Table Of Content.
The Great Basin Naturalist
PublishedatPucno, Uiaii, hv
Bhicham Youxc; Uxaehsity
ISSN 0017-3614
Volume52 December 1992 No. 4
Great Basin Naturalist52i4i,pp.293-299
WINTER NUTRIENT CONTENT AND DEER USE OF
GAMBEL OAK TWIGS IN NORTH CENTRAL UTAH
RosemaiyL. Peiidletoii , FredJ.WagstaFC', and Bruce L. Welch'
—
Abstract. Weexaminedwinternutritional qualitx ot'current-vearbudandstem tissues fromburnedandunburned
stands of Gambeloak (Qiiercu.s oambclii Nutt.). Nutritional analyses werebasedon the amountof forageconsumed In
winteringmuledeer.DeeriLsealongtheUtaJi\'alle\-foothillsa\eraged6.2.5-10.7cmofcurrent-\eargrowth.Ofthetissues
examined,post-Hrebudtissuehadthehighestnutrientcontent,withameanof9.51% crudeprotein,0.19% pliosphonis,
and34.0% in\itrodigestibilit\'. Compositevalues(bud + stem) forunljumedstandswereslightK'higherincnideprotein
andphosphorusandlowerindigcstibilitvthanthosereportedinpre\iousstudies.Nutrientvaluesfrombumcdstandswere
significantlyhigherthanthoseofunbimiedstandsforallthreemeasures.Tannincontentottheburned-arearegrowthwas
alsohigher.OveraOforagevalueofGambeloaktowinteringmuledeerisrelatixcl\ low
Krijwords:Quercusgambelii,Odocoileushemionus,/(iifrieiits.fora<iiii'^hcJifivioi: iitilizafioii.hnncse, winter.
(xanibel oak (Quercus ^auibelii Nutt.) i.s a 17browsespeciesbasedontimespentbiowsing
\aliiable\'ear-roundsourceoffoodandcoverfor andplantweightconsumed.
many wildlife species, including deer, elk, big- Although important to wintering mule deer
horn sheep, small mammals, and a variety of in terms ol forage a\ailabilit\' and jniiatabilitx',
birds (Re\n()lds et al. 1970, Harperet al. 1985, oakranksamongthebottomin nutritional\alue
Tirmenstein 1988). Because of its abundance (Smith 1957, Bunderson et al. 1986). Nutri-
andlocation,oakisanimportantfoodsourcefor tional studies reportwinteroak browseasbeing
winteringmuledeer,pro\idingupto75%ofthe low in essential nutrients and digestibilit\'
available winter browse along the Wasatch (Smidi 1957, Kufeld et al. 1981, Meneelv and
Front (Fern- 1980). Winter use of oak \aries Schenmit/ 1981). Smith and Hubbard (1954)
with location. ])nt it has been report(xl high in described oak as being well liked but of low
some areas along the Wasatch Front, declining forage (|ualit\-. CiuTcntlv, little information is
in the presence of more palatable rosaceous a\ailal)le on the nutrient content of different
shnibs (Smith 1952, julaiuler 1955). Deer use portions of the plant stem oron the selection ol
in western North America ranges from moder- plant parts b\ deer.
ate to hea\A- throughout the year (Kufeld et al. Theeffectof fireon the nutritional status ot
1973 and references therein). In wint(M" prefer- oak brow^seisalsoof some interest toland man-
ence trials. Smith (1950) and Smith and Hub- agers. Fire ma\' proxide an effectixe manage-
bard (1954) ranked oak as 7t]i or lii(j;h(M- out of nuMit tool for opening the canopx of the more
shrubScience.sLahoraton.Iiitc: iiutaiiiRcscaicliStation.L'SD.\ForestSenice,7.3.5North.500East.Pro\().UtaliS4606.
293
L
294 Great Basin Naturalist [Volume52
primariK' ofGaiubel oak and sagebnish (Arte-
misia tridentata ssp. vuscijana), with scattered
patchesofcliffrose {Cowaniastanshitriana)and
bitterbmsh {PumJiia tridentata). Allthree loca-
tions are heavily used by wintering mule deer.
InAugust 1987awildfirebimiedapproximately
1270acresonthesouthwest-facingslopesabove
Orem and Lindon, Utali. Oak present on the
bum showed considerable regrowth two
months followingthe fire. Two studysiteswere
established at the Lindon location, one on the
bum itself, the other in the adjacent unburned
vegetation. Studv sites were also established at
the Pleasant Grove and Hobble Creek Canyon
locations, fora total offour studysites.
DeerutiUzationwas determinedbymeasur-
ingthe length ofmarkedtwigs before andafter
browsincr. In November of 1987, 679 twio;s on
the Lindon bum site and660twigs on the adja-
cent unburned site were marked with colored
plastic tape. Twigs were selected from around
F'ig. I. Location of four oakbrnsh studv sites in Utiili theperiphen'ofmultipleclonestorepresentall
Coimtv,Ut;ili.PG=Ple;is;uitGrove;LB= Lindon-lnimed; directiouiil aspects and a variet}' of heights
\.\J = Lindon-nnhnmed; IIC = HobbleCreekGanvon. accessibleto deer. Twiglengthswere measured
from thetape totheendoftheterminalbud. In
March 1988thetwigswereremeasuredandthe
dense oak thickets to aHow greater herbaceous
growth (Anonymous 1966, Dills 1970, Hallisey numberofcentimetersbrowseddeterminedfor
and Wood 1976, Haiper et al. 1985). Deer use eachtvvig.Theratioofbudtissueandtvvigtissue
of browse species has been found to increase consumed by deer was then calculated. The
followingfireinsome (Honi 1938, Halliseyand procedure was repeated at the Pleasant Grove
Wood 1976), but not all (Kufeld 1983), cases. and Hobble Creek sites the following vear,
The nutrient content of some oak species has where 186twigswere marked and measured at
been reported higher following burning each site.
(Hallisey and Wood 1976, Meneely and Tvvent\-t\vo samples for nutritional analysis
Schemnitz 1981). were collected at mid-winter from 12 burned
The intent of this study was to provide a and 10unbumedoakclonesatthe Lindon loca-
more accurate assessment ofthe nutrient con- tion. Portionsofeach ofthebumedcloneswere
tent ofoak forageconsumed bywinteringnuile fenced in earlyNovembertoensure availabilitv
deer on the Wasatch Front. Specific objectixes ofmid-winter collection material. In late Janu-
were (1) to determine what portion ofGambel an' 200-300 stems were removed from each
oak twigs was used by wintering mule deer in clone, packed in snow, and transported to the
UtiiliValley, (2)todetermine thepercentcrude laboratoi"v. Twigs were collected from all sides
protein, phosphorus, andin\itrodigestibilitx of of theperiphen*ofeach clone toeliminatepos-
terminalbudsandstemsofC^ambeloak,and(3) sible differences due to directional aspect. At
to compare xalues f)btained from adjacent the laboratory, stems from each clone sample
burned and uiiburned stands. were divided into a 1-cm terminal bud portion
and an adjacent lO-cm stem. Theproportion of
current-vear growth sampled (11 cm) was
Materials a.xd Methods approximatelyequaltothatremovedbvwinter-
ing mule deer. Where stem lengths measured
Deerutilizationwasstudiedatlocationsnear less than 11 cm, total current-vear growth was
Lindon, Utali, abo\e Pleasant Gro\e, Utah, and used in the analvsis. Tvvig diameters at 1 and 5
in the mouth ofHobble C:rcek Canyon (Fig. 1). cm from the tipwere also recorded.
Vegetation at these foothill locations consists The ensuing44budandstemtissuesamples
1992] W'lXTEK XLTKIEXTCUXTENT(JFGAMBELO.VK 295
Table L Sumnian-ofdeerutilizationonmarkedtwigsofGambeloakatfourstudysitesinUtahCounty,Utah.
No. h\igs No.twigs Percent Mean
marked browsed browsed utilization(cm)
Lindon-bumed 679 194 28.6 10.7 ± 0.44^'
Lindon-unburned 660 368 5.5.8 10.7± 0.24
HobbleCreek 186 112 60.2 6.3± 0.39
PleasantGrove 186 157 83.9 7.7± 0.33
'Mfaii istaiuiarderror
wereground u.singlicjuidnitrogen andstoredat Percentage data were arcsine transfbrm(>(l
—80 C. Invitro digestibility, cnide protein, and and anaK'zed usingthe General Linear Models
phosphorus were determined for both bud and (GLM) routine available on SAS. The mock'l
stemportions. Thesethree measureswerecon- used was a 2 x 2 factorial design, with burn
sidered sufficient to determine overall nutri- treatment (burned, unbumed) and tissue t\pe
tional qualit\- of oak as the\- are the nutrients (bud, twig) as main effects. Clone was used as
most commonK' deficient in winter diets of the error term for the bum treatment main
range animails (Welch et al. 1986). In vitro effect. Tissue differences were also examined
digestibilit\'wasassessedusingPearsons (1970) separateK' for burned and imbunied areas
modificationoftheTille\-andTerr\' 1963)tech- becauseofasignificantburntreatment x tissue
(
nicjue. This technicjue, while possibK oxeresti- interaction.
niating in \i\'o digestion of cell contents in
tannin-containing forages (Robbins et al. 1987, Results
Nastisand Malechek 1988), reniiiins theeasiest
and most accurate of the in \itro techniques Deer use at the Lin{k)n sites averaged 10.7
(Nastis and Malechek 1988) and is commonly cmforbothbtmiedandunburnedclones (Table
employedinnutritiontdstudiesofrangeforages. 1). Individual twig use \aried wideK; ranging
Inoculum for the digestion trial was obtained from 1.5to33cm.Althoughmeanuseatthetwo
irom aslaughter-house steer. The CO^-injected Lindon siteswasthesame,theburnedareahad
a greater proportion of small bites than the
inoculum was processed within 45 minutes of
remoxal h'om the rumen (Milchunas and Baker unbumed area (Fig. 2). Over 24% of the bites
wereinthe1.5-5cmcategoryatthebumedsite
1982). Studieshaveshownthatinoculaobtained
as compared to 5.7% in this category' at the
from domestic rmuinants can successfulK' unbumed site. Also, a smaller percentage of
approximate digestibilitv' of range forages to marked twi^s was browsed in the bumed area
deer (Palmer and Cowan 1979, Welch et al. (Table 1). Mean use at the Pleasant Grove and
1983). Phosphonis and cnide protein determi- Hobble Creek sites during the milder 1988-89
nationswere madeatthe PlantandSoilAnaKsis winter was somewhat less than at the Lindon
Laboraton atBrighamYoungUniversity'.Crude sites, averaging 7.7 and 6.3 cm, respecti\el\
pAroTteecihnnwiacsonbaAsuetdooAnnaKjlewl.dearli(lTneicthrnoigceonncoInntsetnrtu.- (TabRleesi1d)t.s fro'm the nutrient anaKsis of sam-
ment CoqD., Tarntowii, NY)was used to deter- pled tissues arc gixcn in Table 2. Main effects
mine phosphoms content. To simplify from the anaKsis of \ariance were all highly
comparisons with values reported in the litera- significant. Post-burn sprouts contained more
ture, composite \alues for the complete 11-cm crude protein and phosphorus and were more
sample were calculated as follows: composite digestible than unbumed samples. Bud tissue
\alue = [I0(twig value) -I- bud value]/ll. Bulk exceededstem tissueinall three measures.The
samplesmadeupofonetwigfromeachsampled interaction term was also highly significant for
clone were tested for tannin content. Twigs crude protein and phosphoms (p < .0(X)1 and
were kept frozen at -80 C until us(\ then p = .0021,respecti\ely). Runningseparateanal-
ground under liquid nitrogen. Tannin content \scs for bumed and unburned areas revealed
for each bulk sample was determined at the that the difference between bud and stem
Plant and Soil Analysis Laborator\- using viilueswasgreatest forpost-bum sprouts,creat-
Hagerman's (1987) radial diffusion method. ing the significant interaction term. Bud and
296 Great Basin Naturalist [\'olume 52
Lindon burned Lindon unburned
2 6 8 10 12 14 16 18 20 22 >23
Hobble Creek
8 10 12 14 16 18 20 22 >23 2 4 6 8 to 12 14 16 18 20 22 >23
Twig utilization (cm)
Fiii;.2. I^istrihiitionof stemutilization at FouroaklmislistutK'sitesin UtahCountw Utiili.
TaBLF. 2. Attained significance \alues from anakses of
variancefornutrientcontentofCambeloak.
Sourceof Cnide
variation protein Phosphonis Digestibility-
Bumtreatment
19921 W'lXTEH XlTHlENTCONTFNTOFCiAMHKLOaK 297
Tabll3. .\kaiisami .staiularil (.mtois forpcicfiit cIa matter iiutriciit content oi hudandtwigsaniple.scollected from
b=un.0i0e0d1)anbdetuwnehenumbeuddoliuiikd.tswtiagnd\saglureoswiwnitghnineabruLmintdroena,tmUetnath..CIoxm'tptoerssitfoel\laolwuiensgm=e[aln()s(ti\n\idgic\aatleues)ig+nifbiucdanvtalduief]i/elre1.nces (/;
Cnideprotein Phosphonis Digestibility
Bnmedstands
Hud 9.5 ± 0.36a
Twig
Composite
Inhuineilstand
Hud
Twig
Composite
298 Great Basin Naturalist [\olunie 52
burning. Vegetation at this Colorado location ACKNO\yLEDGMEXTS
consisted of a mixture of mature oak stands,
sagebrush, snowbern- (Sijiuphoricarpos alhiis), Thanks are extended to Len Caipenter,
chokecherrv' {Pntniis vir^iiiidita), and senice- Steve Monsen, and Art Tiedemann for sugges-
bern- {Aniclancliier ainifolia). Burning elimi- tionsonresearchdesign;toWarrenClaiv,Philip
nated big sagebrush plants and decreased Urness, Robert Ferguson, and an anonvmous
production of several other important browse reviewer forcomments on the manuscript; and
species, partially as a result of abnormally dr\' to John Allen and the Pleasant Grove Ranger
weatherconditions. District for access to the Lindon bum site.
We found no evidence for increased use on
the Lindon burned site (Table 1). The mean Literature Cited
number ofcentimeters browsed at the burned
site was identical to that ofadjacent unburned
Anonymous 1966. Conversionofthicketcoverediireasto
stands, even though twigs (sprouts) from the productivegrazinglands.USDA,ForestSeivice,Inter-
burnedstandstendedtobelonger.Also,alower mountainRegion. RangeImprovementNotes II: 1-4.
percentageofmarkedtwigswasbrowsedat the Austin. D. D., and P. J. Urness 1985. X'alues of four
communities for mule deer on riuiges widi limited
burnedsite.Theapparentlyloweruseofburned summer habitat. Journal of Range Management 38:
twigs by deer despite higher nutrient content 167-171.
maybe duetoseveral factors. Oakstands inthe BUNDERSON.E. D., B.L.Welch,AND D.J.Weber 1986.
In vitro digestibilit\'oiJtinipcnis osteospemw (Torn)
area form discrete clones rather than large Littlefrom 17Utahsites. ForestScience.32:8.34-840.
impenetrable thickets. Important browse spe- DeByle. N. v., p. J. Urness. and D. L. Blank 1989.
cies such as sagebrush and bitterbrush present Foragequalitvinburnedandimbumedaspencommu-
on unburned areas were lost as a result ofthe nSittaiteiso.n,USODgdAe,n,ForUteaslti.SeRneisceea,rcIhntPearpmeorunItNaTi-n40R4e.se8arpcph.
fire. Also, a lack ofcover and increased tannin Du.LS, G. G. 1970. Effects ofprescribed burningon deer
content of forage on the bum may have had browse.JournalofWildlifeManagement34:.540-.545.
some effect ondeerpreference. Hacerman.a.E. 1987. Radialdiffusionmethodfordeter-
mining tannin in plant extracts. Journal ofChemical
The oakbrush zone is critical to wintering Ecologv- 13: 437-449.
deer populations along the Wasatch Front. Hallisey, D. M.,and G.W.Wood 1976. Prescribedfire
inscmboakhabitatincentralPenns\Kania. lounialol
Although notthemostpreferredwinterfood,its WildlifeManagement40:.507-.516.
protective cover and sheer abvmdance make it Harper K.T,F.J.Wagstaff,andL.M.Kunzler 198.5.
oneofthemostwidelyused(Smidi 1949, Smith BiologvandmanagementoftheGambeloakvegetative
and Hubbard 1954.'julander 1955). Current tmvopuen:taailinterFaotrureestrevainedw. URaSnDgAe,FEo.rxepsetriSmeennitce,StaItnitoenr.-
emphasis in the Intermountain region is to Ogden.Utidi,GeneralTechnicalReportINT-179.31pp.
managetheoakbrushzoneprimarilyforwildlife Horn. E. E. 1938, Some wildlife-forest relationships.
(Winward 1985). Several management tools Ternacnesa3:ct3i7o6n-s38of0.the North ,\merican \\ildlifi' Confer-
have been suggested, including fire (Haiperet |ulander. O. 1955. Deer and cattU' relations in Utah,
al. 1985,Winward 1985). Burning may result in ForestScience 1: 130-139.
atemporary'improvementin nutritionalquality, KuFE\LeDg,etRa.tiCo.n19t8o3.bRuersnpionng,sessporfaevlikn,gm,ulaenddeecr,hacianttilneganodf
as well as opening the canopy sufficiently to Gambeloakrangekuid.ColoradoDivisionofWildlife,
allow establishment of other shrub and forb FortCollins,TechnicalPublication34.47pp.
species. However,withoutsome formofibllow- KuFELD R. C. M. ,Stevens, and D. C. Bowden 1981.
\\inter variation in nutrient and fibercontentand in
up treatment, the proliferation of oak sprouts vitro digestibilitv ofGambel oak (Qitcrcits t^ainbclii)
mayultimatelyresultindenser,less useableoak andbigsagebnish (Aiicinisiatridcniata)fromdiversi-
forage and reduction of understor\' species fiedsites in('olorado, loumal ofRange Manatiement
34:149-151.
(Harper et al. 1985, Stevens and Da\is 1985, Kufeld, R. C, D. D.Wall.mo, and C, Feddema 1973.
Winward 1985). Moreover, the loss of fire- FoodoftheRockvMountainmuledeer.USD.\,Forest
susceptible browse species such as big sage- SmeernvticeS,tatRioocn,kvFoMrotuCnotllaiinns.F(oCroelsotradaon.dRResaenagrechE.Pxpaepreir-
brush, mountain niiiliogany, and l)itteri)rush RM-111.31pp.
may have serious consequences for wintering -Miaeely. S. C, \nd S. D, S( hkmnitz 1981. Chemical
mule deer (Riggs et al. 1990), outweighing anv cthormepeosvietarisonaltaenrdawinildvfiitrreo.tSlioguetsthiwbielsittveronfNdaetuerralbirsotw2s6e:
possible benefit. 36.5-.374.
. a
19921 Winter \uthik.\tContentofCambelOak 299
Mii.c lll—NAS,D.G.,ANDD.L.Bakkr 1982. Invitrodiges- basedonbrowsingtimeandforageconsumed. |()imial
tion -sourcesofwithin-andbetwecn-triiils\arial)iIitA. ofRange Management8:262-265.
JournalofRange Management35: 199-203. Stevens. R., and N. D.wis 1985. Opportunities for
Nasti.s, a. S., and J. C. Malechek 1988. Estimating improvingforagJ.eproductionintheGambeloaktvpes
digestibilih ofoak browse diets for goats b\ in \itro ofUtiili. Pages37—il in K. L. Johnson, ed., Proceed-
techniques. |ounial ofRange Management 4\: 255- ingsofthethirdUtahslinibecology-workshop.College
258. ofNatural Resources, UtahState Universitv, Logan.
Pai.meh W. L.. and R. L. Chowan 1979. (xiniparison ol In ley M.A.,AND R.A.Terry 1963.Atwo-stagetech-
J.
deerandsheepdigestivecapacities.|oumalotW'ildlitc ni(jue forin vitrodigestionofforagecrops, [ounialof
Management43:V9.S-8()1 die Briti.shGra.s.slandSocietA' 18: 104-111.'
Pearson. H. A. 1970. Digestibilitv triiils: in vitro tccli- TiKMENSTElN. D. 1988. (ptiercus omnlu'lii. In. Fischer,
niques.Pages85-92;/i H.A. Paulsen. E. H. Reid.antl W. C., compiler. The fire effects information svsteni
K. W—. Parder,eds.. Rangeandwildlifehabitatevalua- [data base]. USDA, Forest Senice. Iiitermountain
tion a research svniposium. USDA, Forest Service Research Station, Intermountain Fire Sciences Labo-
Mi.scellaneousPublication 1147. ratorv', Missoula, Montana, magnetictape reels: 1600
Perky. W. M. 1980.Oakbrushmanagementplanandenvi- bpi,.'XSCIIwithGommonLISPpresent.
ronmental assessment report for Pleasant Grove Urness.P D. Neff.andR.K.Watkins 1975.Nutri-
RiingerDistrict.UintaNationalPoorest.8pp.[Mimeo- tive vaJl.u,e ofJ.mule deer forages on ponderosa pine
graphed.] summer range in Arizona. USDA Forest Seivice.
Reynolds. H.G..\\'.P.Glary.andP.F.Ffolliott 1970. Rock-\ Mountain Forest and Range E.xperiment Sta-
Gambeloakforsoudiwesternwildlife.JournalofFor- tion, FortCollins. Colorado, Research Note RM-3()4.
estrv'68:545-^547. 6pp.
Rk;gsE.ffeRc.tsa.,ofRdoJ.meUsrtniecssgo,atasnodnKd.eeAr. wGionntzearilnegz.in1U9t9a0l.i WelcPteldeBr.soL.n,.Ea.ndD. MNc.:AD.rAtVhIlS'r1.986D..'LH.obNbellesGorne.ekJ".—C.
oakbmsh.JournalofRangeManagement43:229-234. superiorselectionJ.oflow-elevationmountainbigsage-
RoBBLNs. C. T, S. Mole, A. E. R\german, and T. .\. biii.sh. USDA, ForestService, Intermouiit;un Research
HanLEY. 1987. Role of tannins in defending plants Station,Ogden,Utali,ResearchPaperINT-370. 10pp.
against mniinants: reduction in dn matterdigestion':^ Welch, B. L., J. C. Pederson. and W. P. Clary, 198.3.
Ecologv 68: 1606-1615. Abilit\'ofdifferentrumeninoculatodigestrangefor-
Smith .-X.D. 1949. DeerforageobservationsinUtiJi. |(>ur- ages.JournalofWildlifeMtmagement47:873-877.
luilofWildlife Management 13;314—315. Winward. A. H. 1985. Perspectives on Gambel oak niiui-
. 1950. Feeding deer on browse species during agement on national forests of the Intermountain
winter. JournalofRange Management3: 130-132. Region. Pages 33^35 in K. L. Johnson, ed.. Proceed-
. 1952. Foodhabitsot muledeerin Utah.Journalof ingsofthethirdUtahshrubecologvworkshop.College
Wildlife Management 16: 148-155. ofNatural Resources, UtahStateUniversitv, Logan.
. 1957. Nutritive value of some browse plants in
winter.JournalofRange Management 10: 162-164.
Smith. A. D.,and R. L. Hubbard. 1954. Preference rat- Received9March 1992
ingsforwinterdeerforagefromnorthernUtahranges Accepted18October1992
