United States 

Department of 

Agriculture 

Forest Service 

Northeastern Forest 

Experiment Station 

Research Paper NE-684 

Survival and Growth of White 

Ash Families and Provenances 

15 Years After Establishment 

in West Virginia 

Thomas M. Schuler 
Abstract 

The survival, growth, and stem form of 45 white ash (Fraxinu americana L.) 

families nested within 22 provenances were evaluated 1 5 years after establishment 

in North Central West Virginia. Geographic family origins encompassed a wide area 

in the eastern and central United States, including locations from Maine in the North 

to Mississippi in the South to Nebraska in the West. Significant differences were 

identified among provenances for survival, stem form, total height, and stem 

diameter and among families within provenances for stem form and total height. 

Latitude was a significant indicator of provenance performance with performance 

declining north and south of the plantation latitude. Performance gains may be 

realized by using proper provenances then identifying superior families within these 

provenances for white ash artificial regeneration activities. 

The Author 

THOMAS M. SCHULER, research forester, received a B.S. 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. 

COVER PHOTO.--A white ash tree from Jackson County, Illinois, included in a 

rangewide provenance and progeny evaluation in West Virginia 15 years after 

establishment. 

Manuscript received for publication 13 September 1993 

USDA FOREST SERVICE 

5 RADNOR CORP CTR STE 200 

PO BOX 6775 

RADNOR PA 19087-8775 

January 1994 
Introduction 

White ash (Fraxinus americana L.) is a species native to a 

wide geographic area in the eastern and central United 

States and Canada. This range extends southward from 

Nova Scotia to Florida in the East and from Minnesota to 

Texas in the West. White ash is known to be genetically 

variable (Wright 1944). To capitalize on the genetic variability 

found in any species, estimates of various levels of genetic 

variation must be developed. Thus, a rangewide white ash 

provenance and progeny test was initiated in 1976 (Clausen 

1984). The white ash tree improvement program included the 

establishment of 22 plantations throughout the eastern 

United States and Canada. This paper reports on the 15-year 

results of the West Virginia plantation that was included in the 

rangewide evaluation. The West Virginia plantation includes 

45 families nested within 22 provenances. Provenance and 

family survival, stem form, total height, and stem diameter at 

breast height (d.b.h.) are described and are related to the 

geographic origin of the planting stock. White ash seed 

collection guidelines are suggested for artificial regeneration 

ventures in the central Appalachian region. 

Methods 

The West Virginia white ash outplanting was established on 

the Fernow Experimental Forest (39.03~ N, 79.67' W), 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 48OF., and 

the length of the frost-free season is approximately 145 days. 

The elevation of the outplanting is 2,700 feet. The soil is a 

Meckesville silt loam (fine-silty, mixed, mesic Typic 

Fragiudults), which is colluvial in origin and has some 

limestone influence. Meckesville soil is well drained, 3 to 4 

feet deep, and is one of the more productive forest soils in 

the region. Northern red oak site index for the plantation is 

between 80 and 85. 

Before plantation establishment, the site was forested with a 

65- to 70-year-old stand of mixed Appalachian hardwoods 

that included a high proportion of black cherry (Prunus 

serotina Ehrh.), yellow-poplar (Liriodendron tulipifera L.), 

northern red oak (Quercus rubra L.), sugar maple (Acer 

saccharum Marsh.), and white ash. In preparation for 

planting, the sawtimber and pulpwood were harvested and all 

remaining stems were felled. The brush and small stems 

were removed from the site to facilitate the planting. The area 

has a northwest aspect with a slope not exceeding 10 

percent. 

The outplanting design is a randomized complete block of 22 

provenances with 45 families laid out in 5 contiguous blocks. 

Trees were planted 10 feet apart, in rows 10 feet apart. Each 

of the 45 families (Table 1) was represented in each of the 

five blocks by a five-tree linear plot. A border row of 

yellow-poplar seedlings was planted around the perimeter of 

the area. The sequence of the family plots on each block was 

randomized. Nursery 1-0 bare-root seedlings from each of 

the 45 families were planted on April 2, 1976. The 

measurements also include growth that occurred during the 

first year in the nursery. Thus, the individual trees have had 

16 growing seasons. Postplanting site treatment has included 

several herbicide applications and weedings that were 

necessary to control competing herbaceous and woody 

vegetation. 

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

the form of the main stem were recorded periodically, most 

recently, in January 1991, 15 years after establishment. Data 

were analyzed by analysis of variance using linear model 

procedures incorporating the randomized block design with 

families nested within provenances. Plot means of the 

surviving trees were used for tree height, d.b.h., and stem 

form analyses (Bresnan and others 1992, Freese 1967, Rink 

and Kung 1991) and were weighted by the number of trees 

they represented to account for unequal variances. 

Differences in survival and stem form were tested for 

following arcsin transformation. 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. 

The 7-year results, as reported by Wendel (1985), were used 

for contrast where appropriate. 

An overall provenance ranking or index was developed using 

the performance criteria where significant differences were 

identified. Multiple-trait selection is not uncommon and has 

been used by others to evaluate two or more traits 

simultaneousiy (Sprague 1966; Bey and others 1974; 

Schuler 1993). To develop the index, the provenances were 

ranked in each category based on overall provenance means 

and the rankings were summed for each provenance. The 

rankings provide a preliminary evaluation of suitability for 

each of the provenances as to their fitness for the growing 

conditions similar to those found on the Fernow Experimental 

Forest. 

Table 1 summarizes the identification information of each 

family, including provenance designation. The families 

originated from 15 states, covering much of the botanical 

range of white ash. 

Results and Discussion 

At the end of 15 years, analysis of variance revealed 

significant differences among mean provenance survival, 

stem form, total height, and d.b.h. Families within 

provenances also differed significantly with respect to stem 

form and total height. 

Survival 

Average provenance survival 16 years after plantation 

establishment was 50 percent (Table 2). Survival after 7 

years was 61 percent (Wendel 1985). Provenance survival 

ranged from 88 percent to 0 percent and accounted for 52 

percent of the variation in survival. Families within 

provenances did not differ significantly and accounted for 

only 4 percent of the total variation. The provenances from 

Mississippi, Louisiana, and Alabama had a mean survival 

rate of approximately 9 percent. In contrast, the two 
Table l .--White ash families and provenances planted in West Virginia 

Family Provenance Latitude Longitude 

number number State County (N) (w) Elevation 

Nemaha 

Williamson 

Jackson 

Jackson 

Forest 

Pike 

Franklin 

Franklin 

Wayne 

Madison 

Madison 

Trigg 

Trigg 

Boone 

Marion 

Marion 

George 

East Baton Rouge 

East Baton 'Rouge 

East Baton Rouge 

Oktibbeha 

Oktibbeha 

Oktibbeha 

Oktibbeha 

Effingham 

Shelby 

Randolph 

Randolph 

Tucker 

Penobscot 

Penobscot 

Cortland 

New Haven 

Washington 

Barbour 

Harrison 

Taylor 

McMinn 

McMinn 

Overton 

Overton 

Overton 

Marshall 

Marshall 

687306 22 TN Marshall 35.4 87.7 800 
Table 2.--White ash mean provenance survival, crop trees, total height, d.b.h., and performance index of 22 

provenances at the end of 15 growing seasons in West Virginia 

Crop Total Performznce 

Provenance Center of Geographic Origina Survival trees Height D.b.h. Index 

Southern IN (Salem) 

North Central WV (Bridgeport) 

North Central WV (Harman) 

Central NY (Cortland) 

Southern IL (Marion) 

Central OH (Wooster) 

West Central IL (Rockport) 

East Central lL (Shelbyville) 

Central CN (New Haven) 

Northeast TN (Livingston) 

Southeast TN (Athens) 

Southwest KY (Golden Pond) 

South Central TN (Lewisburg) 

North Central AR (Harrison) 

Central MS (Starkville) 

Northern ME (Millinochet) 

Southern TN (Winchester) 

Southeast LA (Baton Rouge) 

Southeast NE (Auburn) 

Northeast WI (Crandon) 

Northern AL (Meridianville) 

Southern MS (Lucedale) 

Provenance Mean 50 12 13.4 1.7 

aCenter of geographic origin is represented by a regional description of location within a state and a centrally located 

community within that region. 

provenances from West Virginia had a mean survival rate of 

approximately 79 percent. These results are consistent with a 

region-wide study that found local white ash provenances 

usually performed either at average or above average levels 

for the plantation (Clausen 1984). 

Stem Form 

Overall, plantation stem form was quite poor, although 

provenances and families within provenances were 

significant predictors of stem form. Provenances accounted 

for 18 percent of the total variation in stem quality while 

families nested within provenances accounted for 24 percent 

of the variation. Twelve families did not have any potential 

crop trees. A potential crop tree was defined as a tree that 

could develop a grade 1 or 2 butt log (Lamson and Smith 

1989). Trees with poor stem form characteristics such as a 

low fork, epicormic branches, or a broken top were 

considered unsuitable as potential crop trees. The families 

with the best averages were from Harrison and Randolph 

Counties in West Virginia. However, less than 50 percent of 

the trees from these two families were potential crop trees 

(Table 3). 

Poor stem form characteristics within the plantation may be 

related to the number of provenances and families that 

probably are unsuitable for the growing conditions of the 

Fernow. Poor performance of these families has resulted in a 

longer than usual delay of crown closure. Typically, crown 

closure would occur in 7 to 8 years following overstory 

removal. However, the 1 5-year-old ash plantation has yet to 

reach this stage of development. This is due partially to the 

continued weedings necessary to maintain growing space for 

the slower growing individuals in the plantation. Yet, in the 

absence of crown closure many of the larger trees are 

developing forms more typical of open-grown trees. Lack of 

crown closure also has resulted in the persistence of shrubs. 

The most common shrub within the plantation is blackberry 

(Rubus sp.). This, too, has intensified stem form problems by 

attracting bears that damage plantation trees while feeding 

on the blackberries. Common problems associated with bear 

damage include broken tops and limbs and bark wounds. 
Table 3.--Average crop trees (percent) and total height of 

45 white ash families at the end of 15 growing seasons In 

West Virginia 

Family Family origin Total Crop 

code State County height trees 

Harrison 

Randolph 

Randolph 

Cortland 

Williamson 

Trigg 

Washington 

Taylor 

Barbour 

Jackson 

Shelby 

Marion 

Pike 

Tucker 

Jackson 

Wayne 

Overton 

Overton 

New Haven 

McMinn 

Effingham 

Marshall 

Marshall 

Penobscot 

Nemaha 

Boone 

Madison 

Penobscot 

Marshall 

Overton 

Franklin 

McMinn 

Oktibbeha 

Forest 

Oktibbeha 

Marion 

Franklin 

East Baton Rouge 

Trigg 

Oktibbeha 

East Baton Rouge 

Madison 

Oktibbeha 

East Baton Rouge 

George - 

Family Mean 12.1 11 

Tree Height 

Average tree height differed significantly for both 

provenances and families nested within provenances. 

Provenances accounted for 27 percent of the total variation 

in tree height while families within provenances accounted for 

18 percent of the variation. Mean provenance total height 

was 13.4 feet. The provenance with the best height growth of 

19.8 feet was a local provenance from north central West 

Virginia and was 70 percent greater than the mean for ail 

provenances (Fig. 1). These results clearly illustrate that the 

selection of the correct provenance is critical for height 

growth of white ash in West Virginia. Additional 

improvements in height growth may be possible through 

identification and use of superior families and gains may be 

proportionately larger with increasing age. 

D.B.H. 

Mean d.b.h. for all provenances 15 years after establishment 

was 1.7 inches with the largest diameter of 3.2 inches 

re-corded for the north central West Virginia provenance 

(Table 2). Provenances accounted for 23 percent of the total 

variation in d.b.h. and weresignificant, while families within 

provenances accounted for 11 percent of the variation but 

were not significant--illustrating again, the importance of 

proper provenance selection of white ash in West Virginia. As 

expected, the results for mean provenance d.b.h. were highly 

correlated with the results for mean provenance total height 

(Spearman correlation coefficient r = 0.94). 

Performance Index 

An overall provenance performance index was developed 

based on survival rate, stem form, total height, and d.b.h. 

Including total height and d.b.h, in the index, essentially 

weights the size as 50 percent, survival as 25 percent, and 

stem form as 25 percent of the total. In some cases, more 

than one provenance occupies a particular rank (Table 2). 

Although the basis for the index is subjective, some general 

trends are suggested. Perhaps most evident is that the best 

performing provenances have geographic origins from about 

the same latitude as the test site. The top three provenances 

include two provenances from West Virginia and one from 

Indiana. All are from virtually the same latitude as the test 

site. Two families within these provenances also had the best 

stem form characteristics (Table 3) and were local in 

geographic origin. However, the high ranking of the Indiana 

provenance suggests that longitude may be less critical than 

latitude as a potential performance criteria and that superior 

families may be found in locations not local to the planting 

site or even part of the central Appalachian mountain region. 

Based on the results from the performance index, stepwise 

multiple regression was used to evaluate provenance 

performance as related to latitude, longitude, and elevation of 

the provenance. Latitude-squared and longitude-squared 

terms were included as potential independent variables. 

Latitude and latitude-squared were significantly related to 

survival and total height and accounted for 69 and 68 percent 
Figure 1 .--Range of tree sizes present in white ash plantation in West Virgina 15 years after establishment. 

Tree height differed significantly for both provenances and families within provenances and latitude was a 

significant predictor of provenance performance. 
of the variability for survival and total height, respectively, 

when used as the independent variables in quadratic 

equations (Table 4). Stem form was not related to any of the 

independent variables and d.b.h. was more closely related to 

elevation. The distribution of the survival and total height data 

suggest an optimal latitude for each criterion (Fig. 2), with 

performance declining north and south of the optimum. The 

quadratic equations predict optimal latitudes of 41 .0' N for 

survival and 39.5' N for total height. Higher survival rates of 

slightly more northern provenances may be related to the 

timing of leaf abscission. Clausen (1 982) found dieback of 

the main stem was closely related to the latitude of the seed 

origin and the presence of trees still growing at the time of 

the first killing frost in an Ontario white ash plantation. Thus, 

performance gains through longer leaf retention may be 

achieved at the risk of frost injury. On the Fernow, it appears 

that such risks are too great to realize performance gains by 

using southern seed sources. 

These findings clearly illustrate that white ash seed collection 

zones for mountainous locations in West Virginia should be 

restricted to a narrow latitudinal range. Longitudinal seed 

collection zones appear to be less critical and may provide 

opportunities for identification of superior families and 

realization of performance gains by using superior seedlings 

in artificial regeneration activities. 

Improper selection or use of poorly suited sources of white 

ash in artificial regeneration should be carefully avoided. As 

seen here, the consequences could result in a complete 

regeneration failure of the desired species. The risk of using 

unsuitable sources is increasing in conjunction with the 

increased use of tree shelters and nursery grown planting 

material. As this practice becomes more common, both the 

user and the nursery manager need to be well informed 

about the origin of their sources and develop guidelines for 

localized use. 

Table 4.--Equations describing white ash provenance 

survival rate (S) and total height (H) (feet) as a function of 

latitude and latitude-squared (LS). 

Equation R~ SEa 

S = -1 0.8542 + 0.56293(L) - O.O0687(LS) 0.69 0.15 

H = -246.715 + 13.3245(L) - 0.16867(LS) 0.68 2.91 

a~tandard error of estimate. 

Conclusions 

Preliminary recommendations for planting of white ash in 

West Virginia suggested it was both safe and advantageous 

to plant trees originating 100 to 200 miles south and west of 

the planting site (Clausen 1984). Wendel (1985), reporting on 

the 7-year results of the Fernow white ash plantation, refined 

this recommendation by finding that only families from the 

same general latitude (38'to 40' N) as the test site 

performed well. This translates to a zone of about 70 miles 

north and south of the planting site. However, Rink and Kung 

(1991) found that no significant improvements in height 

growth were observable in West Virginia, although rangewide 

studies of white ash have shown that height growth 

improvements of 25 to 50 percent are possible through 

proper provenance selection. 

The results of this study suggest that white .ash performance 

gains in West Virginia are possible by using proper 

provenances and superior families within these provenances, 

but the provenances and families probably will originate 

within a narrow latitudinal range. Equations developed from 

the provenance survival and total height observations (Table 

4) predict an optimal provenance latitude of 2 degrees north 

of the plantation for survival and virtually the same latitude as 

the outplanting for maximizing height growth. 

Latitude 

Latitude 

Figure 2.--White ash mean provenance survival and total 

height as a function of provenance latitude. Vertical line 

represents latitude of outplanting in West Virginia. 
Literature Cited 

Bey, C. F.; Kung, J. H.; Kucera, J. M. 1974. Multiple-trait 

selection in black walnut (Juglans nigra L.). In: 

Proceedings, Joint AFFRAY Meeting, S.02.04.1-3, Session 

11 1974; Stockholm, Sweden. Stockholm, Sweden: 

Department of Forest Genetics, Royal College of Forestry: 

1 17-1 28. 

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

Rink, George. 1992. Black walnut provenance 

performance in Kansas. Northern Journal of Applied 

Forestry. 9(2): 41 -43. 

Clausen, Knud E. 1982. Variation in frost injury to white 

ash families in an Ontario plantation. Canadian Journal 

of Forest Research. 12(2): 440-443. 

Clausen, Knud E. 1984. Nonlinear regressions and 

contour plots: techniques for selection and transfer of 

white ash provenances. Forest Science. 30: 441 -453. 

Freese, Frank. 1967. Elementary Statistical Methods for 

Foresters. Agric. Handb. 317. Washington, DC: U.S. 

Department of Agriculture, Forest Service. 87 p. 

Lamson, Neil I.; Smith, H. Clay. 1989. Crop-tree release 

increases growth of 12-year-old yellow-poplar and 

black cherry. Res. Pap. NE-622. Broomall, PA: U.S. 

Department of Agriculture, Forest Service, Northeastern 

Forest Experiment Station. 7 p. 

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. 

Schuler, Thomas M. 1993. Survival, growth, and target 

canker infection of black walnut families 15 years after 

establishment in West Virginia. Res. Pap. NE-674. 

Radnor, PA: US. Department of Agriculture, Forest 

Service, Northeastern Forest Experiment Station. 8 p. 

Sprague, G. F. 1966. Quantitative genetics in plant 

improvement. In: Frey, Kenneth J., ed. Plant Breeding. 

Ames, IA: Iowa State University Press: 31 5-354. 

Wendel, G. W. 1985. Performance of white ash progenies 

after 7 years in a West Virginia planting. In: 

Proceedings, 29th Northeastern Forest Tree Improvement 

Conference; 1984 July 18-20; Morgantown, WV. 

Morgantown, WV: West Virginia University: 90-97. 

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Printed an Recycled Paper 

*U. S. GOVERNMENT PRINTING OFFICE: 1994-550- 1 12/80009 
Schuler, Thomas M. 1994. Survival and growth of white ash families and 

provenances 15 years after establishment in West Virginia. Res. Pap. 

NE-684. Radnor, PA: U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station. 7 p. 

The survival, growth, and stem form of 45 white ash (Fraxinus americana L.) 

families nested within 22 provenances were evaluated 15 years after 

establishment in north central West Virginia. Geographic family origins 

encompassed a wide area in the eastern and central United States, including 

locations from Maine in the North to Mississippi in the South to Nebraska in 

the West. Significant differences were identified among provenances for survival, stem form, total height, and stem diameter and among families within 

provenances for stem form and total height. Latitude was a significant 

indicator of provenance performance with performance declining north and 

south of the plantation latitude. Performance gains may be realized by using 

proper provenances and then identifying superior families within these 

provenances for white ash artificial regeneration activities in West Virginia. 

ODC 232.12:176.1 

Keywords: Fraxinus americana, progenies, performance gains, seed collection zones 
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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