Species Hybridization in the Genus Pinus

SPECIES HYBRIDIZATION IN THE GENUS PINUS

BY PETER W. GARRETT

FOREST SERVICE RESEARCH PAPER NE-436 1979 FOREST SERVICE, U.S. DEPARTMENT OF AGRICULTURE NORTHEASTERN FOREST EXPERIMENT STATION 370 REED ROAD, BROOMALL, PA 19008 The Author PETER W. GARRETT is the principal geneticist at the North- eastern Forest Experiment Station's Forestry Sciences Laboratory in Durham, New Hampshire.

.- MANUSCRIPT RECEIVED FOR PlJBLICATION 8 NOVEMBER 1978

Abstract Results of a breeding program in which a large number of pine species were tested indicate that a number of species and hybrids may be useful in the northeastern United States. Austrian black pine x Japanese black pine and hybrids containing Japanese red pine all had good growth rates. While none of the soft pines grew faster than eastern white pine, a number of hybrids grew equally fast and the nonnative component in the crosses may provide pest resistance, which is the number one problem in the Northeast. INTRODUCTION A 6 x 6 foot spacing was used in all plant- T ings at each location. A completely random- IN1947, the genetics project, located at the ized, single-tree plot design was used in some Morris Arboretum near Philadelphia, Penn- plantations while others contained rows from sylvania, began an extensive hybridization a single source or a cornbination of the two. program with a number of genera of trees. All seedlings for the New Jersey planting Reports of species crossability have been pub- were grown in the Washington Crossing State lished by J. W. Wright (1948, 1953, 1958, Nursery. Seedlings were moved to a field lo- 1959). cation adjacent to the nursery when they were This paper reports on the performance of 3 years old (May 1955). the pine hybrids at two locations: Washing- Trees were measured in March, 1967, after ton Crossing State Nursery in west-central 12 field seasons ( 15 years from seed), and the New Jersey (lat. 40°15' N., long. 74"45' W.) following information was recorded: survival, and 'r,Villiamstown, Massachusetts (lat. 42"45' height, diameter, straightness of main stem, N., long. 73"15' W.) . number of branches, multiple stems, forking of main stem or lack of apical dorninance, and MATERIALS AND METHODS presence of cones. Seedlings for the Massachusetts plantings With few exceptions, the trees used as par- were also grown in the New Jersey nursery ents were planted specimens of unknown seed and field planted in May, 1955, and May, origin and were located in the Morris Arbore- 1956, as 2-year-old seedlings. The trees were tum and the surrounding area. Seedlots with measured in October, 1968, after either 13 or numbers preceeded by the letter "N" were 14 field seasons (15 or 16 years from seed), supplied by a cooperator and in most instances and survival, height, diameter, and form were were not of hybrid origin, The exceptions were recorded for each tree. Additional informa- two lots of (Pinus rzigra x Pinus densiflora) tion such as the presence of weevil injury, x open-pollinated seed supplied by Cold heavy fruiting, or unusual branching habits Harbor Laboratory, hng Island, Kew Vork. was noted for individual trees or seedlots. RESULTS nations except for the P. thunbergiana X P. tabulaeformis combination, which produced good height growth. P. syluestris and P. densi- Washington Crossing, New Jersey flora produced good height growth at this lo- Summaries of data for seedlots of section cation and P. banksiana had excellent growth. Pinus, subsections Sylvestres and Contortae In the following list, seedling information are presented in Table 1. Seedlots with similar is from 3- or $-year nursery measurements parentages have been combined. Individual previously published by Wright (1958, 1959). seedlot summaries are included in Table 2. The "tree" infomation was collected in 1967 Summaries for section Strobus, subsection after the seedlings had been outplanted 12 Cernbrae, Strobi, and Cembroides are pre- seasons (15 years from seed). sented in Table 3. Individual seedlot informa- P. thunbergiana x self. Seedlings were 10 tion is presented in Table 4. to 50 percent shorter than crossed P. thun- The soft pines generally had good survival bergiana. Trees were still 20 percent smaller with the possible exception of P. grifithii. and survival was reduced 12 percent. When P. gri@thii was used as the female P. thunbergiana x P. thunbergiana. No parent, survival was lower than that obtained seedling information was available. Trees were using other species; also reduced growth was slightly smaller than most other hybrids in apparent. Growth of P. strobus x P. grifithii the New Jersey planting. hybrids was as good as any other cornbina- P. thunbergiana x P. densiflora. This cross tion, but there was a definite reduction in had good seed set and seedlings were 50 per- growth for the reciprocal cross-P, grifithii x cent taller than P. thunbergiana. This height P. strobus. Survival and growth of selfed lots advantage had shrunk to less than 15 percent of P. strobus were poorer than those of out- by 1967. crossed hybrids. P. thunbergiana x (P. densiflora X P. Survival of the hard pines was fair to good thunbergiana). Trees in New Jersey were 14 with as much variability between seedlots as percent taller than P. thunbergiana or about between species combinations. There was re- the same height as P. thunbergiana x P. duction in survival and growth of selfed lots densiflora. Trees in l'v1assachusetts were 19 (P. densiflora, P. thunbergiana j . percent. taller and 13 percent larger in diarneter than P. thunbergiana. Survival was about Williamstown, Massachusetts the same. Summaries of data for each planting at P. thunbergiana x P. tabulaeformis. Seed- Williamstown are presented in Table 5. All lings were 10 to 30 percent smaller than P. species in these plantings are in section Pinus, thunbergiana. Trees in New Jersey were 20 subsection Sylvestres and Controtae. percent larger, and in &..3:assachusettstrees The reliability of the survival figures is were over 100 percent larger. Survival was again dependent on the number of seedlings also improved by 50 percent. planted. P. thunbergiana x P. yunnanensis. Seed- Seedlots with P. syluestris or P. banksiana lings were about 15 percent larger than P. as the female parent generally had good sur- thunbergiana, and trees in New Jersey main- vival while seedlots with P, nigra, P. thun- tained this advantage. bergiana, or P. contorta as the female parent P. thunbergiana x P. taiwanensis. Seedlings had lower survival. Survival of seedlots con- were 20 percent larger than P, thunbergiana, taining P. densiflora was intermediate re- This advantage had decreased to only a 12 gardless of the pollen parent. The only ex- percent difference by 1967. ception to the above is that survival from P. thunbergiana x P. massoniana. Seed- P. thunbergiana is improved when P. tabulae- lings were 30 to 60 percent taller than P. formis is the pollen parent-an increase of thunbergiana, but the trees were less than 10 nearly 50 percent. percent taller by 1967. Seedlots with P. nigra and P. thunbergiana P. thunbergiana X P. nigra. Seedlings were female parents were shorter than other combi- only slightly bigger than those of P, thunbergiana but the trees were 32 percent taller, percent shorter than P. densiflora, but diam- 51 percent larger in diameter, and had a 36 eters were about the same. Survival was re- percent survival advantage. duced by 40 percent. P. ~hunbergianax (P. thunbergiana x P. P. tabulaeformis x P. sinensis. A compari- densiflora). Seedlings were slightly bigger son with P. tabulaeformis was not possible, than P. thunbergiana, but none was out- but this hybrid was as good as many others in planted. the New Jersey planting. P. thunbergiana x (P. nigra X P. densi- P. nigra X P. thunbergiana. Seedlings were flora). No information is available on seedling slightly larger than P. nigra and trees were performance. Trees were about the same size 10 to 15 percent larger. Survival was 40 per- as P. thunbergiana. cent better. P. densiflora x self. Seedlings were 10 to P. nigra x P. densiflora. Based on seedling 20 percent smaller than crossed P. densiflora. performance, this is one of the best hybrids; By 1967 these trees were 47 percent shorter, however, trees in New Jersey were only 10 60 percent smaller in diameter, and had 30 percent larger than P. nigra. Survival was im- percent less survival than P. densiflora. proved by 60 percent. P. densiflora x P. thunbergiana. Seedlings P. nigra X (P. densiflora x P, thunbergi- were slightly larger than P. densiflora but ana). No seedling information is available on trees were 33 percent shorter and 37 percent this hybrid, but P. nigra X (P. thunbergiana smaller in diameter. x P. densiflora) seedlings were larger than P. P. densiflora x P. sylvestris. Seedlings were nigra. Trees of these two hybrids were about taller than P. densiflora. Trees were slightly the same size, and survival of P. nigra X (P. smaller but survival was equally good. densiflora x P. thunbergiana) was 65 percent P. densiflora x (P. densiflora x P. sylves- better than P. nigra. tris). No information is available on seedling P. nigra x P. tabulaeformis. Seedlings were growth. In New Jersey the trees were 10 to taller than P. nigra, but none was outplanted. 15 percent smaller than P. densiflora, and in P. sylvestris x P. sylvestris. No seedling Massachusetts they were comparable. information. Trees in New Jersey and Massa- P. densiflora x P. nigra. Seedlings were chusetts were about the same size as other P. shorter than P. densiflora, but trees at both syluestris hybrids. locations were about the same size as P. densi- P. sylvestris X P. massoniana. Trees were flora. 14 percent taller and 22 percent larger in P. densiflora x P. massoniana. Seedlings diameter than P. sy lvestris. were taller than P. densiflora. The one surviv- P. syluestris x P. nigra. Seedlings were 25 ing tree was about the same size as P. densi- percent shorter than P. syluestris, but trees in flora. New Jersey were 10 percent taller than P. P. densiflora x (P. densiflora x P. thun- sylvestris. bergiana). No seedling growth information is P. sylvestris X (P. densiflora x P. sylces- available, but trees in Massachusetts were tris). Seedlings were taller than P. syluestris. about the same size as P. densiftora. In New Trees at both locations were approximately Jersey the trees were 17 percent shorter and the same as P. syluestris. 34 percent smaller in diameter. Survival was P. sylvestris X (P. densiflora x P. thunber- about the same. giana). No seedling information is available, (P. densiflora x P. thunbergiana) x P. but the trees in Massachusetts were the same thunbergiana. Trees in New Jersey were 18 size as P. sylvestris. percent shorter and 50 percent smaller in P. banksiana x P. banksiana, P. banksiana diameter than P. densiflora. Survival per- X P. contorta, P. banksiana x (P. contorta centages were similar. x P. ban ksiana). No seedling information is (P. densiflora X P. thunbergiana) x (P. available. Trees in Massachusetts were very densiflora x P. thunbergiana). Trees were 17 similar in height and all very good compared to other hard pine hybrids. Survival was ex- this hybrid was recommended for further cellent. P, banksiana x OP (of questionable evaluation. The trees were 13 percent taller parentage) was 20 percent shorter and 30 per- than P. grifithii, but 30 percent shorter than cent srnaller in diameter than trees of these P. strobus and the reciprocal cross P. strobus three hybrids. x P. grifithii. Information on soft pines was obtained only P, ayacahuite x P. strobus. Seedlings were from the New Jersey plantding. taller than P. strobus and about the same P. strobus x self. No seedling information size as P. strobus x P. ayacahuite. There was is available. Trees u7ere more than 25 percent no P. ayacahuite for comparison but trees of shorter and 50 percent smaller in diameter this hybrid combination u7ere 21 percent than crossed P. strobus. Survival was very shorter and 24 percent srnaller in diameter poor compared to other crosses containing P. than P, strobus. The reciprocal, P, strobus x strobus. P. ayacahuite, was 13 percent taller and 28 P. strobus x P. strobus. Trees were as good percent larger in diameter than P, ayacahuite as any hybrid in the planting. X P. strobus. P. strobus x P. grifithii. Seedlings were P, ayacahuite x P. grifithii. Seedlings were taller and had a larger diameter than P. stro- similar in size to P. ayacahuite x P. strobus bus. This difference was not apparent in trees, and taller than P. strobus. Trees were 20 per- and both the hybrid and P. strobus were the cent smaller than P. strobus and slightly same size. srnaller than P. ayacahuite X P. strobus. Sur- P. strobus x P. (lexilis, Poor seedling vival was the poorest of all P. ayacahuite growth was attributed to a poor pollen parent. hybrids. Trees were still slightly shorter (13 percent) P. ayacahuite x P. flexilis. Seedlings were but this hybrid is not slow growing compared 50 percent srnaller than P. ayacahuite X P. to others in the planting. strobus. Trees were 35 percent shorter and P. strobus x P. koraiensis. The hybridity 23 percent smaller in diameter. Poor growth of this group is questionable. The trees are as may be partially attributed to a poor P. flexi- large as P. strobus, and it is possible that lis pollen parent. this, in fact, is the true parentage of these 14. flexilis X P. grifithii. Seedlings were trees. taller than either P. (lexilis or P. grifithii. No P. strobus x P. ayacahuite. Seedlings were trees of this combillation were found in the taller than P. strobus, but trees were approxi- New Jersey planting; however, the one P. mately the same size. flexilis tree is as large as any other hybrid, P. strobus x P. paruiflora. Seedlings were including those of P. stro bus. shorter than P. strobus. Trees were 39 per- P. peuce x P. strobus. Seedlings were cent shorter and 49 percent smaller in diam- srnaller than P. strobus. There were no trees eter. in the planting; however, P, peuce X 037 is a P. grifithii x self. None of the crosses con- small tree, about one-half the size of P. taining P. grifithii female parents produced strobus, fast-growing trees. Selfing reduced height by 25 percent and diameter by 50 percent. SUMMARY P. grifithii x P. grifithii. This is a small These comments refer only to the species tree in comparison to P. strobus. It may be and hybrids included in the central New Jer- useful in pest resistance work, but it is not sey and northwestern 3lassachusetts plant- suitable for wood production in the New Jer- ings; seed from other sources, species, and hy- sey region. brid combination could possibly outperform or P. grifithii X P. paruiflora. The seedlings underperform this entire group of plants. of this combination were not large and none Pinus resinosa (red pine), P, rigida (pitch was planted. pine), and P. echinata (shortleaf pine) are P. grifithii x P. strsbus. Seedlings were examples of native species that might have taller than either P. stro bus or P. grifithii and been included for comparison puwoses, and many nonnative species were omitted. P, sinensis, which is probably a form of P. Hybrids containing the designation "OP" yunnanensis, did very well in our planting and are of questionable parentage, in some in- should do well in most of the region. Southern stances because the pollen could have come sources of P, tabuEae f ormis apparently grow from several sources including selfing, trees of faster and straighter than northern sources the same species, or trees of a compatible and if this species is as diverse as reported, species. Comments in the Results are, there- parental selection would be worth the effort. fore, limited to hybrids of known parentage, P. yunnanensis, which is a southern species- but all combinations are listed in Tables 2, 4, below 30" N latitude-on mainland China, and 5. apparently does not adversely affect the This group of trees was not produced with growth of hybrid combinations at more north- the intent to study growth rate; in most cases erly latitudes. they were the products of studies to deter- We have not yet found a soft pine that will mine species crossability. The parent trees grow as fast as the native white pine, Pinus were specimens available at the Morris Arbore- strobus, in the Northeast. It appears that the tum and in the surrounding area. They could greatest value of introducing new species or have come from the best or the worst stands using them in hybrid combinations with P. and/or parents. strobus would be to increase resistance to the In spite of these problems, a considerable white pine weevil (Pissodes strobi) or blister amount of information on a large number of rust (Cronartium ribicola) . species and hybrids has been gathered. The The hybrid P. strobus x P. griflithii (syn data from these two locations should provide P. wallichiana A. B. Jacks) looks as good as tree breeders with valuable information on pure P. strobus in our tests. In its native adaptability of these species and hybrids. range, which is restricted to higher elevations A few of the hybrids and species in these in the Himalayan Mountains (below 36" N plantings warrant further investigation. The latitude j, P. grlfithii stands are composed of use of '"elect" rather than "available" par- tall, straight trees obviously unaffected by any ents should result in even better progenies major pests. Progeny tests are currently un- than those represented in this study. derway in New Jersey and Maine to see if Pinus nigra x P. thunbergii and the re- this species has any resistance to the white ciprocal cross looks promising and, because pine weevil. If resistance is observed, hybrids the natural range of both species extends or backcrosses may provide a useful solution above 4.5" N latitude, there is reason to think to the major deterrent to growing soft pine the hybrid would do well in many parts of the in the Northeast. It is already known that Northeast. P. thunbergii is a maritime species, P. griflithii has some resistance to blister rust but it has performed relatively well as a pure and could be used in the North Central species at our two test sites and has done region with P. strobus, and perhaps in the very well in hybrid combination with several West with P. monticola. other species. The other species of soft pine that re- Pinus densiflora extends slightly above 42' sponded well in these trials was Pinus ayaca- N latitude in its native range and in combina- huite. While it did not outgrow P. strobus, tion with both P. nigra and P. sylcestris as this species and hybrids containing P. ayaca- either the male or female parent, it looks very huite grew very well in the region. Consider- promising. Both P. nigra and P. sylvestris are ing that the range of this species does not found at least as far north at 50" N latitude. extend above 20' N latitude in Mexico, we The variation in growth rate and stem form of were surprised by its apparent adaptation to P. sylvestris suggests that the selection of planting sites above 40" N latitude. More in- good parents in this species would yield sub- formation is definitely needed about variation tantial dividends in any hybrid progeny. within the species and its possible pest re- Pinus thunbergii x P. tabulaeformis and x sistance. LITERATURE CITED

Critchfield, W. B., and E. L. Little, Jr. by the Northeastern Forest Experiment Sta- 1966. Geographic distribution of the pines of tion, 1947-1950. U.S. Dep. Agric. For. Serv., the world. U.S. Dept. Agric. For. Serv. Misc. Northeast. For. Exp. Stn. Paper 56. 47 p. Publ. 991. 97 p. Wright, J. W., and W. J. Gabriel. Wright, J. W. 1958. Species hybridization in the hard pines, 1948. Tree breeding at the Arboretum. Morris Series Sylvestres. Silvae Genetics 7:109-115. Arbor. Bull. 4(8):63-64. Wright, J. W. Wright, J. W. 1959. Species hybridization in the white pines. 1953. Summary of tree breeding experiments For. Sci. 5:210-222. Table I.-Washington Crossing, New Jerseydata on trees of section Pinus after combining seedlots of similar parentage from Table 2 Tallest Parentage Number Survival Height Diameter tree 0 8 planted rft) i in) ift j

nigra x nigra nigra x OP nigra X thunb nigra x ithunb x OP) nigra, x densi nigra x idensi x thunb) nigra x sin densi x self densi x densi densi x OP densi x syl densi x syl + OP densi x (densi x syl) densi x nigra densi x mass densi x thunb densi x thunb + OP densi x Cdensi x thunb) (densi x thunb) x thunb (densi x thunb) x OP Cdensi x thunb) x (densi x thunb) tab x OP tab x sin syl x syl syl X OP syl x mass syl x nigra syl x densi + OP syl x (densi x syl) bank x OP bank x (cont x bank) bank x (cont x bank) + OP thunb x self thunb x thunb thunb x thunb + OP thunb x OP thunb x densi thunb x (densi x thunb) thunb x tab thunb x tab + OP thunb x sin thunb x tai thunb x mass thunb x nigra thunb x (nigra x densi) r=, qq~qqqqqqqqggicqqq q PC: ~"j9 qq cq rjqq~?qqqqq~?q rn dririMririri*ddm riM00pi o-l Mri oJ ri rim N mCJmMmMrimriMM

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G coca ~CCCUae c M c-1 cc o wa LC r- ml~di0.1 am w"" r- o CD ma a: OJ CJ CUaV f(M0;O 4 * 4 ri 403 l-i M MCrjM d-10:aC)& C3 0) H Q583 r9 a wwe ~wmPC- CD e ri w CD (DC co c ccm c wo GZZ CD cc LQ a= mZ c ~c. Seedlot Parentage Survival Height Diameter Tallest tree number Q 8 (%> tft) (in) c ft > thunb x self thunb x self thunb x self thunb x self thunb x self thunb x thunb thunb x thunb thunb x thunb thunb x thunb thunb x thunb thunb x thunb thunb x thunb thunb x thunb thunb x thunb + OP thunb x thunb -I- OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x OP thunb x densi thunb x densi thunb x densi thunb x densi thunb x densi qqOOO0 oqoooc OoqocG~Cocc coocc 00 coo 0c;O 0 mr(t-:cncrjtd .d;v&&r-'d &doot-:t-:ujt-..J;cxj &r- loodQ;ct-: &Cj &CjCrj tdcu't-: cd m4riririri mmmmrim mWmmmmdm0-1 rim Noldm WC-l C.tri0-1 riC-lm4

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8g8m00 ~mmm~aommcnim~o~ mm oimao GO mvm oo o mmm mmmmms swmccmmmm mm mmdms rn~rivm ~25 d T-l

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Parentage Nurnber Survival Height Diarneter Tallest tree 9 8 planted eft> (in) v%,) (ft) strobus x self strobus x strobus strobus x strobus + OP strobus x OP strobus x griifithii strobus x griathii + OP strobus x Aexilis strobus x koraiensis strobus x ayacahuite strobus x ayacahuite -t- OP strobus x parviflora grifithii x self grifithii x griftithii griathii x grifithii + OP grifithii x OP grifithii X strobus grifithii x strobus + OP grifithii x koraiensis i- OP ayacahuite x OP ayaeahuite x strobus ayacahuite x strobus + OP ayacahuite x grifithii ayacahuite x flexilis flexilis x flexilis flexilis x OP peuce x OP parviAora x eernbra -t OP =? qqqc qc: ocqqqcqqcc cccoeooci ooqoe qo qcic=:qqqc LO OLnE--* L9C13 800t--Ts;Ct)d'dM3d'~Pf(t-Cl.dlWX;O:0-1t-C33 U=itC3CDV5CXI COW L'363C- 45WOQ rc CC30JiXb;i CC3d dUWrndVdb;ioJG.1W6;1 c.IrnC~o.1dU~~WrimrtmG3r-4 NC-4-i dN&m

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X XXXX XX XXXXXXXXX~XXXXXXXX XXXXX XX XXX XXXX

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lX x X XXX XX xxxxxxx XX Table 5.-Wiliiamstown, Massachusetts-performance of individual seedrots in section Pinus by planting -- Tallest Seedlot Parentage Survival Height Diameter F'orrngL tree number 9 8 C?61 ift) i in) eft>

Nllll nigrax OP N1211 nigra x OP N1009 syl x OP N1033 syl x OP N1128 syl x OP N1215 syl x OP N976 densi x OP N978 densi x OP N980 densi x OP 617 densi x nigra 610 densi x thunb N979 thunb x OP 638 thunb x OP 555 bank x bank 561 bank x bank 563 bank x bank 552 bank x cont 559 bank x cont 562 bank x cont 566 bank x cmt 558 bank x (cont x bank) 565 bank x (cont x bank) 568 bank x icont x bank) 849 cont x OP 850 cont x OP

693 syl x syl 694 syl x syl 698 syl x (densi x syl) 695 syl x idensi x tkmnb) 739 densi x OP 735 densi x (densi x syl) 747 densi x idensi x syl) 737 densi x idensi x thunb) N1261 thunb x OP 705 thunb x thunb 715 thunb x thunb 7 17 thunb x thunb

Continued Tallest Sectdlot Parentage Survival Height Diameter tree number Q 8 ( 721 ift 1 (in) (ft i

102 thunb x t densi x thunb) 707 thunb x idensi x thunb) 7 13 thunb x r densi x thunb) 730 thunb x i densi x thunbi 727 thunb x idensi x thunb) 719 thunb x idensi x thunb) 703 thunb x tab 706 thunb x tab 9 14 thunb x tab 718 tbunb x tab

763 bank x OP 42-353-50 bank x OP 566 bank x (cont x bank) 762 bank x icont x bank) 764 bank x ! cont x bank)

754 nigra x OP N987 nigra x OP N978 densi x OP N980 densi x 0P N1009 syl x OP NlOlO syl x OP N1128 syl x OP N1200 syl X OP "1-best ; 5-poorest. "Largest diameter. APPENDIX List of scientific and common names of species included in this report' Section Strobus Subsection Cembrae P. cembra L. Swiss stone pine P, koraiensis Sieb. & Zucc. Korean pine Subsection Strobi P. ayacahuite Ehrenberb Mexican white pine P. flexilis James Limber pine P. grifithii McClelland Himalayan white pine P. parviflora Sieb. & Zucc. Japanese white pine P. peuce Grisebach. Balkan pine P. strobus L. Eastern white pine

Section Pinus Subsection Pinus P, densiflora Sieb. & Zucc. Japanese red pine P. massoniana Lamb. Masson pine P. nigra Arn. Austrian pine P. resinosa Ait. Norway pine P. sinensis" P. syluestris L. Scots pine P. tabulaeformis Carr. Chinese pine P. taiwanensis Hayata Formosa pine P, thunbergiana Franco. Japanese black pine P. yunnanensis Franchet. Yunnan pine Subsection Australes P. echinata Mill. Shortleaf pine P. rigida Mill. Pitch pine Subsection Contortae P, banksiana Lamb. Jack pine P. contorta Dougl. Lodgepole pine

lcritchfield and Little 1966. Wariety of P. yunnanensis.

U S GOVERNMENT PRINTING OFFICE 1979-603-01117 Head aaaaters of the Northeastern Forest Experiment Station are in Broomal9 , Pa. Field laboratories and research units are maintained at:

@ Beltsville, Maryland. 0 Berea, Kentucky, in cooperation with Berea College. Burlington, Vermont, in cooperation with the University of Vermont. @ Delaware, Ohio. 0 Durham, New Hampshire, in cooperation with the University of New Hampshire. 0 Wamden, Connecticut, in cooperation with Yale University. 0 Kingston, Pennsylvania. @ Morgantown, West Virginia, in cooperation with West Virginia University, Morgantom. 0 Orono, Maine, in cooperation with the University of Maine, Orono. 0 Parsons, West Virginia. 0 Princeton, West Virginia. @ Syracuse, New York, in cooperation with the State University of New York College of Environmental Sciences and Forestry at Syracuse University, Syracuse. @ University Park, Pennsylvania. 0 Warren, Pennsylvania.