Báo cáo khoa học: "Intraspecific variation of growth and adaptive traits in North American oak specie"

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Review article Intraspecific variation of growth and adaptive traits in North American oak species HB Kriebel School of Natural Resources, Division of Forestry, The Ohio State University, Wooster, OH 44691-4096, USA Summary — Variation of growth and adaptive traits has been identified in Quercus rubra L, which has recognizable geographic patterns in replicated test plantations in the central and western parts of the species range. Traits varying geographically include growth rate, drought resistance, cold re- sistance, time of flushing and leaf senescence. Patterns may differ in tests in southern regions. In Quercus falcata Michx, coastal plain sources are superior to others in both central Mississippi and western South Carolina. In 3 partial-range Quercus pagoda Raf tests, local or regional sources out- rank others in growth and adaptability. Both of these species vary widely in cold hardiness. Local trees of Quercus alba L are above the average height of all Indiana trees at age 5 yr in southern In- diana, but local trees of Quercus macrocarpa Michx in Nebraska are not as fast-growing as trees from seed sources 160 km south. Range-wide patterns remain undefined in both of these species. Among western provenances of Quercus nigra L in Louisiana, flushing is latest in trees of the north- ernmost origins. Only fragmentary information is available on variation of growth and adaptive traits in 7 other oaks, all eastern North American species. Quercus / oaks / variation / growth / adaptive traits / hardiness Résumé — Variabilité intraspécifique des caractères d’adaptation et de croissance chez les espèces d’Amérique du Nord. La variabilité des caractères de croissance et d’adaptation a été étu- diée chez Quercus rubra L; des gradients de variation ont clairement pu être établis chez cette es- pèce au vu des résultats obtenus dans des plantations installées dans la partie centrale et occiden- tale de l’aire naturelle. Les caractères, dont la variabilité suit un gradient géographique, sont : le taux de croissance, la résistance à la sécheresse et au froid, la date de débourrement et la sénescence des feuilles. Ces gradients peuvent être différents dans les plantations installées dans la partie méri- dionale de l’aire. En ce qui concerne Q falcata Michx, les origines des plaines côtières sont supé- rieures aux autres dans la partie centrale du Mississippi, et la partie occidentale de la Caroline du Sud. Dans 3 plantations de Q pagoda Raf ne comprenant qu’un échantillon partiel de provenances, les populations locales étaient nettement supérieures aux autres pour la croissance et les carac- tères d’adaptation. Les origines locales de Q alba L ont une meilleure croissance que les autres dans le sud de l’Indiana (à 5 ans); alors que chez Q macrocarpa Michx dans le Nebraska, les ori- gines locales sont moins vigoureuses que celles originaires de 160 km au sud. Les gradients de va- riation au niveau de l’ensemble de l’aire naturelle n’ont pas encore été étudiés pour ces 2 espèces.
En Louisiane, chez Q nigra L, le débourrement est plus tardif chez les provenances les plus nordi- ques. Des données fragmentaires sur la variabilité des caractères de croissance et d’adaptation exis- tent pour 7 autres espèces, toutes issues de l’est des États-Unis. Quercus / chênes / variabilité / croissance / adaptation / résistance INTRODUCTION expected to vary with seed in source ex- perimental plantations. In uniform-environment provenance North America has about 58 species of tests of a geographically variable species, oaks (genus Quercus) of tree size, of extensive provenance sampling covering which about 20 are considered important the entire distribution strengthens the prov- in forest management (Fowells, 1965). enance component of variance in relation Many of the North American oaks are dis- to stand and family components, whereas tributed over a wide range of latitude and range restriction leads to proportionately longitude and over several of the plant har- larger regional and local components (Krie- diness zones used as guidelines in horti- bel, 1965). In several species of Quercus, culture (fig 1, table I). Some are extremely mid-range or confined-latitude sampling in- wide-ranging. Q macrocarpa Michx, one of dicated that, within the region studied, the most widely-distributed species, oc- stand variability was more important than curs from 28-53 °N latitude and 66-105 geographic variability, and geographic pat- °W longitude. Therefore, adaptive traits, terns were not observed (Kriebel, 1965; and perhaps growth rate as well, could be Houston, 1987; Schnabel and Hamrick, 1990). However, this paper demonstrates that results are very different, at least in Q rubra, when samples are more widely dis- persed. Most of the information currently availa- ble on intraspecific variation in the North American oaks is based on population samples covering only parts of the spe- cies distribution. Far more information is available on Q rubra than on any other species. In addition, there have been sev- eral provenance experiments on Q falcata and the closely-related Q pagoda. Report- ed results from research on Q alba and Q nigra are not range-wide and are limit- ed to juvenile material. Some information is available on growth and adaptatibility of Q macrocarpa from one provenance test at age 11 years. Apart from these 6 spe- cies, there is a little information in the liter-
ature on variation of growth and adaptive Variation in growth rate traits in North American oaks. Brief dis- cussions on 7 other species are included Northern red oak (Quercus rubra L) varies in this review. The information is taken with geographic origin in rate of height and from: 1) published research; and 2) un- diameter growth. The geographic pattern published data and reports obtained by was evident in 23-year-old trees in 4 the author. With the exception of experi- range-wide tests in middle latitudes of the mental analysis of one commercially im- species range from eastern Nebraska to portant adaptive trait in Q palustris, the in- northern Ohio (Kriebel et al, 1988), but not formation on these other oaks is based at age 14 years in the same tests (Kriebel on fragmentary data from limited popula- et al, 1976). There was no statistical evi- tion sampling. dence of a pattern in results from limited- area sampling (Kriebel, 1965; Farmer et al, 1981; Houston, 1987; La Farge and Lewis, NORTHERN RED OAK 1987). pattern is follows: The variation as About 25 provenance tests of northern red height growth means are almost always oak (Quercus rubra L) of varying size have highest in trees from provenances be- been established in North America, but tween latitudes 43 and 46°N in an east- some no longer exist and others have not west zone extending from the Mississippi been evaluated. Some are comprehen- River to western Maine. Trees from out- sive, multi-family experiments that are side of this the average, zone are, on range-wide and replicated in several loca- slower-growing. In Ohio, Indiana and tions, while others include only a few pop- Michigan experiments, all but one of the ulation samples or are regional in their provenance samples that exceeded the sampling pattern. The first Q rubra prove- mean annual increment of its age class by nance tests, which were established by of than 1 standard deviation more was Scott Pauley in Massachusetts in 1951 Wisconsin, Michigan, Ontario, New York and 1952, were the most geographically or Maine origin (Kriebel et al, 1988). comprehensive tests of this species in There were indications of a similar pattern North America. They included 80 seed in test of the same material in eastern a sources that sampled most of the natural where the fastest-growing Nebraska, distribution. Unfortunately, the plantations trees were from Wisconsin and extreme were not maintained and the only pub- eastern Minnesota (Schlarbaum and Ba- lished report is a study of cold-hardiness. gley, 1981).These patterns are summar- Nine replicated range-wide tests were ized in table II. planted in the North Central states be- From these evaluations up to age 23 tween 1960 and 1962. Results from 7 of years, we can conclude that at latitudes these have been published. The other in- 40-42°N in the USA significant gains in tensive study was of more than 200 fami- growth of northern red oak can be lies from Tennessee and adjacent areas; achieved by planting trees from seed ori- of 10 outplantings, results from 3 are sum- gins 250-550 km north of the planting lo- marized. Additional information was availa- cality. In addition, since growth varies with ble from 4 other northern red oak studies, stand and family (Kriebel et al, 1988), intra- 2 in the northeastern and 2 in the south- eastern parts of the USA. A summary fol- provenance selection is important for plant- lows. ing in this region.
We do not know whether the same su- in low rainfall regions west of the nances periority of northern over southern origin Mississippi River, near the range limits, are trees of Q rubra applies to plantations in more drought-resistant than those of other other regions. Fragmentary but inconclu- origins. These differences were observed in sive data suggest that it might not apply in provenance test in Kansas, at the south- a regions farther south. In a replicate of the western limits of Q rubra, where mean sum- above experiments that was planted in temperature is highest and mean annu- mer Kansas, tree diameter was inversely corre- precipitation is lowest within the species al lated with seed source latitude, ie, the range. Trees originating from this region, in- southern provenances had the faster- cluding lowa, Kansas and Missouri, had higher survival rates than those from any growing trees. However, data were taken at age 11 years, and the plantation had other provenance (Deneke, 1975). low survival percentages of all seed source Cold hardiness of northern red oak de- samples (Deneke, 1975). A similar trend pends upon geographic origin. Twigs col- was noted in a progeny test in eastern lected from 16- to 18-year-old trees of 38 Tennessee that included families from origins growing in Massachusetts (Pauley Tennessee, Virginia and Kentucky. The and Johnson, 1955) were subjected to shortest 10 families in mean height at age controlled freezing experiments. Cold har- 20 years were from the more northern ori- diness was strongly related to estimated gins (Schlarbaum, 1991).Since all the annual minimum temperature of the mean seed sources were in a narrow latitudinal and to latitude of origin. In all cases, origin range relative to the species distribution, however, cold hardiness was greater than results are not comparable with those of that required by the climate of the origin, the range-wide tests. suggesting that twig hardiness in estab- lished trees is not an important factor in natural selection under contemporary cli- Variation in adaptive traits matic conditions (Flint, 1972). Data of bud-break or leaf flushing of Northern red oak varies geographically in northern red oak depends upon seed source; in the north central region of the drought resistance. Trees from prove-
et al, 1980), with partial replication USA, flushing begins in trees of northwest- tamour in northern Ohio then proceeds ’eastward’ (author’s records). origin, ern through trees of northern origin to trees of northeastern origin, and also ’southward’ Variation in growth rate to trees of central and southern prove- nance, ending in trees of mid-latitude ori- Seed had a strong effect on tree gins from southern Michigan to Pennsylva- source in South Carolina at age 10 years. height nia (Kriebel et al, 1976). This trend is not Southern red oaks that surpassed the local significantly correlated with latitude and it source were from the lower coastal plain of is only weakly correlated with longitude North and South Carolina, southern Missis- (Schlarbaum and Bagley, 1981). In east- sippi, southeastern and north-central Loui- ern Tennessee, the pattern of flushing is siana, southeastern Arkansas and south- very different: the general trend begins in eastern Missouri. Those growing more trees of southern origins and ends in trees slowly were from Piedmont and mountain- of northern origins (Gall and Taft, 1973; ous regions of Virginia and North Carolina, Schlarbaum, 1991). Tennessee, Florida, eastern Texas, Ala- Data of bud-break advances with in- bama, southern Missouri and southern in seed source elevation; in west- crease New Jersey. These provenances are near ern North Carolina, the time spread be- the periphery of the species range. No cli- tween the lowest and highest elevation nal trends were found, nor were there any source was 11 days, regardless of planta- meaningful correlations with latitude, longi- tion elevation (McGee, 1974). tude, mean annual temperature or length Unlike the flushing date, the time of leaf of growing season. in Q rubra is very strongly cor- senescence In central Mississippi, provenance ef- related with the latitude of the seed fects at age 5 years accounted for about source, progressing clinally from north to 70% of total variation, and families within south (Deneke, 1975; Kriebel et al, 1976; stands 20%. There was very little differ- Schlarbaum and Bagley, 1981). ence among stands within provenances. Southern red oaks from seed sources in southeastern Texas and eastern Georgia SOUTHERN RED OAK significantly faster-growing than were those from the other 21 provenances. Two principal studies have been conduct- Farther north, in southeastern Pennsyl- ed on geographic variation in southern red vania, the comparison was made between oak (Quercus falcata Michx). One compris- 2 seed sources of southern red oak that are es two 43-origin, range-wide provenance northern for the species and 2 sources in tests in the Piedmont region of western the southern part of the species range. South Carolina (Schoenike et al, 1982). Progenies from seeds collected in the near- The other is a central Mississippi test of by region of Maryland and Virginia outgrew 112 trees from 43 stands in 23 prove- those of Alabama and Arkansas origins. nances, including most of the natural dis- tribution with the exception of Florida and Variation in adaptive traits areas north of 36°N (Mukewar and Land, 1987). In addition, a few families of 4 prov- enances were tested at the Michaux Quer- Survival of southern red oak in South Car- cetum in southeastern Pennsylvania (San- olina and Mississippi is not source-related.
In southeastern Pennsylvania, it is; trees the tallest of the Tennessee families had a from Mississippi suffer heavy mortality mean height of ≈ 11 m, 2-6 m above the from winter temperatures. Trees of Arkan- means of southern and western trees (Uni- sas sources are less affected, but of the 4 versity of Tennessee, unpublished data). provenances tested, only Virginia and In a more recently initiated study near Maryland trees had high survival rates Mississippi River in extreme northwest- the (Santamour et al, 1980). In Ohio, where ern Kentucky (including provenances from winters are more severe, trees of the Vir- Louisiana, Mississippi, Alabama, Tennes- ginia seedlot that were hardy in Pennsylva- see, Kentucky and Virginia), the trees from nia all died within the first few years after Tennessee and Mississippi were outgrow- planting. Other sources were not tested ing those from other sources at age 5 (author’s records). years. Some were 6 m in height (Rous- seau RT, unpublished data). Farther north, in Indiana, cherrybark CHERRYBARK OAK oaks from the northern extremity of the species range in southern Indiana were Harlow et al (1991) now follow Jensen significantly taller than all others at age 7 (1989) in classifying cherrybark oak as years. The test included 9 seed sources in Quercus pagoda Raf rather than as a form 6 states (M Coggeshall, unpublished anal- of southern red oak (Quercus falcata var ysis and these proceedings). pagodaefolia Ell). The 2 oaks were consid- ered by Ware (1967) and Jensen to be sister species that are incompletely repro- Variation in adaptive traits ductively isolated. In support of species separation, Jensen stated that, "differences between them can be detected consistent- oak is highly variable in cold Cherrybark ly in geographically widespread locales, in- hardiness. In western Tennessee, families dicating that recognition of these taxa as of local origin averaged 92% survival at species is in keeping with the generally ac- age 10 years, while those from Mississippi cepted species concept in oaks." Q falcata and Arkansas averaged 66% (Overton, characteristically occupies a xeric habitat, 1981). In southern Indiana, Coggeshall’s records show that only trees from extreme whereas Q pagoda occurs in mesic habi- southwestern Indiana, the northernmost tats. point in the species range, remained com- pletely healthy after a 10-day period with a low temperature of -31°C. There was a Variation in growth rate high negative correlation between degree of winter injury and seed source latitude. Cherrybark oak has been described as the most rapidly growing southern oak (Ran- WHITE OAK dall, 1972). As in Q falcata, the growth rate of Q pagoda is very dependent upon seed In a western Tennessee test that White oak (Quercus alba L) grows through- source. out the eastern United States, with the ex- included 36 phenotypic selections at age 10 years, trees from Mississippi and Ar- ceptions of northern Maine and the Florida kansas seed sources grew poorly com- peninsula. It also occurs in parts of south- ern Ontario and Quebec. Although it is one pared with those from the Tennessee of the most common and commercially im- sources (Overton, 1981). By age 14 years,
portant oaks in the eastern United States, There was no evidence from any of there is no range-wide provenance test of these limited-sample Q alba experiments the species. The most geographically dis- that survival rate was related to the geo- persed set of population samples is in the graphic origin of the seed. Michaux Quercetum test in southwestern Pennsylvania, which includes small num- bers of trees of 18 families from 9 prove- BUR OAK nances (Santamour et al, 1980). Eight of these provenances are replicated in Bur oak (Quercus macrocarpa Michx) has Wooster, Ohio. There are 2 tests estab- very wide north-south natural distribution a lished by Coggeshall in southern Indiana (table I), extending from Manitoba nearly to containing 63 and 70 families from the Gulf of Mexico. It also has a wide longi- throughout Indiana. Results from an un- tudinal range, extending from New Bruns- published evaluation of 5-year-old in the wick far into the prairies. Bur oak is ex- Indiana tests sent to the author by Cogge- tremely drought-resistant (Fowells, 1965). shall are included in this review. Provenance variation has been studied in eastern Nebraska and on a smaller scale in eastern Pennsylvania and Ohio. Variation in growth rate of variation in No geographic pattern Variation in growth rate growth rate was evident among 24-year- old white oaks of 9 provenances in Penn- test of 50 seed In eastern Nebraska, a sylvania. The sampling dispersion was sampled the species range, but sources from Massachusetts and Virginia in the population sampling was mainly concen- east to Wisconsin and Arkansas in the trated in Kansas and Nebraska. Height at west (Santamour et al, 1980). Results age 11 years was maximum in trees origi- were the same in the Ohio replicate at age nating 160 km south of the plantation at 11 years (author’s data). This absence of 40 °N. There was no observed continuous a pattern is not surprising, considering the geographic pattern of growth rate (Dicke small number of provenances and the vari- and Bagley, 1980). In eastern Pennsylva- ation among seedlots. In southern Indiana, nia, only 2 seedlots each from Kansas, where 5 latitudinal transects were made, South Dakota and Minnesota were tested. including 2 stands per transect, family dif- Height growth tended to increase with de- ferences were about 2.5 times stand differ- creasing latitude of seed origin; Kansas ences. Stand, transect and stand-within- trees averaged about 1.5 times the height transect differences were not significant at of Minnesota trees (Santamour et al, either test location. Growth of trees from 1980). local stands exceeded the test mean at both test sites. These early Indiana results suggested that, in that region, some gain Variation in adaptive traits in growth of white oak may be obtained by using seed sources (or stands) from a lati- tude of up to 2 °N of the planting site, but Date of leaf fall was not related to the lati- tude of the seed origin in Nebraska. In that extreme northern Indiana seed should be avoided for planting in southern India- southeastern Pennsylvania, survival rate of bur oak by age 25 years was 4 times high- na.
trees of more northern origin. A late frost among Kansas trees than among Min- er damaged foliage of local and southern nesota trees. Since the test site has a mild- provenance material, but there was no in- er winter climate and higher precipitation than either the Kansas or Minnesota col- jury to the Arkansas trees (Adams, 1989). In southeastern Pennsylvania, near the lections, the differential survival does not northern range limits, survival of trees of appear to reflect differences in either cold hardiness or drought resistance. Bur oaks provenance was highest, averag- Virginia ing 76%. Survival of Maryland and Missis- from these 2 sources had a high survival sippi trees was lower (Santamour et al, rate in northern Ohio. 1980). Only 3 trees from one source (Vir- ginia) were tested in northern Ohio, a cold- er climate than southeastern Pennsylva- WATER OAK nia. Records show that all 3 were growing at age 11 years. Water oak (Quercus nigra L) is a southern, mild-climate species. At its northernmost limits, mean minimum winter temperature OTHER NORTH AMERICAN OAKS is -18 to -12°C (US Dep Agric, 1960). Most information on geographic variation is exception of one study of Quer- With the available from one 5-year-old provenance palustris Muenchh, information on vari- test in central western Louisiana and from cus ation of growth and adaptive traits in other southeastern Pennsylvania. oaks is based on the testing of open- pollinated progenies at the Michaux Quer- cetum, Longwood Gardens, PA. Seed Variation in growth rate samples from various North American oaks were collected from a small number In Louisiana, 68 water oak families were of trees in each of 2-9 geographically dis- taken from 12 sampling points in the west- persed localities. Trees from each seedlot ern part of the species distribution, consist- were divided into 2 row-plot replicates; the ing of 3 north-south transects, one along total number of trees planted per seedlot the Mississippi River and the others 160 varied from 11 to 42. Analysis of juvenile km east and west of it. At age 5 years, material indicated that there was genetic there was no distinct geographic pattern. variation in growth and phenology, but no Trees from the middle Mississippi River geographic patterns could be identified and middle eastern Mississippi sources (Gabriel, 1958; Santamour and Schreiner, were consistently high in both diameter 1961; Schreiner and Santamour, 1961).An and height growth, but trees of southwest- assessment of growth and survival of all ern Louisiana origin, which averaged 4.2 m oak collections in the Quercetum was at age 5 years, were superior overall in made at age 25 years (Santamour et al, rate of height growth independent of diam- 1980). Three-tree plots of many of the (Adams, 1989). eter Quercetum collections were planted by Kriebel at Wooster, Ohio and measured at age 11 years. The data were not published Variation in adaptative traits and a large part of the test was subse- quently lost due to road construction. Giv- Q nigra trees of northern Arkansas origin en the limitations of these experiments, the results nevertheless report the perfor- flushed 7-10 days later in Louisiana than
the average than trees from the Tennes- of source-documented trees and mance provide some useful information on growth see provenance. and climatic adaptability in relation to prov- enance. Black oak Pin oak There are 11 seed lots of 6 provenances of black oak (Quercus velutina Lam), in the 25-year data from Pennsylvania. Seed ori- Experiments with pin oak (Quercus palus- gins include Alabama, Tennessee, North tris Muenchh) on 19 natural and 2 cultivat- Carolina, Virginia, Illinois and Michigan. ed populations well-distributed throughout The Ohio collections originally included 8 the natural distribution were designed and Q velutina seedlots from the same prove- carried out to evaluate the severity of iron nances and Connecticut. In southeastern chlorosis in solution culture and soil envi- Pennsylvania, seedlot differences in 25- ronments. There were significant differenc- year height growth within provenances es in resistance among the progenies of were small. Although black oaks of Ten- different pin oak parents, but the rankings nessee and Illinois origins were slightly varied considerably among experimental faster-growing than others, averaging environments. There was some indication 16.3 m, North Carolina trees had the high- of a geographic pattern; population sam- est survival rate. In northern Ohio, the ples from northcentral and northwestern mean height of one Michigan family was parts of the species range (Indiana, Illi- 30% greater than the mean of 2 Tennes- nois, Missouri) were consistently among see families. Black oaks of Alabama origin the most resistant populations. One popu- were not winter-hardy in Ohio and there lation from northern Illinois was a particu- larly promising candidate for testing and was dieback of Virginia and North Carolina selection (Berrang and Steiner, 1980). trees. Scarlet oak Shumard oak The Michaux Quercetum plantings include In Pennsylvania, the growth rate of Shu- 4 provenances of scarlet oak (Quercus mard oak (Quercus shumardii Buckl) was coccinea Muenchh) in Pennsylvania and 3 higher in trees of Mississippi provenance in Ohio. In Pennsylvania, trees from Ten- than in trees from Illinois, Tennessee and nessee and Illinois seedlots had a mean Florida sources. Juvenile trees of all height at age 25 years of 15.5 m, a slight sources except Illinois had an extended superiority over the growth of Virginia and growing season and were killed back by Alabama trees. Survival differences were early fall frosts. However, at age 25 years, not source-related. Of the 3 provenances the 70% survival rate of Shumard oaks of represented in Ohio, trees from the one Mississippi origin nearly high was as as that of Illinois trees and growth rate was Virginia provenance were 36% faster- growing at age 11 years than the mean of higher, up to 17.1 m. In the colder climate 2 Tennesee provenances. Scarlet oaks of northern Ohio, the Mississippi and Flori- from Alabama seedlots did not survive in da collections were not winter-hardy. Ohio. Trees of Illinois origin were 17% taller on
Willow oak CONCLUSIONS exception of results from tests of Six seedlots of 4 provenances of willow With the on variation in oak (Quercus phellos L) were tested in Quercus rubra, information Pennsylvania. The provenances were Mis- growth and adaptive traits of the North American oaks is still very limited. Even sissippi, Arkansas, Virginia and Maryland. Willow oaks of Virginia origin were tallest Q rubra is inadequately sampled, consid- with a mean height of nearly 16 m at age ering its abundance and wide distribution. The distribution of Q falcata has been 25 years, but only one-third of the trees survived the winters. The survival rate of more completely sampled than has that of trees in the Arkansas collections was the wider-ranging Q rubra, although the future of the South Carolina tests of about twice that of the others. Only Arkan- sas trees were winter-hardy in Ohio, but southern red oak is uncertain. However, some useful information is now available growth was slow, averaging 1.5 m at age 11 years. about intraspecific variation in growth rate and winter hardiness in this species. Stud- ies of variation in Q pagoda are yielding Shingle oak valuable information, but they do not cov- er the entire species range. Considering the economic importance of Q alba, we Shingle oak (Quercus imbricaria Michx) should be much farther ahead than we represented in both Pennsylvania and was are in our research on this species. Our Ohio by 3 sources (Illinois, Indiana and white oak tests are very limited, both geo- Ohio). There were no clear source-related graphically and in sampling intensity. Be- differences in growth rate or hardiness in cause of its drought-resistance in the prai- either location. At age 25 years in Pennsyl- Q rie regions of North America, vania, trees of Indiana origin averaged merits further research on macrocarpa 15.6 m in height and were taller than those vigor and hardiness. The recently initiated from Illinois and Ohio. effort under way with Q nigra, focusing on the western part of the species range, will provide some information on this previ- Blackjack oak ously untested fast-growing species. Tests of scattered seed sources of 6 other oaks provide fragments of information One progeny of each of 4 provenances of blackjack oak (Quercus marilandica thay may reduce the risk of plantation fail- Muenchh) was tested in southeastern ure and increase productivity in certain re- gions, although they are at best only indic- Pennsylvania. The provenances were in ative and in no case descriptive of Texas, Kansas, Arkansas and New Jersey. Height growth in 25 years varied from a variation of the species as a whole. Indi- cations of variation in resistance to iron mean of 9.4 m for trees of Texas origin to chlorosis in Q palustris may be useful in 7.2 m for trees from the New Jersey seed source. The survival rate of blackjack oak urban forestry. As far as other North varied from 10% in the Texas progeny to American oaks are concerned, including 38% in the progeny from nearby southern the western species, their intraspecific variation is virtually unknown. New Jersey.
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