fire (Harper et JULANDER. O. 1955. Deer and cattle relations in Utah. al. 1985, Winward 1985). Burrung may result in Forest Science 1: 130-139. KUFELD, R. C. 1983. Responsesofelk. mule deer, cattle and a temporary improvement in nutritional quality, vegetation to burning, spraying. and chaining of as well as opening the canopy suffiCiently to Gambel oak rangeland. Colorado Division ofWildlife, allow establishment of other shrub and forb Fort Collins, Technical Publication 34. 47 pp. KUFELD. R. G, M. STEVENS, AND D. C. BoWDEN. 1981. species. However, without some form offollow Winter variation in nutrient and fiber c..'Ontent and in up treatment, the proliferation of oak sprouts vitro digestibility of Gambel oak (Que1UiS ga.mbelit) may ultimately result in denser, less useable oak and big sagebrush (Artemisia tridentota) from diversi 6ed sites in Colorado. Journal of Range Management forage and reduction of understory species 34, 149-151. (Harper et al. 1985, Stevens and Davis 1985, KUFELD, R. c., D. D. WALLMO. AND C. F'EDDEMA. 1973. Winward 1985). Moreover, the loss of fire Foodofthe Rocky Mountain mule deer. USDA, Forest SeIVice. Rocky Mountain Forest and Range Experi susceptible browse species such as big sage ment Station, Fort Collins, Colorado, Hesearc:h Paper brush, mountain mabogany, and bitterbrush RM-lI1. 31 pp. may have serious consequences for wintering MENEELY. S. C., AND S. D, SCHEMNITZ. 1981. Che.mical composition and in vitro digestibility of deer browse mule deer (Riggs et al. 1990), outweighing any three years aftera \'lildf'u:e. Southwestern Natumlist 26: possible benefit. 365-374. - .", ".' ,,'.' ". " " ' , .. ", 1992J WINTER NlYfRIENT CONTENT OF GAMBEL OAK 299 MILCHUNAS, D. G., AND D. L. BAKER. 1982. Invitro diges~ based on browsing time and forage consumed, Journal tion-sources of'Nithin- and between-trials variability. of Range Management 8: 262-265. Journal afRange Management 35: 199-203. STEVENS, R., AND J. N. DAVIS. 1985. Opportunities for NASTIS, A. S" AND J. C. MALECHEK 1988. Estimating improving forage production in the Gambel oak types digestibility of oak browse diets for goats by in vitro of Utah, Pages 37--41 in K. L, Johnson, ed" Proceed techniques. Joumal of Range Management 41: z.55-.-. ings ofthe third Utah shrub ecology workshop, College 258. of Natural Resources, Utah State University, Logan. PALMER, W. L., AND K L. COWAN. 1979, Comparison of TILLEY, J. M. A.,AND R A. TERRY, 1963. A two~stagetech~ deer and sheep digestive capacities. Journal ofWildlife nique for in vitro digestion of forage crops. Journal of Management 43: 798-801. the British Grassland Society 18: 104-111. PEARSON, H. A. 1970. Digestibility trials: in vitro tech TIRMENSTEIN, D. 1988. Quercus gawelii. In: Fischer, niques. Pages 85-92 in H, A. Paulsen, E. H. Reid, and W. C., compiler, The lire effects information system K. W. Parder, eds., Range and mldlife habitat evalua~ [data base]. USDA, Forest Service, Intermountain tion-a research symposium. USDA, Forest Service Research Station, Intermountain Fire Sciences Labo~ Miscellaneous Publication 1147. rato:ry, Missoula, Montana. magnetic tape reels; 1600 PERRY, w: M. 1980. Oakbrush management plan and envi bpi, ASCII with Common LISP present. ronmental assessment report for Pleasant Grove URNESS, P. J" D. J. NEFF, AND R. K. WATKINS. 1975. Nutri Ranger District, Uinta National Forest. 8 pp. [Mimeo tive value of mule deer forages on ponderosa pine graphed.] summer range in Arizona. USDA Forest Service, REYNOLDS, H. G., W. P. CLARY,AND E F. FFOLLIOTT.1970. Rocky Mountain Forest and Range Experiment Staw Gambel oak for southwestern wildlife. Journal ofFor tion, Fort Collins, Colorado, Research Note RM~304, estty 68,545-047. 6pp. RIGGS, R. A., P, J. URNESS, AND K, A. GoNZALEZ. 1990. WELCH, B. L., E. D. McARTHUR, D. L. NELSON, J, C. Effects of domestic goats on deer wintering in Utah PEDERSON, AND J. N, DAVIS. 1986, 'HQbbleCreek'-a oakbrush. Journal of Range Management 43: 229-234. superior selection oflow-elevation mountain big sage ROBBINS, C. T., S. MOLE, A, E. HAGERMAN, AND T. A. brush. USDA, Forest Service, Intermountain Research HANLEY. 1987. Role of tannins in defending plants Station, Ogden, Utah, Research PaperINT-370.10pp. against ruminants: reduction in dry matter digestion? WELCH, B. L" J. C. PEDERSON, AND W. P. CLARY. 1983. Ecology 68, 1606-1615. Ability of different rumen inocula to digest range for SMITH, A. D. 1949. Deer forage observations in Utah. Jour ages. Journal ofWildlife Management 47: 873-877. nal ofWildlife Management 13: 314-315. WINWARD, A. H. 1985. Perspectives on Gumbel oak man~ . 1950. Feeding deer on browse species during agement on national forests of the Intermountain --Wlc;·nter. Journal of Range Management3: 130-132. Region. Pages 33--35 in K. L. Johnson, ed., Proceed _-=' 1952. Food habits of mule deer in Utah. Journal of ingsofthethird Utah shrub ecology workshop. College Wildlife Management 16: 148-155. ofNatural Resources, Utah State University, Logan. ---c' 1957. Nutritive value of some browse plants in winter. Journal ofRange Management 10: 162-164. SMITH, A. D., AND R. L. HUBBARD, 1954. Preference rat~ Received 9 March 1992 iogs for winter deer forage from northern Utah ranges Accepwd 18 October 1992