United States 

Department of 

Agriculture 

Survival, Growth, and Juvenile- Mature 

Forest Service Correlations in a West Virginia 

Northeastern Forest 

Experiment Station 

Sugar Maple Provenance Test 

Research Paper NE-689 

25 Years After Establishment 

Thomas M. Schuler 
Abstract 

Survival, total height, diameter at breast height (d.b.h.), and stem quality of sugar 

maple trees of different provenances were compared 25 years after establishment 

in north-central West Virginia. Provenances were from Michigan, Minnesota, West 

Virginia, Massachusetts, New Hampshire, Vermont, Maine, and Quebec, Canada. 

There were significant differences between provenances for all traits except stem 

quality. By provenance; total tree height ranged from about 49 to 37 feet; d.b.h. 

from 6.7 to 3.6 inches; and survival from 100 to 15 percent. The predictability of 

total tree height 25 years after establishment based on mean provenance height at 

age 2, 6, 10, and 15 years is discussed. Results suggest that juvenile height 

growth may be a good predictor of mature height performance, thus decreasing 

the need for rotation-length trials. 

The Author 

THOMAS M. SCHULER, research forester, received a B.S.F. degree in forestry 

from Purdue University and an M.S. degree in silviculture from Colorado State 

University. In 1992 he joined the forest management research project at the 

Northeastern Forest Experiment Station's Timber and Watershed Laboratory at 

Parsons, West Virginia. 

Manuscript received for publication 11 March 1994 

Cover photo: Sugar maple provenance outplanting in north-central West 

Virginia 25 years after establishment 

USDA FOREST SERVICE 

5 RADNOR CORP CTR STE 200 

PO BOX 6775 

RADNOR PA 19087-8775 

August 1994 
introduction 

Sugar maple (Acer saccharurn Marsh.) is an important 

hardwood species of the eastern, central, and northern 

hardwood forests of the United States and the southern 

hardwood forests of Canada. The wood of sugar maple is 

highly valued for furniture and flooring and the sap is used 

extensively for the production of maple syrup. Sugar 

maple represents approximately 6 percent of the total 

hardwood volume of the United States and in most 

regions, the growing-stock volume is increasing (Godman 

and others 1990). Yet, while data are available on the 

genetic variability of other important hardwood species 

such as white ash, black walnut, and black cherry, there is 

relatively little information on the genetic variability of sugar 

maple as related to survival and growth (Kriebel 1975). 

This paper reports on the West Virginia outplanting of a 

Northeastern Forest Experiment Station rangewide study 

of sugar maple. As part of this rangewide study, a sugar 

maple provenance test was established in north-central 

West Virginia in 1968. Seed sources were from throughout 

the Northeastern and north-central United States and 

Southern Canada. In part, the objectives of this study 

were to determine the nature and extent of genetic 

variation between provenances and to compare 

characteristics of juvenile and mature growth. At the 

inception of this study, characteristics of mature growth 

were expected to be observed in about 40 years. This 

paper reports 25-year growth, survival, and characteristics 

of stem form for sugar maple trees of different provenances, 

and examines the potential for predicting characteristics of 

mature trees irom juvenile traits. High correlations 

between juvenile and mature traits imply that shortened 

breeding cycles and increased gain in genetic improvement 

per unit of time are possible through the selection of 

superior genotypes at juvenile ages. 

Methods 

The West Virginia sugar maple plantation was established on 

the Fernow Experimental Forest (39.0°N, 79.7"W) in 1968 

near Parsons, West Virginia. Mean annual precipitation on 

the Fernow is about 58 inches and is distributed evenly 

throughout the year. Mean annual temperature is 48°F 

and the length of the frost-free season is approximately 

145 days. 

The study site was previously occupied by second-growth 

hardwoods consisting of northern red oak (Quercus rubra 

L.), yellow-poplar (Liriodendron tulipifera L.), sugar maple, 

black birch (Betula lenta L.), white ash (Fraxinus 

arnericana L.), and other hardwood species. The site 

index for the area averaged about 70 for red oaks. Soils in 

the area are mapped as Dekalb channery loam (coarseloamy, mixed, mesic Typic Dystrochrept). The area has an 

easterly aspect and a slope of 5 to 15 percent. The 

elevation of the study area is about 2,800 feet. 

In preparation for planting, the sawtimber and pulpwood 

were harvested and all remaining stems were felled. Brush 

and small stems were pushed off to the side to facilitate 

planting. Surface disturbance was kept at a minimum. 

Five contiguous blocks, each 130 by 45 feet, were laid out 

with the long dimension roughly parallel with the contour. 

Originally, 2-year-old seedlings from 14 provenances each 

represented by progeny from seven or eight parent trees in 

two-tree plots were planted at a spacing of 5 by 5 feet, 

bringing the total number of trees in the initial outplanting to 

1,109. 

This outplanting included provenances from Michigan, 

Maine, Massachusetts, Minnesota, New Hampshire, 

Vermont, New York, and Quebec, Canada (Table 1). Two 

naturally occurring sugar maple seedlings of about the 

same size as the planting stock were selected in the area 

immediately adjacent to each block and included as 

controls, bringing the total number of provenances to 15. 

However, their location outside of the plantation grid, 

unknown age, and fewer trees per provenance make their 

value as a control group questionable. This problem was 

exacerbated in 1982 when the rest of the plantation was 

thinned but the local source was not included in the 

thinning due to the fewer trees in the provenance relative 

to the other provenances. The thinning removed the 

poorest tree in each of the two-tree plots. When only one 

tree of a pair had survived, the surviving tree was not cut. 

Survival percentages reported here are not adjusted for the 

thinning due to the inconsistent application. 

The latitude of the 15 provenances ranged from 39"N in 

West Virginia to 47"N in Minnesota. Longitude ranged 

from 70°W in Maine to 94"W in Minnesota. The elevation 

of the source material ranged from 350 feet in Quebec to 

2,800 feet in West Virginia. The planting stock was grown 

in a USDA Forest Service nursery in Burlington, Vermont, 

for 2 years prior to being shipped to West Virginia before 

the beginning of the third growing season. After planting, 

each tree was mulched with a sheet of black plastic, 30 by 

30 inches. A screen cylinder of 1/4-inch hardware cloth, 6 

inches in diameter and 18 inches long was placed around 

each tree in the summer of 1968 to protect the stems from 

rodents. The protective screens were removed in 1973. 

Total tree height, d.b.h., survival, and remarks pertaining to 

the form of the main stem were recorded periodically. Height 

measurements began after the second growing season; the 

most recent measurements were recorded at the end of the 

25th growing season in November 1992. Stems with 

desirable form and dominant or codominant relative crown 

position were designated as potential crop trees at the time 

of the 1992 measurement. Potential crop trees were defined 

as dominant or codominant trees with no forks below 17 

feet, and no undesirable stem form characteristics such as 

cankers, excessive crook, butt-log rot, birdpeck, or other 

characteristics that would reduce the potential grade of the 

butt log. 

Data were analyzed by analysis of variance using linear 

model procedures incorporating the randomized block 

design with seed sources nested within provenances. Plot 

means of the surviving trees were used for tree height, 

d.b.h., and analyses of stem form (Bresnan and others 

1992; Freese 1967; Rink and Kung 1 WI), and were 
Table 1.-Identification and geographic origin of the 15 provenances included in 

north-central West Virginia sugar maple plantation established in 1968 

Provenance County and state Latitude ON Longitude OW Elevation 

Ingham, MI 

Berkshire, MA 

Sultivan, NH 

Addison, VT 

Lewis, NY 

Franklin, ME 

Iron, MI 

Quebec, Canada 

Cass, MN 

Mille Lacs, MN 

Franklin, VT 

Chittenden, VT 

Rutland, VT 

Bennington, VT 

Tucker, WV 

Feet 

600-1 400 

1500 

400 

900-1 000 

900 

560 

1550 

350 

1300 

1300 

600 

700-800 

1550-1 650 

900-1 1 00 

2800 

weighted by the number of trees they represented to 

account for unequal variances. Differences in survival and 

stem form were tested for after arcsin transformation of 

these variables. The West Virginia provenance was not 

included in the survival analysis since it was not thinned 

while the other provenances were thinned. Regression 

analysis was used to estimate relationships between 

performance factors and provenance latitude, longitude, and 

elevation. Data were analyzed and tested for significance at 

the 5 percent level unless otherwise noted. The 6- and 10- 

year results were reported by Wendel and Gabriel (1974, 

1980) and were used for contrast where appropriate. 

Results and Discussion 

At the end of 25 years, analysis of variance revealed 

significant differences in provenance survival, total height, 

and d.b.h. Stem quality also was evaluated but there were 

no significant differences among provenances. Table 2 

reports the provenance means for survival, d.b.h., total 

height, and stem quality. Latitude, longitude, and elevation 

of provenances were not correlated with provenance 

performance. 

Seed sources nested within provenances were not 

significantly different for any of the traits evaluated and 

reflect a trend in declining variation between families 

within provenances. After 6 years, average height was 

significantly different between families within eight 

provenances. After 10 growing seasons, there were 

significant differences between families within only four of 

the provenances (Wendel and Gabriel 1980). 

Quality 

Stem quality, as in previous remeasurements, did not 

show significant differences related to geographic origin. 

Overall, stem form was fair and appeared to be improving 

from earlier observations, though direct comparisons are 

difficult due to the increased size of the trees. At age 25, 

approximately 27 percent of the trees qualified as crop 

trees. The range of values in this category included 

several provenances above 40 percent and several at 10 

percent or less. The most common problem with stem 

quality was low forking. At the time of the outplanting it 

was noted that many of the seedlings had multiple stems, 

and early plantation cleanings may have promoted the 

persistence of these multiple stems. However, stem form 

was not recorded and it is impossible to determine if there 

was a seed-source component to the observed 

characteristics of stem quality until these observations 

were initiated in 1978. It should be noted that the seed 

sources in this study were selected based on sap-sugar 

characteristics. Observed stem-form or growth 

characteristics should not be interpreted as potential 

performance characteristics of a rangewide assessment 

of superior sugar maple trees with respect to form or 

growth rate. 
Table 2.-Mean total height, d.b.h., survival, and stem quality measured as percent of crop trees for all sugar maple 

provenances in north-central West Virginia plantation 25 years after establishment 

Location 

(origin) 

Percent Percent Percent Percent 

of mean of mean of mean Crop of mean 

Height height D.b.h. d.b.h. Survival survival trees crop trees 

Chittenden, VT 

Lewis, NY 

Sullivan, NH 

Berkshire, MA 

Bennington, VT 

Franklin, VT 

Iron, MI 

Addison, VT 

Franklin, ME 

Rutland, VT 

Quebec, Canada 

Cass, MN 

Ingham, MI 

Mille Lacs, MN 

Tucker, WV 

Mean 

Survival 

Feet 

48.6 

48.6 

47.2 

46.9 

46.8 

46.3 

46.1 

45.5 

44.8 

44.7 

44.3 

42.4 

42.1 

37.6 

33.6 

Inches 

6.2 

5.8 

5.8 

5.6 

4.8 

5.3 

6.0 

5.1 

5.1 

5.5 

5.4 

5.4 

4.8 

3.6 

4.0 

Percent 

41 

34 

42 

34 

30 

39 

31 

39 

23 

41 

30 

15 

43 

17 

100 

Percent 

44.2 

40.8 

36.2 

30.2 

33.4 

37.8 

7.4 

22.0 

45.0 

24.4 

13.4 

30.6 

18.8 

6.6 

10.0 

Mean provenance survival was 37 percent. Survival 

ranged from 100 percent for the unthinned West Virginia 

provenance which was excluded from the analysis of 

variance of survival to 15 percent for the provenance from 

Cass County, Minnesota. The highest survival rate of the 

14 thinned provenances was from lngham County, 

Michigan. This provenance had a survival rate of 43 

percent, 16 percent greater than the mean for all 

provenances. Perhaps the most noteworthy aspect of 

the differences in provenance survival is that both of the 

most westerly provenances from Minnesota also had the 

poorest survival. 

Because full site occupancy does not occur until after 

crown closure, survival is not a good predictor of future 

performance until competitive interaction occurs between 

individual trees. Significant differences in provenance 

survival rates did not occur until 15 years after plantation 

establishment. At age 25, the provenance effect 

accounted for 60 percent of the total variation in survival 

in the analysis of variance. This contrasts with the 2-, 6-, 

and 10-year results where survival was not significantly 

different between provenances and overall survival was 

above 90 percent. A closer initial spacing would decrease 

the time required until competitive interaction occurs and 

likely would magnify differences in fitness at a much 

earlier age than seen here. 

The differences in survival have been biased downward 

by the thinning that occurred after 15 years. Those 

sources with higher survival were thinned more heavily 

because they had a greater percentage of two-tree plots. 

Stem Diameter 

Mean provenance d.b.h. 25 years after establishment was 

5.2 inches, with the largest trees originating from 

Chittenden County, Vermont (Table 2). This source 
averaged 6.2 inches and was 19 percent greater than the 

mean for all provenances. Provenance differences for 

stem diameter were significant and accounted for 44 

percent of the total variation in this tract. The source from 

Mille Lacs County, Minnesota, had the smallest mean 

d.b.h. of 3.6 inches and seems poorly suited for the 

growing conditions of the central Appalachian region as 

reflected by both poor survival and growth. As expected, 

the results for mean provenance d.b.h. and the results 

for mean provenance total height were highly correlated 

(r = 0.78). 

Total Height 

Overall provenance average height was 44.4 feet at age 

25. Average tree height varied significantly between 

provenances and the provenance effect accounted for 56 

percent of the total variation in tree height. The tallest 

trees in the plantation originated from seed collected in 

Chittenden County, Vermont, and from Lewis County, 

New York. These trees averaged nearly 49 feet in height 

at age 25 and were nearly 10 percent taller than the mean 

for all provenances (Table 2). Average height for the local 

seed source was about 34 feet and was only about 75 

percent of the mean total height for all provenances. 

These results suggest that significant improvements in 

height growth of sugar maple might be possible using 

superior trees from the correctly identified provenances. 

However, it is difficult to determine whether the relatively 

poor performance of the local provenance is due to 

meaningful genetic differences or to differences in 

experimental design previously addressed. 

Phenotypic Age-Age Correlations 

In this study, there were significant differences in provenance 

tree height at each measurement period (2, 6, 10, 15, and 

25 years after establishment; Table 3). The measurements 

at age 2 are significantly and positively correlated with the 

results from age 25 (p .: 0.01). A correlation matrix of 

provenance tree height reveals a highly significant but 

declining correlation as time lengthens from the first 

measurement period (Table 3). A matrix of Spearman 

correlation coefficients, which evaluates relative rank rather 

than absolute value, provides similar results. It appears 

that genetic differences in sugar maple are exhibited in 

seedling and sapling stages and persist for at least 25 

years, an age that is characterized by pole-size trees. 

Although remaining significant, the correlations have 

declined consistently with age and it cannot be determined 

if this relationship will remain significant for an additional 

50 to 75 years or until such time as end-of-rotation 

characteristics are realized. However, the objective is to 

identify only the superior provenances; the correlation of 

juvenile and mature characteristics of all of the provenances 

is not necessary. 

To further evaluate the potential for seedling and sapling 

demonstration of genetic superiority, the provenances 

were stratified into rankings based on the first height 

measurements taken 2 years after establishment. At that 

time, significant provenance differences accounted for 

51 percent of the total height variation, similar to the 

Table 3.-Correlation matrix of sugar maple mean 

provenance total height for 15 provenances included in 

a north-central West Virginia plantation established in 

1968 (all relationships significant at 0.01 level) 

provenance effect 25 years after establishment. The 

provenances were stratified into three groups based on 

the top 20 percent (n = 3), the bottom 20 percent, and the 

middle 60 percent (n = 9). Although subjective, these 

groups were used to characterize the best and worst 

performers after graphical analysis of the 2-year results. 

The relationship between these three groups has not 

changed over time (Fig. 1). The provenances from 

Chittenden County, Vermont, and Lewis County, New 

York, were ranked second and third tallest after 2 years 

and have remained in the top category at every 

remeasurement period. The provenance from Berkshire 

County, Massachusetts, was ranked first in total height 

after 2 years and is currently ranked fourth in total height. 

Further, after 10 growing seasons, the top three provenances 

have remained the same without exception. Thus, while 

the correlation of all the provenances continues to decline 

with respect to total height, the tallest provenances just 2 years 

after establishment generally remain the tallest 25 years 

after establishment. 

Conclusions 

Sugar maple, an important species of the eastern 

hardwood forest, exhibits genetic variation in total height, 

d.b.h., and survival. Much of this variation appears to be 

related to geographic origin. Such variation could be used 

to enhance desirable traits through breeding programs. 

For such programs to be most effective, the time between 

subsequent generations and selection should be minimized. The minimal duration of a breeding cycle must be 

at least as long as the time required for establishment of 

consistent genetic differences among seed sources and 

the onset of flowering for sexual reproduction. Sugar 

maple usually does not flower before 22 years of age 

(Olson and Gabriel 1974). This factor appears to be the 

most limiting for minimizing the breeding cycle interval as 

sustained genetic differentiation in height seems to occur 

at a much earlier age. While early selection of breeding 

traits is bound to be less than perfect, the gain per unit of 

time may greatly accelerate the breeding process. 
Age (years) 

Top 20% Mid 60% Bottom 20% 

-8- + 

Figure 1 .-Relationship between mean provenance total height and year of 

remeasurement. The classifications used are based solely on height measurements 

taken 2 years after establishment. 

Literature Cited 

Bresnan, Daniel F.; Geyer, Wayne A.; Lynch, Keith D.; 

Rink, George. 1992. Black walnut provenance 

performance in Kansas. Northern Journal of Applied 

Forestry. 9(2): 41-43. 

Freese, Frank. 1967. Elementary statistical methods for 

foresters. Agric. Handb. 317. Washington, DC: US. 

Department of Agriculture, Forest Service. 87 p. 

Godman, Richard M.; Yawney, Harry W.; Tubbs, Carl H. 

1990. In: Burns, Russell M.; Honkala, Barbara H., 

tech. coords. Silvics of North America: Volume 2. 

Hardwoods. Agric. Handb. 654. Washington, DC: US. 

Department of Agriculture, Forest Service: 78-91. 

Kriebel, H. B. 1975. Twenty-year survival and growth of 

sugar maple in Ohio seed source tests. Res. Circ. 

206. Wooster, OH: Ohio Agricultural Research and 

Development Center. 9 p. 

Olson, David F., Jr.; Gabriel, W. J. 1974. Acer L. Maple. 

In: Schopmeyer, C. S., tech. coord. Seeds of woody 

plants in the United States. Agric. Handb. 450. 

Washington, DC: U. S. Department of Agriculture, 

Forest Service: 1 87-1 94. 

Rink, George; Kung, Fan H. 1991. Height and diameter 

variation in twelve white ash provenancelprogeny 

tests in eastern United States. In: Proceedings, 8th 

central hardwood forest conference; 1991 March 3-6; 

University Park, PA. Gen. Tech. Rep. NE-148. Radnor, 

PA: U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station: 179-1 87. 

Wendel, G. W.; Gabriel, W. J. 1974. Sugar maple 

provenance study: West Virginia outplanting 6-year 

results. In: Garrett, Peter W., ed. 22nd northeastern 

forest tree improvement conference: proceedings of a 

symposium; 1974 August 7-9; Syracuse, NY. Syracuse, 

NY: State University of New York: 163-1 71. 

Wendel, G. W.; Gabriel, W. J. 1980. Sugar maple 

provenance study: West Virginia outplanting-10- 

year results. Res. Pap. NE-460. Broomall, PA: US. 

Department of Agriculture, Forest Service, Northeastern 

Forest Experiment Station. 5 p. 

fiU. S. GOVERNMENT PRINTING OFFICE 1994-550- 1 1 Z/8OOZ 1 
Schuler, Thomas M. 1994. Survival, growth, and juvenile-mature correlations in 

a West Virginia sugar maple provenance test 25 years after establishment. 

Res. Pap. NE-689. Radnor, PA: U.S. Department ofAgriculture, Forest Service, 

Northeastern Forest Experiment Station. 5 p. 

Survival, total height, diameter at breast height (d.b.h.), and stem quality of sugar 

maple trees of different provenances were compared 25 years after establishment in 

north-central West Virginia. Provenances were from Michigan, Minnesota, West 

Virginia, Massachusetts, New Hampshire, Vermont, Maine, and Quebec, Canada. 

There were significant differences between provenances for all traits except stem 

quality. By provenance, total tree height ranged from about 49 to 37 feet; d.b.h. 

from 6.7 to 3.6 inches; and survival from I00 to 15 percent. The predictability of 

total tree height 25 years after establishment based on mean provenance height at 

age 2,6, 10, and 15 years is discussed. Results suggest that juvenile height growth 

may be a good predictor of mature height performance, thus decreasing the need for 

rotation-length trials. 

Keywords: Acersaccharum, genetic variation, seed sources, phenotypic age-age 

correlations. 

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Headquarters of the Northeastern Forest Experiment station is in Radnor, 

Pennsylvania. Field laboratories are maintained at: 

Amherst, Massachusetts, in cooperation with the University of Massachusetts 

Burlington, Vermont, in cooperation with the University of Vermont 

Delaware, Ohio 

Durham, New Hampshire, in cooperation with the University of New Hampshire 

Hamden, Connecticut, in cooperation with Yale University 

Morgantown, West Virginia, in cooperation with West Virginia University 

Orono, Maine, in cooperation with the University of Maine 

Parsons, West Virginia 

Princeton, West Virginia 

Syracuse, New York, in cooperation with the State University of New York, 

College of Environmental Sciences and Forestry at Syracuse University 

University Park, Pennsylvania, in cooperation with The Pennsylvania State 

University 

Warren, Pennsylvania 

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