White Spruce Above And Beyond Treeline In The Arrigetch .

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ARCTICVOL. 39, NO. 3 (SEPTEMBER 1986) P. 247-252White Spruce above and beyond Treeline in theArrigetch Peaks Region, Brooks Range, AlaskaDAVID J. COOPER’(Received 20 December 1983; accepted in revised form12 November 1985)ABSTRACT. White spruce trees occur in the Arrigetch Creek valley and its tributaries at great distances above and beyond current treeline, which is at760 m elevation. The highest tree foundis at 1465 m elevation on a south-facing limestone slope. Treesalso occur up to 5.0 km beyond treeline ongranitic parent rock. Thesetrees appear to occur at the highest elevationof thenorthArctic Circle in North America and include some of the highest treesin Alaska.Key words: Brooks Range, Alaska, treeline, white spruce, seed dispersalRESUME. On trouve des CpicCas blancs dans la vallte du ruisseau Amgetch et de ses tributaires i des distances bienau-deli de la limite forestiereau sud. Certains arbres seactuelle, quise trouvei une Clbvationde 760 m.L’arbrele plusClevC est i 1465 m d’CMvationsurun versant de calcaire exposemere granitique. Ces arbres semblent croitrei leur plus hautetrouvent aussijusqu’i 5.0 km au-dessus de la limite forestibre, croissant dans de la rocheClbvation au nord ducercle arctique en Amdrique du Nord et comprennent peut-&tre les arbres les plusClevCs en Alaska.Mots clCs: chaine de Brooks, Alaska, limite forestiere, dpicda blanc, dispersion desp i n e sTraduit pour le journal par Maurice Guibord.INTRODUCTIONare relicts of a much warmer interval. Elliott (1979a, 1979b,1979c)hasshownthatsexualreproductionrarely, if ever,occurs inspruce at treelineinKeewatinbutdoes occur inWhile conducting researchon arctic-alpine tundra ecosystems in Labrador-Ungava.the Anigetch Peaksregion(Fig. 1) from1978 to 1980, theThe continental divide in the Anigetch Peaks region north and south, separating the Alatna Riverdrainage on the east[Moench] Voss) trees hundreds of mabove and several kmfrom the Kobuk River drainage to the west(Fig. 1). Treeline inbeyond the last cone-bearingtrees. Reports of treeline stabilitythe Alatna River-Arrigetch Creek region occursat 700-760 min Alaska are varied. As earlyas the 1920s, Mertie (1925) waselevation (Murray, 1974; Cooper, 1983), while in the Kobuksurprised to find treesat 1060 m elevationin the BrooksRange,valley treeline occurs at680 m elevationon north-facing slopesat higher elevation than in the Alaska or Coast ranges. Elseand 550 m on south-facing slopes(Goldstein, 1981). Moist airwhere inthe Brooks Rangeseedlingsabove treeline are reported masses from the Pacific Ocean and Bering-Chukchiseas enterby Garfinkel and Brubaker(1980), Goldstein (1981) and Odasz the Brooks Range from the west and southwest and are fun(1983). HoweverDrewandShanks(1965) andDensmorenelleduptheeast-west trendingKobuk and NoatakRiver(1980) showed treelineto be in equilibrium. In western Alaskavalleys. Atthe heads of these valleys the Arrigetch and IgikpakHopkins (1972) has found that treeline has advanced since theplutons (Brosge and Pessel, 1977; Nelson andGrybeck, 1978)1940s. In interiorAlaska, Marchand (1976), Haugen and Brown ceated extensive highlands, which impede the eastward move(1978) andViereck(1979)allreport that treeline is notinment of air masses.equilibrium withthecurrentclimate. In the Alaska RangeSome botanical differences between the vegetationof theDentonandKarlen(1977) report treeremains 76 m aboveKobukandAlatnaRiversides of the Anigetch Peaksarepresent treeline but no recent change in treeline. In southernsignificant. The ferns Cystopteris montanu (Lam.) Bernh. andAlaska Griggs (1934) reportedtreeline to be advancing.Dryopteris dilututu (Hoffm.) Beauv. ssp. americuna (Fisch.)The twentieth-century warming trend in Alaska documented Hult. and the grass Culamagrostis canadensis(Michx.) Beauv.by Hamilton (1965a, 1965b), Blasting and Fritts (1973) andare locally common forestinopenings and lowelevation arcticGarfinkel and Brubaker (1980) may be responsible for whatalpine tundra around Walker Lake and KaluluktokCreek in thetreeline changes are occurring. The pollen recordindicates thatKobuk drainage. In the Alatna Riverdrainage the twoferns arewhitesprucewasestablishedsouth oftheBrooksRangevery rare and restricted to springs, and the grass occurs at the5000-6000 years ago, became established in the Alatna Valleyon disturbed sites. This appears to indicateborder of ponds and1500 years ago and may still be slowly advancing northwardon the Alatna Riverside of theincreasing continental influences(Brubaker et a l . , 1983).Arrigetch Peaks. This wasalso shown by Ellis etal. (1981) andIn Canada, Nichols(1976)hypothesizedthat the modemcan influence the elevation oftreeline (Wardle, 1974).warming is too smallto advance treeline and present reproducThis paper documents interesting occurrences ofsprucewhitetion shouldbe looked for within the forest zone. Larsen (1965,at high elevationsnorth of the ArcticCircle and thuscontributes1974, 1980) and Savile (1972) feel that tree islands at treelinesubstantially to treeline ecology and distribution.‘Thome Ecological Institute,4860 Riverbend Road, Boulder, Colorado80301, U.S.A.@The Arctic Institute ofNorth America

D.J. COOPER24815w.FIG.1. Sketchl54W15O'wmap of northern Alaska.METHODSWhen trees were found, their positions were recorded bytaking compassreadings to high peaksin the area. Height of thetree was measured and its elevation, habitat and health noted.Where possible a minimum age of the tree was estimated bycounting the bud scale scars on the trunk. Trees would have tobe cut down toaccurately determine their age, and this was notdone. Regional vegetation wasclassified according to standardBraun-Blanquet methods (Westhoff and Maarel, 1978) and ispresentedinCooper(1983). NomenclaturefollowsthatofHulten(1974) for vascular plants andThomson(1979) forlichens. Bedrock types are from the mapof Brosge and .S. SurveyPass 1:250 OOO scale topographic map.RESULTSTree data is presented in Table 1, and the locations of thesetrees are shown in Figure 2. The trees occurred on all threethe ArrigetchCreek valley:bedrock types thatoutcropingranite, limestone and shale. Some trees showed dieback on afew branches, but no dead trees were found. While bud scale

TABLE 1. Tree and habitat dataht(cm)45249WHITE ationtY Pe8845679601111298146526.51067245.33.7 7.01.6.o3.52.6-- WW-870 .6 li1.o.5- Casc-Droc-SflatflatNSliSliSES cshdist km distance from the nearest cone-bearing trees.circum cm circumference of bole, 1 cm above the ground surface.Cate-Clst Alliance Cassiope tetragona-Cladonia stellaris.Casc-Vaul Association Carex scirpoidea-Vaccinium uliginosum.Casc-Droc Association Carex scirpoidea-Dryas octopetala ssp. octopetala.Heal-Elin Alliance Hedysarum alpinum ssp. americanum-Elymusinnovatus.Talus broken rock.gr granite and gneiss.li limestone and marble.sh shale.scar counts were not doneof all the trees, it is clear that thesetrees do not represent theeffects of just a few warm summers.Tree number 9(Figs. 3 and4) occurred in a coarse limestoneblockfield at 1465 m, the highest in elevation. Only scatteredvascular plants werepresent, and thisparticular tree occurred inapatchof Salix arctica. The highesttrees all occurredonsouth-facing slopes and were in protectedhabitats. Trees 2, 11and 12occurred in shallow snowbeds. Trees 10, 11 and 12occurred onor very near ridgetops. Tree 10 is shown in Figure5. The broad, sloping, south-facing limestone andshale uplandsin the Arrigetch valley havesparse and openvegetation, and treeseedlings probably have less severe competition than in more(Picea marianadensevegetation.Althoughblackspruce[Mill.] B.S.P.) occurs at treeline, no seedlings above treelinewere found. Occasionalplantsof alder (Alnuscrispa [Ait.]Pursh ssp. crispa) were found at elevations above 1100 m onrocky slopes.Trees 1 , 2 , 3 and 4 occurred inor on the edge of thick lichenmats dominated byCladonia srellaris(Opiz) Pouz Vezda, C.rangiferina (L.) Web. and C. arbuscula (Wallr.) Rabenh. ongranitic parentmaterial (Fig. 6). Because all tributaries ofArrigetch Creek head in granite, the trees found well beyondtreeline in the valley bottom alloccur on granite. Trees 3 and 4were approximately 3.7 and 5 .O km beyond and 100- 150 mabove treeline in the unnamed valley south of the ArrigetchCreek valley. Some treesare within 2 km of cirques. Brubakeret al. (1983) also report whitespruce at great distances beyondtreeline at Redondo Lakein the Alatna Rivervalley. DISCUSSIONWhile regional climate differences between the Alatna andKobuk River valleys may control the smalldifferences in theirestablished treelines, microenvironmental differences appear tohave greater influence on tree establishment and growth today.Growth and survival Piceaof is known to be enhanced by warmgrowingseasontemperatures (Tranquillini, 1979; Black andBliss, 1980). Protected, south-facing and valley-bottom sitesprovide warmer habitats. If present climate ameliorates andthese trees above and beyondtreeline live to reproductive age, aseed source will be availablefor expansion of forests. Whetheror not the individualtrees reported herelive to reproductive age,they indicate that rapid and dramatic changes in treeline asreported by Kearney and Luckman(1983) for Alberta can occurin the southern BrooksRange.These trees also indicate that seed dispersal in the BrooksRange can be considerably greater thanthe 60-90 m from theparent tree under normalconditionsor 300 m with a strong windreported by Zasada (197 1) or within a horizontal distance ofapproximately twice the heightof the tree reported by ViereckandSchandelmeier (1980). EvenMarshall’s 1930 estimate(Marshall, 1970) of seeds blowingup to 1000-1200 feet(305-366 m) is anunderestimate. Wind in the Arrigetch regionis generally light, especially in winter, when intense temperature inversions dominate. Winter snowpack, as seen in Marchand April 1981, is loose and without wind crusts. Althoughsome spruceseeds were seenon the snowsurface in 1981, seedtransport by blowingacross a hard-packed snowsurface probably occurs rarely. Occasional very strong winds were experienced, especially in late summer andearly fall, which is whenseeds are disseminated.These trees in the Arrigetch Creek valleyinclude what appearto be the highest in elevation reported to date for any regionnorth of the ArcticCircle in North America, as well as some ofthe highest in Alaska.

D .J . COOPER250FIG.2. Section of U.S. Geological Survey, Survey Pass Topographical Map1:250 OOO scale, showing locationsof trees cited.NOTE ADDEDDuring the summer of 1985, after this paper was submittedfor publication, the author found white spruceat elevationsup to1495-1510 m on Fireweed Mountain, in the Wrangell Mountains near McCarthy, Alaska. McCarthy is approximately 200miles southof the ArcticCircle. The exact location is61’27’Nlongitude, 143’2’E latitude, in the west central portion ofsection 11, Township 5South, Range 13 East on the McCarthy(B-6) Quadrangle, U.S. GeologicalSurvey 1:63 360 seriestopographic map.The highest cone-bearing trees in thisregionare found at 10601130 m elevation. Trees were notfound in theshrub tundra zone immediatelyabove the forest zone but werescatteredthroughoutthearctic-alpinezoneabove the shrubtundra. The highesttreesoccurredinashalloweast-facingsnowbed on volcanic parent material. Most trees are somewhatgnarled, with dead branches. Ages range from approximately10-13years for small treesto individuals witht r u n k s up to 7 cmin diameter that appearto be older than 50 rien providedfieldassistanceduring the summers in the Brooks Range.I would like to thank both ofthem. This research was funded in part by a contract from the U.S.National Park Service, an Explorer’s Club grant and two grants fromthe University of Colorado.

25 1WHITE SPRUCEFIG. 3. Arrigetch Creekvalley, showing locations of five trees above and beyondtreeline. White spruce are visible as darkobjects along creek. The farthest treeup this valley is number 5 . The established treeline is where the creek makes a thesharp bend to the right. Valley faces northeast.FIG. S. Tree number 10 is seenjust to the leftof the number 10, elevation 1067 m.Treeline is in the valley bottomto the right. Tree number 12 is just off the edge ofridge. South is to the right.at 1465 m elevation. The tape is cm on the rightand inches on the left. The plant above and belowthis tree is Salk arctica.FIG. 6 . TreeFIG. 4. Tree number 9,locatednumber4 , 5 km beyond treeline.Tape is in cm. Light-colored plantsare Cladonia spp. lichens, Grass is Fesruca altaica.

D.J. COOPER252HOPKINS, D.M. 1972. The paleogeography and climatic history of Beringiaduring Cenozoic time. Inter-nord 12:121-150.HULTEN, E. 1974. Flora of Alaska and Neighboring Territories. A manualofthe vascular plants. Stanford Stanford University Press. 1008 p.BLACK R.A., and BLISS,L.C. 1980. Reproductive ecology Piceaof rnarianaKEARNEY, M.S., and LUCKMAN,B.H. 1983. Holocene timberlinefluctua(Mill.) BSP, at tree line near Inuvick, NorthwestTemtory, Canada. Ecologitions in Jasper National Park, Alberta. Science 221:261-263.cal Monographs 50:331-354.BLASTING, T.J., and FRI'ITS, H.C. 1973.Pastclimate ofAlaskaandLARSEN, J.A. rritories: studies in subarctic and Arctic bioclimatology. Ecological Mononorthwestern Canada as reconstructed from tree rings. In: Weller, G., andgraphs 35:37-59.Bowling, S., eds. Climate of the Arctic. 24th Alaskan Science Conference.-.1974. Ecologyof the northern continental forest border.In: Ives, J.D.,Fairbanks, Alaska: Universityof Alaska. 48-58.andBarry, R.G., eds. Arctic and Alpine Environments. New York: Methuen.BROSGE, W.P., and PESSEL, G.H. 1977. Preliminary reconnaissance geo341-369.logic map of the Survey Pass quadrangle, Alaska.U.S.G.S. Open File report-.1980. The boreal ecosystem.New York: Academic Press. 500 p.71-56 1:250,000.MARCHAND, P.J. 1976. Growth and population structure of white spruce inBRUBAKER, L.B., GARFINKEL, H.L., and EDWARDS, M.E. 1983. lateAWisconsin and Holocene vegetation history from the central Brooks Range: the forest-tundraecotone,TwinMountainarea,Alaska. In:Young, S.B., ka.Wolcott,implications for Alaskan palaeoecology. Quaternary Research 20: ernStudies,No.9:COOPER, D.J. 1983. Arctic-alpine tundra ecosystemsof the Anigetch Creek97-123.valley, central Brooks Range, Alaska.Ph.D. thesis, University of gtheCentralBrooksBoulder. 827 p. tables.Range. Berkeley: University of California Press. 173 p.DENSMORE, D. 1980. Vegetation and forest dynamics of the upper DietrichRiver valley, Alaska.M.S. thesis, North Carolina State University, Raleigh. MERTIE, J.B. 1925. Geology and gold placers of the Chandalar Lake quadrangle. U.S. Geological Survey Bulletin 773. 221p.183p.MURRAY, D.M. 1974. Notes on the botany at selected localities in the e,Alaska.FinalReport,variation in the White River valley and Skolai Pass, Alaska and YukonContract No. CX-9000-3-0125 from National Park Service to University ofTemtory. Quaternary Research 7:63-111.Alaska. 218 p.DREW, J.V., and SHANKS, R.E. 1965. Landscape relationsof the soils andNELSON, S.W., and GRYBECK, D. 1978. The Anigetch Peaks and Mountvegetation in the forest-tundra ecotone, upper Firth River valley, AlaskaIgikpak plutons, Survey Pass Quadrangle, Alaska.U.S. Geological SurveyCanada. Ecological Monographs 35285-306.Circular 772-B: B7-B9.ELLIOTT, D.L. ICHOLS,H.1976.Historicalaspectsof thenorthernCanadiantreeline.current regenerative capacity. Ph.D. thesis, University of Colorado, BoulArctic 29:38-47.der. 192 p.ODASZ, A. 1983. Vegetation patterns at the treelimit ecotone in the upper-.1979b. The current regenerative capacity of the northern Canadiantrees, Keewatin, Northwest Territories, Canada: some preliminary observa- Alatna River Drainageof the Central Brooks Range, Alaska. Ph.D. thesis,University of Colorado, Boulder, Colorado. 224p.tions. Arctic and Alpine Research 11:243-251.SAVILE, D.B.O. 1972. Arctic adaptations in plants. Canadian tinversusLabradorAgriculture Research Branch Monograph 6. 81 p.Ungava. Proceedings, Eighth Annual Eastern Canadian Arctic Workshop,THOMSON, J.W. 1979.Lichens of theAlaskanArcticSlope.Toronto:INSTAAR. University of Colorado, Boulder, Colorado. Abstract.University of Toronto Press. 314 p.ELLIS, J.M., HAMILTON, T.D., and CALKIN, P.E. 1981. Holocene glaciaTRANQUILLINI, W. 1979. Physiological ecology of the alpine timberline.tion of the Arrigetch Peaks, Brooks Range, Alaska. Arctic 34:158-168.Berlin: Springer-Verlag. 137p.GARFINKEL, H.L., andBRUBAKER, L.B. 1980.Modemclimate-treegrowth relationships and climatic reconstruction in subarctic Alaska. antcommunitiesinAlaska. Holarctic Ecology 2:228-238.286 872-874.-and SCHANDELMEIER, L.A. 1980. Effects of fire in Alaska andGOLDSTEIN, G. 1981. Ecophysiological and demographic studies of whitespruce Picea glauca (Moench) Voss at treeline in the Central Brooks Range adjacentCanada - aliteraturereview.BureauofLandManagement,Alaska. Technical Report 6.of Alaska. Ph.D. thesis, University of Washington, Seattle.207 p.WARDLE, P. 1974. Alpine timberlines. In: Ives,J.D., and Barry, R.G., eds.GRIGGS, R.F. 1934. The edge of the forest in Alaska and the reasons for itsArctic and Alpine Environments. New York: Methuen. 371-402.position. Ecology 1580-96.WESTHOFF,V., andMAAREL, E. vander.1978.TheBraun-BlanquetHAMILTON, T.D. 1965a. Alaskan temperature fluctuations and trends: anapproach. In:Whittaker, R.H., ed. Ordination and Classification of Commuanalysis of recorded data. Arctic 18:104-117.Junk.nities. b. Comparative glacier photographs from northern Alaska. Jour617-126.nal of Glaciology5479-485.ZASADA, J.C. 1971. Natural regeneration of interior Alaska forests-seed,HAUGEN, R.K., and BROWN, J. 1978. Climatic and dendroclimatic indicesseedbed and vegetative reproduction considerations.In: Fire in the Northernin the discontinuous permafrost zone ofthe central Alaska uplands.In: 3rdEnvironment. Pacific Northwest Forest and Range Experimental on,Alberta,Canada.Portland, Oregon. 231-246.392-398.REFERENCES'

beyond the last cone-bearing trees. Reports of treeline stability in Alaska are varied. As early as the 1920s, Mertie (1925) was surprised to find trees at 1060 m elevation in the Brooks Range, at higher elevation than in the Alaska or Coast ranges. Else- where in the Brooks Range seedlings above treeline are reported

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