ABSTRACT 15 - YEAR PERFORMANCE OF EUROPEAN BLACK PINE IN NORTH CENTRAL UNITED STATES BY Nicholas Collins Wheeler A provenance test of European black pine (Pinus nigra Arn.) representing 25 native stands and 2 plantations was established at 4 test sites in the north central United states in 1961. Average plantation mortality varied from 18 to 42% among the 4 sites with the greatest mortality occurring within 2 years after planting, probably a result of tranSplant shock. There were large differences among seedlots in tree height and diameter. Generally, the same seedlots grew fastest at all test locations. The slowest growing seedlots were from warm, mild cli- mates; the fastest growing seedlots were distributed essentially at random throughout the range. Natural infections of twig and needle blights in two plantations made it possible to evaluate genetic resistance to these diseases. Dothistroma pini caused serious defoliation of current and yearbold foliage on trees grown in Nebraska. In Michigan, Cenangium fer— ruginosum infections resulted in the death of the entire upper third of many trees. One seedlot each from Yugoslavia, Austria and northern Greece were quite resistant, whereas seedlots from Corsica, southern Greece and Italy were most susceptible to both diseases; the differences Nicholas Collins Wheeler .4 Q) (27 were significant. Insect damage also occurred but was not severe. ")4 fl)" ‘u’ Genetic differences in resistance to attack were found for black- headed pine Sawfly (Neodiprion sertifer), Zimmerman moth (Dioryctria zimmermani) and European pine shoot moth (Rhyacionia buoliana). Many seedlots which appeared resistant to Dothistroma and Cenaggium were damaged by insect attacks. In addition to growth and pest resistance data, over #0 morph- ological, anatomical and physiological characters were measured in this provenance test. Two seedlots from Spain were similar enough in several traits to be considered a separate variety (var. pyrenaica 3 so were three seed- lots from Corsica (var. poiretiana). Trees from the Peloponnesian Peninsula of southern Greece were similar to each other in growth and pest resistance performance. Elsewhere in the range, there were no consistent geographic trends, differences among seedlots from neighboring ' areas were often as large as those among seedlots obtained from widely separated areas. Hence it is concluded that most taxonomically des- cribed varieties or subSpecies are invalid. Many distinctive characteristics of the Corsican variety can be explained in terms of isolation and natural selection favoring sur- vival in a warm habitat. Genetic drift is a possible cause of the seemingly random variation found elsewhere. Diseases have often been the limiting factor to growth of EurOpean black pine in our plantations. The three most resistant seed- lots are, therefore, the most highly recommended for commercial plant- ing even though they are slightly inferior in some traits (e.g. height and diameter growth) to other seedlots. 15 - YEAR PERFORMANCE OF EUROPEAN BLACK PINE IN NORTH CENTRAL UNITED STATES BY Nicholas Collins Wheeler A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Forestry 1974 ACKNOWLEDGMENTS I wish to extend my sincere gratitude to Dr. J. W. Wright. His instruction and guidance have been invaluable. I am also in- debted to Carl Lee and a number of others for the use of certain data which they collected. I would especially like to thank Don Dickmann, Don.DeHayes, Bruce Bongarten, Kim Steiner and Bonnie Pietila for many stimulating discussions and their endless encourage- ment. I am extremely grateful to my good friend Pamala Pease for the considerable time she spent helping to prepare and type the manuscript. Finally, I would like to thank my parents, Otto and Dorothy Wheeler, for their unfailing support, advice and friendship. 11 LIST OF TABLE 0 O O C O 0 LIST OF FIGURES . . . . . INTRODUCTION . . . . . . MATERIAL AND METHODS . . RESULTS AND DISCUSSION . Mortality O o o o a Growth Rate . . . . Disease Damage . . . Insect Damage . . . Geographic and Evolutionary Considerations Taxonomy 0 o o o o o PRACTICAL RECOMMENDATIONS LIST OF REFERENCES . . . APPENDICES . . . . . . . TABLE OF CONTENTS iii 10 13 18 20 22 21+ 26 28 Page AppendiXA-FIELDDATAOCI000000000000...28 Appendix B - NEEDLE AND BRANCH CHARACTERS . . . . . . . . 46 iv TABLE 1. 10. 11. LIST OF TABLES Relative heights of Pinus nigra seedlots in Ohio, Nebraska and MiChigan o o o o o o o o o Damage by diseases and insects to Pinus nigra at plantations in Michigan, Nebraska and Ohio Number of trees planted, current mortality and height growth of Pinus nigra at Kellogg Forest, Michigan (MSFGP 5-61). . . . . . . . . Floristic and vegetative characteristics of Pinus nigra at Kellogg Forest, Michigan WSFGP SE) 0 O O O O C I O O O O 0 O O C O O Disease and insect damage in Pinus nigra at Kellogg Forest, Michigan (MSFGP 5-61). . . . . Anatomical and wood property characteristics of Pinus nigra at Kellogg Forest, Michigan (I'ISF‘GP5'61)0000000coo-coo... Result of analyses of Variance. Kellogg Forest, Michigan (MSFSP 5-61) . . . . . . . . Plantation establishment and growth char- acteristics for Pinus nigra at Russ Forest, MiChigan (I‘ISFGP 6-61;. 0 o o o o o o o o o o o Foliar and insect damage characteristics for Pinus nigra at Russ Forest, Michigan (MSFCpé-61)ooooooool00000000 Flowering and insect and disease damage characteristics of Pinus nigra at Russ ForeSt, MiChiga-n (MSFGP 6" l o o o o o o 0 Result of analyses of variance. Russ Forest, Michigan (MSFGP 6-61) . . . . . . . . 1h 28 29 30 31 32 35 36 37 38 TABLE 12. 13. 14. 15. 16. 17. Plantation establishment, growth and winter injury data for Pinus nigra at Dunbar Forest, northern Michigan (MSFCP 13-61). . Plantation establishment, mortality and growth characteristics of Pinus nigra at Plattsmouth, Horning Farm, Nebraska (MSFGP63-61)00000000000000 Foliar and fruiting characteristics of Pinus nigra at Plattsmouth, Horning Farm Nebraska (MSFGP 63-61) . . . . . . . . . . Plantation establishment, growth and foliar characteristics of Pinus nigra at Wooster, Ohio (MSFGP 66-61) . . . . . . Foliar, insect damage and fruiting characteristics of Pinus nigra at Wooster, Ohio (MSFGP 66.61) . . . . . . . Plantation establishment and growth characteristics of Pinus nigra in Mason County, Illinois . . . . . . . . . . . . . vi #0 41 1+2 “3 45 FIGURE 1. LIST OF FIGURES Natural range of Pinus nigra Arnold . . . . . . . . . Distribution of native stands of Pinus nigra represented in the present provenanCGSLUdYo....o...o.o....... vii 6 INTRODUCTION European black pine is a widesPread tree Species of central and southern Europe, Asia Minor and northern Africa. It is known by sev- eral common names such as Austrian, black, Corsican, and Calabrian pine. Strictly Speaking, the name EurOpean black pine (Pinus nigra Arnold) refers to the entire Species; the other common names typically refer to trees from Specific geographic locations such as Corsican pine (P, nigra var. poiretiana) from the Island of Corsica. Consid- erable taxonomic confusion exists in the description of this Species. Rehder (19#0) presents a detailed list of the more than 80 different Latin names proposed for all or parts of the Species. European black pine from Austria was one of the earliest tree introductions into the United States, arriving in 1759. It is pop- ular as an ornamental and windbreak in many parts of north central United States. In recent years, Pinus nigra has proved one of the best trees for roadside planting because of its high salt tolerance. It is particularly well adapted to calcareous soils and will grow as rapidly as native pines when planted for timber on such sites, but the acreage of such plantations is small. European black pine is grown on a relatively small scale for Christmas trees in the U. S. because of difficulties encountered in shipping it in large quantities; the branches are brittle and apt to break if trees are piled high. How- ever, the tree re5ponds nicely to shearing and is preferred by many people. As noted, the natural range of European black pine is large. The range is markedly discontinuous with isolated stands occurring from Algeria and Spain in the south and west to Austria and the Crimea SSR, in the north and east. It has a latitudinal range of 130 (35a to 48° N) and a longitudinal range of 48° (60 W to 420 E) (Fig. 1). Very few genetic studies of EurOpean black pine have been re- ported. It has been the subject of some small unreplicated provenance trials conducted in Belgium, France, New Zealand and eastern United States. In addition, there are two previous replicated provenance tests. The first one, of about the same size and age as the present study, was conducted in West Germany and was described by Rghrig (1966). Prof. Dr. RShrig (personal communication) stated that his plantations were so heavily attacked by a Species of Scleroderris (insect) that they have been abandoned. The second study, established in the late 1960's in France was described by Arbez and Millier (1971). Details of these studies are summarized in Mirko Vidakovic's review paper (in press) on the genetics of EurOpean black pine. They have shown the presence of considerable genetic variability in traits such as height, foliage color, resistance to cold and other characters. The present paper is a summary of the first 15 years' results of a replicated 4-p1antation provenance test of European black pine con- ducted in north central United States as part of the NC-99 (formerly NC-5l) project entitled "Improvement of Trees through Selection and Breeding". Previous papers on this NC-99 study have been published by Wright and Bull (1962), Lee (1968, 1970, 1971, 1972). and Peterson and Read (1971). Figure 1. Natural range of Pinus nigra Arnold. (from Lee, Ph. D. Thesis, Michigan State University) O. can .0» - oON .0“ on. O. c on W... L .\.. 0 fl ..\ .\. Q o o 0 m on“ ' Q o . O .\ o. as .ooo . D 090 o o a .e. u/ , 0 //f C o x 0 /.on o . k a. 11. it. a r a fi c. .. s i. a ‘b .. ... — its 1...! J .w I“ ‘. l .\.. r.\o .. . o . .a. . c 1 P. . 400' f. x. ‘ ’ qa .o. D. . .. K..." \x/l i \ I..’.. .... r a...” v s W ‘3 .2 \\..ln. s x . . . 23L. ,. c \ no | o r..r\n..\ ~ (31.x. ‘ ”I \..\/.{\. w 1‘1‘ nv\ , . . K. .. c . x . x. $41 a r; .1-.. ._ A . : . p. x m r!...\: it! v f/ A on? 00' con oOn on“ .0“ on. .0. on .0 on .0. MATERIALS AND METHODS Seed from 10 average trees in each of 25 native stands and 2 plant- ations was obtained from cooperators by Wright in 1958 (Fig. 2). The seed was sown in an East Lansing, Michigan nursery in the Spring of 1959 following a randomized complete block design and grown for 2 years. De- tailed methods and results of the nursery plantings are given by Wright and Bull (1962). Fourteen field plantations were established in the Spring of 1961 in several north central states. Due to problems in shipping, mor- tality of the seedlings was high. Only four of the original plantations had sufficient survival to afford useful data. All plantations follow a randomized complete block design with 4-tree linear plots. The number of replications per plantation varies from five to ten and individual tree Spacing varies from 7 x 7 feet to 10 x 10 feet. Trees were planted in furrows and chemical weed control (simazine, amino-triazole) was applied to each of the plantations. Growth rate was monitored from time of planting to age 15. A number of additional characters (e.g. pest resistance, leaf morphology and anatomy etc.) were also measured during this period although seldom more than once. Disease resistance was interpreted as a function of pathogen frequency. In Nebraska, Dothistroma pini damage was evaluated by the percentage of infected needles obtained from one shoot from each tree in a seedlot. A Cenangium infection was defined as a lateral branch or 5. Figure 2. Distribution of native stands of Pinus nigra represented in the present pro- venance study. 45- 40' 35' 30' 20' IO' 0. I0“ 40' 35'. 20' o. terminal leader that was partly or wholly destroyed. Individual plantation details are as follow: W. K. Kellogg Forest near Augusta, Kalamazoo County, Michigan. Rolling hills, 0-25% slope on Oshtemo loamy sands, glacial derivation; average mortality 17%: 1973 mean height 15-6 ft. Degrees of freedom: seed16t-26, block-9, error-225. Fred Russ Forest near Dowagiac, Cass County, Michigan. On flat, heavy clay soils; average mortality 47%; 1974 mean height 14.5 ft. Degrees of freedom: seedlot-ZO, block-6, error-101. Apple Creek State Hospital near Wooster, Wayne County, Ohio. Level, clay-loam soil. Fertilized in 1962, 1967 and 1968. Average mortality 34%; 1967 mean height 5.3 ft. Degrees of freedom: seedlot-22, block-6, error-122. Rorning State Farm near Plattsmouth, eastern Nebraska. 0n 6% east slope, Silt loam of loessial origin; average mortality 18%; 1973 mean height 17.3 ft. Degrees of freedom: seed- lot-ZO, block-4, error-78. RESULTS AND DISCUSSION Mortality Average mortality for all seedlots varied from 18 to 47% among the four plantations. Most mortality occurred within two years of planting, probably as the result of tran5p1anting Shock. There was a weak geographic trend to mortality of’Piggg‘giggg origins. Corsican and other French seedlots suffered noticeable win- ter injury while in the nursery as well as in the permanent plantations. Mortality was highest in these seedlots, averaging 49%. Mortality in the remaining nonuCorsican and French seedlots averaged 25% in all planta- tions. 10 Growth Rate With few exceptions, those seedlots which grew most rapidly at one site were among the tallest at other test locations (Table 1). Were it not for the diseases which are now damaging some of the plan- tations, there would be good reason to believe that the relative height differences shown in Table 1 would continue to exist in the future. In plantations such as the one at Kellogg Forest which have been measured at frequent intervals there were few changes in relative performance from 1965 to 1973. Twenty-four of the 27 seedlots which were below average or above average in the nursery at age 2 remained so at age 15. There are serious disease problems in two of the test plantations, however. Entire tops of trees are dying, resulting in decreased height growth. As of 1973, the severity of damage was still so low as to not appreciably influence height growth, but if the damage continues, some seedlots can be eXpected to fall far behind their present rate of growth. The three Corsican seedlots grew most Sloyly, as was similarly reported in Rghrig's (1966) West German study. \This was probably a consequence of mild climate trees being grown under severe northern conditions, for Corsican pine has proved very successful in areas of New Zealand and Great Britain with mild climates. Considering the other, non-Corsican seedlots , it is difficult to make any generalizations about geographic trends in height per- formance. For example, Greek seedlots were among the tallest and shortest; CHE-22 and GEE-24 from the same part of Macedonia differed in height by 23%. In that respect, EurOpean black pine differs from 11 .Table 1. Relative heights of Pinus nigra seedlots in Ohio, Nebraska and Michigan. ‘Relative height, as a percent of plantation mean, (a) when_p1anted at Country Wooster Plattsmouth Kellogg Forest Russ Forest and Ohio Nebraska Michigan Michigan Seedlot Age 8 Age 15 Age 15 Age 15 SPA-2 113 98 103 101 SPA-3 97 87 89 97 ITA-7 131 109 114 121 FHA-10(b) 122 -- 114 95 FHA-14 89 -- 89 85 con-11(b) 55 A-- 89 85 COR-12 -- 84 86 7O COR-13 -- -- 84 70 CHI-8 113 107 114 109 YUG-15 111 111 106 102 ADS-23 105 105 106 103 CHE-16 -- -- 68 -- ORE-17 101 110 108 - 98 GRE-18 107 105 105 115 ORE-19 109 106 111 113 ORE-20 95 106 111 113 CRE-21 76 85 84 8? CHE-22 95 106 108 113 GRE-24 95 80 86 87 CHE-25 116 -- 110 102 ORE-26 95 91 97 104 CHE-27 -- 106 105 99 GRE-28 101 104 105 97 CHE-2 9 II- n.- 81 87 TUR-l 91 96 95 86 TUR-4 95 97 111 10 TUR15 951, 95 103 99 Plantation mean (ft) 5.4 17.3 15.6 14.3 F-value (seedlot) 7.16*** 7.88*** 5.05*** 2.19** L.S.D. (.05) 18.0 11.0 17.00 19.0 (a)AUStria, CORsica, CRImea, FRAnce, GREece, ITAly, SPAin, TURkey, YUGoslavia. (b)Parenta1 stand was a plantation of the presumed origin shown. **, ***)Indicates significance at the 1 and .1% levels, respectively. 12 Scotch pine, another south Eurasian Species. Wright gt El (1966) found that Scotch pines from Spain were consistently slower growing than southern French Scotch pines, which in turn differed consistently from Italian, Yugoslav, Greek, Turkish, etc. Scotch pines. These same two Species exhibited considerable differences in trunk diameter growth. When eXpressed as a proportion of height (height/diameter ratio), diameter in European black pine is nearly 50% greater than in Scotch pine. In absolute terms, this means a 15 foot tall black pine would have a diameter approximately 2 inches greater than a Scotch pine of similar height. Peculiarly, those Scotch pine seedlots from Spain that put on the least height growth had the largest trunks. The situation was reversed in black pine, with Spanish trees having the smallest trunks relative to trees from the rest of the range. 13 Disease Damage Considerable disease damage has occurred at two of the European black pine test plantations. In Plattsmouth, Nebraska an epidemic needle blight, Dothistroma pini, began in 1967. More recently, a serious twig blight infection of Cenangium ferruginosum was observed at Kellogg Forest in Michigan. The extensive damage at these test sites made it possible to evaluate genetic resistance to both diseases. Dothistroma_pini has caused extensive losses to Pinus nigra plantings in several Great Plains states. It has been particularly damaging to shelterbelts of European black pine in Nebraska (Peterson, 1967). The disease has also been observed on Pinus nigra in Great Britain and a number of European countries and on Monterey pine in New Zealand, Chile and Africa. Dothistroma is a disease that infects current and year-old foliage, resulting in browning and death of the needles. At the Nebraska plantation, defoliation became so severe at the peak of.inp faction that few trees were believed capable of surviving. -The disease was controlled shortly thereafter with a fungicide but once more reached epidemic levels in 1971. Very large and significant differences in resistance to the disease were observed between seedlots during both periods of infection (Table 2). The highly resistant YUG-15 and slightly less resistant AUS-23 and ORE-18 seedlots are of great interest. On trees from those Seedlots, only 3, l6 and 22% of the needles were infected, reSpectively. Only three other seedlots had infection rates of less than 50%. In contrast, 14 Table 2. Damage by diseases and insects to Pinus nigra at plantations in Michigan, Nebraska and Ohio. Country Disease damage Insect damage and Dothistroma Cenangium Zimmerman Black-headed Seedlot pini ferruginosum __pine moth _pine sawf1y_ Zrneedles infections ---% trees attacked -- infected per tree SPA-2 76 1.0 15 30 SPA-3 '88 1.4 26 40 ITA-7 48 3.0 21 50 FHA-10 -- 2.3 35 32 FRA‘IL" "'" 1 04 17 20 COR-11 "" 30 8 19 3 COR-13 -- 3.6 29 0 CRI-8 51 .8 19 3 YUG-l5 3 .l 4 26 GEE-16 -" 300 5 . 8 GRE-l? 30 .6 10 30 GRE-20 31 .5 8 21 CHE-21 98 2.0 9 l CHE-22 -- .4 11 13 GRE-25 -- .4 12 5 GRE-27 61 1.4 11 3 CRE-z 8 55 .4 13 19 ORE-29 -- 2.5 9 2 TUR-4 57 1.4 16 14 Mean 50 1:5 14 16 F-value (seedlot)(§) 3.93*** 1.60* 94.39*** L.S.D. (.05) (a) 1&0 25.0 23.0 *, **, ***) Indicates significance at the 5, 1 and .1% levels, resPectively. (a)lnformation not available. 15 75 - 98% of the needles were infected on Spanish and southern Greek trees, resulting in nearly complete defoliation. Cenangium ferruginosum has had considerable esthetic and biological impact on European black pine. Commonly known as "pruning disease" be- cause of its selective branch removal through infection, Cenangium has previously been reported to cause severe damage over large planted areas of Pinus nigra in Hungary and Germany (Laubert, 1926; Lengyel, 1963; Lorenz, 1967). In 1973, the disease caused extensive damage in the black pine plantation at Kellogg Forest in Michigan. At this site, disease expression was variable. 0n trees with slight infections, damage was characterized by a twig blight of vigorous lateral and terminal shoots, resulting in the death of the current year's twigs. More severe infections, however, killed the previous two or three year's growth. In such cases, the entire top third of a tree died. Over 44% of the 900 living trees in the plantation were infected. Half of the infected trees lost their leaders Although mortality due to the disease was low in 1973 (3 dead trees), many trees appeared so weakened that they might not survive the following season. There were large and statistically significant (f=3.93, 26 and 225 degrees of freedom for seedlot and error, resPectively) differences among seedlots in resistance to Cenaggium (Table 2). The YUG-15 and AUS-23 seedlots were once again, as with Dothistroma, clearly superior in resistance. The few infections observed on trees of these seedlots were generally confined to the new year's growth on single lateral branches. They were the only seedlots not to suffer leader damage. The most severely damaged seedlots were of Corsican and southern Greek origin. In these, l6 nearly 50% of the trees had dead leaders and half-dozen or more branches were simultaneously infected. However, there were individual disease-free trees in each of these seedlots. It is important to note that the same two seedlots -- YUG-15 and AUS-23 were most resistant to both diseases and that ITA-7, COR-12 and ORE-21 were among the most susceptible to both diseases. However, there were some seedlots (e.g. TUR-l and ORE-28) which showed more resistance to one disease than to the other. This indicated that two different diseases were involved, and that Specific factors were responsible for resistance to each. 0f additional interest is the confusion that exists concerning the true pathogenicity of Cenaggium. The disease is considered by some (Lorenz, 1967) to be primarily saprophytic while others feel it is a facultative parasite on trees predisPosed to disease by unfavor- able conditions. The bulk of the literature on Cenangium indicated the latter case may be more common. Lengyel (1963), in an extensive study of climatic variables, found that an outbreak of pruning disease in 7200 ha. of Pinus nigra in Hungary was strongly related to certain climatic factors, expecially drought, which may have resulted in a physiological weakening of the trees. Lukonski (1968) and Kobayashi and Maniya (1963) also found Cenaggium to be a problem on weakened pines in Poland and Japan. An infection on a ponderosa pine plantation in British Columbia, Canada resulted in 25% of the 300 trees exhibiting flag- ging of vigorous branches in the upper crown. That infection was preceded by two years of drought (Molnar, 195“). 17 In Michigan, it did not appear that a climatic variable was reSponsible for physiological weakening of trees but rather that cli- matic conditions may have been optimal for the Spread of this particular disease. The proper set of environmental conditions for Spread of Cenaggium is not know, however. The disease apparently occurs in serious proportions only every 10 to 15 years in Michigan. Such was the case in 1973 when seemingly healthy trees were damaged in much of central and southern Michigan. In addition to the Pinus nigra infection previously mentioned, severe damage was noted in plantations of Scotch pine Christmas trees and in pine hybrids of Japanese red pine and European black pine. The same hybrid pine was damaged by Cenangium in Fort Wayne, Indiana in 1973. 18 Insect Damage Damage due to insect pests has been moderate in the black pine test plantations. At least one infestation of the black-headed pine sawfly, Zimmerman pine moth or the European pine shoot tip moth has been observed at every plantation with the exception of Plattsmouth, Nebraska. All three insect pests were observed at the Kellogg Forest plantation. The most commonly occurring insect has been the black-headed pine sawfly (Neodiprion sertifer) which in the larva stage, feeds on year-old needles. The sawfly has been noted to occur in both Michigan plantations and in Ohio, the latter infestation being the most severe. Nearly 37% of all the trees in the Ohio plantation had sawfly larvae in 1968, while fewer than 10% of the trees were attacked at the Mich- igan test Sites (Table 2). The differences in susceptibility to sawfly were large and statistically significant among seedlots. Peculiar- ly, some of the seedlots which were most resistant to the sawfly (e.g. CHE-21, CHE-29, COR-12, and COR-13) were among the most susceptible to the previously described diseases. The sawfly damage does not appear to have markedly influenced survival or growth rate in the infested planta- tions, but has made the trees unsightly. The Zimmerman pine moth (Dioryctria zimmermani) is a phloem-borer whose presence can be recognized by large pitch exudations, normally at the main Stem nodes. The insect has been seen only at the two Michigan sites, where approximately 14% of all trees were attacked. Potentially, the Zimmerman moth is probably more capable of inflicting restricted growth, poor form and increased mortality damage 19 than is the pine sawfly. However, infestations to date have been relatively light, both on an individual-tree and plantation basis. Differences among seedlots in susceptibility to the Zimmerman moth were significant (5% level, Table 2). The most resistant seedlots were widely scattered over the Species' range and included COR-12, YUG-l5 and TUR-l. There appeared to be no apparent relationship between a particular seed- lot's resistance to Zimmerman moth and its resistance to the other insect and disease pests previously described. The European pine shoot moth (Rhyacionia buoliana) was observed only once in the test plantations, at Kellogg Forest in the winter of 1974. Damage at that time resulted primarily in death of the lateral, 1973 growth, although death of a few leaders was attributed to the insect. Approximately 200 individual attacks were noted in the plantation. Differences in susceptibility occurred and were significant (5% level). Those seedlots suffering the fewest attacks were YUG-l5, COR-11, ADS-23 and ORE-25. The economic importance of this insect on Pigp§.glg£§ in Michigan has not yet been fully assessed because of its short period of infesta- tion. At the time of this writing (summer, 1974) however, a rather extensive infestation of shoot moth was observed on Pinus densiflora x.P.‘pig£§ hybrids at the Michigan State Tree Research Center in southern Michigan. Some trees lost up to 50% of their crown. Holst and Heimburger (1955), in a review of resistance to European pine shoot moth in hard pine Species, reported that resistance was largely a function of resin production. Pinus densiflora is not very resinous and therefore quite susceptible to attack. They also reported that European black pine from Austria and Corsica were resinous and highly resistant which coincides with our data. 20 Geographic and Evolutionary Considerations It is extremely difficult to identify any major trends in variation in response to geographical, elevational or climatological variables for most of the characters previously described in this study. Almost without exception, characters varied as much within regions as they did between regions. This was repeatedly illustrated in the 13 Greek seed- lots, eSpecially GRE-22, CHE-24 and ORE-25. These showed large differences in height, diameter, branch size and resistance to Cenaggium twig blight, even though the parental stands were within 50 miles of one another. Although there seems to be a lack of range-wide trends, a few Specific geographic areas are worthy of note. Heavy winter damage, high initial mortality and poor growth were common to all three Corsican seedlots. The seedlots from the Peloponnesian Peninsula of southern Greece were also below average in growth rate. In general, these deficiencies were most noticeable during the early years of the experi- ment. As the plantations grew older, winter damage lessened and relative growth rate of these 6 southern Greek trees were extremely susceptible to disease but relatively resistant to insect attacks. It would be valuable to be able to predict areas that would yield trees of superior growth or pest resistance. For the most part, this is not possible. Fast growing seedlots occurred from southern Italy to the Crimea. Strong resistance to insect attack was found in Corsica, southern Greece and northern Turkey. Perhaps of greatest interest were the disease resistance results. The most resistant seedlots to both Dothistroma and Gena ium, YUG-15 and AUS-23, were the only representatives of the northern extension of the Pinus nigra range in Europe. Whether 21 strong resistance to the disease is an inherent character of these two northerly seedlots or whether it occurred merely by chance can not be determined solely on the observations reported in this study. The great amount of within-region variability over the rest of the range suggests the latter case may be true, although eXperience with Scotch pine gave different results. Wright gt El (1966) found considerable regional uniformity in pest resistance, as well as large between- region differences. From an evolutionary standpoint, European black pine poses an important question-~what evolutionary mechanism has had the greatest influence on the Species? Strong selection pressures and isolation surely have resulted in the adaptation of Corsican and southern Greek seedlots to warm, mild climates. These trees are uniform among them- selves, yet as a group, differ considerably from the remainder of the seedlots. Elsewhere European black pine also has a discontinuous range and tends to be a relatively uncommon tree. It is possible that some of the genetic differentiation occurred as the result of random gene fixation and could be interpreted as genetic drift. 22 Taxonomy The taxonomic status of European black pine is extremely com- plicated. As previously mentioned, over 80 Latin names for Species, subspecies, varieties and forms have been proposed for all or parts of the black pine range. Many of the names are mere synonyms resulting from improper publication procedure, but the entire synonomy cannot be so dismissed. The most recent taxonomic treatments 0 European black pine consider it to be one Species, Pinus nigra Arnold, consisting of a small number of lesser taxa. Gaussen 33 El (1964) and BleCic (1967) divided Pigug piggg into 5 subsPecies while Lee (1968), recognizing different groupings, Split the Species into 5 varieties. The latter work was based on the combined data of 40 growth, anatomical and chemical char- acters measured in a replicated provenance test. Even though these recent studies proposed a much more Simplified taxonomy for Pinus nigra, their practical application is still question- able. Although varietal or subSpecies differentiation may be justified from a strictly taxonomic standpoint, the characters utilized as a basis for the decision are often so obscure (e.g. number of hypodermal layers) as to be of little practical significance. Indeed, in our study only seedlots from Corsica and Spain were easily recognized. Lee (Ph.D. thesis) stated that although it is possible to differentiate populations-of Pinus nigra after statistical study based on dozens of Specimens, it is practically impossible to definitely assign any single Specimen to a variety on the basis of its morphology. Even after an extensive study of 40 characters he found the only positive way of identifying a Specimen was to know the geographic location of the parent tree. 23 If, in fact, such difficulty is commonly encountered in identifi- cation and classification of suDSpecies or varieties, then the question of what constitutes a valid intraSpecific taxon needs to be given greater consideration. Justification for the recognition of intraSpecific taxa should be based, at least in part, on the following points: 1. Geographical or ecological distribution. Theoretically, the entity in question should possess an identifiable range, the boundaries of which are geographical or ecological in nature. 2. Genetic similarity. A taxonomic entity should possess a basic genetic component common to all members. 3. Identifying characters. The ability to identify a common set of phenotypic characters among Specimens is the foundation for taxonomic grouping. 4. Usefulness. The taxonomic classification should be applicable, not only under elaborate laboratory conditions or in genetics planta- tions, but under field conditions as well. With these requirements in mind, a reassessment of the taxonomic status of Pinus nigra is certainly worth undertaking. From a strictly practical standpoint (e.g. improvement through breeding, ornamental planting), knowledge of individual stand performance is much more valuable than the taxonomic classification system thus far proposed. For purposes of identification in our study, only Corsican (var. pOiretiana) and Spanish (var. pvrenaica) seedlots warrant separate varietal status. PRACTICAL RECOMMENDATIONS Growth rate, susceptibility to winter damage and insect and disease resistance are the most important considerations when selecting Pinus nigra seed for ornamental, timber or Christmas tree purposes. Differences among seedlots were large for all these characters and constitute a basis for improvement through selection of the proper seed source. Corsican and southern Greek seedlots were the least satisfactory under northern United States conditions and their use is not recommended. Variation patterns in the remaining seedlots do not follow any major geographical or climatological trends, thus recommendations by region are virtually useless. That is, the performance of one popu- lation cannot necessarily be predicted by knowing the performance of a neighboring population. The data obtained in this study indicated that to insure superior growth and pest resistance, it is necessary to return to Specific stands from which the seed was originally obtained. Two seedlots, YUG-l5 and AUS-23, were superior in all~around per- formance at the four test sites. They grew rapidly and had high resistance to diseases and most insects. A third seedlot, GRE-25, was also quite successful and can also be recommended as one of the best geographic origins for planting in north central and northeastern United States. Listed below are the locations from which seed for the present study was obtained. The seedlots are listed in descending order of superiority. YUG-l5. Latitude 43°52' N, longitude 19°32' E. Seed collected 24 25 from natural stand, 900 meters elevation, near Sumska Sekcija, Kremma, Yugoslavia. AUS-23. Latitude 48°10' N. longitude 16°15' E. Seed collected by Franz Kluger at 500 meters, near Vienna, Austria. ORE-25. Latitude 41" 22' N. longitude 21f25' E. Seed collected from natural stands on Mt. Rhodopi near Paranestion, near Drama, in Macedonia, Greece, at elevation of 900 to 1000 meters. Further improvement in Pinus nigra may be obtained by increasing the number of geographic origins tested, by large scale testing of individual tree progenies from within the superior stands and by cros- sing selected trees chosen for their desirable traits. The trees in the NC-91 experiment are now beginning to produce seed, but not in enough quantity to meet ornamental or Christmas tree planta- tion requirements. One way to insure a source of genetically superior seed for future needs would be to establish a seed orchard containing the most desirable seedlots. This has not yet been done. For now, the grower must rely on other means of obtaining seed. If seed is being purchased from commercial seed dealers, the grower should specifically request that it be collected from the areas recommended, namely YUG-l5, AUS-23 and ORE-25. LIST OF REFERENCES 26 LIST OF REFERENCES Arbez, M. et.Millier, C. 1971. Contribution a l'étude de la variabilite’geographique de Pinus nigra Arn. Ann. Sci. Forest. 28: 23-49. Bleéié, V. 1967. GymnOSpermae in Catalogue Florae Jugoslaviae, 1-2: 8-9, Ljubljana. Gaussen, H., V. H. Heywood, and A. U. Chater. 1964. Pinus nigra Arn. in Flora EurOpa, Tutin, T. G., V. H. Heywood, N. R. Bruges, D. H. Valentine, S. M. Walters and D. R. Wess (eds). Vol. 1, Cambridge Press. Holst, M. and C. Heimburger. 1955. The breeding of hard pine types resistant to the European pine shoot moth (Rhyacionia buoliana Schiff). Forestry Chron. 31: 162-169. Kobayashi, T. and Y. Mamiya. 1963. A Cenangium causing dieback of —-—-———H———— Japanese pines. Bull. For. Exp. Sta. Meguro, 161: 123-150. Laubert, R. 1926. Beobachtungen und Bemerkungen fiber das seuchenhafte diesjahre 'Zweigspitzen sterben' der Kiefern. Laudw. Zeit., 43: 543-544. Lee, C. H. 1966. Anatomical characters and chemical composition of European black pine needles as influenced by geographic origins and nitrogen fertilization. Michigan State University, Ph. D. thesis. Lee, C. H. 1968. Geographic variation in European black pine. Silvae Genetica 17: 165-172. Lee, C. H. 1970. Response of different European black pine provenances to nitrogen fertilization. Silvae Genetica 19: 122-123. Lee, C. H. 1971. Trunkwood-branchwood Specific gravity and tracheid length relationships in Pinus nigra. Forest Sci. 17: 62-63. Lee, C. H. 1972. Statistical efficiency in wood quality study based on a black pine plantation. Silvae Genetica 21: 249-250. Lengyel, G. 1963. Dieback of Pinus nigra var. austriaca in Hungary, 1960 - 1962. Erdesz. Kutat, 59: 55-75. 2? Lorentz, Irene. 1967. Untersuchungen zur Biologic and PathogenitSt von Cenangium ferruginosum. Arch. PflSchutz, 3: 143-153. Lukonski, S. 1968. Badania nad biologia szkodliwoscia grzyba Cenangium ferruginosum. Pr. Badaw. Inst. Badaw. Lesnij 196: 351-353 0 Molnar, R. 1954. A disease causing flagging on Ponderosa pine. Bi-mo Progro Rep. DiVO For. 3101., D31). AgriC. Can., 10: 3-7. Peterson, Glenn W. 1967. Dothistroma needle blight of Austrian and Ponderosa pines: epidemiology and control. Phytopathology 57: 437-441. Peterson, Glenn W. and R. H. Read. 1971. Resistance to Dothistroma Eini within geographic sources of Pinus nigra. Phytopathology 1: 149-150. Rehder, Alfred. 1940. Manual of cultivated trees and shrubs. 2nd ed. The MacMillan Company, New York. R3hrig, Von E. 1966. Die Schwarzkiefer (Pinus nigga Arnold) und ihre formen II. Erste ergebnisse von provenienzversuchen. Silvae Genetica 15: 21-26. Vidakovic, Mirko., (in press). The genetics of European black pine (Pinus nigra Arn.). Wright, Jonathan W. and W. Ira Bull. 1962. Geographic variation in EurOpean black pine - two year results. Forest Sci. 8: 32-42. wright, Jonathan w., s. s. Pauley, R. B. Polk, J. J. Jokela and R. A. Read. 1966. Performance of Scotch pine varieties in the north central region. Silvae Genetica 15: 101-110. APPENDICES APPENDIX A Table 3 o 28 APPENDIX A FIELD DATA Number of trees planted, current mortality and height growth of Pinus nigra at Kellogg Forest, Michigan (MSFGP 5-61); Country & Trees Mortal- Height, total Height, total sayi-_3em__yr _____________________ Age 15 15 5 6 7 10 15 5 6' 7 10 15 Year 73 73 63 64 65 68 73 63 64 65 68 73 COR_11 48' i 31-747nug$er1577E42 73" °77m886 '92 COR-12 40 18 31 41+ 60 146 411 73 76 72 80 86 COR-13 40 15 30 46 67 152 404 70 79 81 84 84 SPA- 2 40 3 43 61 86 182 495 101 108 103 100 103 SPA-'3 36 7 46 64v 88 181427 106110 106 99 89 ITA- 7 36 2 43 66 95 214 549 101 114 116 114 114 FHA-10 “0 7 42 63 89 198 549 99 108 109 109 114 FHA-“ 40 9 44 59 77 166 427 101 102 93 91 89 Ans-23 “0 4 42 57 80 181 511 99 98 96 99 106 rue-15 40 4 48 66 92 202 511 117 114 110 110 106 633-16 40 29 36 49 65 145 328 85 84 78 80 68 CHE-17 1+0 6 43 62 84 193 518 100 106 101 106 108 GRE-18 40 7 46 61 83 192 503 107 104 100 106 105 GEE-19 4o 3 46 64 94 206 526 107 108 113 113 110 CHE-20 “0 1 40 58 75 163 533 90 100 90 90 111 GRE_21 40 10 34 51 69 155 404 79 87 83 85 84 GRE-22 36 2 48 64 86 187 518 113 109 103 103 108 GRE_24 36 2 50 64 87 162 411 116 109 105 89 86 GRE_25 4o 10 61 80 106 218 526 142 137 127 120 110 CHE-26 1+0 1 41 57 80 166 415 97 97 97 91 97 ens-27 40 6 45 66 93 182 503 104 113 112 100 105 GRE-28 40 7 42 65 89 206 503 99 111 107 113 105 car-29 ‘ “0 14 1+2 58 78 :79 389 98 99 9+ 98 81 TUR‘ 1 (+0 2 [4'5 61 82 170 (+57 106 105 99 93 95 Tun-1+ 40 2 46 66 92 198 533 106 113 111 109 111 TUB- 5 36 1 43 61 83 180 495 101 105 100 99 103 CHI- 8 40 0 50 70 100 214 549 117 120 120 117 114 Mean 7' 43’ 58 83 182 477 100 100 100 100 100 F value (seedlot) 7.3 6.3 3.9 29 Table it. Floristic and vegetative characteristics of Pinus nigra at Kellogg Forest, Michigan (MSFGP 5-61). Country & WI Branch Br. Cones Male Date of Deg. of Segdlot ______ 8. ggle_ _dlafl. _____ 3 £020_ 2010*Sh0_ _L£0_D§_V_|_ _ Age 5 6 10 10 15 15 15 15 Year 63 64 68 68 73 73 73 73 Char.~ 11 13 37 38 62 51 52 53. T%' 0 mm no. #T _ 58 57 39 7.6 34 5 8 12.9 gg§_1; 72 49 38 6.6 13 0 - 10.3 COR_13 68 52 38 6.5 0 3 11 10.0 SPA- 2 8 62 38 10.7 0 4 21 7.0 SPA- 3 8 61 36 9.2 1 3 4 7.7 ITA- 7 28 59 37 8.2 9 8 20 12.7 FRA-lO 10 60 37 9.4 43 4 - 14.0 FHA-14 35 52 35 8.0 9 11 32 7.0 ADS-23 o 64 34 10.4 36 19 11 12.0 YUG-l5 0 64 37 9.4 9 6 5 10.8 CHE-16 0 63 40 8.0 9 2 - 13.8 GEE-17 2 61 36 9.3 0 16 10 13J+ ORE-18 0 63 37 9.6 0 7 6 13.6 CHE-19 2 60 33 9.3 0 13 13 13.6 GRE-20 2 66 39 8.9 4 3 5 12.3 CHE-21 0 55 35 7.6 0 3 4 13.1 GEE-22 o 60 36 8.9 13 10 9 12.5 GEE-24 0 62 39 7.4 l 26 12 16.0 CHE-25 0 60 33 9.5 17 20 13 12.8 GRE-26 2 55 34 8.0 0 16 12 13.8 CHE-27 2 66 40 8.8 0 11 18 12.7 GRE-28 0 66 39 1 .4 4 14 15 13.7 CHE-29 0 67 35 8. 0 9 9 13.3 TUR- l o 59 39 8.2 0 6 11 13.5 TUR- 4 0 64 35 9.8 0 24 18 14.8 TUR- 5 2 65 43 8.4 0 7 8 13.4 CR1- 8 o 67 39 10.6 0 22 13 .15.4 Mean 11 61 37 8.7 8 ll 17 15.9 F value 61 3.8 3.8 1.0 2.2 for seedlot. WI a Winter injury, Male stob a male strobili, Date of pollen shed, and Degree of leaf development. 30 Table 5.. Disease and insect damage in Pinus gigr§_at Kellogg Forest, Michigan (MSF‘GP 5-61) . Country & Tr. with Inf./ Leader Sht.Moth Zim.Motthucosma BHP saw éesdioi _ _ .Csnensium_Tres .1111“. _ 141:..- - .121". .. _. BIL- ._ .Iaf... .. Age 15 l5 15 15 15 ' ' 11 11 Year 73 73 73 73 73 69 69 Char.i 57 58 59L_ 64 65, 4; 43 % no. % no. no. % trees infected- COR-ll 45 3.8 35 2 3 10 14 COerZ 45 2.9 25 4 2 0 4 COR-13 42 3.6 32 4 0 0 0 SPA- 2 38 1.0 10 5 1 5 8 SPA- 3 32 1.4 18 10 1 6 6 ITA- 7 60 3.0 32 13 1 20 34 FRA914 28 1.4 18 6 1 3 0 AUS-23 22 .3 0 3 5 8 0 YUG-15 10 .1 O 2 1 ‘ 8 13 ORE-16 28 3.0 22 19 2 25 8 GEE-17 30 .6 2 5 0 5 37 CRIB-18 25 .5 5 13 l 5 13 GEE-19 58 1.9 22 7 l 3 13 GEE-20 3O .5 2 10 2 l3 l3 GEE-21 50 2.0 25 36 1 l6 3 GEE-22 32 .4 7 12 1 10 3 GEE-24 28 1.6 20 13 O 10 5 GEE-25 l5 .4 2 3 0 3 7 GEE-26 58 1.4 30 26 0 25 10 GEE-27 40 1.4 15 29 l 10 O GEE-28 20 .5 5 8 2 8 8 GEE-29 50 2.5 32 26 2 40 3 TUR- 1 30 .6 5 12 0 11+ 3 TUR- 4 48 1.5 18 11 3 l3 5 TUR- 5 ' 50 1.8 22 6 l 10 0 CR1- 8 40 ..§, 10 5 Q, .10 3 Mean 37 1.5 16 11 1.4 11 9 F value for seedlot 3.13 5.12 3.90 1.40 1.09 2.53 “.39 Columns 1,2 and 3 - Damage by Cenan ium ferruginosum ZR Sht. Moth - European pine shoot moth hygcionia buoliana Zim. Moth - Zimmerman moth (Dior ctria zimmermani BHP saw - Black headed pine sawfly Neodigrion sertifer) 31 Table (3. Anatomical and wood pr0perty characteristics of Pinus niggg at Kellogg Forest, Michigan (MSFGP 5-61). Country & Needle Scler. Sp. Gr. Tr. Between eeedloi _ _ _ Eminent. _ _. .LeyeL‘S- 1ren.c.h.. Jen. _ iundie_disi._ _ Age 6 6 ll 12 6 Year 64 64 69 70 64 Char. 28.29 26 39a 40a 27 Len. wid. L/w -- microns -- # x10“ mm microns 003-11 778 395 1.96 .4 42831 .93 85.6 con-12 728 372 1.96 .5 37942 .93 82.0 SPA- 2 738 375 1.97 .9 39+64 .94 77.2 SPA“ 3 799 393 2.03 . .9 39962 .92 80.0 ITA- 7 770 397 1.94 .7 41955 .95 70.4 FHA-10 776 392 1.98 .7 42123 .97 78.4 FHA-14 732 384 1.91 ;.9 39266 .93 73.6 AUS-23 838 400 2.10 .8 40707 .94 81.2 GEE-16 765 394 1.9+ 1.0 42113 .97 72.0 GEE-17 747 389 1.92 1.0 41537 .96 74.4 ORE-18 744 367 2.03 .9 42143 .97 85.2 GEE-19 722 379 1.91 .9 40762 .97 62.0 GEE-20 731 368 1.99 .9 41525 .96 70.4 GEE-21 763 380 2.01 .9 39476 .96 76.0 GEE-22 720 366 1.97 1.0 39733 .96 70 .4 GRE -24 792 414 1 . 91 1 .0 41270 . 97 76 . 0 GEE-25 795 402 1 . 98 1 . 0 40592 . 96 70 .4 ORE-26 776 398 1.98 1.0 38635 .97 81.6 GEE-27 857 444 1.93 1.0 38572 .98 87.6 GRE-28 781 401 1.95 1.0 43588 .96 86.0 TUR- l 762 381 2.00 1.0 40514 1.00 73.2 TUR- 4 . 814 418 1.95 1.0 40635 .99 76.4 TUR- 5 _ 843 413 2.04 1.0 41063 .99 78.4 (1121- 8 846 441 1.92 1.9 41526 199 78.9 Kean 772 393 1.97 .89 196 76.5 P value 4.9 4.2 1.6 7.41 3.48 1.32 1.32 for seedlot Scler- Sclerenchyma layers. Sp. Gr. - Specific gravity. Tr len - Tracheid length. 32 Table 7. Result nf analyses nf variance. Kellogg Farest, Michigan MSFGP 5-61. Character Degrees of Mean Squares F values freedom S R SR S R SR S R 393. Br. wood Sp. Gr. 26 9 234 .0026 .0163 .0007 3.48**21.66** 40a. Tracheid length. 26 9 234 48. 1972 Ht. 26 9 23% 19.64 3.733 5.263** 52. Date of Pol. Shed 23 9 110 9.37 .54 17.35** 53. Deg. of Lf. Dev. 26 9 221 49.77 3.12 15.95** 58. Cenangium total Inf.26 9 233 170 43.2 3.93** 57. Cenangium ,# Tr.Inf.26 9 225 2.77 .855 3.13** 59. Cenangium, # Leader 26 9 225 2.04 .525 3.90** 63. 1973 Ht. 25 9 130 286.8 56.8 5.05** 64. Pine shoot math 26 9 233 7.40 5.29 1.39 65. Zimmerman moth 26 9 233 .185 .169 1.094 S s Seedlot R a Replicate SR = Brrnr 33 CHARACTER DESCRIpTTON A description of the characters measured at the Pinus nigra plan- tation at Kellogg Forest, Michigan (MSFGP 5-61). 1. 11 15/61. Dead tree count. WAL, Jun. 2. 4 62. Replaced dead trees. No count made. WAL. 3. 9/25 62. Dead tree count. WAL. 4,5. 6 25/63. 3 fascicles of 1962 leaves collected for anatomical study. Carl Lee 6. 7/21/63. 10 fascicles collected pr tree in reps l to 4. Time 5 hours. J.W. Andresen, Heidi H. Schlosser. 7. 11/21/63. Height, half-inches. Carl Lee. 8, 11/21/63. Dead tree count. Carl Lee. No insect damage. south ends of reps 2,3,4, need weed control. 9. 4/10/64. Needle length, eighths-of-inches. Reps 1,2, 1963 nddeles. Carl Lee. 10. 4/10/64. Bud color, reps 1,2, Carl Lee. Measurements discarded. 11. 4/10/64. Winter damage, No. of trees per 4 trees having brown needles. Carl Lee. 12. 6/3/64. Count of spray-damaged trees. Carl Lee. 360 trees whitened, all recoved later same year. 13. 9/15/64; Branch angle. Measured to nearest 5 degrees of flat- test 1963 branch with vertical. Carl Lee. 14. 9/25/64. Needle length. Collected 1 fascicle per tree. cm/4 C.L. 15. 9/25/64. Height, half-inches. A few 1ammas shoots. C.L. 16. 9/25/64. Needle width, mm/4. C.L. Fert + unfert. separately 17. 9/25/64. Needle serrations per 2 mm. Carl Lee. Fert. + unfert. separately. 18, 19. 9/25/64. No. of rows of stomata on ventral and dorsal side reSpectively. Carl Lee. Fert. + unfert. reSpect. sep. 20. Nov. 1964. Leaf color, judged from ground samples of leaves col- lected 9/15/64, ground for mineral analysis. 2 samples per origin graged. One sample was reps l-lO ert., one sample was reps. l-lO unfert. Color scoring by Wright and Lee. Grade l=yellow~green=Munsell 5.016/6. Grade l7=blue-green- 5.016/8. 21. No. or resin canals. Measurements by Carl Lee 22. Position of resin canals. on the l-needle-per-tree 23. No. of hypodermal layers, ventral side samples collected 9/25/6. 24. No. of hypodermal layers, corner. 25. No. of hypodermal layers, dorsal side. 26. No. of sclerenchyma layers. 27. Distance between fibrovascular bundles, in .04=mm units.. 28. Length of endoderm, 104-mm. units. 29. Width of endoderm, 104-mm. units. 30. 1965 height, 1/2-inch units. Carl Lee. Nov. 1965 31. 1963 height, l/2-inch units. " " " " 32. 1964 growth . 1964 - 1963 height, 1/2 inch units. C.L. 33. Dead tree count, 8/22/68. C.L. Heavy weed competition reSponsible for increased mortality in reps 4 to 7. 34. 8/22/68. No. of trees with flowers or cones. None were seen. 35. 8/23/68. Height, fortieth-of-foot units. C.L. 36. 8/24/68. Number of insect-damaged trees. Damage on 1964 to 1967 whorls. Apical portion of twig dead. Eucosma damage? 34 Character description (continued) 37. 380 8/24/68. Branch angle, 5 degree units. Angle with vertical of flattest branch of 1968 whorl. Carl Lee. Branch diameter, southernmost branch of 1968 whorl, measured by steel calipers in mm/4 units. Branch severred, taken to Stevens Point for study. Carl Lee. 39, 40. 7/23/69. Height, eighth-of-foot units. Measured fertilized 41. 42. 43. L140 45. 46.. 47. 48. 49. 50. 510 52. 53. and unfertilized trees reSpectively. No significant diff., d data combined. JWW. Corrected heights by a 5-tree running average. Correction reduced error term only slightly. 7/23/69. Trees per plot with Eucosma damage. 1-2 twigs per damaged tree 7/23/89. Trees per plot with cones. wa. 7/23/69. Trees per plot with black-headed pine sawfly damage Damage per tree slight. Deviations from variety mean. JWW. 7/23 height. Correction to 7/23 height on basis of 5-tree running average. 7/23/69. Trees per plot which are Scotch pine. 7/23/69. Corrected 7/23 height on basis of 5-tree running average. Dead trees, by position in plot. 5/10/73. Kim Steiner. Tree, per 4-tree plot, with 2 year cones. 5/10 73. K.S. Trees, per 4-tree plot, with l-year cones. 5 10/73. K.S. Trees, per 4-tree plot, with male strobili. 5/10/73. K.S. Date of pollen shed. 1=May 26, 2=May 29, 3=June l, 4=June 4, 5=June 7- K08. . Leaf phenology grades-sum for 4 trees. K.S. 6/4/73. 1=needle still in sheaths, 2=need1es just barely out, 3=need1es %“ out of sheaths, 4=needles %" out of sheaths, 5=needles %-1" out of sh. 54.to 56. Preliminary data taken on disease. Discarded. 57. 580 59. 60. 61. 62. 63. 64. 64a. 65. No. of trees infected in each plot. Disease is Cenangium fer- faginosum. Some damage was confounded with shoot moth and was sub- sequently' removed in character summary, 64a. Nicholas Wheeler 1/17/74. Total no. of branches + terminal infected per 4-tree plot. N.W. No. of leaders infected per plot. N.W. 1/17/74. No. of whorls where every branch has been infected. N.W. 1/17/74. Square root transformation of no. of cones observed per plot. Nicholas Wheeler 1/17/74. Dead trees, by position in plot. 1/17/74. N.W. Ht., in t feet. Mean of two tallest trees in plot. N.W. Pine shoot tip moth-total no. of shoots infected per'4-tree plot. N.w. 3/30/74. Revised Cenangium damage to remove confounded shoot tip moth. N.W. 4/28/74. Zimmerman moth-total no. of trees infected per 4-tree plot. N.W. 4/13/74. 35 Table 8. Plantation establishment and growth characteristics for Pinus nigra at Russ Fhrest, Michigan (MSFUP 6~61). Country & Trees Mortal- Height,total Height, total Diameter 86291": __ _ Elan: _. -117 _______________________ Age 2 15 3 5 8 15 3 5 8 15 15 Year 61 73 62 64 67 73 62 64 67 73 73 Char. 31 6 12 23 29 6 12 23 29 35 no. no. centimeters -- 7 of mean- in. CnRall 28 25 24 39 98 408 78 90 93 94 5.0 CnR-12 28 27 28 38 130 305 93 88 122 70 3.0 CnR-lB 28 24 21 35 90 305 74 82 83 70 3.6 824- 2 24 4 32 47 115 442 113 108 110 101 5.4 $24- 3 28 13 34 55 120 424 119 127 114 97 5.4 113-.7 28 5 29 48 129 527 101 110 122 121 5.9 FRA-10 28 16 34 47 128 410 118 108 121 95 4.8 FRA-14 28 12 25 38 93 372 86 88 87 85 4.6 AUS-23 28 9 34 53 114 451 117 121 108 103 5.] YUGblS 28 6 32 50 123 445 111 116 117 102 4.9 GRE-lo 28 27 28 48 109 -- 98 111 104 —- _. GRE-l? 28 16 27 43 96 430 96 100 91 98 5.4 (RB-18 28 6 28 48 118 503 98 110 113 115 6.0 (RE-d9 28 10 3O 43 104 446 105 99 99 102 5.8 (RE-20 28 8 28 45 102 491 99 103 96 113 5.8 CHE-21 28 9 25 33'91 378 86 80 87 87 5.0 (RE-22 24 5 30 42 104 491 105 98 99 113 5.4 (PB-24 24 10 30 43 110 378 105 99 105 87 5.2 CHE-25 28 17 35 44 106 445 124 101 101 102 5.4 (RE-26 28 10 26 36 85 454 94 84 81 104 5.9 (RB-27 28 8 23 36 95 433 80 84 90 99 5.4 (RB-28 28 18 28 43 116 424 99 99 109 97 5.4 (RE-29 24 18 34 43 93 382 107 99 89 87 5.2 TUR- l 28 9 23 35 79 376 80 82 71 86 4.8 TUR- 4 28 ll 26 41 101 451 90 94 96 103 5.8 TUR- 5 28 24 28 42 101 408 97 97 96 94 5.6 CR1. 8 28 7 27 43 98 476 92 100 94 109 5.8 Mean 13 29 43 105 100 100 100 100 5.3 r=va1ues, Sig. * ** ** *w * ** it ** 36 Table '9. Foliar and insect damage characteristics for Pinus nigra at Russ Forest, Michigan (MSECP 6-61). Country & Crown Foliage Zimmer La Lf WI BA LA BHP Frost $241.03 _ _ .. -diag’hx-jelgr- 3239.0: ._ ._ _ .19.. .. .. _ _. .. _ 2a! iniurv Age 11 11 11 3 5 5 6 5 6 6 Year 69 69 69 62 64 64 65 64 65 65 Char. 26 27 25 7 13 14 15 16 20 21 cm/cm grade % of Tr. T% mm 1% o o T% LF% COR-11 .64 6.0 25 0 74 10 49 55 25 74 COR-12 .63 5.0 -- O 95 56 57 —- -- -- COR-13 .58 6.2 -- O 85 56 57 48 O 26 SPA- 2 .65 5.8 28 8 96 13 51 35 15 21 SPA- 3 .68 5.6 53 19 85 28 48 38 48 6 ITA— 7 .57 7.1 30 5 78 10 52 50 28 49 IRA-10 .57 6.6 58 6 80 28 49 49 23 34 FWA-l4 .73 4.9 28 0 95 20 58 45 6 55 ADS-23 .62 4.8 15 20 72 10 56 55 19 36 “JG-15 .57 4.8 8 O 55 13 56 56 9 48 GRE-16 .67 6.5 -- 0 73 O -- —- -- -- ORB-17 .62 5.6 12 7 50 10 58 64 O 56 GRB-18 .62 5.8 21 5 54 20 54 61 17 51 ORE-19 .64 4.3 35 O 72 3 54 64 20 63 GRE-ZO .63 6.2 8 5 54 28 52 73 14 47 GRE-Zl .70 4.6 15 10 55 3 51 63 0 76 GRE-24 .85 4.1 14 12 73 3 55 66 14 57 GRE-25 .58 6.0 35 O 51 O 55 59 0 44 GRB-26 .60 4.6 4 14 72 7 51 63 O 66 GRB-27 .64 4.6 25 5 79 O 58 56 10 86 (RE-28 .82 5.4 31 12 60 10 56 61 9 78 (RB-29 .71 5.0 25 O 64 17 51 62 O 45 TUR- 1 .68 3.9 12 10 64 10 55 61 5 66 TUR- 4 .63 6.6 21 10 73 O 51 56 5 83 TUR- 5 - .67 4.7 50 14 73 0 56 61 14 62 CR1. 8 .60 5.2 25 O 74 10 56 61 O 75 Mean .63 5.2 23.2 4 70 13 54 56 12 54 T1 = Tree percent, Lf% = Leaf percent LA, LA = Leaf angle, WW a Winter injury, BA = Branch angle, La = Lammas shoots, ‘ 37 Table 10. Flowering and insect and disease damage characteristics of Pinus nigra at Russ Forest, Michigan (MSFGP 6-61). Country 8: Zimmer. Male Fernale Diplodia Pine shoot gee-glgt _____ biolh_ _ _ _F_];oy_ _ flower _______ 1:1th ______ Age 16 16 16 16 16 Year 74 74 74 74 74 Char. 3O 28 32 33 34 (a) (b) (c) (a) (a) CnR-ll «. 8 6 0 0 l CnR-lz O O O 28 O COR-13 20 O O O 7 SPA- 2 5 0 O O 2 SPA- 3 9 O 0 O 0 ITA- 7 11 3 O 6 O ERA-10 13 l O 9 l FRA-14 7 1 0 0 1 AUS-23 3 6 2 O 0 YUG-lS 0 3 2 0 0 (1213.17 6 3 3 0 0 (1113-18 5 7 4 0 O (RB-.19 7 9 2 6 O (RE-20 3 6 5 0 O (RB-21 3 2 O 1 0 (RB-22 2 9 1 O O GUS-24 2 7 3 2 0 ORE-25 2 7 3 2 O CRIB-26 5 10 2 0 0 CRIS-27 2 8 O 0 0 (PE-28 2 5 l O 0 0213-29 0 1 1 O O TUR- l 0 3 3 0 l TUR- 4 7 8 2 8 0 NR. 5 l4 0 0 6 O CRT.- 8 11 8 5 O 1 Mean 4 4 1 2 .5 P value 1.98* 3.0* 1.4 Tab 1e 11 . (MSFGP 6-61). 38 Result of analyses of variance. {uss Forest, Michigan Character Degrees of Mean squares F values freedom 8 R sn“ 3 R SP 3 R 6. fit. cm 1962. 20 6 113 91.8 170 46.0 2.0* 3.72** 12. fit. cm 1964. 2. 6 113 219.5 459 96.9 2.27**4.64** 23. Ht. cm 1967. 24 6 120 1068 5640 469 2.24**1l.8** 7. Lam. growth, trees 25 9 164 .259 .446 .210 1.23 2.13* 13. Lf. len, mm 2 6 106 1312 521 138 8.94**3.50** 14. Win. Inj. T2 20 6 106 656 2285 458 1.40 4.98** 15. Br angle 20 20 6 106 1519 1638 2040 .75 .80 16. Lf angle 20 6 106 12180 4660 2026 6.00**2.29** 17. Lf Stiffness 20 o 106 .451 '.091 .232 1.94* .39 18. Bud color 20 6 106 .0476 .032 .052 .92 .61 20. Bhp sawfly, trees 25 9 164 .431 .177 .270 1.60 .61 21. Frost injury 25 9 164 3450 1150 730 4.73** 1.57 28. Male flowers 20 6 101 1.44 .19 .47 3.03** .40 29. Ht. 1/4 ft, 1974 20 6 101 216 400 98.6 2.20** 4.18** 30. Zimmerman noth 20 6 101 1.94 .66 .98 1.98* .678 32. Female flowers 20 6 101 .39 .57 .27 1.433 2.09 ** 35. Stem diameter 20 6 101 116 169 75 1.547 2.253* 37. Needle angle 20 6 101 .98 .44 2.22** 38. Branch size 20 6 101 .81 .68 1.19 39 Character description, Russ Forest, Michigan (HSFCP 6-61). bU-JNH .000 U1 o 7. ll. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 6/13/61. Dead tree count, J. L. Ruby and J. u. Wright. 6/13/61. Count of trees which have started growth. Ruby, Wright. 12/5/61. Dead tree count. Bright. 4/26/62. Count of trees replaced. Soil dry, temp. 76 F., windy. Bright, hollenbect. Dead tree count, Bright, Wright. Killed one 3-foot rattler. In June Bright sprayed amino-triazole around trees. June weed control ineffective, now 100% weed cover. 8/13/62. Height, cm., wa, JNB. 8/13/62. Trees per plot with 1ammas growth. JHW, JND. 3/23/64. Dead tree count. Carl Lee. " Height, half-inches. Carl Lee. Leaf length, eighths of inches. C.L. Trees/4 trees with winter injury, brown needles. C.L. Angle of 1963 branches with vertical, to nearest 5 degrees. Angle of 1963 needles with leader, to nearest 5 degrees. " Needle stiffness, 1 = stiff, 2 = less stiff. No diff. C.L. Bud color. No differences. C.L. 6/9/64. Dead tree count. JND. ” Trees/4 trees with black-headed pine sawfly attack. J.B. Frost damage, spring 1964. X of levs. brown, to nearest 5%. 1/9/67. Dead tree count. hanover and Wright. " “eight, inches. hanover and Wright. 6/1/74. Male flowers, 0 = none, 1 = l to 100, 2 = more than 100. Nicholas Wheeler/ height in 1/4 feet of two tallest trees in the plOt. N.U Zimmerman moth, no. of trees infected in 4 treeplot. N.H Dead tree count by position. Female flowers 0 = none, 1 = 1 to 25, 2 = more than 25. 6/2/74. Diplodia, number of branches infected per 4 tree plot. ” Shoot tip moth, number of branches infected per 4 tree plot. NM. Diameter at 2nd internode in 1/10 inches. Color, 1 = yellow, 4 = green. Data discarded. Needle angle, 1 = closely appressed, 4 = widely diver- gent from stem. N.N. Branch size, 1 = fine, 4 = robust. 40 Table 12. Plantation establishment, growth and winter injury data for Pinus nigra at Dunbar Forest, northern Michigan (MSFGP 13-61). Country & Trees Mortal- Height Winter Injury 5.69.4.1“: _ .. 313v; - it! ______________________ Age 3 7 7 9 7 9 Year 61 65 65 67 65 67 no. no. --cm--- --- grade(a)- COR-11 4O 23 21 48 96 8 COR-12 4O 22 23 58 88 8 CnR-13 4O 28 20 53 72 8 SEA. 2 40 16 42 61 43 8 SPA- 3 36 8 43 85 54 8 TTA- 7 36 14 32 81 74 6 FRA-lO 4O 26 32 79 54 8 [WA-l4 36 36 -- -- -- - AUS-23 40 5 45 147 30 1 YUG-ls 32 6 47 137 24 0 GRE-l? 4O 12 36 116 44 1 GRE-18 36 6 45 147 33 4 GRE-l9 36 16 43 -- 46 - GRE-20 40 7 35 119 48 4 GRE-Zl 40 31 29 -- 67 - GRE-ZZ 40 6 41 119 49 3 GRE-24 40 25 30 91 68 8 GRE-26 4O 11 30 68 51 4 ORB-27 36 20 31 96 64 8 GRB-28 40 7 41 144 45 2 GRB-29 40 21 21 94 68 8 TUR- l 40 10 3O 79 59 2 TUR- 4 ' 40 17 35 137 54 4 TUR- 5 36 18 29 76 65 l CRI— 8 40 15 37 94 51 3 Mean - 4F% 35 98 57 4.7 (a) Grade, age 7: 0 low, 100 = very high age 9: 0 low, 8 = severe 41 Table 13. Plantation establishment, mortality and growth character— istics of Pinus nigra at Plattsmouth, Morning Farm, Nebraska (HSFGP 63-61). Country & Tr. Tr. 10631 height Total height 5%fl%_w__fl;fl9fl ____________________________ Age 2 4 6 7 8 10 11 13 15 6 10 11 13 15 Year 61 63 64 65 66 68 69 71 73 64 §8_*69 71 73 no. no. --~ centimeters -------- -- Z of mean ---- EUR-11 -- -- -- -- -- -- -- -- -- -- -- “- COR-12 25 6 27 37 67 149 214 329 442 51 66 69 78 84 con—13 25 12 -- r— -- -— -- —- -- —- -— -— -- -- SPA. 2 25 2 58 77 127 250 310 402 518 107 101 101 96 98 SPA- 3 25 3 47 69 114 202 256 368 460 87 82: 83 88 87 ITA_ 7 23 2 57 90 147 278 342 448 573 104 113 111 107.109 FRA-10 25 _- —- -- -- _- _- __ -_ __ __ _— _- -- FRA_14 ’4 4 -- __ —— -- _- __ __ -_ __ _- -- __ Aus-23 25 2 72 117 171 290 352 451 554 133 117 115 107 105 YUC-15 25 2 66 103 165 288 370 490 585 121 117 120 117 111 ORE-19 0 -_ __ -- _ _- -2 -2 __ __ __ __ _- __ 688_1 25. o 74 117 169 296 357 475 579 136 120 116 113 110 can 18 25 1 63 105 158 280 340 466 554 117 112 110 111 105 ORE-19 2) o 64 102 155 280 346 466 559 118 112 112 111 106 ORE-20 25 2 54 88 137 252 318 439 559 99 102 104 104 106 GRE-Zl 25 7 33 57 100 185 238 353 448 61 75 77 84 85 GRE-ZZ 8 1 -- -- 139 -~ 294 -- —— —- -_ 91 -- -— CRE424 25 6 46 77 115 208 258 353 424 86 84 84 84 80 CRE-ZS 0 -— -- —— —— -_ -- __ _- __ _- _- _- __ 'GRE-26 25 5 53 85 129 230 280 378 482 97 93 91 90 91 688—27 25 3 51 80 123 242 308 445 557 95 98 100 105 106 GRh—28 25 0 57 95 140 266 326 426 548 104 107 106 101 104 ORE-29 25 3 -- -- -- -— -- —- —— -_ -- -- -- —- TUR— 1 25 1 54 89 136 241 304 414 509 99 98 99 99 96 TUR- 4 2 3 47 73 118 228 284 384 512 87 92 90 91 97 TUR~ 5 25 0 48 77 128 236 289 396 502 89 95 94 94 95 CR1- 8 25 1 54 91 140 260 328 445 566 .992105._136_196_1o7 Mean 16% 54 87 135 247 308 100100 100 100 100 Table 14. Plattsmouth. Horning farm, Nebraska 42 Foliar and fruiting characteristics of Pinus nigra at (MSFGP 63-61). Country & Ndl. 1n. Ndl. dia. Ndl. 1n. Ndl. dia. Cones WI 5888 _______________________________ Age 15 15 15 15 10 15 6 Year 73 73 73 73 68 73 64 mm mm -—-- % of mean -- no. no/ gr. P10 (10116.11 -- -- -- -- -- -— -- CnR-IZ 151 1.59 110 96 0 3O 1 SEA. 2 163 1.44 119 87 .0 155 1 SPA- 3 157 1.30 115 78 0 40 l TIA- 7 149 1.66 109 100 O 235 l FHA-10 -- -- -- -- -- -- - Flu-14 -- -- -- -- .. «II-- '- AUS-23 136 1.82 99 110 11 220 0 YUGblS 129 1.67 ‘94 101 0 75 0 GRB-l6 -- -- -- -- -- -- - GRB-17 133 1.65 ”97 99 0 125 O GRB-18 139 1.70 101 102 0 120 0 GRE-l9 125 1.57 91 95 0 75 0 GRE-ZO 121 1.56 88 94 l 105 O GRB-Zl 111 1.63 L81 98 O 95 l GRE-Zz cu- -- ..- -- -- u- - ORE-24 126 1.84 92 111 l 110 0 ORB-25 -- -- -- -- -- -- - (RE-26 128 1.74 93 105 0 105 0 (RE-27 148 1.70 108 102 0 75 O GRE-28 127 1.55 93 93 O 130 O GRE-29 -- -- -- -- -- -- - TUR- l 135 1.79 98 108 3 190 0 TUR- 4 131 1.66 96 100 0 100 0 TUR- 5 133 1.74 97 105 0 65 0 CR1- 8 152 1.80 111 108 0 125 0 Mean 136 1.65 100 100 .8 94 N81. 8 Needle, 1n. = length, dia. = diameter, grade: 0 = none, 1 = Some. WI = Winter injury, 43 Table 15. Plantation establisMent, growth and foliar characteristics of Pings nigra at Wooster, flhio (MSFGP 66-61). Country 81 -'I‘rees Trs. Total Ht. - Total Ht. Leaf Len. $24192 _ - _ £12933- Read ______________________ Age 3+ 3. 6 7 8 6 7 8 6 6 Year 62 62 65 66 67 65 66 67 65 65 no. no. ---- cm --- -- % of mean mm fimean COR-11 4O 40 43 64 88 70 53 55 79 81 COR-12 4O 39 SO -- -- 82 -- -- 9O 93 COR-l3 O -- -- -- -- -- -- -- -- -- SRA- 2 4O 32 63 134 180 108 112 113 133 137 SEA- 3 4O 29 57 116 155 93 97 97 111 114 I04- 7 4O 31 67 152 210 111 127 131 121 125 ERA-10 40 34 63 131 195 104 109 122 84 87 FRA-l4 40 34 55 110 143 90 92 89 125 129 AUS-23 4O 22 62 119 168 101 99 105 97 100 YUG-IS ' 4O 18 68 134 177 115 112 111 89 92 688-17 40 29 56 119 162 92 99 101 91 94 GRE-18 4O 31 62 128 171 191 107 107 95 98 CHE-19 40 28 61 131 174 100 109 109 86 89 CPR-20 4O 25 55 116 152 89 97 95 79 81 (PE.21 40 38 43 88 122 71 73 7 80 82 (RE-22 4O 22 58 119 152 95 99 95 86 89 (RE-24 4O 30 58 119 152 95 99 95 103 106 (RE-25 4O 30 67 143 186 110 119 116 91 94 (RE-26 40 35 51 116 152 84 97 95 94 97 (RE-27 0 -- -- -- -- -- «- —- -- -- CHE-28 40 27 60 122 162 98 102 101 94 97 (nu’Zo 40 4(7 0-- .0. a- an- o.- u... _. ...-- TUR- l 40 31 55 110 146 91 92 91 99 102 Tug- 4 40 35 54 116 152 89 97 95 92 95 TUR- 5 4O 29 54 113 152 88 94 95 97 100 CR1. 8 40 26 64 137 180 105 114 113 105__108 Mean 77% 61 120 160 100 100 100 97 100 P value 12.1 7.38 7.15 10.6 Note: Mortality, as noted above, is not a true indication of permanent plantation representation. Many trees were replaced the second year. 44 Table 16. Foliar, insect damage and fruiting characteristics of Pinus nigra at Wooster, Ohio (MSFGP 66~61). Country & Foliage Color BHP Sawfly Female Strob. éesdins. _______________________________ Age 9 , 9 9 Year 68 68 68 gtade(a) # trs. inf. trs/plot with COR-ll 4.0 6 1 COR-12 —- .. .. CnR-13 -- .. -- SPA- 2 5.0 22 l SPA- 3 4.0 23 O ITA- 7 6.0 27 5 FRA-lO 6.0 21 2’ FRA-14 4.5 18 2 ADS-.23 4.0 1 5 YUG—lS 4.5 17 . 3 0118-16 -- _. .. CRIS-17 4.5 14. 5 (PE-18 4.0 9 2 (RE-19 4.0 7 1 (RE-20 4.0 11 5 (RE-21 3.5 O l (RE-22 3.5 S l (RE-24 4.5 1 1 ($113.25 4.0 2 3 CPR-26 4.5 15 0 (RE-27 -- -- —- (RB-28 4.0 13 0 (BR-29 —- -- -- 'IUR- 4 4.5 10 0 111R- 5 3.5 5 2 CR1. 8 4.5 6 6 Mean 4.3 10.5 2.6 P value 3.19* 5.54* 1.67 (a) grade: 0 = yellow, 10 = bane-green 95 Table 17. Plantation establishment and grr-wth characteristics of Pinus nigra in MaSon County, Illinois. Country & irees irees Trees Total height §e£dlot _ __ __ __ Elgnteg __ _Dgad __ _ glivgfg) _____________ Age 3 3 5 5 5 __ Year 61 62 64 64 64 no. no. no. em % mean CflR-ll 24 23 4 18 62 C0R-12 24 22 1 ,9 31 CflR-l3 24 12 2 18 64 SPA- 2 24 12 14 33 114 SPA... 3‘ 24 15 14 28 98 TIA- 7 24 21 4 21 74 IRA-10 24 13 5 32 111 IMA~14 24 18 4 27 96 ANS-23 24 4 20 26 89 ‘HJG—lS 24 8 19 32 ~lll GRE-16 0 _- —- _- -- 098-17 24 12 11 28 99 CRE~18 24 ll 14 29 102 GRE-19 24 15 11 36 127 GRH-20 24 15 15 25 86 ORE-21 24 15 10 25 88 GREeZZ 0 -~ —- -- -- (RB~24 24 11 10 26 89 GRB~25 t0 -- -- —- —» GRE-26 24 5 13 27 94 QUE-£7 24 9 13 33 115 GRE—28 24 -- —- -- _- (PE-29 24 16 8 31 106 109- l 24 1? 15 27 95 10R” 4 24 11 12 33 114 TUR- 5 24 14 12 31 107 CRT-.8 24 21 ‘_fl‘~__2 24 85 Mean 58% 42% 28.6 100 (a) Number of trees remaining alive after replacement of original mortality two years prior. APPENDIX B APPENDIX B NEEDLE AND BRANCH CHARACTERS Since the establishment of the geographic origin tests in.§i22§ ni ra, a number of morphological, physiological and anatomical characters have been measured by Carl Lee and others. host of this information was obtained from the Kellogg Forest Plantation because it had the lowest mortality and most complete representation of seedlots. Lee (1968) measured 16 different needle characteristics in Pinus nigra, 10 of which exhibited statistically significant differences among seedlots. In addition, several branch characters were observed including branch- wood Specific gravity and tracheid length (Lee, unpublished data). This information is valuable in the identification of genetic variabil- ity patterns and more Specifically, identification of individual seed- lots that may provide a basis for improvement through breeding. A few generalizations may be made concerning patterns of genetic variability in leaf and branch characters of European black pine. Lee (1968) noted that Western seedlots (Spain, France, Corsica and Italy) were characterized by long, fine needles with a blue-green tinge. The needles were often closely appressed to the stem. With the exception of the Spanish seedlots, western trees had relatively fine branches. Some individual western seedlots could be distinquished relatively easily because of their particular combination of characters. Spanish trees had an airy appearance due to their long needles, short needle retention and very robust branches. Corsican seedlots had fine, orange 47 branches and long, curly needles. Anatomically, western seedlots had leaves with more resin canals and fewer hypodermal and sclerenchyma layers than did trees from the remainder of the Species' range. In contrast, eastern seedlots (Yugoslavia, Austria, Greece, Turkey, and Crimea) normally had short, stout needles on robust branches. Although large between-seedlot differences existed in most characters measured, it was essentially impossible to identify Specific eastern seedlots. Considerable within-region variation existed. Two important wood properties, tracheid length and Specific gravity, were also found to differ considerably among seedlots. Although statistically significant (1% level), the absolute differences in tracheid length were so small as to be of little practical importance. Specific gravity was relatively uniform over much of the Species' range with the single exception of YUG-lS which had a far higher Specific gravity than the other seedlots (Specific gravity = .44 for YUG-lS and .41 for all others, approximately). Perhaps the most valuable informa- tion obtained about Specific gravity was its relationship with tree height. Although weak (correlation coefficient r = .37, 25 degrees of freedom), the positive relationship between 1968 height and branchwood Specific gravity implied that selection for faster growing individuals does not necessarily result in selection for wood of lower specific gravity. Ill1|llllllHHIIIlllllllll11111111111111 111111 3105502334