United States 

Department of 

Agriculture 

Forest Setvice 

Adaptation of Eastern 

White Pine Provenances 

Northeastern Forest 

Experiment Station to Planting Sites 

Research Paper NE-703 

Maurice E. Demeritt, Jr. 

Peter W. Garrett 
Abstract 

Eastern white pine provenances from the extreme limits of its natural range are 

changing from above- and below-average stability to average stability for height 

growth with increasing age. The regression method is useful for evaluating the 

stability of provenance to planting sites. The same general conclusions are reached 

for the performance of provenances at different planting sites from the regression 

method as from the relative difference method. 

--- 

The Authors 

MAURICE E. DEMERIT, JR. (deceased) was a research plant geneticist with the 

Northeastern Forest Experiment Station at Durham. New Hampshire, when this 

study was completed. Most recently he was a quality assurance specialist with the 

Northeastern Station's Forest Monitoring Program at Radnor, Pennsylvania. 

PETER W. GARRETT, research geneticist and Project Leader with the Northeastern 

Forest Experiment Station at Duham, New Hampshire, received a B.S. degree from 

Michigan State University and MS. and Ph.D. degrees from the University of 

Michigan. He joined the USDA Forest Service in 1966. 

Manuscript received for publication 6 November 1995 

Published by: For additional copies: 

USDA FOREST SERVICE USDA Forest Service 

5 RADNOR CORP CTR SUITE 200 Publications Distribution 

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Delaware. OH 43015 

October 1996 
Introduction 

Eastern white pine (Pinus strobus L.) is one of the most 

important timber trees in Northeastern United States and 

Eastern Canada. Provenance and/or genetic variation 

studies of this species have been underway since 1959 

(Garrett et al. 1973; Demerin and Kettlewood 1976. Funk et 

ai. 1975; Thor 1975; Kriebel et al. 1974). 

The ability to identify provenances that are adapted to a wide 

range of environments would be particulady useful to forest 

managers. One method of obtaining information on the 

adaptability of provenances is from the genotypeenvironment interaction from the analysis of variance of 

provenances over locations. A second method is a 

regression technique used by Finlay and Wilkinson (1963) to 

evaluate the phenotypic stability of 277 barley varieties. The 

regression technique consists of computing the linear 

regression of yield on the mean yield of all varieties for each 

site. The regression coefficient (b) was used as an indicator 

of stability. Average stability would be: b = 1 .O, perfect 

stability b = 0 and below average stability b > 1 .O. 

Phenotypic stability using the Finlay-Wilkinson technique 

was reported for jack pine (Pinus banksiana Lamb.) by 

Morgenstern and Teich (1969) and for white spruce (Picea 

glauca [Moench] Voss) by Nienstaedt (1969). 

We report on the stability or adaptability of 29 eastern white 

pine provenances in 12 plantations in the Northeastem 

United States using 10-year height and 16-year height and 

diameter with the regression technique. There are no 

previous reports on the adaptability of eastern white pine 

provenances to planting sites. 

Materials and Methods 

In 1955, the USDA Forest Service initiated a rangewide 

provenance trial of eastern white pine. Cones from 10 trees 

chosen at random in good stands of natural origin were 

cdlected by cooperators. Initially, 32 sources were included in 

the study, but two provenances (17 and 26) failed to produce 

seedlings in the nursery, and a third seed source (32) was not 

included in the northeastern plantings (Fig. 1, Table 1). 

Nursery trials of this material were conducted in New York, 

New Jersey, Maryland. North Carolina, and Wisconsin. 

Plantations were then established by the USDA Forest 

Service's Northeastem, Southeastern, Lake States, and 

Central States Forest Experiment Stations, and by the 

Southern Research Station of the Ontario Department of 

Lands and Forests. Seedlings within sources were bulked 

when lifted from the nursery beds and then selected at 

random for field trials.The Northeastern Station's plantings in 

1959 (plantings 7 to 11) were established with 2-0 stock 

grown in the Maryland State Forest Nursery. The 1960 

plantings (1 to 6) were established with 3-0 stock grown in 

the New Jersey and New York State Forest Nurseries. 

Planting locatiins are shown in Figure 2 andTable 2. Three 

Figure 1 .-Seed sources 

used in the provenance 

tests (shaded areas 

indicate the natural range 

of eastern white pine; 

numbers indicate seed 

collection points, Table 1). 
Table 1.--Seed sources for eastern white pine provenance study 

Union County,GA 

Transylvania County, NC 

Greene County, TN 

Garrett County, MD 

Greenbrier County, WV 

Monroe County, PA 

Monroe County, PA 

Clearfield County, PA 

Clearfield County, PA 

Ulster County, NY 

Ulster County, NY 

Franklin County, MA 

Worcester County, MA 

Penobscot County, ME 

Allamakee County, IA 

Ashland County, OH 

Forest County, WI 

Cass County, MN 

Lunenburg County, NS 

Sunbury County, NB 

Quebec County, PQ 

Pontiac County, PQ 

Norfolk County, ON 

Algoma District, ON 

Carroll County, NH 

Lake County, MN 

Houghton County, MI 

Palaski County, VA 

Sauk County. WI 

Source Location Latitude ~ongiude Elevation 

Feet 

Figure 2.-White pine provenance planting 

sites (Table 2). 
Table 2.-Location and design of plantings with mean plantation 10- and 16-year height and 16 year diameter 

Plantation Location Design Latitude Longitude Elevation Mean Mean Mean 

no. 10-year 16-year 16-year 

height height diameter 

Feet Feet Feet Inches 

1 Orono, ME ill Me53'N 68"39'W 100 4.1 15.7 2.70 

2 Alfred, ME I 43O32'N 70°40'W 300 6.8 16.8 4.21 

2 Alfred. ME II 4832'N 70°40'W 300 6.6 18.7 4.32 

2 Alfred, ME 111 43*32'N 70°40'W 300 7.4 19.5 4.54 

3 Essex Junction, VT Ill 44@28'N 73"09'W 327 6.5 17.3 3.67 

4 Paul Smiths, NY I MC26'N 74"13'W 1,815 5.3 14.3 3.04 

6 Warren, PA I 4I050'N 79"15'W 1,180 8.4 22.2 4.64 

7 Standing Stone, PA Ill 4O037'N 7a055'W 960 5.8 17.3 3.25 

8 Kennett Square, PA I 3g052'N 75"41'W 400 7.1 20.5 4.53 

9 Savage River State Ill 3gm40'N 79"15'W 2.740 9.0 16.1 2.98 

Forest, MD 

10 Horseshoe Run, WV I 39"1I7N 7g035'W 1.720 13.1 29.1 5.89 

11 Rison. MD lI 36"30'N 77"20'W 1M) 4.8 26.7 4.14 

planting designs, each a randomized complete-block, are 

included in this report (Table 2): 

Design I: Asingle tree from each provenance located 

randomly in each of 24 blocks. Spacing was 10 feet within 

rows and 10 feet between rows. 

Design II: Two-tree row plots of each provenance located 

randomly in each of 24 blocks. Provenance trees were planted 

at a spacing of 7 feet in rows and 14 feet between provenance 

rows. Additional seedlings were planted between provenance 

rows to achieve a final spacing of 7 x 7 feet. 

Design Ill: Four-tree row plots of each provenance located 

randomly in each of 12 blocks. Provenance trees were 

planted at the same spacing as those in Design II with 

addiiional seedlings between rows to again achieve a final 

spacing of 7 x 7 feet. 

The added (nonstudy) trees were removed from all Design II 

and Ill plantings 1 to 3 years before the 16-year 

measurements were taken. Total height was measured on 

each provenance tree after the 1966 growing season (10 

years from seed), and total height and diameter were 

recorded for each tree after the 1972 growing season (16 

years from seed), except for three plantings in Alfred, Maine. 

Those plantings were remeasured after the 1973 growing 

season (17 years from seed) and the data were wmected for 

the extra year's growih. Not all provenances were represented 

in evely planting; this accounts for the different number of 

plantings in the test for each provenance (Tables 3 -4). 

Simple linear regression and correlation coefficients for 10- 

and 16year height and 16year diameter were computed for 

each provenance across all plantings. Provenance means 

were the dependent variables (y) and planting means for all 

provenances at the planting were the independent variables 

(x). 

Simple linear correlation coefficients of provenances over 

planting locations were positive and high for 10- and 16-year 

diameters (Table 3). High correlation coefficients indicate 

that regression coefficients (b) for provenances over 

plantings are good measures of provenance stability with 

respect to the environments tested. The planting mean of the 

trait measured was used as an indication of the productivity 

of that environment. A relatively low planting mean would 

indicate a poor environment and a relatively high mean 

would indicate a good environment (Table 2). 

The average relative difference for each provenance was 

determined by the following formula: 

Average Relative Difference (%) = 

(for each provenance) 

sum provenance means at each plantation 

Sum plantation means. 

x 100 

This statistic was used to compare the performance of a 

provenance to all provenances across all planting sites (Table 

4). No confidence intervals were constructed for average 

relative differences in height and diameter measurements. 

Confidence intervals (95%) were constructed around each 

provenance's regression coefficients (Steel and Torrie 1960) 

to assign provenances to stability classes. 

Provenances were assigned to one of three stability classes 

for 10- and 16-year height and 16-year diameter based on 

their respective regression awfficients (Table 5). The three 

stability classes are: 

Above Average = b significantly less than 1. 

Average = b not significantly different from 1. 

Below Average = b significantly greater than 1. 

Stability classes were divided into two performance classes: 

provenances with above- and beiow-average relative 

differences when evaluated across all plantations (Table 5). 
e Table 3.-Llnear regression, standard error, and correlation coefficients of seed-source mean height at 10 and 16 years, and mean diameter 

at 16 years at each of the plantations on their respective mean heights or diameters of all seed sources at the plantations 

Seed 10-year height 16-year height 1Gyear diameter 

Location N b SE of reg. r b SE of reg. r b SE of reg. r 

source coef. coef. coef. 

1 Georgia 

2 North Carolina 

3 Tennessee 

4 Maryland 

5 West Virginia 

6 Pennsylvania 

7 Pennsylvania 

8 Pennsylvania 

9 Pennsylvania 

10 New York 

11 New York 

12 New York 

13 Massachusetts 

14 Maine 

15 Iowa 

16 Ohio 

18 Wisconsin 

19 Minnesota 

20 Nova Scotia 

21 New Brunswick 

22 Quebec 

23 Quebec 

23 Quebec 

25 Ontario 

27 New Hampshire 

28 Minnesota 

29 Michigan 

30 Virginia 

31 Wisconsin 
Table 4.-Average 10- and 16-year helght and 16-year diameter of each seed source as a 

percentage of all plantatlons 

Seed source Location N 10-year height 16-year height 16-year diameter 

Geomia 12 110 105 115 

~orthCarolina 

Tennessee 

Maryland 

West Virginia 

Pennsylvania 

Pennsylvania 

Pennsylvania 

Pennsylvania 

New York 

New York 

New York 

Massachusetts 

Maine 

Iowa 

Ohio 

Wisconsin 

Minnesota 

Nova Scotia 

New Brunswick 

Quebec 

Quebec 

Ontario 

Ontario 

New Hampshire 

Minnesota 

Michigan 

Virginia 

Wisconsin 

Table 5.-Assignment of seed sources by source number (Table 1) to stablllty and performance classes for 10 and 

16-year helgM and 16-year dlameter 

Stability Class Performance Class 10-year height 16-year height 16-year diameter 

Specifically adapted to 1,2,3,6,9, 1,5,9 2, 9, 13 

Below average favorable environments 13.24 

(source above average) 

Specifically adapted to 5 none 5 

Below average favorable environments 

(source below average) 

Well adapted to all 4,7,10,11, 3,4,6,7,8,10,11, 1.3,6, 7,8,10, 

Average environments in the tests 12, 16,27,31 12,13,20,24,27,31 11,12,16,24,27,31 

Poorly adapted to all 8. 18, 19,20, 2, 15, 18, 19.21, 4, 15, 18, 19, 20, 

Average environments in the tests 25, 30 22,23,25,28,29,30 23,25,26,29,30 

Specifically adapted to none 16 none 

Above average poor environments 

(source above average) 

Specifically adapted to 14, 15.21,22, 14 14,21,22 

Above average poor environments 23,28.29 

(source below average) 
Results and Discussion 

In the Northeastern United States, eastern white pine 

provenances are becoming more stable for height growth with 

increasing age. At 10 years of age, of the 29 provenances 

tested, 8 had below-average stability and 7 had aboveaverage stability. At 16 years, only three of the initial eight 

provenances with below-average stability at 10 years were still 

below average, and only one of the seven with above-average 

stability still was above average. One provenance (Ohio 16) 

was stable at 10 years and above average stability at 16 

years. The assignment of provenances 7 and 31 to the 

average-stability class should be interpreted with caution. 

Provenance 7 was planted only in northern plantings (less 

favorable environment) and provenance 31 only in southem 

plantings (more favorable environment). 

The values for provenance stability may have been 

influenced by how far from their origin the provenance was 

planted. Central provenances were moved only a maximum 

of 4" latitude north or south, while southern provenances 

were moved a maximum of 9" latitude north and northern 

provenances a maximum of 10" latitude south. 

When 10-year height measurements are used, there is a 

tendency for southern Appalachian provenances to have 

below-average stability. This also is true for some 

provenances from the central part of the eastern white pine 

range. Below-average stability implies that the provenance 

at 16 years. Using the same data and the relalive difference 

method. Demerin an0 Kettlewood (1976) found that 

provenances were becoming more similar in growth from 

age 10 to age 16. Also, Funk et al. (1975) found that in the 

Central United States, southern Appalachian sources were 

losing their superiority by age 15. 

When diameter is used as the measurement, provenance 

stability at age 16 is nearly identical to that using 16-year 

height growth. The lone difference with the 16-year height 

data'is that Provenance 13 (Massachusetts) has belowaverage rather than average stability, and Provenances 21 

(New Brunswick) and 22 (Quebec) have above-average 

rather than average stability. Demeritt and Kettlewood (1976) 

generally found the same good correlation between 16-year 

height and diameter data for eastern white pine provenances. 

The use of performance classes within stability classes only 

serve as indicators of the performance of provenances in 

their stability class. However, only three provenances (5, 

West Virginia). (8, Pennsylvania), (20, Nova Scotia) changed 

from below- to above-average performance between age 10 

and 16 for height growth. Only provenance 2 (North 

Carolina) changed from above- to below-average for height 

growth during the same period. In no case did any change in 

average relative difference in height growth exceed 5 

percent (Table 4). Overall, there is good agreement for 

assigning provenances to stability-adaptation classes for 10- 

and 16-year height and lgyear diameter. Three were some 

willgrow well in go& environments. Garrett et al. (1973) proven&e shifis when stability and performance classes 

found that 10-year-old trees from southem Appalachia, and were evaluated simultaneously. 

some sources from as far north as central Pennsylvania, 

were preferred for southem plantings. Funk (1971) reported 

that 10-year-old trees from the southern Appalachian 

sources were superior in height growth in midwestern 

plantings. Provenance 24, a source from southern Ontario, 

has below-average stability and thus might grow well in good 

environments. This may be the result of modifying effects of 

Lake Erie on its evolution. 

There is a tendency for northern provenances to have 

above-average stability for 10-year height growth. This 

implies that these provenances will be specifically adapted 

(grow best) in less favorable environments (northern 

locations). Some provenances from the central portion of the 

species natural range also fall into this category. On the 

basis of 10-year height growth, Garrett et ai. (1973) found 

that these sources were preferred for northern plantings. 

About half of the provenances were assigned to the 

average-stability class and thus were adapted (well or 

poorly) to all environments based on 10-year data. Because 

most of these were from the central portion of the natural 

range of eastern white pine, they were moved a relatively 

short distance to any planting site in this test. 

Using height data at 16 years, no provenance could be 

associated with stability class with any degree of reliability. 

Literature Cited 

Demeritt, M.E.; Kettlewood, H.C. 1976. Eastern white pine 

seed source variation in northeastern Unlted States: 

16-year results. Gen. Tech. Rep. NC-26. St. Paul, MN: 

U.S. Department of Agriculture, Forest Se~ice, North 

Central Forest Experiment Station: 80-87. 

Finlay, K.W.; Wilkinson, G.H. 1963. The analysls of 

adaptatlon in a plant-breeding programme. Aust. J 

Agric. Res. 14: 742-754. 

Funk. D.T. 1971. Eastern white pine seed source trials: 

Ten-year results from three midweetern plantations. 

Res. Note NC-113. St. Paul, MN: U.S. Department of 

Agriculture. Forest Sewice, North Central Forest 

Experiment Station. 4 p. 

Funk, D.T.;Allen, R; Williams, R.D. 1975. Fifteen-year 

performance of eastern white pine seed source tests 

In the Lower Ohlo Valley. Proceedings of the Central 

States Forest Tree lmprovment Conference. 9: 153-158. 

Garrett. P.W.; Schreiner, E.J; Kettlewood, H.C. 1973. 

Geographic varlation of eastern white pine in the 

Northeast. Res. Pap. NE-274. Upper Darby, PA: U.S. 

Provenance 16 (Ohio) was the only sourw that changed Department of Agriculture, Forest Service, Northeastern 

from average stability at 10 years to above-average stability Forest Experiment Station. 14 p. 
Kriebel, H.B.; Roberds, J.H.; Cox, R.V. 1974. Genetic 

variation in vigor in a whlte pine Incomplete diallel 

cross experiment at age 6. Proceedings of the Central 

States Forest Tree lmprovment Conference. 8: 40-42. 

Morgenstem, EK; Teich, kH. 1969. Phenotypic stability 

of height growth of jack plne provenances. Canadian 

Joumal of Genetics and Cytology. 11: 110-117. 

Nienstaedt, H. 1969. White spruce seed source variation 

and adaptatlon to 14 planting sites In northeastern 

United States and Canada. Proceedings of the 

Commission on Forest Tree Breeding in Canada. 11: 183- 

Steel, R.G.D.; Torrie. J.H. 1960. Principles and 

procedures of statistics. New York McGraw-Hill Book 

Co. 481 p. 

Thor, E. 1975. White pine from the southern 

Appalachlans. Proceedings of the Central States Forest 

Tree Improvement Conference. 9: 145-152. 
Demeritt, Maurice E.. Jr.; Garrett, Peter W. 1996. Adaptatlon of eastern white 

pine provenances to planting sites. Res. Pap. NE-703. Radnor, PA: US. 

Department of Agriculture, Forest Sewice, Northeastern Forest Experiment 

Station. 7 p. 

Eastern white pine provenances from the extreme limits of the natural range of this 

species are changing from above- and below-average stability to average stability 

for height growth with increasing age. The regression method is useful for 

evaluating the stability of provenance to planting sites. The same general 

conclusions are reached for the performance at different planting sites from the 

regression method as from the relative difference method. 

Keywords: Pinus strobus, provenances, adaptation 

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College of Environmental Sciences and Forestry at Syracuse Unlversity 

Warren, Pennsylvania 

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