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Height and Growth Rings of Salix Lanata Ssp. Richardsonii Along the Coastal Temperature Gradient of Northern Alaska

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Height and Growth Rings of Salix Lanata Ssp. Richardsonii Along the Coastal Temperature Gradient of Northern Alaska Height and growth rings of Salix lanata ssp. richardsonii along the coastal temperature gradient of northern Alaska D. A. WALKER Institute of Arctic and Alpine Research, UniversiQ of Colorado, Boulder, CO, U.S.A. 80309 Received March 14, 1986 WALKER,D. A. 1987. Height and growth rings of Salix lanata ssp. richardsonii along the coastal temperature gradient of northern Alaska. Can. J. Bot. 65: 988-993. Salix lanata ssp. richardsonii from open-tundra and streamside populations were studied at seven sites along a 100-km north-south transect following the Sagavanirktok River from the Alaskan arctic coast to the foothills of the Brooks Range. Mean July temperatures along this transect vary from 2.6 at the coast to 10°C at the base of the foothills. Mean maximum heights of the sampled open-tundra willows increased from 10 + 2 at the coast to 37 + 8 cm at the southern end of the transect. Mean maximum heights of sampled streamside willows increased from 0 at the coast to 147 i 25 cm. The mean maximum height of willows in both habitats showed very strong correlations with thawing degree-days. Mean growth-ring widths increased from 92 + 20 at the coast to 188 i 57 pm at the southern end of the transect and were also highly correlated with the temperature gradient. The results are discussed in light of other arctic studies of willow growth rings and Cantlon's system of vegetation subdivisions within the Alaskan arctic. WALKER,D. A. 1987. Height and growth rings of Salix lanata ssp. richardsonii along the coastal temperature gradient of northern Alaska. Can. J. Bot. 65 : 988-993. Le Salk lanata ssp. richardsonii, provenant des tundras exposCes et des populations situCes pres des ruisseaux, a CtC CtudiC i sept sites le long d'un transecte nord-sud de 100 km longeant la Riviere Sagavanirktok, de la cBte arctique de 1'Alaska aux contreforts de la chaine de montagne Brooks. Les tempkratures moyennes de juillet le long de ce transecte varient de 2.6 i la c6te i 10°C i la base des contreforts. La moyenne des hauteurs maximums des saules des tundras exposCes augmente de 10 i 2 i la cBte i 37 + 8 cm i 1'extrCmitC sud du transecte. La moyenne des hauteurs maximums des saules riverains augmente de 0 i la cBte i 147 5 25 cm. La moyenne des hauteurs maximums des saules des deux habitats montre de trks fortes corrClations avec des degrCs-jours de dCgel. La largeur moyenne des anneaux de croissance passe de 92 + 20 i la cBte i 188 i 57 pm i llextrCmitC sud du transecte et Ctait Cgalement en Ctroite corrClation avec le gradient de temperature. Ces rksultats sont discutCs i la lumibre d'autres Ctudes arctiques sur les anneaux de croissance du saule et du systbme de Cantlon sur les subdivisions de la v6gCtation i 1'intCrieur de l'arctique de 1'Alaska. [Traduit par la revue] Introduction Shanks 1968; Young 1971). Most of the growth-form changes The stature and productivity of shrubs are worldwide criteria along this north-south transect are on a scale of a few centi- for dividing the Arctic into vegetation subzones (SOrenson metres. The major exception to this is the height of riparian 1941; Polunin 1951; Bocher 1954; Andreev 1966; Alexan- willows, which increase from prostrate forms at the coast to drova 1970; Bliss 1981). For example, Nicolas Polunin (1951) nearly tree-sized shrubs at the northern edge of the foothills. Less pronounced changes to shrub physiognomy occur on the commented that " . .the vegetable productivity on land increases more markedly than the totality of species as we open tundra. travel further south. ." He further stated that this increased In this study, I examine the height and growth rings of the productivity is due to the greater importance of shrubs and most common erect willos, Salix lanata L. ssp. richardsonii dwarf shrubs. Although biogeographers have long emphasized (Hook.) A. Skvortz., along the coastal temperature gradient the importance of shrub physiognomy for defining arctic and relate the results to recognizable vegetation zones on the vegetation zones, there are few data that quantitatively show coastal plain. The study transect follows the Dalton Highway how the shrub heights and growth rings respond to temperature along the Sagavanirktok River for 100 km southward from the along continuous latitudinal transects. arctic coast (70'15' N, 148'20' W) to the base of the arctic The central arctic coastal plain of northern Alaska (Fig. 1) is foothills of the Brooks Range (69'25' N, 148'35' W) (Fig. 1). a good area to examine the vegetation effects of a steep coastal I chose Salix lanata for three principal reasons: (i) it is a temperature gradient because the vegetation zonation is highly woody species with a multiyear growth record; (ii) it occurs compressed in this region. Three of Young's (1971) four arctic abundantly along the entire transect; and (iii) it exhibits nearly floristic zones occur within 50 km of the Beaufort Sea coast in its full range of height growth potential along the Sagavanirk- the vicinity of Prudhoe Bay. The only other area of the Arctic tok River transect. where Young (1971) shows this to occur is a region east of the Methods Kolyma River in the USSR. The central Alaskan arctic coastal plain near the Sagavanirktok River is extraordinarily flat with Fieldwork most environmental factors other than temperature varying I measured the heights and growth rings of S. lanata at eight loca- narrowly so that it is possible to study the vegetation effects of tions along the Sagavanirktok River (Fig. l). Five of these locations a steep coastal temperature gradient with minimal confounding "East Dock," "Drill Site 9," "Deadhorse," "Mile 350," and "Pipeline Intersection," were selected within 40 km of the coast to effects from other variables. measure willow growth in the steepest part of the temperature Along the gradient, inland from the coast there is an increase gradient. The two stations at the southern end of the transect, "Pump in the total number of plant species and subtle changes to the 2, Coastal Plain," and "Pump 2, Foothills," were established within overall tundra vegetation physiognomy (Clebsch 1957; 2 km of each other to determine whether the better drained environ- Cantlon 1961; Wiggins and Thomas 1962; Clebsch and ment on the upland affects the height of open-tundra willows. Printed in Canada 1 IrnpnrnC au Canada WALKER WILLOW +TRANSECT - - .-- 0 0 5% E w '7 L gz 2 w Willow k U m Collection 1 N N Z .- Sites aa - fQ v 5s E .-g g zg an L a I no I. I Brooks Range '- '- Coastal Plain '. Foothills I Latitude I I I I I I I I I 68" 69" 70" Distance to I I I I I I Arctic Coast (krn) 200 100 0 FIG. 1. Location of temperature stations and willow collection sites. At each location the heights of 50 of the tallest willows were AGE-CLASSES OF WILLOW COLLECTIONS measured in a streamside site and 50 on a nearby open-tundra site. I selected the tallest willows because I wanted representatives of the willow's full growth potential at a given site. I also haphazardly SITE3 collected 50 willows without their roots from each open-tundra loca- tion (except the East Dock) for analysis of growth rings. The willows were cut at ground level with a pocket knife. Growth-ring analysis MILE The willows were sectioned and mounted according to procedures 350 outlined by Jensen (1962). The stems were dehydrated for 2 weeks in -'n alcohol. They were then sectioned in 10 pm thick slices, mounted, fixed by Mayer's albumin, and allowed to dry for 24 h. They were stained with safranin and fast green, cleared in clove oil, and mounted I I - I I in Permount medium. O FRANKLIN lo. BLUFFS I I The slides were studied by projecting them on a wall, using a Leitz I X 2. microscope-slide projector. The annual rings were marked on long 5 I - I i I I strips of paper along two radii for each section. The two radii were PUMP2 10 I I compared to locate rings that were missing on one or the other (COASTAL PUlNl Him+, sample. Mean ring width of an individual willow was calculated by I I I dividing the mean radius by the number of growth rings. The 25 PUMP 2 10 I ~FOOTHILLS~ willows with the clearest growth records were selected from each 2 station for the final analysis. Most of the sectioned willows, 107 of 175, were in the 16- to - 30-year age-class (Fig. 2). This subset was used for regression analy- Number of Growth kings - sis of mean increment widths to compare roughly equivalent age FIG. 2. Age-class of willow collections. The 107 willows between groups. The mean ring widths were calculated at each station for all the ages of 16 and 30 (broken lines) were used for one portion of the 25 willows and for the willows in the 16- to 30-year age-class. The regression analysis. data were divided into two subsets. The first consisted of data from all 175 willows, and the second consisted of data from the 107 willows in have shown that the best-fit equations are obtained by measuring the 16- to 30-year age-class. the distance to the coast along the primary wind vector (N 75" E). Data analysis Both distances were tried in these correlation analyses, and there were Height and growth-ring widths were regressed against distance only slight differences in the results; therefore, for clarity of presenta- from the coast, predicted mean July temperature, and predicted tion, only the results using the shortest distance to the coast are pre- thawing degree-day s (TDDs), using linear and exponential regression sented here.
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