Forest Vegetation Related to Elevation in The White Mountains of

by William E3. Leak and Raymond E. Graber

USDA FOREST SERVIC EARCH PAPER NE-299

&ORTHEASTERN FOREST BFYERIMENT STATION FOREST SERVICE, U.S. DEPARTMENT OF AGRICULTURE 6816 MARKET STREET, UPPER DARBY, PA. 19082 WARREN T. DOOLITTLE, DIRECTOR The Authors WILLIAM B. LEAK joined the Northc;istcrn I"ortlst Expcrirnent Station in 1956 and now is principal silvi- culturist at the Station's Forclstry Scicnctbs 1,:it)or:ttory at Durham, N. H.

RAYMOND E. (;RABER is :1 plant ecologist :it the Northe;tstern Fortst Experimcr~t P;t:ltion9s Forchst ry Sciences L:ibor:~tory ;it Durh:lm, N. H. Hcl is it grad- uate in forcst maniigcmcnt of thch Univclrsity of 'L"Ii;isl.l- ington and has studied forcst ecology itt Orclgon St:itt~ University and botany :it the Univcxrsity of New H;lmpshire.

MAXI'SCRIPTRECEIVFD FOR P17NLICtSTION 8 E'EJ<~<~AKY I!f7 1 Forest Vegetation ated to E in The White Mountains of New Hampshire

ABSTRACT Maximum tree size and species composition are related to eleva- tion on (disturbed by logging) and -Mount Whiteface (uncut) in the White Mountains of New Hampshire. Species migrational trends and differences between the two moun- tains in species elevational limits indicate that both hardwoods and softwoods will move to higher elevations in areas where cutting and heavy disturbance are eliminated.

I N THE MOUNTAINOUS areas of New other mountain systems in to- England, elevation above sea level is one of pography, geology, and past use. the most important factors affecting the species composition and structure of forest vegeta- tion. Because these forest characteristics relate AREAS AND METHODS closely to resource values and management Our research dealt with two mountains in possibilities, there is need for specific infoma- New Hampshire: Mount WBshington, which tion about the influence of elevation. is 1,917 meters (6,288 feet) high, and Mount Much of the past research on mountain Whiteface, 1,215 meters (3,985 feet) high. ecology in New England has dealt with the Mount Washington is a heavily used recrea- alpine zone above timberline (Bliss 1963 and tional area, whose lower slopes up to about 11966, Harries 1961, Griggs 1946). Information 914 meters (3,000 feet) were irregularly about elevz,tional relationships is available for logged up until the late 1800s. The study area the Adirondacks (Wolway and Scott 1969) on Mount Whiteface, in the Bowl Research arid the of Vemont (Sic- Natural Area, has never been logged. cama 1968). However, outside of the detailed In the summer of 1971, a trensect was laid vegetational work on the Hubbard Broofr out on the southeastern slope of Mount Wash- watersheds (Bormann et al. 1970), little spe- ington, beginning at an elevation of 632 meters cific information is available for the White (2,073 feet) near the Appalachian Mountain Mountain range, which differs somewhat from Club's Pinkham Notch base camp and ending at 1,373 meters (4,505 feet) at the base of Mount IYashington before the turn of the cen- Nelson's Crag. At every 10-meter rise in tury. For the typical northern hardwoods- elevation, measured with an aneroid barom- beech, yellow birch, and sugar maple-no eter, a series of nested plots was set out. substantial diminution in tree size was evi- On the main plot, which was 3 x 20 meters, dent on Mount Whiteface up to 900 meters. laid out along the contour, the diameter above For the spruce-fir components-red spruce, root swell was measured on all trees of at balsam-fir, and paper birch-tree size remains least 50 millimeters diameter. Ages of the large up to 1,100 meters on Mount Whiteface largest and smallest stems (including those and 1,200 meters on Mount Washington. less than 50 mm) were determined by boring, For those who use diameter nieasurements sectioning, or counting terminal bud scars. at breast height, dbh in mrn can he estimated On a 3 x 3-meter subplot, diameter was rneas- from diameter above root swell by: ured on all stems that were between 1.372 Hardwoods: Dbh (mm) = -11.68 + 96 meters tall (4.5 feet) and 50 mm diameter. (dia. rnm, root swell) Smaller vegetation was measured on a 1 m' It' = .997 circular subplot. Diameters were measured S,,= 111.04 just above root swell fright at the point of i90ft~coods: Dbh (mm) == --17.80 + -95 curvature for the root swell on the uphill side (dia. mm, root swell) of the tree) because this point can be identi- R2= .996 fied on all sizes of trees and it is approximately S,,= 10.43 the point used in making increment borings. Dbh proves useless on young trees and on stunted trees near timberline. Basa6 Area and Species Compositisn During the summer of 1972, a compass line On both Mount Washington and Mount was run up the northeast side of Mount Whiteface, basal areas per acre increased mod- miteface, beginning at an elevation of 610 erately with elevation as the proportion sf meters (2,000 feet) at the softwoods increased. Then basal area seemed in the base of the Bowl and ending at 1,201 to decline at 1,100 meters on Mount nThite- meters (3,940 feet) near the mountain Lop. face and at 1,300 meters on Mount Washing- At every 50-foot (15.24 rn) rise in elevation, ton. Percent species composition differed basal areas by species were accumulated with somewhat between the two areas, partly be- a 10-factor p~sm.The largest tree per species cause of the differences in past land use. in the prism plot was measured for diameter In general, Mount Washington has a wider Just above the root swell and aged with an range of species than the late-successional increment borer. The smallest stems per spe- stands on Mount Whiteface (table 2)- How- cies on the prism plot or a I x 10-meter sub- ever, on both areas the typical northern hard- plot also were measured and aged. woods are most abundant in the 600 to 700 On both transects, elevations were double- meter elevation classes. Red spruce appears in checked at several points, and the errors were greatest abundance in the 800 to 1,000 meter distributed. These corrected plot elevations elevation classes. Balsam-fir first becomes an were used in analyzing the data. important component in the 800-meter class and continues to increase in relative abun- dance with higher elevation. On ,Mount mrash- RESULTS AND DISCUSSION ington, paper birch is abundant from the 700- through 1,100-meter class, while on &4ount Maximum Tree Diameter FVhiteface it is evident mainly in the 1,000 to 1,108 Maxirnum tree diameters commonly ran meter classes. eonsidesably larger on Mount Whiteface than on Mount Washington except at elevations 'pecies' Ranges above 94f9 TP+~^.S.; {tahh j I), This pattern Based on the occurrence of stems more than probably resulted from the logging activity on I year old, elevational ranges of species were Table 1 .-Maximum tree diameters, by species and 706-meter elevation classes (600 to 696, efc.1 [In mm, above root swell] Elevatian class Red Paper Mt. Striped Mt. Red Balsam Eastern Black (meters) BrP"'l %::? 2 maple birch maple maple ash Aspen spruce fir hemlock spruce

MOUNT WASHINGTON 179 370 301 66 126 -

Table 2 .-Average species composition, by species and 7 00-meter elevation classes [In percent of total basal area 1 ------Elevation class ~~~~h Yellow Sugar Red Paper Mt. Striped Mt. Red Balsam Eastern Black (meters) birch maple maple birch maple maple ash spruce fir hemlock spruce MOUNT WHITEFACE 600 29 25 46 ------700 28 32 15 - 6 - 2 5 - 10 2 - - 800 - 18 4 - 8 - 4 2 - 42 22 - - 900 - 2 - - 4 - - - - 61 33 - - 1,000 - - - - 15 - - 2 - 37 46 - - 1,100 - - - - 38 - - 3 - 14 45 - - 1,200 ------24 76 - - MOtTNT WASH1NGTON 2 68 4 17 4 1 1 - - - - 45 12 - 32 - 1 - 7 2 - 17 - - 12 - - 3 - 19 - - - - 14 - 1 - 38 - - - - 15 - - - - 37 - - - - 13 - - - - 16 - - - - 5 - - 3 Figure 1 .-Erevationat ranges of woody species on Mount Whiteface and Mount Washington, based on occurrence of stems over 1 year old. Transects began at 610 and 632 meters above sea level and ended a3201 and 1,373 meters, respectively.

HOBBLEBUSH

MT. ASH

BALSAM FIR

BLACK SPRUCE

RED SPRUCE

PAPER BIRCH I

YELLOW BIRCH

BEECH

MT. MAPLE

STRIPED MAPLE m MOUNT WHITEFACE

RED MAPLE !1 0MOUNT WASHINGTON SUGAR MAPLE

600 700 800 900 1,000 1,100 1,200 1,300 1,400

ELEVATION, METERS

graphed for each area (fig. 1).Occasional out- remain as a permanent resident in the spruce- liers were ignored. Northern hardwoods gen- fir association. erally occur below 850 meters, although yel- The same conclusions can be made from low birch is found up to 950 meters. The soft- preliminary information about the detection woods (and paper birch) are found throughout of species' migration. When maximum and a much broader range although, as noted minimum ages are plotted over elevation (or earlier, they become predominant above 800 distance), it becomes possible to detect ad- meters. vancing, stable, or retreating species fronh. Note that the elevational ranges of the The data from Mount WashingLon, already hardwoods are consistently narrower in the published, indicated that the hardwoods gen- disturbed Mount Washington stands than in erally were advancing uphill, while paper birch the uncut Mount Whiteface stands. The op- and spruce-fir were retreating. In contrast, the posite is true with softwoods: softwood (and more recent data from Whiteface indicate that paper birch) ranges are narrower on Mount the northern harrdwmds such as beech (fig. 2) whiteface than on Mount Washington. and sugar maple have attained almost per- It seems l&ely that, when cutting is elimi- fectly stable up"bll fronts. Yellow birch (fig. nated, northern hardwood ranges will expand 3) still seems to be advancing uphill, while to their upper linrnt---850 do 950 meters--- the lower end of its elevational range is re- while softwood ranges will retreat. The end treating uphill drastically (although the shape result wmld be a general reduction in lower- of the lower front is nod well defined by the altitude softwoods and a fairly st~r'i;line of data), The softwoods and papr birch (fig, 4) dmarcation between the northern hardwood seem to be retreating only sporadically and And spruce-fir associations. Paper birch would slmi3y uphill. 700 800 900 1000 1100 1200 1300

ELEVATION, METERS

Figure 3,-Maximum and minimum age for yellow birch over elevation, showing approximate shape of migra- tional fronts. Mount Whiteface.

ELEVATION, METER Figure 4.-Maximum and minimum age for paper birch over elevation, showing approximate shape of migra- tional fronts. In this case, the curves are drgwn as en- velope curves, ignoring some of the irregularities in plotted points. Mount Whiteface.

ELEVATION, METER

IN CONCLUSION This information on species development Bliss, L. C. 1963. ALPINE PLANT COMMUNITIES OF THE PRESf- related to elevation may be useful in forest DENTIAL ZANGE, NEW HAMPSHIRE.Ecology 44:678- land-use planning. The infomation from 697- Mount Whiteface should be especially useful B1g6$-giANT ,, ,,,, ,,,,- in evaluating the potential species and sizes VIRONMENTS ON MT. WASHINGTON,NEW HAMP- that can develop at specific elevations. The Eco'- 36(2) :125-155. Bormann I?. H., T. 6. Siccama, 6, E. Likens, ad infomation in this paper cannot be extended R. w. ~&i~~a,~. beyond the White :\;lountains, although 1970. THE ~~UBBAKWBROOK ECOSYSTEM STvDY: COMPOSIa7ON AND DYNAMICS OF THE TREE STRATUM. similar conditions may exist at similar lati- ~,,1. ~~~~g.- 40:373-3=, tudes in , , and New Uork. Griggs, R. F. 1946. THETIMBmLINES OF NORTHERN AM~ICAAND evidence is given On changes ,N,,,ETA,,N. Ecology 27: 275-289. in species elevational relationships in dis- Harries, H. turbed and uncut areas. Without logging, t::kE, ~~&",",,",N~l~~o~~~~~o$P hardwoods apparently would progress, and com~,AND MOVTNG $nm.Eeol.. Soc. Amer. Bull. softwoods recede, to a point of stability at 42:108- higher elevations; and the line of demarca- H7\zy'$~~~~kI~!E$,","O~I;I:,","~tt. AT tion between the two associations tend WBITGFACE~IOUNTMN IN THE ADIRONDACMS.Sbte Univ. N. Y. Atmss. Sci. Res. Gent. Rep. 92. Al- to become more distinct. bany. 236 p. Siccarna, T. G. 1968. ALTITUMNALDZSTRIBUTZON OF FOREST VE@E;TA- TION IN RELATION TO SOIL AND CLIMATE ON THE; SLOPES OF THE GREENMOUNTAINS. Ph.D. thesis, Univ. Vt., Burlington. 618 p. Scientific Name American beech Fagus grandkfo Eia E hrh. Yellow birch BeluEa ulteghaniensis Britton Paper birch i ineluding mountain Betuta papyrifera Marsh., in- paper birch ) eluding B.ccrrdifolia Reg. Sugar maple Acer saceharum Marsh. Red rnaple Aeer rubrum L. Red Spruce Pieea rubens Sarg. Balsam-fir Abies balsamea ( L,)Mill. E;-\stern hemlock Tsuga canadensis ( I;.) Carr. Black spruce Picea mariana iMill. ) B.S.P. Hobblebush Viburnum alnifolium hlarsh. Mountain rnaple Aeer spicalum Lam. Striped maple Aeer pensy lr ianicum L. Mountain-ash Sorbl~snmeriearza Marsh. THE FOREST SERVICE of the U. S. Depart- ment of Agriculture is dedicated to the principle of multiple use management of the Nation's forest re- sources for sustained yields of wood, water, forage, wildlife, and recreation, Through forestry research, cooperation with the States and private forest owners, and management of the National Forests and National Grasslands, it strives - as directed by Congress - to provide increasingly greater service to a growing Nation.