PINUS L. Pine by Stanley L

Home , Pinus canariensis, Pinus edulis, Pinus heldreichii, Pinus leiophylla, Pinus merkusii, Pinus peuce

PINAS

Pinaceae-Pine family PINUS L. Pine by Stanley L. Krugman 1 and James L. Jenkinson 2

Growth habit, occurrence, and use.-The ge- Zealand; P. canariensis in North Africa and nus Pinus, one of the largest and most important South Africa; P. cari.bea in South Africa and of the coniferous genera, comprises about 95 Australia; P. halepereszs in South America; P. species and numerous varieties and hybrids. muricata in New Zealand and Australia; P. Pines are widely distributed, mostly in the sgluestris, P, strobus, P. contorta, and P. ni'gra Northern Hemisphere from sea level (Pi'nus in Europe; P. merkusii in Borneo and Java 128, contorta var. contorta) to timberline (P. albi- 152, 169, 266). cantl;i,s). They range from Alaska to Nicaragua, The pines are evergreen trees of various from Scandinavia to North Africa. and from heights,-often very tall but occasionally shrubby Siberia to Sumatra. Some species, such as P. (table 3). Some species, such as P.lnmbertionn, syluestris, are widely distributed-from Scot- P. monticola, P. ponderosa, antd. P. strobtr's, grow land to Siberia-while other species have re- to more than 200 feet tall, while others, as P. stricted natural ranges. Pinus canariensis, for cembroides and P. Ttumila, may not exceed 30 example, is found naturally only on the Canary feet at maturity. Islands, and P. torreyana numbers only a few Pines provide some of the most valuable tim- thousand individuals in two California localities ber and are also widely used to protect water- (table 1) (4e). sheds, to provide habitats for wildlife, and to Forty-one species of pines are native to the construct shelterbelts. The bulk of naval stores United States. These species are also the most still comes from pines, and the seeds of some widely planted in the United States. Planting species are a valuable food source. Increasingly, has extended the range of a number of them, pines are planted to improve man's environ- including P. strobus, P. banksinna, P. rad'inta, ment. P. pond,eroso var. scopulorum, P. resi,nosa, and, Sixty-five species and varieties which are now P. oirginiana (71, 266). Many of the native being planted or which have a potential in the eastern pines, especially those from the southern United States are listed in table 1; included are States. do not do well in the western States. The 40 species and varieties native to the United same appears to be true for many western pines States, one to Mexico, one to the Caribbean when they are planted in the eastern States region, 11 to Europe, Africa and the near east, (120,201). and 12 to Asia. Many introduced pines have been planted and Geographic races and hybrids.-The impor- grown successfully in the United States. Four tance of planting seeds or seedlings from the of these-P. sglaestris, P. thunbergi,ana, P. proper source cannot be stressed to strongly. densiflora, and P. nigra-have become natural- Seed origin is extremely important in determin- ized in parts of New England and the Lake ing the ability of a species to grow and succeed States (261r, 270). in a given environment. Many pines with an Many other pine species have been suce.ess- extensive range, as well as some of limited fully planted outside their native range. These natural range, have developed geographic races include P. patuln in South Africa; P. radinta in that are morphologically and physiologically South Africa, New Zealand, Australia, and distinct (gz). These differences make each race South America ; P. insularzs in East Africa; P. best suited for growing in certain environments. etliottii var. elliottii, P. taedn, P. pinaster, and, As a general rule, seeds from sources in moist P. palustris in South Africa, New Zealand, and regions are smaller and produce faster growing Australia; P. ponderoso in Australia and New and less deeply rooted seedlings than seeds from sources in drier regions. Southern seed sources . 'Timber Management Research, USDA Forest Serv- commonly differ from northern sources by being rce. faster growing, capable of growing longer in a 'Pacific Southwest Forest and Range Exp. Stn. season, more susceptible to damage by winter

598 PINUS freezes, less susceptible to late spring and early ern origins require a longer stratification period autumn frosts; and having seed dormancy that (3 wee[s or more) than seeds of southern origin can often de overcome with a shorter presowing (less than 3 weeks). Seedlings from northern treatment (122, 218, 260, 266). sources tend to be more frost resistant than For many of the pines, detailed data about those from southern sources. These differences geographic rdces are not available. In some among sources appear to be clinal and reflect cases our understanding of races of pines native the latitudinal disliibution of the species (175). to the United States comes from plantings in Pinu,s balfouriana-Seeds of northern origin other countries. When appropriate, this infor- (Lake Mountain, California) are longer than mation is included in the following species sum- those of southern origin (Mineral King, Cali- maries: fornia) and have persistent seed wings com- Pinus attenuata-From morphological and pared to the detachable wings of southern nursery studies, knobcone pine in California can sources (154.). be separated into two major groups, one north Pinus banksiana-Sources differ in seed size, and the other south of the Monterey-San Luis growth, form, and susceptibility to insect and Obispo County line. Seed weight tends to in- disease damage. Seeds tend to be larger from crease from north to south. and seeds of north- the warmer parts of the range (71) . In Minne-

TAsLp l.-Pinus: nomenclature, occutrence, and uses; data comTtilers Data compilers Scientific names and synonyms Common names Occurrence IJses' for the species

P. albicaulic Engelm. whitebark pine Subalpine in the northern T,W,E - R. J. Steinhoff. Rocky Mountains and Coast Mountains of British Columbia through the Cascade Range to the southern Sierra Nevada. P. arietata Engelm. ... bristlecone pine, Small, scattered subalpine H,E G. H, Schubert. P. balfouriana var. foxtail pine, areas in California, aristata (Engelm.) Engelm. hickory pine. Nevada, Utah, Arizona, and New Mexico. P. armandii Franeh. Armandpine Moderate-highelevations T,E P. O. Rudolf. from central and southwestern China to northern Burma and southeastern Tibet; Taiwan and Hainan. P. artcnua'a Lemm. knobcone pine Rocky slopes and ridges w,E S. L. Krugman. P. tuberculatc Gord. from southwestern Oregon and California to northwestern Baja California. P. balfouriana Grev. & Balf. ------foxtail pine, Subalpine in Klamath H, W Do. Balfour pine. Mountains and southern Sierra Nevada, California. P. bankeiana Lamb. jack pine, Canada, from the Mackenzie T, H, W, D. H. Dawson. P. diaaricata (Ait.) Sudw. scrub nine. River and Alberta to S, E. banksiana Nova Scotia and south pine, black into the Great Lakes pine, gray region and New England pine. States. P. brutia Ten. Calabrian pine Crete, Cyprus, and Lebanon S,E S. L. Krugman. P. halepensis vat. brutia north through Turkey. (Ten.) Elwes & Henry P. canariendt C. Smith Canary Island Dry, exposed slopes of the T,W,E P. O. Rudolf. prne, central and western Canary pine. Canary Islands. P. caribaea Morelet Caribbean pine Bahama Islands, western T,E S. L. Krugman. P. bahamensis Griseb. Cuba, Isle of Pines, and P, hondurensis Loock Central America from British Honduras to Nicaragua. 599 PINAS

Tnnln l.-Pi,nus: nomenclature, occu,rrence, and uses; data compilers-Continued

compilers Scientific names and Occurrence Uses' Data synonyms Common names for the species

P. cembra L.- - --- Swiss stone High elevations in the Alps T,W,E P. O. Rudolf. P, montana Lam. pine, cembran and Carpathian Moun- pine, arolla tains of central Europe. prne. P. cembroidet Zucc.- Mexican pinyon, Semiarid mountain regions w, E___-,,,. S, L. Krugman. nut pine, in Mexico. southeastern plnyon. Arizona. southwestern New Mexico. and southwestern Texas. P. clauta (Chapm.) Vasey sand pine, Sandy plains of central and T,H L. Jones. spruce pine. coastal Florida and the southern tip of Alabama. P. con,orta Dougl. var. shore pine, Low elevations on the w,E S. L. Krugman. con,orta lodgepole Pacific Coast from pine, beach southeastern Alaska to pine, coast California. prne. P. contorta vat. latilolia Rocky Mountain Rocky Mountains, from T,H,W,S J. E. Lotan. Engelm. lodgepole pine. Canada to Utah and Colorado, and the Black Hills. P. contorla yat. nrurrayana Sierra Nevada High elevations in the T,H,W S. L. Krugman. (Grev. & Balf.) Engelm. lodgepole pine. Cascade Range-Sierra Nevada chain from southwestern Washing- ton to Baja California. P. couhcri D. Don.. Coulter pine, Coastal mountains from H,E Do. nut pine, central California south big-cone pine. to northern Baja California. P. densiflora Sieb. & Zucc.------Japanese Midelevation in the moun- E S. L. Krugman red pine. tains of Japan and and P. O. Korea north to eastern Rudolf, Manchuria and adjacent u.s.s.R. P. ecfrinnra Mill. shortleaf pine, Coastal plain of south- T,H,W J. M. McGilvray. southern eastern United States to yellow pine. eastern Oklahoma and eastern Texas. P. edulis Engelm. pinyon, Semiarid regions in Utah, T,H,E G. H. Schubert. P. cembroides var. edulis Colorado Colorado, Arizona, and (Engelm.) Voss pinyon pine, New Mexico. nrr+ nina P. elliottii var. densa Little South Florida Southern and central T,E R. L. Barnes. & Dorman. slash pine. Florida, and Lower Florida Keys. P. elliottii Engelm. var. elliottii- slash pine, Coastal Plains from South T,H,E Do. P. caribaea Morelet swamp pine, Carolina to eentral pitch pine, Florida and southeastern yellow slash Louisiana. pine. P. engelntannii Carr. Apache pine, Mountains of southeastern T,E G. H. Schubert. P.Iatifolia Sarg. Arizona Arizona and south- P. apacheca Lemm. longleaf pine. western New Mexico south through the Sierra Madre Occidental, Mexico. P. /e.tifis James limber pine, Subalpine in the Rocky T,W,S,E R. J. Steinhoff. Rocky Mountains and southern Mountain Sierra Nevada. white pine. P. gerardiana Wall. chilgoza pine, Mountains of eastern T,H,E P. O. Rudolf. P. aucklandii Lodd. Gerard pine. AfEhanistan, northern P. chilghoza Ehh. Pakistan. Kashmir. and northern India. P. glnhra Walt. spruce pine, Coastal Plains from South T.H J. P. Barnett, cedar pine. Carolina to northern Florida and west to southeastern Louisiana.

600 PINAS

and, uses; data compilers-Conlinued

t Data compilers Scientific names and synonyms Common names Occurrence IJses for the species

Aleppo pine, Mediterranean region, from T, W' E P. O. Rudolf. P. halependc Mill, - .---. to P . alepensis Poir. Jerusalem Spain and Morocco pine. Turkey and Jordan. P. helilreichii Christ Heldreich pine, High elevations from cen- W' E Do. P. heldreiehii' var. leucodermis Balkan pine, tral Yugoslavia to (Ant.) Markgr. Bosnian pine, northern Greece and in P.leucodermis Ant, graybark pine. southern Italy. P. nigra'rar, Ieucodermis ( Ant. ) Rehd. pine,-, High elevations from east- S' E S. L. Krugman. P. insularis Endl...-.--- . Khasi P. khasaa Royle ern India to South Vietnam. and mountains of northern Luzon, Philippines. Do, Grev. & B?lf_. ----- Jefrrey .. -- Mountains of southwestern T, H, W' E'- P. ietrregi . Pine P. ponderoso' var. jefiregi Oregon, California, (Grev. western Nevada and & Balf.) Engelm. northern Baja California. P. O. Rudolf. P. koraiensis Sieb. & Zttcc'------Korean pine, Mountains from southeast- T eedar Pine. ern Siberia to South Korea; central Honshu, Japan. S. L. Krugman. P.lantbertiano Dougl. sugar pine, Mountains from western T' W pino real. Oregon through Cali- fornia to western Nevada and northern Baja California. W. J. Rietveld. P. leiophylla var. chihuahuana Chihuahua pine, Mountains of central T, E (Engelrn) Shaw yellow pine, Arizona and southwest- Engelm. prno real, ern New Mexico south P, chihuahuana throush the Sierra Madre Occidental in northern Mexico. T, E - S. L. Krugman. P. merkusii Jungh. & de Vriese ----- Merkus pine, Mountains of southeastern . Tenasserim Burma to North and pine. South Vietnam; rvestern Sumatra: Luzon and Mindoro in the PhiliP- pines, H, E A. L. Roe. P. monophvlla Torr. & Fr6m. ------singleleaf Semiarid mountains from W' P. cembroides var, pinyon, nut southeastern Idaho monophglia (Torr. & Fr6m.) pine, pinyon. tbrough the Great Basin Voss rangei to central and soufhern Calif ornia and northern Baja California. R. T. Bingham. P. rrronticolo Dougl. western white Southern British Columbia T' W pine, Idaho to western Montana, white pine, northern Idaho and silver pine. northeastern Oregon; south in the Cascade Ranse. Klamath Moun- tainJ and Sierra Nevada to central California. in the Pyrenees, W' E P. O. Rudolf. P. ntugo Turra- -- Swiss mountain Subalpine P. montana Mill. pine, Mugho Alns. Carpathian and pine, dwarf Baikan Mbuntains of inountain pine. central and southern Europe. E S. L. Krugman. P. muricaaa D. Don bishop pine, Coastal California and H' P. remorata Mason prickle-cone northern Ba.ja California ; pine, Santa Santa Rosa, Santa Cruz' Cruz Island and Cedros Islands. pine. P. O. Rudolf. P. nigra Arnold-, Austrian pine, Southern Europe, northern T' W' S, E P.laricio Poir. black pine, Morocco, SicilY, and European Cyprus. black pine. 601 PINAS

T.qslr l.-Pinus: nomenelature, occztrrence, and uses; data compilers-Continued

Scientific names and synonyms Common names Occunence Uses t

P. palu*ric Mill, ____ longleaf pine, Coastal plains from south- T. H. E J. M. McGilvray. P. australh Michx. f. southein pine, eastern Virginia to cen- longstraw pine tral Florida and west to eastern Texas. P. paruiflora Sieb. & Zucc.-----___---____ Japanese Mountains throughout E -. P. O. Rudolf. P_. pe.ntaphyllc Mayr ivhite pine. .fapan from iofittre"tt F. htnekomatsu Kyushu north to south_ Miyabe and Kudo ern Hokkaido; Korean Island of Utsuryo. P. patula Schiede and Deppe Mexican Eastern Mexico from T, W- S. L. Krugman. weeping pine. Tamaulipas south to Oaxaca. P, peuce Griseb...- -., Balkan pine, High mountains of western T, E. P. O. Rudolf. P. encelsa ltan". peuce Macedonian Eulgaria, southern Yugo- (Griseb.) Beissn. pine, Greek slavia, eastern Albania, stone pine. and northern Greece. P. pinaster Ait.- - maritime pine, Coastal areas in Morocco. T,W,S Do. P, maritima Poir. cluster pine, Portugal, Spain, southern pinaster pine. France, western Italy, Corsica, Sardinia, and northeastern Algeria. P. pinea L. ,- Italian stone Portugal, Spain, and north T,H,S,E Do. pine, umbrella shores of the Mediter- pine, stone ranean Sea to Lebanon: prne. northeastern Turkev: Ibiza,-S;;di;i;;si.iiJ, Crete, and Cyprus. P. ponderotavat, arizonica Arizona pine, Extreme southwestern T,H.W W. J. Rietveld. (Engelm.) Shaw Arizona pon- New Mexico and south- P. arizonica Engelm. derosa pine, eastern Arizona through Arizona the Sierra Madre Occi- yellow pine. dental to Durango. P. pon.derosa Laws. var. ponderota ponderosa pine, Southern British Columbia T,S,E R. J. Boyd. bull pine, south to central ldaho, rock pine, and through the Cascade western Range, Coast Ranges, yellow pine, and Sierra Nevada to blackjack pine. southern California. P. ponilerota var. rcopulorunt Rocky Mountain Rocky Mountains from T, H, W, J. L. VanDeusen. Engelm. ponderosa Montana south to New s, E. prne, western Mexico, Trans-Pecos yellow pine, Texas and northern blackjack pine. Mexico: west to eastern Nevada and east to central Nebraska. P. purnita Regel Japanese Eastern Siberia south tc, S. L. Krugman. P. cembra iat. pum;ta pitl. stone pine, northern Mongolia, Man- dwarf Siberian churia. Korea. and pine. through the Kuril Islands to central Honshu, Japan. P. pungent Lamb. - Table Mountain Appalachian Mountain T,H,W,E S. Little. pine, hickory region from Pennsylvania pine, mountain to northeastern leorgia. pine. P. quodrilolia Parl. Parry pinyon, Semiarid mountains of W,E,H S. L. Krugman. P. cembroides yar. nut pine, extreme southern Cali- parryana Yoss prnyon. fornia and northern Baja California. P. radiata D. Don Monterey pine, Coastal central California T,S,E Do. P. insignis Dougl. rad.iata pine, and northern Guadaluoe rnsrgnrs prne. Island, Mexico. P. reeinosa Ait. red pine, Great Lakes region from T, H, W, D. H. Dawson. Norway pine, southeastern Manitoba E, S. hard pine, and Minnesota east to pitch pine. Newfoundland, Nova Scotia and New Jersey.

602 PINAS

TASLp L-Pinus: yromenclature, occurrence, and uses; data compilers-Continued Data compilers Scientific names and synonyms Common names Occurrence Uses' for the species

P. risida Mill. pitch pine, Northeastern United States T, H, W' E S. Little. hard pine, and the Appalachian bull pine. Mountain region, from Lake Ontario and Maine to western KentuckY and northern Georgia. P. roxburghii Sarg.. Chir pine, Monsoon belt of the outer T, S, E P. O. Rudolf. P.Iongifolia Roxb. longleaf Himalayas, from north- Indian pine, eastern West Pakistan to Bhutan. P. rabiniana Dougl. Digger pine, Low-elevation dry sites T' H S. L. Krugman. gray prne. in the Coast Ranges and Sierra Nevada, California. P. eerotina Michx. pond pine, Coastal Plains from south- T B. Benson. P . rigida var. serotina marsh pine. ern New JerseY to centrai (Michx.) Loud. Florida and west to central Alabama. P, sibirica Du Tour Siberian Ural Mountains east T, E P. O. Rudolf. P. cembra var. sibirica stone pine. through western and Loud. central Siberia to northern Mongolia. of southwestern T, H, W W. J. Rietveld. P. urobilormis Engelm. . . . southwestern Mountains P. flerilis var. refl'era Engelm. white pine, Colorado, Arizona, and P. reflera (Engelm.) Engelm. border limber New Mexico south in the pine, Mexican Sierra Madre Occidental white pine. and Oriental to central Mexico, P. ilrobus L. eastern white Newfoundland and Nova T, H, W, R. E. Graber. pine, northern Scotia west to south- S, E. white pine, eastern Manitoba, south white pine, throuEh the Great soft pine, Lakes and Appalachian Weyinouth Mountain regions to pine. Iowa, western KentuckY, and northern Georgia. R. S. Walters. P, ryhtestria L. Scotch pine, Eurasia, from Scotland and T,S,E Scots pine. Spain to Finland and Tirrkev. across U. S.S.R. to the Sea of Okhotsk and Manchuria. P. taeda L. loblolly pine, Coastal Plains and Pied- T,H,W B. F. Mclemore. oldfield pine. mont from Delaware to central Florida to eastern Texas, southern Arkansas, and southern Tennessee. P. thunbergiana Franco Japanese Maritime in South Korea T,W,S,E P. O. Rudolf. P. tlrunbergiiParl. black pine. and in Japan from northern Honshu south to northern Ryukyu Islands. S. L. Krugman. P. toneyana Parry-- - -.. Torrey pine, Santa Rosa Island and E Soledad pine, low coastal bluffs in San Del Mar pine. Diego Co., California. S. Little. P. drginianaMill. Virginia pine, Aooalachian'Piedmont Mountain and T,W scrub pine, regions, from Jersey pine. LonE Island south and wesf to central Alabama, v'estern Tennessee, and southern Indiana. P. wallichiana A. B. Jacks. blue pine, Mid- and hiEh-elevations T,E P. O. Rudolf. P. excelsaWall. Bhutan pine, in the Himalayas, u'est P. g rifr.tlLii McClelland Himalayan to eastern Afghanistan P. nepalensis de Chambr. pine. and east to southern China and northern Burma. forestry' 1 T: timber production, H: habitat or food for wildlife, W: watershed, S: shelterbelt, E: environmental 603 PINAS

sota there is a change in cone serotiny from Mendocino White Plains-and has serotinous closed cones in the north to predominantiy open cones like those of the Rocky Mountain variety, cones in the south (19f). Winter colorafion of var. Iatifolia. "Ihe other varieties have cones needles is less in seedlings from lower latitudes which open at or soon after maturity; the open tlran in those from higher latitudes (ZeZ). In cones of var. contorfo persist indefinitely while Canada, height growth was greater for sources those of var. lnumo,yana do not (lL\.In tests in from areas with longer growing seasons; selec- northern Europe, seedlings of var. contorto tions moved north made bettei treigtrt- growth grew faster, and were more branchy and less than selections moved south (i04). winter hardy than those of var. lntifolia from Pi,tr,us brutia-several varieties have been interior British Columbia and the Rocky Moun- described in the Black Sea region. The var. tains (6y'). The var. mul.raAarLa has the largest pitltyusa is found along the nortliern and north- seeds. At temperatures of 50' to 68" F. the var. eastern shores of the Black Sea. and var. eI- Iatifolta germinates twice as fast as coastal darica is found in the central Transcaucasus sources (44). (152).In northern California, an Afghanistan Differences between sources also are found source related to var. eldarica is out-growing within a given variety. Seeds from high eleva- var. pithyusa and appears to be both fiost and tions in central British Columbia germinated drought hardy and of good form (90, 1 ZZ) . faster at 68' F. than at other temperatures, and Pinus canariensis-This pine is found natu- seed from low elevations germinated faster at rally only in the Canary Islands where it ranges temperatures above 68'F. (8/). Commonly, the from-.2,100 to 7,200 feet above sea level (15-Z). southern sources grow faster than the northern Seedlings of several sources from the various sources of a given variety. islands and elevations, when exposed to low Pintts cot.tlteri-This pine is endemic to Cali- winter temperatures in the Institute of Forest fornia and Baja California. No races have been Genetics' nursery at Placerville, California, described, even though it grows in isolated showed marked differences in cold hardiness. stands from 500 to 7,000 feet and on fertile to Seedlings from 4,000 feet showed more cold very poor soil types. Mount Diablo sources are damage than seedlings from 6,200 feet on considered to have the poorest form, with Tenerife Island. Seedlings from 6,200 feet on .qreater branching than any other source (Z7g\. Palma Island were badly damaged, suggesting Pinus densiflora-A number of cultivars have that sources from different islands rn'iy als6 been described (150) . This pine is widely differ in susceptibility to low temperature"(122). planted and hardy in the Lake States, New Pintts caribaea-Three geographic variants England, and southern Ontario (Z6t+). are recognized. The var. caribaeaiative to Cuba Pinus echin,o,ta-Ten-year results in a range- and the Isle of Pines has persistent seed wings, wide test suggest that in the southern Uniied while the other two varie[ies do not. In testsln States the southernmost sources from east of South Africa, var. caribaea showed better the Mississippi River are superior to northern growth than var. hondut'ensis from the mainland sources in both survival and growth (260). of Central America. The var. hondur.ensts has Seeds from northern sources should be used in the largest seeds; var. bahamensis from the the northern extremities of the range (1llr, Bahama Islands has the smallest (150). 260, 262). In an Oklahoma test, an Arkansas Pinus cembro-A number of cultivars of this seed source showed the best performance, even spe_cies have been reported (51, 1g0). surpassing a local source (189\. Pitttts cembroides-Two varieties have been Pinus edulis-A single leaf variety, var. described. The var. remota found on the fallar, is found in the mountains at 6,0b0 feet Edwards Plateau in southwestern Texas has in central and eastern Arizona, very and in the Grand thin seedcoats. The var. bicolor- is found in Canyon and parts of New Mexico. Its seeds tend southwestern New Mexico and southeastern to be larger (137\. and have a thicker seedcoat than Arizona the more common variety, var. edulis. In con- Phtus clattsa-ln central Florida, this small trast to var. edulis, var. fallar seldom produces pine has closed cones. But in western Florida it seeds in quantity (1Sf). has an open-cone variety, vat. immuoinata- Pinus elliottii-Two varieties are recognized. Choctawhatchee sand pine, whose cones"ripen in The var. densa is found only in south Florida. September and shed seeds during October (es, Compared to typical slash pine, var. elliottii, the 138). var., germinates Pintts dettsa faster, has a grasslike cotttorta-Four varieties are recog- seedling stage with crowded needles. and has only Lized'r'he -thouSh three are listed in the table"s. heavy wood with very wide summer rings (lgg, fourth, var. bolanderi, has a restricted 110, 216). Sources range of var. elliottii in north- on the northern California coast-the eastern Florida appear to be the poorest, as they 604 PINUS are susceptible to ice damage and are less origins (217). Seeds from northern Idaho are drought resistant than northern and western il;tt"" ihan'those from washington and cali- sources (216,260). fornia---Pir*; (295). Pinus flerilis-seedlingq from--- southern mugo-A number of horticultural stands are'faster growinglhan seedlings from varieties hav6 been described. They vary from a northern stands (219). snrawlins shrub, vat' pumilio, to a small tree' Pi,nus halepensis-In Israel at least two ele- v-ai. iostFata Q8:D. Varieties also diffe-r in seed and others can size and germination capacity (1p7\.-Seedlings vational ecotypes are recognized, high be expected in other parts of the range (1I8, of io*-"t6uation origins-are not hardy at 152). elevations (256). Pinus heldreichii-Pinus tt'eldreichid' var. Pinus muricata-Trees from origins north a timber- and south of Fort Ross, California, differ in leucodermis, considered by sorne to be I1 line tree, commonly is found on drier sites and g"o*ttt form, foliage color, and cone shape' often on soils formed on limestones (51). The triito*i", tiees fiom northern sources tend var. heldreichiiforms open forests in the moun- to grow larger with fuller and more com-pact tains at elevations between 3,000 and 5,000 feet (a-r )l In tests in Australia, the northern (180). ""oiunssouices maintained better growth rate and form ' tests in northern Rho- than the southern sources (69). Pi,nus insulnris-In and desia, trees from seed of Philippine origin were ptnus nigru-Several distinct varieties more vigorous and had better form than those numerous cultivars are recognized. The var' from India, Burma, or Vietnam origins (152, iiia*iin" from the Crimea, Turkey, and 2oo). Cyprus tends to have the largest seeds (17-,500 Pinu.s ieffreui-Evidence for distinct ceq to'ZO,SOO per pound) ; the var. c,ors,i9a.na frorn graphic rices ii lacking, but differences in seed Corsica tras ttr6 smallest seeds (28,000 to 36'000 trees from per pound) (152, 187). Wood in the Corsican origin are still important. Generally, in seed collected east of the Sierra Nevada are irari6tv is' considerably better than -that slower growing, more drought resistant, and typicai Austrian pine, var. austrinca, !.rom the less susceptible to cold damage (85). Also trees ddtt peninsula- and eastern Alps. Stands of from high-elevation sources tend to be slower iir. iitiarfco"n planted in Belgium are considered growing than those from lower elevations (93) ' one of the more cold hardy varieties. Other Siberia, China, and varieties are var. cebennensis from the Pyrenees Pinus koraiensis-The hispanica ftom Spain, Korea sources most likely should be considered in southern France, vat' geographic that in Japan. and var. mauritanica from Morocco and Algeria. a race distinct from general can be made for dis- Some statements Pinus leiophylla var. chihunhunnn-Two various geographic sources (152): tinct forms have been found' One has poor form with a short, crooked bole and many branches; Western Groups: Sources in France and the other has good form and grows to heights Spain are often proven to be 4rought re- of 80 feet (152), siitant and are indifferent to soil type. Pinu.s merkusii-At least two distinct races Central Groups:-and Sources in Corsica and Italy are recognized, one on the Asian mainland and grow well have good form. But-they the other on the island of Sumatra. Seeds of need a high humidity and grow poorly on mainland origin are larger than those from limestone soils. Sumatra. Trees from mainland origins pass Eastern Group;: Sources in the Balkan and through a grasslike stage and tend to be uni- Crimea are6s appear to do well on the nodal with a straight, cylindrical bole, but thgy poorer limestone soils. do not grow as tall as those from Sumatra. The Pinus palustris-Seedlings of different geo- Sumatra trees may reach 200 feet in height and graphic origin differ in height, growth, cold tend to be sinuous in gron'th (40)' Some con- iesi-stance, and survival (71, 260'). Southeastern sider the two races as separate species; P. mer' and central Louisiana sources did poorly; south- kusii on Sumatra, and P. merkusiana, on the ern Florida sources failed outside of their area Asian mainland (41). of origin. Central Gulf Coast sources grow well Pinus monophglla-No varieties are recog- throughout the Gulf Coast (260). nized, but there are variants which have partly Piius p ara ifl or a-Several horticultural f orms or mostly two needles in a fascicle, rather than are cultivated, (123) . Pinu,s paraifl'ora is thought just one (137). to consist of two geographical varieties which (49). P inus m o ntic ol,o-Local vari ati ons associated intergrade in central Honshu, Japan with elevation and site are recognized. Pro- Piitrs patula-Because of its rapid growth, genies from high-elevation origins grow faster this species has become one of the most im- at high elevation than those from low-elevation nortant sources of wood in the summer rainfall 605 PINUS

areas of South Africa (148).It also grows well Sierra Nevada are faster growing but less frost in East Africa, New Zealand, and Australia resistant than sources east of the crest (122). (128, 152). A variety, var. longepeduneulata, Closely related to P. ponderosa in appearance from the States of Oaxaca and Chiapas, Mexico, is P. toasltoensfs. This rare pine, which is often has been reported. The cones of this variety wrongly identified as P. ponderosa var. ponder- open quickly at maturity, thus differing from oso, is known to be in only three areas: east var. patuln which has closed cones. The seeds of slope of Mount Rose, Nevada; southern Warner var. Iongepedunculata are black with brown Mountains, northeast California; and Bald marks, while those of var. patuln are pure black Mountain Range in the eastern Sierra Nevada. (1 t8) . Both its male and developing, second-year fe- Pinus peuce-Il Europe seeds from the Rila male cones are purple to purplish black, unlike and Pirin Mountains in Bulgaria are considered ponderosa pine. Washoe pine flowers in July; to produce the best trees (77I). its cones mature in August and September and Pinus pinaster-This species is highly vari- open in September. Cones and seeds are handled in the same manner as var. ponderosa, and its able, and at Ieast four main races are recognized germinate (152). The French or Atlantic race, which is seeds promptly after a 60-day cold itself quite variable, stratification (47, 122). is commonly found on the ptmgens-Seed coastal sands. The Portuguese race is also found Pinus weight, cone length and on coastal sands but is superior to the French width appear to decrease as elevation increases race in growth, form, and drought resistance; and as latitude decreases. Cone serotiny is most has done common in the southern part of the range. No it well in tests in South Africa and (27[t). western Australia, and appears to have some distinct varieties are recognized seed dormancy (105, 266). The Corsican race is Pinus radinta-Monterey pine occurs natu- most commonly rally in four relatively small and well separated found in the mountains. The populations. Moroccan race shows differences between moun- The var. binata occurs on San tain and near coastal origins, Guadalupe Island and is slower growing than in that sources of populations (Monterey, mountain origin fail when planted on the coast. the three mainland The mountain sources are thought to be frost Cambria, and Afro Nuevo Point). The Cambria resistant (152). population has the largest corr€s orrd seeds, and Pittus ponderoso-The the Monterey population has the smallest (70). three main varieties In tests in Australia, trees of origin are var. ponderosa, var. arizoniea, and var. Cambria seopulort+m. did not do as well as those from the Afro Nuevo These varieties differ in seed and Point and Monterey populations (69). cone size, needle length and number per fascicle, germination, P'intts 1'psi11ssa-lhis species is considered speed of rate of growth, form, re- one pines, sistance of the least variable of the and no to drought and low temperatures, and subspecies or varieties are recognized, (74,266). survival (31, 71, 218, 257, 259). Within the varieties However, some height growth, form, and wood there are also source differences in quality differences may among populations some of these characteristics. germina- exist Speed of in the Lake States, New England States, and tion at different temperatures was found to West vary seeds Virginia. In northern sources, seeds are for from the Pacific Northwest, often smaller, lammas frequency is generally Rocky Mountains, and the Southwest. South- less, and west sources frost resistance is higher than in reached maximum germination at southern sources (74). the lowest temperature (91).In tests in Oregon Pinus rigida-No distinct geographic sources and Washington, growth rate generally in- are known, creased but there are variations in form and with seed origins from east to west, and development between populations. Throughout from south to north in the eastern part of the most of the range, range; slowest grou'th trees consistently open cones was shown by sources and shed seeds soon after maturity; in southern from eastern and southeastern parts of the New range (218\. Jersey the vast majority bear cones that In tests in California, growth rate remain closed at maturity and only open at generally increased as seed source elevation irregular (6,71). decreased (33). intervals The latter trees are In tests in northern Arizona, characteristic of areas with a history of wild eastern and southeastern sources did well, but (71). northern fires and western sources failed-as did Pittus sabin,iana-Seeds from populations in the southernmost source (195\. Northern mild climate areas germinated quickly sources of var. scopttlorum more are characterized after stratification than those from the colder by a relatively good growth rate and frost re- areas. sistance (257). Seedlings of southern origin grew longer Southern sources of this variety than those of northern origin. are slower growing Larger cones were but frost resistant. In Cali- more frequent in the northwest part of the fornia, sources of var. ponderosa west of the range than in the Sierra Nevada (82). 606 PINUS

Pinus serotina-Cone serotiny is greater in localities, introduced sources have produced southern and coastal populations than in north- better trees than local sources' ern and Piedmont populations (211t)' Pi,nus strobus-Seeds from the western part the P inus si,b i.r i c a-Di sti n ct diff erence s in gr owth of the range are lighter than seeds from rate, branching habit and fat content of seeds eastern paft, and seeds of southern origin re- exist between certain populations. No varieties qui." u iottg"" stratification period than those are recognized, but several forms have been of northern origin (72, 162, 264). Field tests described. Pinus sibiri,cu f. humistrata is a fru"" A"-onstrafed that southern sources, such dwarf form that grows on mountain summits u* tftot" from the southern Appalachians, tend P'itttts si'biricaf. coronans has a wide io gro* faster and to continue shoot elongation and ridges. (73, and dense crown, is reasonably drought resist- lon"ger in the fall than do northern sources ant, and is found from sea level to 6,600 feet' iosi. ln artificial freezing studies and field Pimts si,birica f . turf osa grows on peat (185). observations in the northern Lake States, south- Pinus syluestris-Scotch pine, the most ern sources are more sensitive to low tempera- eastern widely plan-ted introduced species i1 tlre United tures (162,'had'blueigreen 196). In fall, seedlings of Statei, js probably the most intensively studied oiigin foliage compared to the of all pin-es. The first cornparative pine seed yellow-green foliage of northwestern sources source trials involved this species. It is the pine (267).' with the greatest natural range' and it grows Piitu,s foeda-Numerous field studies have in many different ecological situations (266, demonstrated definite geographic variations in 268). Numerous varieties, forms, and ecotypes growth rate, drought and cold hardiness, dis-ease have been described. A conservative estimate of iesistance, and survival. Seedlings of Maryland the number of geographic varieties ranges from origin tend to grorv less than those of other 21 to 52 (268). There is also considerable varia- oriEin when pianted in different locations tion within named varieties. Sources differ in (260,261). Soulhern sources outgrow northern many characteristics including seed size, ger- .ou"".. in South Africa (210).In many of the mination, dormancy, seed and cone color, tree tests, local seed sources were best. Sources west form, growth rate, structure of root system, of the Mississippi River are more drought and flowering characteristics, needle color, and sus- disease resistant than most of the eastern ceptibility to heat, cold, or drought (28, 18-6, sources. And southern sources tend to be more gZ-0, ZOO, 263). Seed size decreases from the susceptible to low temperature damage than south (44,200 per pound in Spain) to the north northern sources (260). (101). formed trees (127,000 per pbund-decrease in Lapland) Growth Pintts thunbergiana-The better rate tends to from southern to north- are from inland sources (1/+5). ern sources. In tests in Sweden, southern Pi,nus torreyan,a-In a common planting on sources grew faster and later in the autumn the California mainland, trees of mainland than did northern sources (117) . Similar results origin grew taller and had a single trunk, while were found in Michigan tests, where certain trees from a Santa Rosa Island source were French sources grew three times as tall as slower growing, bushy, and freely branched. northern Finnish and northern Siberian sources The islald souice produced larger cones (86). (266, 268). The more southern sources, however, to low temperatures than Numerous pine hybrids have been described. were more susceptible least 200 first- northern sources. Under Canadian prairie condi- As a conservative estimate, at tions, Russian and Finnish sources survived generation and second-generation hybrids as sources (tnZ). tn well as backcrosses, crosses between varieties better than more southerly three Michigan, needles of Spanish, southern France, of the same species, and crosses involving Balkans, and Asia Minor origins remained or more diffeient species either occur naturally of or have been produced artificially (122,1/t1, 169, green during the winter, while those Siberian provide yield and Scandinavian origins turned yellow (266, 266). No attempt is made here to 268).In tests conducted in Sweden, seed origin statistics for the numerous hybrids, since such germination rate at a given tempera- data are highlv variable. The number of sound influenced and indi- ture. Sources from northern latitudes or high seeds produced depends on the species germinated well over a wider range viduaitrees involved, as well as on the environ- elevations is made. of temperatures than those of southern latitudes mental conditions under which the cross or low elevations (117). Sources in the extreme Some natural hybrids are relatively common; northern parts of the range and certain sources e.g., P. palustris I taeda, known as Sonder- from Turkey and Greece show the greatest seed egger pine, which occurs in Louisiana and else- dormancy (101). Lack of a fully developed em- where in the South (36), and P' contorta y bryo accounts for part of the dormancy of bartksiann, in central Alberta and southwestern northern sources (116). In some European Mackenzie (272). Most other hybrids occasion- 607 PINAS

ally occur where the two parent species are torta; in others they do not form until the trees naturally associated. Examples of this in Cali- are much olCer-e.g., P. lnmbertiana and. P, fornia are P. ponderosa y jeffreEi, P. jeffreyi resinosa (table 3). Pines are monoecious with 7 cotilteri, P. radiata )( attenuata (48, 169). male (microsporangiate) and female (mega- In the South are P. taeda X echinata in east sporangiate) strobili borne separately on the Texas, and P. taeda y seroti,na throughout their same tree. Male strobili predominate on the common range (tnS, 27L, 266). The hybrid basal part of new shoots, mostly on older lateral between P. densiflora and P. thmtberginna branches in the lower crown. Female strobili occurs naturally in Japan but has also been are found most often in the upper crown, pri- produced spontaneously in plantations of these marily at the apical end of the main branches in species in Michigan (26.1). In Europe, P. syl- the position of subterminal or lateral buds. But aestri.s occasionally crosses naturally with both frequent exceptions will be found to this general P. nigra and P. mltgo where these species are scheme. For example, P. banksiana, P. clausa, planted near one another (266). These are but P. rad'iata, or P. attenttata are multinodal in the a few of the many hybrids which have been bud, and female strobili are found occasionally reported to occur naturally. at a secondary whorl position (71, 122). Pintn Several pine hybrids have been produced in attenuata, P. radiata, and P. airginiana fre- relatively large numbers by controlled pollina- quently produce female strobili in all parts of tion methods. These include P. rigida \ taeda, the crown (71,122). In temperate climates, the which is an important hybrid in the reforesta- earliest stages of male and female strobili can tion program in Korea, and P. attenuata y be detected in the developing buds during the rad'iata, which is being planted in California and summer or fall; the male develops l" to several Oregon (83, 106). Many other hybrids are being weeks before the female strobilus (76, 122, 169) . produced in smaller numbers and tested for Male and female strobili of the southern and their suitability in various parts of the United tropical pines emerge from buds in late winter; States. e.g. P. elliottii var. densa, P. elli,ottii var. el- Flowering and fruiting.-In certain species, Iiottii, P. glabra, and P. palustris. Strobili of reproductive structures are first formed when other pines emerge from the bud in early spring, the trees are only 5 to 10 years old-e.g., P. or late spring and early summer (table 2). The atten.uata, P. banksiana, P. clausa, and P. con- male strobili are arranged in indistinct spirals

Tasl,n 2.-Pinus: phenology of flotuering andfruiting

Species Location Flowering Cone ripening Seed dispersal Data dates dates dates source P. albicaulis--. ,,,- California - JulY August-September not shed lqQ qt.n P. aristata Arizona July-August September-October September-October 181, 206 P. armandii--, .---.-.. California , April-May- ,-- August August-September - 100 P. attenuata------, do April January-February--- closed cone', - .- , 100 P. balfouriana-, do July-August September-October September-October 1 00 00E P. banksiana ,- , Lake States May-June September September P.bru,tia ., ., California March-May January-March closed cone t oo P.canariensis-,.--,-. do April-May September September-October 100 P. caribaea Cuba - January-Feb- July-August Septdmber 150 ruary. P. cembra -. Germanv May August-October not shed 194 P, cembroides . Califorriia May-June November-December November-December- 122 P. clausa Florida September-De- September September " 29, 235 cember. P. contorta vat. contorta California May-June September-October Fall' 122,235 oary var. Iatifolia Rocky Moun- June-July August-September September-October " tains. YAT, mUrrA,AAnd, Caliiornia MayJune September-October do l qo P. coulteri do do August-September- October n 122,235 P. densiflora do Anril do September-October 122 P, ecltinata., South Carolina M-arch-April October-November October-November P. edulis Arizona June September September-October 132,133,13/+ P. elliottii vat. densa Southern January-April August-September September-November 30 Florida. var. elliottii Florida January-Feb- September-October- October 59 ruary. P.engelmannii, Arizona May November-December November-February- - 62,235 P. flerilis California, June-July August-September . September-October, 122 Montana.

508 PINUS

o h \n SN lo sr F. qsl \t 6r s E a srGas\.t Sco -9rS(oor*Scn \oo \sl oo\ AStSq)ooqiqclcosnSoosloo\\\\\\stoo \\ \\\.8 \ N G\r .Cr \o s - q s ortosi s s Gt '+\tb st ct GaGa \sl G{ \ :\ro\\ q oo -+cec! oa oo sl \or6ci sl srNE \ ^ct \E k)\ ea N ro\o\rov) \ \ \st\ \ \ B Gast\ \ \\k) \i \o\ , ltr i tr ,F }-c i> ta .. k E .E a s t{ FF -B 3 i =s f: + E .P = €B+ lr rE.E.iET,!t€!E€: i.;:E i fi i'S i6o t OF* iiEH?.€ Qao BF a ;! iE:i i a;*,EEl 3, >H- EF:a s; i; :aEa v) flE ;f;EUf E:s.E;il.* ab q !; Fgi,XBEp:6fE.f FE? ;iiEi.Escr?E;iEo.tlEsi? _l: ;s 3* -o,a i : a) k 3p5'eLo.dFr3€Eiii iSiB 4E6a ar iiliHEIi;in 5Ei Er F Q.! l.e A ;#si t;3f;* i=.E.Et #:Er:E!=5 p:* F FF" #g -'-- n lr =Ei ;: € : t=E'iE,"'i E"B€; 4 to i : tu= >, 's.\ -a;i +) o) .2 1 ri. lro_a E q)9EI Y o ^ts{ o g,a 15 o r I r P o. 'lr I E !^o - ! aa a) -a : ; *a tr -c ! c c * '=/ : A^^ q) ;^ Ud P iib 6 € idli6^^'Ex- i^ ti o O.O O o q) O o I O qr O olqJ o H-o dh !tr a . !! F ! ! |l lr a) H A. A^A^ r^ A^ b0€

I s -l .* \o cn HFIsH ca qi st (o q-txlAr G.t \ C.TNN -+6r N q6l\r h\ G{\\O \ H: Nao S) Ge E a-\O qQ \ N tiNN q \ oi .+rO N ra: cC \ \ b- 1..\ \\ f.- \ \\Sa \ N \ sistF \hstsl c3l q qvl Y P r6N i (o

-+ sa \o c) \o € ts s\o O\orosfoAAoJ G! a! 06 q$ocqtooa:gJE st r€ q \6t \6fN-\\\ I b \t - S N c) sts -+q^S Gr c)- q.s oo cr qsa$-to\o s s to gb.+co oJ a s\q sl ai oo\ Gt q a! .+stst a: a! Gn $!i\ ot OJ\\ Bq \ \ ca\ H \\ \- -\ ^\ \ -\--\\ \ tsal- - \\ CA -Gas-\tso--6t-slo..-qosloo- G6 \O \SQ qJ \ NoO\OoOB B Sf qetq \ \ \ :+qsi G\ \ -\ B\6t s.F 0) ktN \6\ \ N NHE\\Q N \ E\\ \o N N NA\ E \o\tb) \a \aJ \

a qOO O@ O tO o@ s LC66.l6rl6ii ic\ cn I i | 6t ca c\ t'- rico !$ iilc\l tJlrol ttl : F > c\t H 6.1 c\ c\ 6.1 6\ l

.l 61 il rot- 6.1 fl* (o<> N I(i co c\

Fl € + q!? N!? o Ooots-c.l @ <1 @O l-- 0O @O)c0 Oroco * Fl I l-Hdill .. hts ^,Hl lliil Hi I ti I tltl d lo llrJ h | | lc.n u) o llo | | ttl r- oN oi c6 h ro ocai rNrooc60co H C{O!O 6.1 ro @ ro c{to r @

tdB .j .sS' .c'e i a g S oiS *Fo 3 b !E s ;is €a 3t F :: :o *,F S EoS a irl HlE FFS * E 3s€is € dLQ, Eis$' ,o .nF Ss!.: !:ds s .c i*s s s * rlrrt cddP S $:B Eb E € S SS333 *Pc!!> S ! E*rrPvv>>w \ s5e{s.-IE a.: o. o" o.tua"a"tu && & \ * o"tu a. o" o"a"a.d 610 PINAS

c)ssq) P R R3 R s\dsc6dser\odld\aqs oo stsl $ co a! \a c) ao S st \ er .+ \o S oo 9 oo 9f q or \ stsa q.t n \ si oo \ oo st oj \ sf \ oo \ co \ ea \ \ B\ \ \ \ \ \ - \ ct- \ S S sr s q)oo N- - \sl qJ S\ 6 q si oa oi.+N- si \ \ 6i €co 6t \ Gr sr qJsa o! oo q \O\ \ \tGt \ \ \ \ \\\ B lO \a \\i \ \\ \ \r \ 5N \ Sl\ \ \ \

;. Er;,e' E E f,€ l;iF ;,;iii: i i :E i€;Ef ir E,:i,E,,i,i,i i ' '6 e;gg ;g'ii;€i'i'iiai$;g'; ; €tu;iial:; ;:iiiE€;a€*i;Ei:r:

oFlF3"€F oo o o B9 g E gi 6 Eh F : eio YV : Y 3 .c 3l-E !e $; tZ; Pe .2 tr o +AAFFIFFIIFF €p.i'le +-B x:x=:6E€€;3;;:E Ei' o 'rr € aS €S **? oF 'os kktr.:Lo, trt ! !8frd.8Eae-bg k! o L " 8E-Ek! ! tub0tu>bo b! 0.= b0 bo tub4 .d b! b!tu br

(o oo \o GT st \o-NN-+vJ \ -+ \a sJ o:Ct A \ ch sa.. s6 \ 6t N \o 0o\ GN .+ slG55 .+ G.6st \ \ q $St ts -+ct Sl \o \ \ st q\oE sr si \E \ s)\ N \ \

\o ui <4 6t 6J t-- m 16 n*t'- O)

\o ro Ga\ oo cbs a sr o: lo-ro st roa e6oO-\Acnc) ^\ G! vJ() \O c) Nojsc!\OqS 9) V) Gt Gbq qJ \ \ro - Gt\ ct B 9r^\qG-rsf\q N - \ - ro -sl \ S .\ \oss\o q (o s - s a \o a\os s ^ - er q6qSq\t6to:qG,) S\ \ S9! q qr a: st g)\q ao e, \ OO\ Gt \ A! \ aOGt 9) c) Gf or-^\\N\\-\-\\(o \V! \ . E\ B \ \S (o ts S .oo - q) \ . qQ \o<)-s6\ -qr co tsN \ \ q oO.$\ \ \ ao 9r \ -+oO Sa slsa\.+\\\\q Sr\ \ \OG\f -+^ \O \O \ cO\\O laj 16 B \ \i 6a\ \ ST E N \ TOVJ N KJ \

oro @ oo t- |rJ al c\ o .L -! 1l)0 .dl h d<{Nic!iiHH N iilN

Hr oo i o @ o r H m fl-3E co t-

Ioc\I @ Oro o

q) \srooo t sr\stct-qce$ st st \s()q^s a -ooqlqrsoo ro-B\\\\oo\ -(o\os-sa ooof\\BN\a\ob$co\st

0) o d *r i- € tr ; q3,:E E r* a; A € ! tr triE€ x s? (J ii,; a I q I :ieE L ; liiFc s i: a) t S ir5;E?*:;s*ttg: I " g"oE CA E'E'= E" ,E s c6 a) q) \ 'aii Ocd -! ,rE .s @: O+i o ar_= L tr o t3 6 € d^ p.E6': o € !o!!o o Ro > o b0'trtuhD

>1 a 0) a) ts Cr jrOBi kt.$c! a) o- a S\}\ sn G\ Gn ooftJ\ lGtstst d;

h& ( ce :dc\l !Fi 3c 9 L I I sf + j H F t3q) q

a) I ( R rl trI b0 I 9!Ss) dl ( a \9)\ b0 l -B-\oGn I s6.\60raq5 Ir q6B.+G.rciN !l I EENts.\ a) ol.dl ooqi\oro\N\€\+\EoOttc) I €l

a) EI v) al +O rO O O € rO O SH H d I S; t J J J ; \F

a) l'

I I :N co d :6.r d i il -'rtl lro |\6 o o oc\l @@o) bh ,1 E

Fi .d. 'aN B*: o .? 'F'F i ! o iitaY a "F{PSdsis'€ c €F g S *:)d"Co':aqq$$ee * { \ \ \ \ A. 612 PINUS in clusters 0.5 to 2 inches long (57, 787, 209, and the seeds are rapidly dispersed (table 2). 225). Before ripening they can be green or Drying causes the cone scales to separate owing yellow to reddish purple, but are light brown to differential contraction of two tissue systems: to brown at the time of pollen shed; in most woody strands of short, thick-walled, tracheid- species they fall soon after ripening. Female like cells extending from the cone axis to the strobili emerge from the winter bud shortly tip of the cone scales, and thick-walled scleron- after the male strobili and are green or red to chyma cells in the abaxial zone of the scale. (2, purple (51, 71, 181, 225). At the time of pollina- 229). In a few species with massive cones, the tion they are nearly erect, and 0.4 to 1.5 inches scales separate slowly, and seeds are shed over long and sometimes longer. After pollination, periods of several months; e.g., P. coulteri, P. scales of the female strobili close, and the rorbtrt'ghii, P. sabiniana, and P. tomeEana strobili begin a slow development. At the end of (225, 233) . the first growing season they are about one- In some species part or all of the mature cones eishth to one-fifth the lengbh of mature cones. remain closed from several to many years or Where temperatures are favorable, development open on the tree only at irregular intervals; e.g., continues through the winter as in P. elliottii P. attenuata, P. banksiana, P. brut'ia, P. elausa, var. elliottii in Florida, and in P. attenuato and P. contorta var. latifolia, P. ltalepensis, P. P. ponderoso at low elevations in the Sierra mtrlicata, P. pi,nastet', P. ptmgens, P. radiata, Nevada (122,25I). Fertilization takes place in and P. t'igida (51 , 71, 225) . In addition to their spring or early summer about 13 months after closed-cone habit (serotinous cones), P. bank- pollination, and the cones begin to grow rapidly. siana, P. clausa, P. r'igida, and P. contortahave Growth of a new shoot leaves the developing forms whose cones open prornptly at maturity cone in a lateral position. As the cones mature (29,71, 196, 197). The closed-cone habit is the they gradually turn from green, purple, or result of three factors: (a) extremely strong violet purple to yellow, light brown, reddish adhesion bet'*'een adjacent, overlapping cone brown, or dark brou'n (table 3). scales beyond the ends of the winged seeds (738, Cones and seeds of most snecies mature 126); (b) cone structure; and (c) the nature rapidly during late summer and fall of the of the two tissue systems in the scales described second year (table 2). Cones of a few species above. The scales apparently are held together mature during late winter of the second year or by a resinous substance. The melting point of early spring of the third year; e.g., P. attenuata, the resin seal for P. contorta var. latifolia is P. brutia, and P. merkusii (25, 10,722). Seeds between 113" and 722" F. (ltl). Heat, especially of P. attemnta and P. bruti,a are mature during that from fire, melts the resin and permits the the fall, about 16 to 18 months after pollination, cones to open. Stiil other species shed partly but the cones are not fully developed until late opened or unopened cones, and the seeds are winter (25, 122). Pittus leiophAlla, P.leiopltUlla dispersed only when the cones have disinte- var. ehihtnhuan.a, and P. pinea require a full 3 grated on the ground; e.g., P. albicuulis, P. years for their seeds and cones to ripen (5-l , eembra, P. pu,mila, and P. sibirica (51, 169, 185, 136). The requirements of P. torreyana are still 225) . unclear. This species has been described as need- Commonly, cones that opened on the tree are ing 3 years to mature its seed, but there is shed rvithin a few months to a year after the evidence that the seeds mature in 2 vears. with seeds are dispersed. In some species, however, the cones requiring 3 years to open tiZZ). opened cones may remain on the trees for up to The interval between large cone crops is 5 years or indefinitely; e.g., P. attenuata, P. variable and depends on the species and environ- eontorta, var.Iatifolia, and P. rigida (71,225). mental factors. Some species consistently pro- Mature seeds vary widely in size, shape, and duce a large crop every year, while others show color (2t+7u) (fig. 2). They range in length from a cyclic pattern of 2 to 10 years between Iarge one-sixteenth to one-tenth inch (2 Lo 3 mm.) for cone crops (table 3). P. banltsi,anlc, to more than three-fourths inch Mature cones (fig. 1) vary widely in size and (19 mm.) for P. sabin,iana. They are ellipsoid weight. Those of P. rnugo are 1to 2 inches long ( P. radiata ), p ear -shap ed ( P. pu,mila ), cylindric and weigh about 0.06 ounce, while those of P. (P. gerardian,a), more or less triangular (P, lambertiana may be 12 to 25 inches long and 'pttngens), ovoid (P. petrce), or convex on the weigh about 1 to 2 pounds. Those of P. sabiniana inner and flattened on the outer side (P. pinea) and P. cotilteri often weigh more than 2 pounds (51, 180). The seedcoat, which may be reddish, (51,180,225). purplish, greyish, brown, or black, and is often The mature cone consists of overlapping mottled, can be rather thin and papery to hard woody scales, each of which bears two seeds at and even stony (5/, 209,225). the base on the upper surface. The cones of most In most species a membranous wing is at- species open on the tree shortly after ripening, tached to the seed, but in some species the wings

613 PINAS

P. albicaulis whitebark pine

P. attenuata knobcone pine

P. cembroides Mexican pinyon

P. banksiana P. clausa jack pine sand pine

..& t.rt'" t\ tt rl

i"\\4'

P. engelmanii P. flexilis P. leiophylla var. chihuahuana Apache pine limber pine chihuahua pine

Frcunn l.-P'inus: mature cones collected before seed dispersal, /2 y. 614 PINAS

P. monophylla singleleaf pinyon

P. ponderosa var. arizonica P. ponderosa var. scopulorum Arizona pine Rocky Mountain ponderosa pine

-il .* iw * {,' P. rigida t'l pitch pine ffirF

P. pungens P. serotina Table Mountain pine pond pine

P. strobiformis P. sylvestris P. virginiana southwestern white pine Scotch pine Virginia pine

FIcuRE l.-Pinus: mature cones collected before seed dispersal, % X-Continued. 615 PINUS

q-p (*"** .$F &n ,r,'r,.4b P. albicaulis P. aristata P. armandii P. attenuata P. balfouriana whitebark pine bristlecone pine Armand pine knobcone pine foxtail pine

','J flp

P. banksiana P. brutia P. bungeana P. cembroides P. clausa jack pine Calabrian pine lacebark pine Mexican oine sand pine

"p & t

P, contorta var. murrayana P. coulteri P. densiflora P. echinata Sierra Nevada lodgepole pine Coulter pine Japanese red pine shortleaf pine p

P. edulis P. elliottii var. elliottii P. engelmannii P. flexilis P, gerardiana pinyon slash pine Apache pine limber pine chilgoza pine

&;-* ' w

P. glabra P. halepensis P. heldreichii P. insularis P. jeffreyi spruce pine Aleppo pine Heldreich pine Khasi pine Jeffrey pine & t ffi

P. koraiensis P. lambertiana P. leiophylla var. chihuahuana P. merkusii P. monophylla Korean pine sugar pine Chihuahua pine Merkus pine singleleaf pinyon

FrcuRo Z.-Pinus: seeds as shed naturally from their cones, 1 X: some are wingless when shed. 616 PINUS

are absent or rudimentary; e.g., P. rnlbi,cauli,s, Ripe cones can be collected from standing P. armandii, P. cembro,, P. flerilis, P. gerar- trees, from newly felled trees, and from animal d'iana, P. koraietrcis, P. pumila, P. sibirica, and caches. To avoid large yields of immature seed, P. strobiformis (209, 225, 233, 2rt7a) (fiq. 2). collection from animal caches should not begin In others the wing or modified "wings" may until late fall, when the seeds are definitely remain attached to the cone scales when the mature (205) . For most species, cone collections seeds are shed; e.g., P. cembroides, P. edulis, P. from standing trees should start as soon as the gerardiana, P. monophulla, and P. qzutdrifolia cones are ripe and just cracking, since most (225, 233). The seed wings are easily detachable seeds are shed promptly from opening cones. from the seed of most hard pines except P. pinea, For closed-cone species, collections can be de- P. ronbttrghii, and P. can,ariensis,' those of the layed without loss of seed, and frequently delay soft pines are firmly attached except for P. is even desirable. Although seeds may be ma- aristata and certain sources of P. balfouriana ture in the fall, the closed cones are very difficult (169,209,233). to open at that time; additional maturation on The mature seed consists of a seedcoat which the tree facilitates both cone opening and seed encloses an embryo imbedded in a food-storage extraction (25,122). tissue, the endosperm (female gametophyte). To avoid extensive collections of immature Attached at the micropylar end of the whitish or empty seeds, it is advisable to first check endosperm is a brown papery cap, the remnant ripeness of seeds in small samples of cones from of the nucellus. The endosperm and papery cap individual trees. A mature seed has a firm white are covered by a thin, brown, membranous ma- to yellow, or cream-colored "endosperm" and a terial-the remnant of the inner layer of the white to yellow embryo which nearly fills the ovules integument (fig. 3). endosperm cavity. With experience this visual check is very useful, and with some species it may be essential. Ripeness for some species can be estimated by changes in cone color. Colors of green and ripe cones for most species are listed in table 3. For example, P. Ttond.erosa var. ponderosa seed coa t cones are mature when the color changes from green or yellow green to brownish green, yellow endos perm brown, or russet brown; P. resinoscl. cones turn from green to purplish with reddish brown on cotyl edons the scale tips; P. strobrts cones turn from green to yellow green with brown on the scale tips or to light brown. For some species such color hypocotyl change comes too late to be a useful index; e.g., in P. palttsti'is the ripe cones are still green in radicle color and may have already started to open before turning brown (25It). L. For species in which changes in cone color may not be useful, flotation tests of cone specific gravity be. These FIGURE 3.-Pinus ponderosa, ponderosa pine: longitudi- may tests are based on the nai section through a seed, 6 X. fresh weight of the cone. Species for which cone specific gravity has been related to seed maturity and their appropriate tests are shown in table 4. Collection of cones.-Cones should be collected The easiest way to determine if cones have from trees superior in growth and form char- reached a desired specific gravity range is to acteristics. Larger cones generally contain more see if samples of freshly picked cones will sink s-eeds, but normaily all cones are collected except or float in liquids with known specific gravities. those with obvious disease and insect damage. Pitttts Ttonderosa seeds are ripe when the cones Widely spaced, dominant trees with full crowns will float in kerosene, P. strobu.s is ripe when produce the most seeds per cone, provided ade- the cones just float in linseed oil, and P. glabra quate pollen from other trees is available. When is ripe when the cones just float in SAE 20 trees are isolated and pollen from other trees is motor oil. Cones from standing or felled trees Iimited, seed yield tends to be low. In dense, should be tested only when fresh, since excessive young stands most species usually produce litile drying will lead to erroneous conclusions as to seed. Those species which form fire thickets are maturity. exceptions i e.g., P. attenuata, P. banksinno,, P. Collecting from felled trees should only take clausa,, P. rigida, and P. serotina (71). place after the seeds mature. Otherwise there 618 PINAS

TAsLp .-Pitttts: speci,fi,c grauity of ri.pe cones und liquids used for testing ri.peness ba flotati,on Specific gravity Flotation Data Species of ripe cones test liquid' source P. aristata 0.59-0.80 kerosene -. p06 P. contorta var. latifolia 0.43-0.89 1l,g P, densiflora, 1.1-0 P. echinata 0.88 Seff zo ,r"t"i"if, oil-part kerosene to 4 parts lin;;ed ;lt 36s1, P. edulis 0.80-0.86 kerosene z0l, P. elliotti.i vat. denso, <0.89 SAE 20 motor oil !t4 vzr. elliottii <0.90 SAE 20 motor oil 251, P. glabra, 0.88 SAE 20 motor oil P. jeffreui 0.81-0.86 903 P.Iambertianrt, 0.70-0.80 203 P. palustris 0.80-0.89 SAE 20 motor oil 25 lt P. ponderosa valf. arlzonrca 0.88-0.97 191 vat. ponderosa 0.80-0.86 kerosene 71 vat. scopulorum <0.85 kerosene P. rad.iata <1 water_ 130 P, resinosa 0.80-0.94 ketosene " 71 P. serotinq, 0.88 11tr,251, P. strobif ormis 0.85-0.95 sic" eih"iot 191 P. strobus 0.92-0.97 linseed oiI oa< P. sylaestris 0.88-1.00 66, 131 P. taeda 0.88 SAE 20 motor oil, or 1 part keroie". io a pa.ts ti"seea olt P. airginiana <1.00 68 ' Test should be made as soon after picking as possible to prevent excessive drying; the liquids have the following specific gravities: kerosene 0.80, 95 percent ethanol 0.82, SAE 20 motor oil 0.88, linseed oil 0.93. Five or more freshly picked cones should float before crop is considered ripe. 'Red pine cones which float in a 50-50 mixture of linseed oil and kerosene are within 10 days of being ripe.

is a risk of harvesting immature seeds. In some immediately spreading the fresh cones in thin species, e.g., P. taeda and P. echinata, nearly layers on a dry surface in the sun, or on trays mature cones can ripen in the crown on felled in a well-ventilated building, or by placing trees, but in other species they may not (251). them in sacks hung from an overhead rack With some species, slightly immature seeds can protected from rain (205, 221, 223, 25/+). The be successfully ripened in the cones after re- cones should dry slowly to prevent "case hard- moval from the tree-(P. elliottii var. elliottii) ening." After initial drying, the cones can be (23, 25L); in cold moist storage-(P. lam- stored temporarily in well-ventilated bags or bertiatta) (121) ; or in proionged cold dry stor- trays. For many species, ripe cones open satis- age in closed gsntainsrs-(P. ui,rginiana) (37). factorily under the above conditions, but cones Such methods should only be attempted if of some species may require additional heat in completely mature seeds cannot be collected. either a cone drying kiln or a heated shed. Prop- Cones most often are hand-picked. either erly air-dried cones of a few species may open from ladders or by climbing the trees. For satisfactorily after a few hours in a kiln, but some species, hand cutters oi a cutting hook those of other species may require several days. must be used to detach the cones, and hooks may Cones of most species can be opened at kiln be needed to bring the cone-laden branches to temperatures not exceeding 130' F. and a the picker. With certain species, mechanical humidity of about 20 percent; but cones of a tree shakers have been used for the rapid har- few specieS, €.g., P. banksiana, P. clattsa, P. vesting of cones; e.g., P. elliottii var. ell;iottii, ponderoso" var. scopulorum, require higher P. palustris, and P. taeda (119). With a few temperatures (205, 221,223, 235) (table 5). others; e.g., P. edulis, P. monophylla, and P. Cones stored long enough in containers to quadrifolia, large amounts of seeds are col- have dried without opening or cones dried under lected by shaking the tree or beating the crown cool conditions may not open properly during to extract the seeds, and then gathering them kiln drying. In such cases, the cones must be from the ground (204,235). soaked in water for 72 to 24 hours, and then Cone processing and seed extraction.-Cones kiln dried before they will open satisfactorily should be dried immediately after collection to (223\. avoid mold development and excessive internal Serotinous cones have beerr opened by dipping heating, which lead to rapid seed deterioration. them in boiling water for 10 to 120 seconds. Drying can be accomplished in 2 to 60 days by Immersion times up to 10 minutes, however,

619 PINUS

Tanr,p 5.-Pinus: cone processing sched,ules and ai,able periods for seed,s in cold, storage

Cone processing schedule Viable period for seeds Kiln drying in cold storage 2 Species Time in Air- boiling drying Mean Data water time 1 Time uemper- source Time Data source ature Seconds Days Hours "F. Years P. albicaulis" -, 0 15-30 0 I 122,166 P. aristata- .. 0 2-8 0 206 I 166,204 P, armandii - 0 15 0 P. attenuata- 15-30 1-3 48 r20 16 isz, pog P. balfouriana 0 2-8 0 122 16 122,202 P. banksiana 0 2-4 16ii 126 10 122,213 10*30 3-10 0 126 P. brutia 0 3-20 0 78,122 s iip, soe 0 10 130 78,122 P, canariensis 0 z-to 0 I q.) is iip, sos P. caribaea,-. - 3 150 P. cembra"----.- 1+ 1/e P. cembroides " 0 2-8 0 '| liz P, clausa 10-30 2-4 t45 11 4, 138 iis P. contorta var. contorta 0 2-20 0 63, 729 77 122,202 0 96 r20 63, 122 var. Iatifolia " 30-60 2-30 0 14s 7+ 202, 242 0 6-8 t40 23s vat. muffaaand, 0 2-20 0 63, 722 tt1n iip, zop P. coulteri 0-120 tJ--t- D 0 100 D 122,766 0 72 t20 100 P. densiflora 0-30 z-a 0 196,264 2-5 s8, lrr, P. echinata 0 48 105 159, 25[ 35 159, 255 P. edulis' 0 .) 0 204 P. elkottii vat, densa 0 8-10 120 oa 0 A 0 qr) var. elliottii 0 8-10 rto 254 35 113,296 0 iz 0 25t, P, engelmannii 0 60 1io oq< P. fleri,Iis 0 lb-to 0 10q 5 122,219 P, gerardiana 0 15 0 P. glabra 0 48 100 1+ ii, to P. halepensis 0 10 130 10 78,112 0 3-10 0 118,122 P. h.eldreichii 0 5-20 0 100 P. insularis 0 5-20 0 122,146 P. jeffreyi 0 oA t20 7qo 18 166,122 0 5-7 0 laa P.Iambertiana 0 qA 120 2L 202,2/t6 0 o-l 0 100 P. merkusii. . 0 o-l 0 2+ ia P. monophylla n 0 2-3 0 P, monticola 0 1A 110 122, 2.13 20 ioz, st o 0 0 P. mugo 0 48 rto 122,171, 5 112,174 P. muricata 0 48 120 63 P. nigra 0 24 -T ID io+ s8 0 3-10 0 7AO P. palustris 0 48 100 254 5-10 1Ir, 157 P. paruiflora 0 D_I') 0 700 P. patula. 15-30 l-2 48 I.LD 2l 122, 202 P. pinaster 0 IID 178,17/, 11 112,166 0 4-io 0 119 16)6) P. pinea 18 soe ponderosa P. . val', arxzon1.ca 0 60 110 235 vat. ponderosa 0 120 26,722,235 18 26,202 0 4-ri 0 26, 122, 235 var. scopulorum 0 .) 16i o/ 15+ 6, 0 4-t2 0 100 P. purnila" P. pungens 0 72 120 3 0 30 0 .D

620 PINAS

T.q.sln 5.-Pittus: cone processing schedules and uiuble periods lor seeds in cold storage-Continued

Cone processing schedule Viable period for seeds Kiln drying in cold storage' Species Time in Air- boiling drying Mean Data water time' Time temper- source Time Data source ature Seconds Days Hours Years P. quadrif olia n 0 2-8 0 104 P. radiata 60-120 0 48-72 L20 9lr6 it 712,202 60-120 o-t 0 2l|6 P, resinosa, 0 I 130 221, 238 30 98, 213 , 248 P. rigida u 11 52, 98 P. rosburghii 0 4J- 56, 112 P. sabiniana 0 48 L,O 1qo 5 P. serotinn, 0 48 105 ql P. sibirica" - 238 P. strobif ormis 0 ii 0 101 -so, P, strobus 0 4-t2 130 10 ss P. sylaestris 0 10-16 120 176, 212 15 98,115 0 d-t^; 0 7 0o P. taeda- -, , 0 48 105 159, 25.1 9+ 159,235 P. tlrunbergiana 0-30 5-20 0 10a 1ry1 11 202 P. torreyana 0 5-20 0 6 122, 166 P. airginiana 0 ,. 170 5+ 112,235 'Air-drying times are for a temperature range of 60" to 90' F. When kiln-drying is used, it should be p_receded by an air-drying period that was not reported for most species. A period of 3 to ? days is recomntended rvhere no times are listed. 'Period after which at least 50 percent of the seeds germinated. Storage temperature ranges for most species were either 0' to 5' F. or 33' to 41' F. The iower range is preferred. Seed moisture contents betrveen 5 and 10 percent' were satisfactory. ^ P. albicaulis, P. cembra, P. pumila, arrd P. sibirico.' Cones must be broken up to release the seeds. n P. cetnbroid4s, P. edulis, P.-monophylla, and P. quadrifolia; An alternate procedure is to shake the tree to release the seeds and collect them on a cloth spread on the ground. 'P. contorta var. latif olia.' Time required in boiling rvater is estimated. Reported treatment rvas 5 to 10 minutes in water at 148' F. or higher (l/9). n P. rigida: Cones were soaked in water overnight and dried in a rvarm room (52). have been needed to open some lots of serotinous After completing the dewinging and cleaning cones (122). This procedure, by melting the processes, empty seeds of many spec,ies can resins between the ione scales and by wetting be separated from the sound seeds by flotation the woody cone, produces maximum scale re- in a liquid having a suitable specifi-c gravity. flexing (ISA, lpo)-. This procedure has been used on the species After the cones are opened they are shaken listed below' to remove the seeds. Seeds normally are ex- Flot.atiort. IiqtLicl tracted by placing the cones in a large mechan- .f or separating Data e trtpt 11 seetis source icai tumbler or shaker, or in a small manual Species shaker for small lots. Seeds are then dewinged P. brutia rvater 118 by machines of various types, by being flailed P. coulteri water 129 P. ech.inata 959f ethanol 159 in sack, rubbing. Dewinging of a few -water a or by P. echinata )5 ^l species can be simplified by first wetting the P. elliottiivar. rvater 2 5lr seeds, then ietting them wings are loosened eiliottii. dry; glabra be (223, P. 951 ethanoi 15 by this method and can then fanned oat P. halepensis 959/. ethanol 118 25/t). Care must be exercised in the use of F. nigra 95/.ethanoi 118,17/t mechanical dewingers, since they may injure P. palustris pentane 158 the seed. Seeds of three species-.P. aristata P. pinaster 95!f ethanol 118,17t+ P. pinea rvater 77,118 (206), P. palustris (25tt), and P. syluestris P. sabinian.a rvater .122 (115) especially susceptible to mechanical P. strobus i00!'. ethanol 221 damage-are and must be dewinged very carefully. P. syluestri.s petroleum ether .127 The seeds are cleaned by using mechanicai P. taecla rvater 159,25I clipper cleaners, fanning mills, screens, or Viability may be reduced after seeds have gravity separators to remove the mixture of been immersed in an organic liquid such as broken seed wings, pieces of cone scale, and ethanol, pentane, or petroieum ether. The re- other impurities. duction, however, can be minimized by using

627 PINAS

oo a3 o: N =k S\o 9aNYroo.Sq)bstto V)ta ooqkr qolooqN Gi -Q oO rO :iA:+si sl 9i \ A \i \o q cb cJ.+ cJ eJ ca eJ $i qJqi c. \ S N\o \tG\6c ca6.c$ S ^=a stG\\o\\c.l\\\ot\c{ sisr (o(ocasrci stsl\slG\l\\eJ\(oGt(oca \

a t- o $ -L-l- -]- + € cn

(-) $ U \3 OdootilOOtOrO0OcaHO @ coo oo ldr c6 0) t- @ r cliO)HOINOi c\ro6INNO(OOt-(D(OO oNoilro 6ldc)oooc€)L.-ro q) q) 6It'-irO

dY ;j; AT r) roh a GC \Gb Sl s st NSt N eJ sr*i \o *i ,o oi-t doo \o sl 'sr sa Ct On\ CfGACStOC \t\ oo -N 6t eSel-\6 q \GA I Sa ro6t loloG.l\c{ Sl\B \ I Si (c)Ga

q b T.Q) *rd li i t.. lo) 6.t@ Nco6im l-' 'd!:tr r o€<{ o -jd o'dc; .i6.j .4 C\ C\f - a) I ttttl il lr cO IrJ c\l b .dr ro co Hrorrit o .J o' doooo c;qJp

1lr) o !Q6IAO0O Oi ltro tF.. q 6'j '-.: ! oi oi o; es ; ; a9Rs6rFi ,l ^i.; HFO oc.i.j

I I I t.- l@o ttr- UrZ c\ .$ a eq coi!Qb- 166, i rTo !b I T TT + :r boby;d la* 6.t m

i b0 I a) E€ I rv ,^ I i E-E' l{* 'Eo'E'E.E dd i f - ': i 6 -r i':'P h tr - E ;': Hj .iteoo-' e * X;*iX!- H h E .i Fl i : Y Cqi* o I i: ! E'gEfEs s 5 s e:lr Ef E€ ;;!!cdd6grJOn gG<(JQOUF;6 : dE€EGf r F6* lG6? ; < fE( a ='q

q q

oo \o kr 8n qJ o) s $ R sr Ga Gn 6a \o ,o I rcili qro rooj\ CE 6f sJ ooqqo snn*$Ex$ R*$:$st-$*sr N\6a B\sl\ RS SRR

-]- -L rt -r -f (a@

9-- e4e: OeOC oo n3?f^??39 o.?@ <1e.noq6f61 6.-o-\. q6-- --E6-5- tr)vd i\qD(ocoliic6c\r HO coN; c\(o{r6l ro Hi iiH;c6;rjco-lclcQos rj rl)

ooc\ ioo@oo(o<) oot-- EEFo^qm- o-61\o-c!EEFEE co^€o-SgE .1o-o-o^3883 .{-_83 6ror3t3 HC\dOO)t'-

to st *+s st

GT ,.i sl Gb qr \+ b-rs\o toN \\ON N Gr kJ 6i Gi .+ql\ \ Nb9i:+ orci oisiLi ci \. srQ - !: st\ st -N bqr \r ^\ 9r ^Gr ^6r c6oONc6(o\O- 6 rr^^qca-5-\a\\- q\o si c)9:+ - !59lro Gb qcrro s\ OO blo\so\\ N \\oca oi .bc\lo .c.l \sib \\

\oqocc!6:q gq4 .c?oqoq lir r.-r--co 60 @ t$Ooodo qr'-i 1 ttl '1 d,oqo c; c.i*cj t-C6 .d *d * @N I 'jc; c; c;ocjJj "l+

Q.$OH h caoo) r-i on cri rj ?qqqqqqqooq6? lttl ry+rj "d ddJJ d;cdc.i m cd N c.6 061 -i c.i c.i c.i

QlaA (crocc \ci icacoro<{c6 **"1 c.<1 ltl I l.o oN 6c'1FY*T C! m co 6J

!E - ie+-a a a cdd.3 cdii -l'.j3 ! 'e , iq/9! Fdd a '&c @or F.I o >'t I a; c.9:i.S 'FZE- Qa E6 5?a * z H .:: !'F'Fs b: - csq Y u!'F : o i.ll '664>;: r5;t.: .I E H.i r€#.s X< i.I I BE€E€€ t'3 5!: J.dcB.da"'=da-?!!-s l'- :H 'A #i !s F8s* t f'l < -V9-A -V a ca

624 PINAS

cold strati'f'ca.- prolonged cold stratification for 6 to 9 months is Tlsr,u 1.-Pinus : recommended scarifica- -'i;; (33"-41' F. in a mo.'i'st medi'um) iiruch irore effective (100, 121). Acid Tteriod's ' seeds of pines and,' oth'er pregermination treatments- tion is not recommended for Continued (122).' Seeds of P. cembra, P.ltorainsi,s, P. paruifl'ora, Recommended and P. si,birica are suspected of having imma- cold strati- ture embryos at the fime of collection. Ger- fication period Data by placing the Species sources mination has been increased Fresh Stored seeds first in a warm moist environment for seed seed several months and then in cold stratification for several more months (table 7, footnote 2) Days Days (8,9,173,238). P. pu,mila_ 120*150 120-150 8,97,722 P.pungens, - 00 Germination tests.-For reliable tests of seed P. quadrif olia 0 0-30 viability the seed is allowed to germinate under P. radiata 0-7 7-20 108, 122 97,122,193 near-optimum conditions of aeration, moisture, P. resinosa 060 ex- P. rigida , 0 0-30 52,235 tempeiature, and light' On the basis of P. ronburghii 00 1q.) tensive experience and experimentation, stand- o 122,165,235 P. sabiniana 60-120 60-120 ardized sebd tests for a number of pine species P. serotina 0 0-30 tt4 238 been established by the Association of P. sibirica " 60-90 60-90 have P. strobif ormis 60-120 60-120 /, 191 Official Seed Analysts (70), the International P. strobus 60 60 122,193,235 Association (108), and certain re- 91, 101, 235 Seed Testing P. sylaestris 0 15-90 gional such as the Western Forest P. taedo 30-60 30-60 161 organizations P. thunbergiana 0 30-60 18,99 tree Seed Council (263). Recommendations and P. toneyana 30-90 30-90 122, 235 procedures given by these orgariz-ations and P. rirginiana 0-30 30 97, 239 are summarized in table 8. 56, 99 bthers who test seedi P. wallichiana 0-15 15-90 In addition, the results of seed tests conducted given for 61 species 'P. cembra: cold stratification period has. been re- under known conditions are duced to 90 days by first soaking the seeds rn concen- and varieties. ii ti""i" "t"ia"tot 3 to 5 hours, or by mechanically germination pine seeds can be effec- iit't"a is The of .^."iiivits Tt ieeds,(729, 1g/*), b:ut acid treatment tested in any medium or container that not now recommended," tively '*; i;: ;";b;;',"p. koiaiensis, P. paruiflora, P. sibirica: provides good aeration and holds adequate In some lots embryos may be immature and requre a moisture. For a number of species, light, com- ;;"* ;.i.t stratification period (2 months-at 70" to monly supplied by a cool white fluorescent l-a-p' g0; ir.ip;uceding the cold-stratification period (9, 99, oary\ is required for reliable tests. When light is '"iij. (1-52). and pinea usual exposure is 8 hours in each -Cooaedulis (204), P. pinaster ^P' necess-ary, the usii, ELrmlnation'has been obtain-ed after soak- 24-hour period. Different temperatures are with no i"i-J""aii" E"ta iao' F.) water for 24 hours employed-in seed testing; constant 68' F. and su-bsequent-'; stratifi cation. com- p.-iiiii"t a cold stratification pe,riod.of 60- days was alternating 86'/68" F. regimes are most sufficient when seeds were first soaked in sulturtc acro mon. When alternating temperatures are used, i""'lO-*i"nies (9.q, 122, 23s), but acid treatment is the higher temperature ordinarily is for 8 hours not now recommended' 16. The duration of most '*; is speede.d bv cracking.the and tfie lower-is for i;: ;'"ii;;iio :- germination 400 to 1'000 *rlcf<' seedcoats, t-etore stratification (122, 165, 235\' tests is from 3 to 4 weeks. Usually seeds per test are adequate. Germination is epigeal (fig. a). may exhibit extreme Cutting tests are commonly uesd for rough Seeds of some species quality. Such tests can e.g., those that requile. more than determinations of seed dormancy; also provide information on soundness and can 6O duy.'of str'atincation (table 7-) ' The dor- physiological or physical be used as an emergency guide in fall sowing of tnu".v may be due to dormancy. X-ray A pretreatment may be, needed to over- fresh seeds with embryo factois. methods supply good information on sound- come a phisiological block in the embryo; e'g', too p. i"mUnitiana* (121), or effect a phvsical ness. Estimates of viability from the above to make it permeable to tests are the most subject to error, since the change in the seedcoat actually germinated (221 223, watei; e.g., P. sabini'ana (82, 121). The dor- seeds are not , anatom- 2sL) (Chapter 7). mancy can also be more complex; -an ically immature embryo with a physiologi-cal Biochemical methods employing a rapid vi- be coupled with an impermeable ability indicator such as one of the tetrazolium block may gen- seedcoat as in P. cimbra. Acid scarification of compounds can also be used, but are not seedcoats has been used with several species; erally recommended. The results are highly cle- e.g., P. cerlbra', P. peuce, and P. sabiniana,b:ut pendent on the analyst's experience, and the 625 PINUS

d8 €k (o00cncosisSt\cncodj q\bq\\ AX A Stootoo -oi -OaS .+Cl 9r -h -coco - - cicoN arqJoo\* o q ooqqn\ s \t f, <) c\ N ei s q \ ro c) \i f, q N s oi q c) cn s ca ei at :) ei s s ci oo f, ei \o Q s oiStAf..anGnSOoiqco \ SA \ St c\l \ St \ \ St str r \ \ \ \ \ Sl G.l \ 6l H St St (o \ (o \ \ 6t \ \ St \ \ St GaGa\oosaq\\sl\\ a l- + l a e 3: m

tr :Y tu Rto nqoNLot-o)ro<{(l)N t-otr 6o oooc6roro r b-mt$ oo (o Nt-- o)

IroOOroo>ot- I F-@ lloro l t- F- Ftr Fe) !O i :to:o)or

13 tr 5 Caro{'Nc)oo{r00moi @oo@oo @oiNNOcomic>ooco@t-co(c 6.100@(o(o(oiit--oo q .J Glq.ci6tcacaN! Nm6l C\l o) co c{ crl c6 N n 6i (o (o c.o N at N c6 (o c6 N N N c{ i c6 c\ c! G.l cI H c\l 61 6t c6 co o ac6i I $

() @ o ca @ ca 0o oo co o oo N oo co co @ c{ h @ 0o @ b- r @ o) co oo oo oo Io co N aI co o aI 6rco6lc\o@@c!@o . (O (O (g (O (t (O cg (O cO cO (O (.o (.o qO (O (o (o (,o $ Fr . n'.O @ € t'- r @ <) r rO IO t-- h cg t-- r C- @ t- r r (c F F- t- b- t- u) li\\\\l\\\\ -Na tr-@ O) @ @@t- eO @ oO@ @@ g) a h o o Q s. € F l@t-(ocl) I t l

' r :\ r>AA Fr>g H E Frpq Qr> mi-l-{*i -L i - oi&gi ^/i 4r@H*n -4FAAAAA ?,4 i?? a aFttu ,V FVXfu HT aFts

h+! q, ++ -T- -r S € o O.d o$ @co oO oo oco @ oo o o @co 0oo@oooo@@ooo@co@oo €(o@@

-r,€€ -LIIIIIIIII I t+++l I | +r++r t++t t+l i +++ lll+r++r+t

I i

ai s.s a *i *q a I \..,1 i: .e* sis i= .: .d i\ l- F. s.e '€ oFa ,:c io bS s .3i{ "d* Q$ I F- *+ .: ! a € 's' $ Ei il s s : sil ll Pcn cg cdp i$ ,o : : :i6 aD R, AA \ \ ry q [l\ tu ry&a' a' 626 PINUS

cOiYiC! H loo roF{o)o + oout

roo oobN@ 11. ;ico Io FoJ(O

lt-o lt'-

jHo looo c\ t-

cc F C)iilOO N F m cO ; sr o n ooo <1 oi o s € oo gQq b cc Q N c0 (!) i o c. c{ di j m cr N N ot c': r N c' N N 6r 6lc\61 @cit c6

1I: u: @@ 6NN@il

(t 0|o 00 @ lco

: .l

i H 2r lr1 H + +l t*& i cq+&Pa + -t-!: '^aa a.uaQ AFlEi H aAa tui aa>a

+-+++ o€@o@ocoootd ooco@ooococo --*oc)Jo * ooooo-liocoooo@ooSoooggoocioococooo6o

-1- -rl l++l I l+l l+ ++l I I l++l l++l+++++ | I I I I I l++l I I I l+++l I++++l

l ll tl I i il l i t- -! iriii.lil rl i tr i .-r ' , F g: -':-QO r- i: l,i si3 EHT i a-O I 'S p- € i E "cP i'iE 3q q!AL B o+i 'd€R E i L ,o "Ei' S -!Lo;; d s *> > B) s EE s: 5it =a.a.o;o'.{dda. a. vpacc)fl PINUS

lrd o I st lo) s3 oG6CtO:GD ia \qgjaior lo HO oo - -co - ooNf."6tGtqioo\ r - st(o q -eJt.Li a!\o \ t;. ^>a q6 eJ o! oO 6f G.a qC GD oo q cl qi eJ GD oo S 6 (o a O si S S I S S \ \ I A S \ <) c| O l9 ts 9r \ G.f \ q Ci \ St \ \ Sl G\ \ N \ \ N oo \ ai 6l \ \ Sl \ B \ N B e.t G.l \ \ l4 t+ trl.i I a o) rl) crt H @ (O o) rO ro c/: i t il rLI a b.- l€ .aJ t6! d nl t.- (J ! 6l ,(o icb1.6t- a (ot--t--Al(0 o) 6a b- 0o co la @ @

ut = oF{ co co oo co @ co co 0o d N c\ (0 d @ 6 6 € ol @ co @ @ 6J @ 1(: o r!) @ 6I co oo o qQ s (O (O (O (O (O (O (€) (O (O (0 (c' (O (O (O (O u . @

i 6 te1.9 ile .tr q) i -..r H FPI IU lq A i q) rtE nc! I i O - .tr I m mm^, -+mmm^ o..^ o.mm I .3 co si E-i d tr FrFiFi ui a 3 u ,FiFp! l- -r-r +++ +++ +++ i9 90dE u cOco@oo@ocoooo $ coo o o o @ @ @ €6 <{co@ (o @ @ oo co@o co o o li .. E-{ cd 6! Fr AI6IH l>. =dFgt fl a;5 & \r l- Nv+)o cd | ! '3!.9€ ul l;|i, ERrEo"F^ io 1.. aE9.EL! # td i v d a9o/ +tl r r+r I l++l+t i l+++l | | | | | | l++t | | | I iil .EEl: b it >It*FsSi F l;?.:o.do>l H,r $-:oj6 a HEE Iif:iFEg lSIaboh'iiad6FE Eg!(6o i.-i:i .38 fr a i:.i S.s3o;Ls lgE€ ilit:q ,* Fg o ts6 rB .s.: | fl e F.s rE.E:s .T;:: .:S '; .5s ':nF ': !S E s{ SO) ti q=: i€ a iS SE | ;18 f,E;v qqQts$ ts "' FiEg$#fr57 a : L14 | '': ^^ .. a; {d o"da"tun: | fl;;;;etsJ" E" 628 PINUS age of the seed. The viability estimates often The seeds can be either drill-sown or broad- are much higher than the germination capaci- cast by hand or machine, but most large ties obtained from germination tests (Zpg') nurseries drill-sow in beds because it is more (Chapter 7). economical. The amount of seed sown per unit Nursery and field practice.-Pines are suc- area and the sowing density vary with the cessfully nursery grown in most parts of the species, seed size and germination capacity, and United States and in virtually all soil types. the desired density. The stock density influences The soil should be fertile, and of good drainage the vigor and size of the seedlings and trans- and aeration. In most large nurseries the soil plants produced. The stock density will depend is fumigated in the fall or spring before sowing on the species, the length of time seedlings will to control soil-borne diseases, insects, nema- rerrain in the nursery bed, and whether they todes, and weed seeds. Nursery practices are will be transplanted. summarized for 35 species and varieties in Seeds are sown at densities selected to pro- table 9. duce from 75 to 75 seedlings per square fbot. Higher tree survival factors are obtained when \ .i:' j medium-to-low sowing densities are used. Most nurseries seeds \ \r{ ,)r ,,' ,' sow at a slightly higher density '\ l if the-seedlings are to be placed in transplanl xtL beds for one or more years. A lesser density is desired for 2-0 than 1-0 seedlings. Depend- iti ing on the species, seed lot, and nursery, sowing ,titV:';,\ densities range from 2 to 20 ounces of seed per ,, il 100 square feet of bed. One northwestern nurs- ,, ery drill-sows P. monticola at 35 seeds per ''ii lineal foot in rows spaced 3.5 inches aparf to rlrilco obtain a density of 120 seedling per square iii foot for 2-0 seedlinSs (Zttg\. A second wesrern ro nursery drill-sows P. monticola at 18 seeds per |i lineal foot with 6 inches between rows to obt-ain ..;r i\ i a density of 35 seedlings per square foot for d\i i, il All a\\ c, its 2-0 seedlings (916). Ultimately, then, ex- { I tj,, perience determines the correct sowing density I Lt for agiven species and nursery for a particular )ll planting situation. Average nursery-germina- !i tion has ranged from 20 to 85 perient of the germin_ation capacities found ln laboratory tests. Of the seeds that germinate, as little as FIcURE A,-Pinus res,inosa, red pine: seedling develop- 19 and as much as 90 percent produce useable ment at 1, 7, and 30 days after germination. seedlings; the average has been about 5b percent. In temperate regions, pine seeds can be sown At the time of sowing, seeds are drilled or in the fall or spring. It is now common practice pressed firmlv into the soil. then uniformlv to spring-sow nondormant seeds. Dormant seeds covered withlA 16 ri inch soil, sand. or mulch. too may be spring-sown, but they must be pre- Fall-sown seeds should be placed slighily deeper treated. Some nurseries pretreat dormant seeds than, spring-sown seeds t-o protecfthem frbm of all species in the same manner; however, wind erosion and frost action. Large-seeded this is not to be recommended. The pregermina- species, as P. albieaulis, P. lambertinna, and p. tion treatment used on each specles seed lot mon,oph11lla, are covered to a depth s1 r/2 inch. should be that which achieves best germination Smaller for seeds require the least coverlng. fhe that lot. Seeds with embryo dormancy can southern pines, P. echinata, P. elliottii, p. palu,s- be sown in fall without a pretreatment. Com- tris, P. taed.a, pared seedlings produced and P. airgintatta, are pressed to by spring-sown into the soil surface seeds, those from fall-sown seeds are commonly and covered with burlap or chopped pine Iarger and better developed after one season. straw. Such materials protect With fall-sown seeds, however, sowing must be the seeds from birds or displacement by rain Iate enough to avoid fall germination, so that and help maintain soil moisture. Pinus corttorta seedlings are not subject to winter freeze dam- (var. cdntorta and. var. latif olia), p. densifl,ora, age and mortality. Fall-sown beds also are more and P. tlttmbergiana are sown /s inch deep; p. subject to losses from rodent damage. banksiana, P. eanatiensis, P. edulis, and p.

629 PINUS

ql h \o\CO \t cB \ sa s.l €! se ct Sf-b\G!\t st st N ;ss*""RNB $ *$$e N$qN* (oca - Log

6\l d i I d i* o6lc\lc! pNf li- J F c!6ic!j-ij "\ "!J cd i. I $S e{,tF- d c\JJ"lJ"lH .1 ^d oeojd'jod:ddHo-! tl rr | | | lt t lL | | | | +c{-Nl 6l c6 6tc! m c\ 6l c.tc.l co citco c.l i N ; A l^^ odiodood oo docidodo'id d od d .i oC) l\N | | ttrtttl tl llltlltl lt lrtlrllll I lt | | ii\i H H6lHi6tdc\ Hd niFN6lCAiC\ di ic\IHFn6.1 c\IiN H iAl ts i + c>ooo cooroo ro roro (o cO (0@ oo9cooutoro [email protected] o oH o OYoF @ Ir) F-

i\N l-N\a \N \N q) iH- i\ i\ tsssss i ,S Sls S I S a I I ltttl Ittl o \i sssss $s S +

i'i€ \ I ,O , a. | 1A i ': : iF '5 'P v) E * li L s *lF:Sg \ o iYF }X a o ,. r 3 i F- k-, F aaz ::f.:4 ii > p oo Eb! @):. _,.d^AA^ _ H'6E " =bE t;.FE* q I € X ]XHEEpE .aag a tr F'loti'o i*F" t€! gF€geE Fr is&3*a# o.d a 'a E ;E;fr S E'at 'a E

I I tt\a b! s\i\i\i\r@ r$ r* rfl i- - sssssls Ll .#sss"F#** is ssssx#tst8s a3 Lss I o I I I 'E*t ro oo | | | I I I i to IoL a o gn gs ns 5s a Fl sgo oo sssss o- ss sss o- oI o k hk hkFr Fr .F f-. + I rob OorOooo16o roo <00roc6vio1l) .d,99 S'oo lo o(O ro2 a H:l s 6trm I c6irroN1()

ir; i r ]oobll b0 tx I i rtrtr ooo oo ooooooooraOO @ooohoooooOooo o$oo q@ 610?9(0 (oco <)ooo1()<1 + c\.1 c6 <1 m co(o F 6l _c @o ro @ 6Jo@ !A o n No ro .4 a c\co d al <{<1 c\r fr N c6 c6 i FE rupg F*= ci9o 'si aaVz iiiiir i l4/ I i P ) !l* .Eoc r; I S.s 'a Fi i tra ! iS; s I .Es: :q90 a s F'$ l"iF. : o ts - eSi s.i p r FE i d'tr R.€ o s6. :shE.6 = F5sc5ss :.li P! gI P !Qd a Fs F H q\S o o S I $ :i !s5€EFF !6P d3 g .:.P,SsFss': iiiii lt rui; * Ei E aaE a;d{{a; \\\{\\&* ddd d{dda;o" t. l.d d { 630 PINAS nxonticola are commonly sown /a inch deep. Literature and Other f)ata Care should be taken not to sow the seed too Sources deep. Cited Germination is complete for most species (1) Albert, R. from 10 50 Data filed 19?0. USDA Forest Serv., Region to days after spring sowing. But 3, Coronado Natl. Forest. some lots dormant seed species, even of after (2) Allen, R., and Wardrop, A. B. pregermination treatment, may continue to 1964. The opening and shedding mecha,rism germinate several months to a year after sow- of the female cones of Pin.us racliata. Aust. ing; e.g., P. albicau,lis, P. cembt'a, P. peuce, and J. Bot. 12: L25-134. P. strobif ormis (122). (3) Allison, A. C. Correspondence, 1969. Virginia Dep. Nat. In most nurseries, fungicides are used to con- Resour. trol damping-off, and sprays are used during (4) Altman, P. L., and Dittmer, D. S. the season to control insects and other diseases. 1962. Biological handbook of growth. 608 p. fn southern nurseries the fusiform rusts on P. Fed. Am. Soc. Exp. Biol., Wash., D. C. taeda and P . elliottii and brown spot on P . palus- (5) Andreev, V. N. tris, and, in Lake States nurseries the sweetfern 1925. Pintrs peuce Griseb., 1n [Dendrology, vol. I, Gymnosperms p. 96.1 Ukraine State blister rust on P. banksiana, P. sylaesf.r'is, and Printing 0ffice. (In Russian.) P. Ttonder"oso are controlled by repeated applica- (6) Andresen, J. W. tions of fungicides. 1963. Germination characteristics of Pintts Generally, transplants or older age classes are rigida seed borne in serotinous cones. Broteria 32 (3-4) : 151-178. recommended for difficult sites. In the Lake (7) ----- States and Prairie Plains, for example, P. 1965. Stratification of seed of Pinus strobi- banksiana is grown as 1-0 or 11/z-0 stock for fornis. Tree Plant. Notes no. 72, p. 5-7, easy to average sites, 1-1 or 2-0 for difficult (8) Asakawa, S. sites, 1957. IStudies on hastening the germination and 2-1, l-2, or 2-2 for windbreaks. Most of the seeds of the five-leaved pines.] Gov. of the white pines are grown as 2-0 and 3-0 Forest Exp. Stn., Meguro (Tokyo) Bull. or as transplants 2-1 and 2-2. Tn specialized 100: 41-54. (In Japanese.) nurseries, pines are routinely grown as con- (e) Correspondence, June 17, 1969. Minist. Agric. tainer plants. Either young seedlings or seeds (Tokyo) planted For., Meguro, Japan. are directly in containers with a soil (10) Association- of Official Seed Analysts. mix. Depending on the nursery, partial shade 1965. Rules for testing seeds. Proc. -t-ssoc. is provided during the germination and seedling Off. Seed Anal.54(2): 1-112, illus. establishment phases. Normally, container- (11) Baker. L. A. grown pines are cultured 1 to 5 years before Correspondence, 1969. Oreg. State Forest. Dep., Dwight L. Phipps Forest Nursery, outplanting. Care must be taken not to grow Elkton, Oreg. pines in too small a container for too long a (12) Railey, L. H. time since they will become container-bound or 1939. The standard cyclopedia of horticul- rootbound. ture. 3 vol., 3,639 p. The Macmillan Co., New York. Methods of mulching. watering, shading, pest (13) Barnett, J. P. control, lifting, and other nursery operations Data filed 1968. USDA Forest Serv., South. have been described in several regional hand- Forest Exp. Stn., Nerv Orleans, La. books (95, 107, (14) 221, 223, 25/t). Each one con- 1969. Long-term storage of longleaf pine tains much useful information on nursery seeds. Tree Plant. Notes 20(2\: 22-25. production of pine seedlings. trDt -- pine species can prop- -1970. Flotation in ethanol affects storability Ali be vegetatively of spruce pine seeds. Tree Plant. Notes agated either by rooting or grafting (122, 178, 21 (4) : 18-19. 231). However, rooting success for most species (16) ---- and Mclemore, B. F. decreases rapidly when scions are taken from 1967. Improving storage of spruce pine seed. trees older than 5 years. PitttLs radiata, P. at- Tree Plant. Notes 18(2): 16. (17) -- s1l Mclemore, B. F. tenuata, P. den.siflora, and P. thunbergiana 1967. Study of spruce pine cone maturity and are relatively easy to root. But only P. radiata seed yield. Tree Plant. Notes 18(2): 18 is extensively propagated by rooting cuttings (18) Barton, L. V. under nursery and greenhouse conditions (230). 1930. Hastening the germination of some coniferous seeds. Am. J. Bot. 17: 88-1.15. Selected trees of many other species are cloned (19) Bates, C. G. by rooting of cuttings. Grafting also is rou- 1930. The production, extraction. and germi- tinely used to propagate rare material or to nation of lodgepole pine seed. U.S. Dep. clone individual plants, seed pro- Agric. Tech. Bull. 191, 92 p. as in orchard (20) Bennett, M. grams designed to produce genetically improved Communication, 1969. Arizona Cypress Gar- forest tree seed (122). dens, Sedona, Ariz.

631 PINUS

(21) Benson, D. A. (40) Cooling, E. N. G. Data filed 1969. USDA Forest Serv.. Eastern 1968. Fast growing timber trees of the low- Tree Seed Lab., Macon, Ga. land tropics. Pinus merkusii. No. 4. L69 p. (22) _ Commonw. For. Inst., Dep. For., Oxford Data filed 1970, USDA Forest Serv., Eastern Univ. Tree Seed Lab., Macon, Ga, (41) (23) Bevege, D. I. and Gaussen, H. -- 1970. In Indo China: Pinus m,erkusiana sn. 1965. An investigation of cone and seed nov. et non P. merkuszli maturity of slash pine in southern Jungh. et de Vriese. Queens- Trav. Lab. For. Toulouse Tome 1 Vol. 8. land. Aust. For.29(3): 135-148. p. (24) Bonninghausen, R. A. Art. T, 8 Communication, 1968. Florida Forest Serv., (42) Cnm, W. H., and Brack, C. G. E. Tallahassee. 1953. Performance of Scotch pine races under (25) Boskok, T. E. prairie conditions. For. Chron. 29G\: 1970. ISeed maturation period in Pinus aqi rAo brutia, Picea orientolis and Abies born- (43) Critchfield, H. M. muelleriana.l Orm. Arast. Enst. Tek. Correspondence, October 9, 1969. Glass Butll. 42: 64 p. (In Turkish.) Mountain Tree Farm and Nursery, St. (26) Boyd, R. J., Jr. Helena, Calif. Data filed 1970. USDA Forest Serv., Intermt. (44) Critchfield, W. B Forest and Range Exp. Stn., Moscow, Idaho. 195 ?. ileographic variatio n in P inus contor ta. (27) Britton, N. L., and Shafer, J. A. Maria Moors Cabot Found. Publ. 3, 118 p. 1908. North American trees. 849 p. Henry (45) __ Holt and Co., Nev'York. 1963. Hybridization of the southern pines in (28) Brown, J. H. California. South. Forest Tree Imnrov. 1969. Variation in roots of greenhouse grown Comm. Publ. 22: 40-48. seedlings of difrerent Scotch pine prove- (46) __ nances. Silvae Genet. 4(4\: 111-112, 1966. Phenological notes on Latin American (29) Burns. R. M. Pinus and. Abies, J. Arnold Arbor. 47(4\: Data filed 1968. USDA Forest Serv., South- 313-318. east. Forest Exp. Sta., Marianna, Fla. (47) and Allenbaugh, G. L. (30) Byrd, R. E. 1965. Washoe pine on the Bald Mountain Communication, 1968. Labelle. Florida. (31) Range, California. Madroiio 18(2): 68-64. Callaham, R. Z. (48) - and Krugman, S. L. 1962. Geographic variability in growth of 1967. Crossing the western pines at Placer- forest trees. In Tree Erowth. T, T. Koz- ville, California. Univ. Wash. Arbor. Bull. lowski (ed.), p. 311-325. Ronald Press Co., 30(4): New York. - 78*81,92. (32) (49) and Little, E. L.. Jr. --- 1963. Provenance research: investigation of 196-6. Geographic distribution of the pines of genetic diversity associated with geog- the world. U.S. Dep. Agric. Misc. Publ. raphy. Unasylva 18(2-B) :78-74. - 991, 97 p. (50) D. (33) and Liddicoei, A. R. Curtis. J. 1961. 1955. Effects of origin and storage method Altitudinal variation at 20 years in germinative ponderosa and Jeffrey pines. J. For. on the capacity of ponderosa - 59(11): 814-820. pine seed. USDA Forest Serv.f Intermt. (34) Forest and Range Exp. Stn. Res. Note Carlisle, H., and Brown. A. H. F. 26,5 p. 1968. Biological flora of the British Isles. (51) Dallimore, W., and Jackson, Pinus sylaestris L. J. Ecol. b6(1): 269_ A. B. 307. 1967. A handbook of Coniferae and GinkEo- (35) Cerepnin, V. L. aceae. Ed. 4, rev. by S. G. Harrison, 229 1964. [The importance of Pinus syluestris p. St. Martin's Press, New York. seed origin, weight, and colour in selec- (52) Dansbury, C. tion. I Sel. Drev. Porod er Vost. Sibiri. n. Correspondence, 1968. Washington Crossing 58-68. (In Russian.) State Forest Nursery, Titusville, New (36) Chapman, H. H. Jersey. 1922. A new hybrid pine (Pinus palustris X (53) Day, R. J. Pinus taeda). J. For. 20t 729_784. 196?. Whitebark pine in the Rocky Moun- (37) Church, T. W., Jr., and Sucoff. E. I. tains of Alberta. For. Chron. 4B(B) | Z7g_ 1960. Virginia pine seed viable two months 282. before natural cone opening. USDA Forest (54) Debazac, E. F. Serv., Northeast. Forest Exp. Stn. Res. 1964. Manuel des conifdres. 172 p. Ecole Nat. Note 102, 4 p. Eaux For6ts, Nancy. (38) Claveria, J. R. (55) Delevoy, G. 1953. Growing Benguet pine (pinus insu- 1935. Note preliminaire sur I'influence de laris) in Cebu Province.-philippine J. For. I'origine des graines chez le pin maritime. 9:57-76. Bull. Soc. Cent. For. Belg. 42(g): 9?_105. (39) Cocozza, M. A. (56) Dent, T. V. 1961. Osservazioni sul circlo riproduttivo di 1.947. Seed storage particular Pin.us with reference heldreicftii Christ. var. leucorlermis to the storage of seed of Indian forest Ant. del monte pollino. Accad. Ital. Sci. plants. Indian Forest Rec. (N.S.) Silvic. For. 10: 9T-110. (In Italian.) 7(1): 1-134. 632 PINUS

(52; Derr, H. J. (74) and Lester, D. 'f. 1955. Seedbed density affects longleaf pine 1970. Genetics of red pine. USDA Forest survival and grorvth. Tree Plant. Notes Serv. Res. Pap. WO-8, 13 p. no. 20, p.28-29. (75) Fritts. H. C. (58) Dimitroff, I. 1969. Bristlecone pine in the White Moun- 1926. [Study of the seed material of Pinus tains of California, Growth and ring-width peuce.l Ann. Univ. Sofia. Fac. Agric. 4: characteristics. Tree-Ring Pap. 4, 44 p. 259-306. (In Bulgarian.) Univ, Ariz. Press, Tucson. (5e) Dorman, K. W., and Barber, J. C. (76) Gifford, E. M., and Mirov, N. T. 1956. Time of flowering and seed ripening in 1960. Initiation and ontogeny of the ovulate southern pines. USDA Forest Serv., strobilus in ponderosa pine. Forest Sci. Southeast. Forest Exp. Stn. Pap,72,75 p. 6(1); 19-25. (60) Duff, C. E. (77 ) Goor, A. Y. 1928. The varieties and geographical forms 1955. Tree planting practices for arid areas. of Pinus pinaster Soland., in Europe and FAO Forest. Dev, Pap. No. 6, 126 p. South Africa, with notes on the silvi- (?8) and Barney, C.'W. culture of the species. S. Afr. Dep. For. 1968. Forest tree planting in arid zones. 409 Bull. 22-a, 55 p. p. Ronald Press Co., New York. ( 61) Duffield, J. W. (7e) Graber, R. E. 1951. Interrelationships of the California 1965. Germination of eastern white pine seed closed-cone pines with reference to Pinus as influenced by stratification. USDA muricata D. Don. Ph.D. Thesis Univ. Calif. Forest Serv. Res. Pap. NE-36, 9 p. Berkeley. 114 p. (80) Graber, R. E. (62) Data fiIed 1967. USDA Forest Serv., North- 1953. Pine pollen collections dates-annual east. Forest Exp. Stn., Upper Darby, and geographic variation. USDA Forest Penn. Serv., Calif. Forest and Range Exp. Stn. (81) R,es. Note 85, I p. Data fi]ed 1970. USDA Forest Serv., North- (63) Eden. J. east. Forest Exp. Sta., Upper Darby, Correspondence, 1969. Calif. Div. Forest., Penn. Davis State Forest Nursery, Davis, Calif. (g2) Griffin, J. R. (64) Edwards, M. U. 1962. Intraspecific variation in Pirzas sobi- 1954-55. A summary of information on niano Dotgl PhD thesis, 274 p. Univ. Pinus contorfa. For. Abstr. 15: 389-396; Ca1if., Berkeley. 16:3-13. (83) (65) Eliason, E. and Conkle, M. T. J. 1967. Early performance of knobcone and 1942. Data from cone collections of various Monterey pine hybrids on marginal timber species in New York. Ner.v York Conserv. p. sites. USDA Forest Serv. Res. Note PSW- Dep. Notes Forest Invest. 39, 1 156, 10 p. (66) and Hill, J. 1954. Specific gravity as a test for cone ripe- (84) Haasis, F. W., and Thrupp, A. C. ness with red pine. Tree Plant. Notes no. 1931. Temperature relations of lodgepole 17. n. l-4. pine seed germination. Ecol. 12: 728-744. (67 ) Ellis, R. W. (85) Haller, J. R. 1969. Correspondence, August 20, 1969. 1957. Taxonomy, hybridization and evolution USDA Forest Serv., Mt. Sopris Nursery, in Pinus ponderosa and Pinus jeffregi. Carbondale, Colorado. PhD thesis 169 D. Univ. Calif.. Los (68) Fenton, R. H., and Sucoff, E. I. AnEeles. 1965. Effects of storage treatments on the (86) ripening and viability of Virginia pine 1967. A comnarison of the mainland and seed. USDA Forest Serv. Res. Note NE- islan

633 PINAS

(92) Heidmann, L. J. (ed.), p. 98-109. Akad. Nauk SSSR Sibirs- Data filed 1969. USDA Forest Serv., Rocky koe Otdel, Vol. 62 [Engl. Transl. TT65- Mountain Forest and Range Exp. Stn,, 50123, CFSTI, U. S. Dep. Commercel. Flagstaff, Ariz. (110) Lebkov, V. F., and Cherednikova, Yu. S. (93) Heit, C. E. 1963. Fruit bearing of stone pine forests of 1958. The effect of light and temperature the Lena-Ilim interfluvial area. .In Fruit- on germination of certain hard pines and - ing of the Siberian stone pine in east suggested methods for laboratory testing, Siberia. A. P. Shimanyuk (ed.), p. 35-79. Proc. Assoc. Off. Seed Anal. 48: 111-117, Akad. Nauk SSSR Sibirskoe Otdel, Vol. (94) __ 62. IEngl. Transl. TT65-50123, CFSTI, 1963. Balkan pine-promising new exotic U. S, Dep. Commercel. conifer. Am. Nurseryman 118(12) : 10, 11, (111) Jacaline, D. V., and Lizardo, L. 32, 34, 36. 1958. Silvieal characteristics of Benguet pine (e5) _ (Pinus insularis Endl.). Philippines Bur. 1964. Tips on growing healthy, vigorous coni- For. Silvical Leafl. 2, 32 p. fer seedlings and transplants. N. Y. (112) Jones, L. Christmas Tree Grow. Bull. 2(1) : 5 p. 1962. Recommendation for successful storage [also Am. Christmas Tree J., Feb. 1966]. of tree seed. Tree Plant. Notes no. 55, p. (96) _ 9-20. (113) 196?. Laboratory germination studies and 'What suggested testing methods for 1.0 less com- 1966. Storing pine seed: are best mon and exotic Pi,nus species. (Unpub- moisture and temperature conditions? lished. ) - Georgia Forest Res. Counc., Res. Pap. (97) _ 42, 8 p. 1967. Propagation from seed. Part 9. FaIl (114) _ and Benson. D. sowing of conifer seeds. Am. Nurseryman Data filed 1969. USDA Forest Serv.. South- 126(6): 10-11. 60-69. east. Forest Exp. Stn., Macon, Ga. (98) _ (115) Kamra, S. K. 1967. Propagation from seed. Part 10. Stor- 196?. Studies on storage of mechanically age method for conifer seeds. Am. damaged seed of Scots pine (Pinus Nurseryman 126(8): 14-54 (not inclu- syluestris L.). Stud. For. Suecica 42, 19 p. sive). (116) _ (99) _ 1969. Investigations on the suitable germi- 1968. Propagation from seed. Part 12. Grow- nation duration for Pinus syloestris and ing choice, less common pines. Am. Nurs- Picea abies seed. Stud. For. Suecica 73, eryman 127 (2\: L4-15, 712-120. 16 p. (100) (117) __ and Simak. M. 1968. Thirty-five year's testing of tree and 1968. Germination studies on Scots pine -- shrub seeds. J. For. 66(8): 632-634. (Pinus sylaestris L.) seed of different (101) provenances under alternating and con- 1969. Propagation from seed. Part 19. Test- stant temperatures. Stud. For. Suecica 62, ing and growing Scotch pine seeds from 14 p. - difrerent sources. Am. Nurseryman 129(7) : (118) Karschon, R. 10-15,110-118. L961. Studies in nursery practice for pines. (102) _ La-Yaaran 11(1): 1-12. Correspondence, 1969. New York Agric. Exp. (119) Kmecza, N. S. Stn., Geneva, N. Y. 1970. Using tree shakers for pine cone col- (103) and Eliason, E. J. lections in Region 8. Tree Plant. Notes 1940. Coniferous tree seed testing and factors 21(1): 9-11. affecting germination and seed quality, (120) Kraus, J. F. - N. Y. State Agric. Exp. Stn. Tech. Bull. 1963. The Olustee arboretum. USDA Forest 255, 45 p. Serv., Res. Pap. SE-4,47 p. (104) Holst, M. (121) Krugman, S. L. 1962. Seed selection and tree breeding in 1966. Artificial ripening of sugar pine seeds. Canada. Can. Dep. For., Forest Res. Rr. USDA Forest Serv. Res. Pap. PSW-32' Tech. Note 115, 20 p. 7p. (105) Hopkins, E. R. (t22\ 1960. Germination stimulation in Pinus pi- Data filed 1969. USDA Forest Serv., Pac. naster. West. Aust. For, Dep. Bull. 66, Southwest Forest and Range Exp. Sta., 1o p. - Institute of Forest Genetics, Placerville, (106) Hyun, S. K. Calif. 1962. Improvement of pines through hybrid- (123) ization. IUFRO Proc. 13, Vol. 1 (2), 2 p. Krtissmann. G. (107) 1960. Die Nadelgehiilze. Ed' 2, 335 p. Paul International Crop Improvement Association. Hamburg. 1963. Minimum seed certification standards. Parey, Berlin and Int. Crop Improv, Assoc. Publ. 20,128 p. (124) Lamb, G. N. (108) International Seed Testing Association. 1915. A calendar of the leafing, flowering and 1966. International rules for seed testing. seeding of the common trees of the'Weather east- Proc. Int. Seed Test. Assoc. 1966: 1-152. ern United States. U. S, Mon. (109) Iroshnikov. A. I. Rev., Suppl. 2, p. 3-19. 1963. [Fruit bearing of stone pine forests in (125) Larson, M. M. the northwestern part of the eastern 1966. Racial variation in ponderosa pine at Sayan.l In Fruiting of the Siberian stone Fort Valley, Arizona. USDA Forest Serv. pine in east Siberia. A. P. Shimanyuk Res. Note RM-73. 7 D. 634 PINUS

(126) LeBanon, R. K., and Roe, E. I. USDA Forest Serv. Res. Pap. NE-134, 1945, Hastening the extraction of jack pine 16 p. sceds. J. For. 43: 820-821. (145) Littlefield, E. W. (127) Lebrun, C. Data filed 1932. New York State Dep. Con- 196?. [Separation of (full and empty) seeds serv. by specific gravity measurement through (146) Lizardo, L. immersion in liquids.] Rev. For. Franc. 1950. Benguet pine (Pinus insr,rlaris Endl.) 19(11): 786-789. (In French.) as a reforestation crop, Philippine J, For. (128) Leloup, M. 7(1-4): 43-60. 1956. Tree planting practices in tropical (14?) Loiseau, J. Africa. FAO Forest. Develop. Pap. 8, 306 1945. Les arbres et ]a for6t. Vol. 1, 204 p. p. Vigot freres, Paris. (129) Letourneux, C. (148) Loock, E. E. M. 1957. Tree planting practices in tropical 1950. The pines of Mexico and British Hon- Asia. FAO Forest. Develop. Pap. 17, 172 duras. Union S. Afr. Dep. For. Bull. 35, p. 244 p. (130) Libbv, W. J. (149) Lotan, J. E. Correspondence, February 26, 1968. Sch. Data filed 1969. USDA Forest Serv., Intermt. For. and Conserv., Univ. Calif., Berkeley. Forest and Range Exp. Stn., Bozeman, (131) Lindquist, C. H. Mont. 1962. Maturity of Scots pine seed. Can. Dep. (150) Luckhotr, H. A. Agric., Res. Br., Summary report for the 1964. Natural distribution, growth, and Forest Nursery Station, Indian Head, botanical variation of Pinus caribaea and Saskatchewan, p. 20-21. its cultivation in South Africa. Ann. Univ. ( 132) Little, E. L., Jr. Stellinbosch vol. 39, ser. A, no. 1, 160 p, 1938. The earliest stages of pifryon cones. (151) Magini, E. USDA Forest Serv., Southwest. Forest 1955. IConditions of germination of Aleppo and Range Exp. Stn. Res. Note 46, 4 p. and Italian stone pines.l Ital. For. Mont. ( 133) _ 10(3): 106-124. (In Italian.) 1938. Stages of growth of pinyons in 1938. (152) and Tulstrup, N. P. USDA Forest Serv., Southwest. Forest 1955. Tree seed notes. FAO For. Develon. and Range Exp. Stn. Res. Note 50,4 p. Pap. 5, 354 p. (134) (153) -Malac, B. F. 1940. Suggestions for selection cutting of 1960. More on stratification of pine seed in pifryon trees. USDA Forest Serv.. South- polyethylene bags. Tree Plant. Notes no, - west. Forest and Range Exp. Stn. Res. 42, p.7-9. Note 90, 3 p. (154) Mastrogiuseppe, R. J. (135) 1968. Geographic variation in foxtail pine. -- 1941. Managing woodlands for pifiyon nuts. USDA Forest Serv., Pac. Southwest For- Chron. Bot. 16: 348-349. est and Range Exp. Stn., Progress Rep., (136) _ 15 p. Inst. Forest Genet., Placerville, Calif. 1950. Southwsslsln frsgs-a guide to the (155) Mclntyre, A. C. native species of New Mexico and Arizona. 1929. A cone and seed study of the mountain U.S. Dep. Agric., Agric. Handb. 9, 109 p. pine (Pir'nrs pungens Lambert). Am. J. (137) Bot. 16: 402-406. 1968. Tu'o new pinyon varieties from Ari- (156) Mclemore, B. F. zona. Phytologia 17(4) :329-342. 1961. Hila of full and empty longleaf pine (138) - and Dorman, K. W. seeds are distinsuishable. Forest Sci. 7: 1952. Geographic differences in cone open- 246. ing in sand pine. J. For. 50(3): 204-205. (157) (139) -_ and Dorman. K. W. 1961. Storage of longleaf pine seed. Tree 1952. Slash pine (Pinus elltottiil, its nomen- Plant. Notes no. 47, p. 15-19. clature and varieties. J, For. 50: 918-923. ( 158 ) - (140) and Dorman, K. W. --- 1965. Pentane flotation for separating full 1954. Slash pine (Pinus eiliottii) including and empty longleaf pine seeds. Forest Sci. South Florida slash pine. USDA Forest 17:242-243. - Serv., Southeast. For-est Exp. Stn. Pap. (15e) 36, 82 p. Data filed 1968. USDA Forest Serv.. South. (141) and Righter, F. L Forest Exp. Stn., New Orleans, La. 1965. Botanical descriotions of fortv arti- - pine and Barnett. J. P. ficial hybrids. U.S. Dep. Agric. Tech. 1967. Bull. 1345, 47 p. Effective stratification of spruce - seed. Tree Plant. Notes 18(2): 17-18. Little, S. 1941. Calendar of seasonal aspects for New (161) and Czabator, F. J. Jersey forest trees. Forest Leaves 31(4): 1961. Length of stratification and Eermina- t-2, t3-14. tion of loblolly pine seed. J. For. 58: 267- ( 143) - 269. 1959. Silvical characteristics of pitch pine (162) Mersen, F. (Pinus risida). USDA Forest Serv., 1963. Ecotypic variation in Pinus strobus L. Northeast. Forest Exp. Stn. Pap. 119,22 p. Ecol.44: 716-72'1. Mikhalevskaya, O. B. 1969. Local seed sources recommended for 1960. [The biology of Pinus pumila Rgl. in loblolly pine in Maryland and shortleaf Kamchatka.l Nauch. Dokl. Vyssh. Shkoly, pine in New Jersey and Pennsylvania. Biolog. Nauk.3: 136-141. (In Russian.)

635 PINAS

(164) Miller, H., and Lemmon, P. E. (185) Pravdin, L. F. 1943, Processing cones of ponderosa pine to 1963. Selection and seed production of the extract, dewing, and clean seed. J. For. Siberian stone pine. /ro Fruiting of the 4L(r2\: 889-894. Siberian stone pine in east Siberia. A. P. (165) Mirov, N. T. Shimanyuk (ed.). Akad. Nauk SSSR 1936. A note on germination methods for Sibirskoe Otdel Vol. 62, p. 1-20. [EnSl. coniferous species. J, For. 34(7'l 719- Transl. TT65-50123, CFSTI, U. S. Dep. 723. Commerce.] (166) (186) _ -- 1946. Viability of pine seed after prolonged 1964. Scots pine variation, intraspecific cold storage. J. For.44(3): 193-195. taxonomy and selection. 208 p. Acad. (16?) Nauk SSSR [Engl. Transl. TT69-55066, 1956. Photoperiod and flowering of pine. CFSTI, U.S. Dep. Commerce.l Forest Sci. 2: 328-332. (18?) Rafn, J. (168) -_ 1915. The testing of forest seeds during 25 1962. Phenology of tropical pines. J. Arnold years, 1887-1912. 91 p. Langkjaers Bog- Arbor. 43(2') : 218-2L9. trykkeri, Copenhagen. (Printed for private (169) cireulation. ) 1967. The genus Pinna. 602 p, Ronald Press (188) Read, A. D. Co., New York. 1932, Notes on Arizona pine and Apache (170) -Muller, C. A. pine. J. For. 30: 1013-14. Data filed 1968. Ala. Div. For.. Edward A. (189) Read, R. Hauss Nursery, Atmore, Ala. Personal communication. 1969. USDA (171) Miiller, K. M. Forest Serv., Rocky Mountain Forest and 7932. Pinus peuce, the Macedonian white pine Range Exp. Stn,, Lincoln, Nebr. as a substitute fot Pinus strobus. Blister (190) Rehder, A. Rust News 16(3) Suppl.:62-70, 1940. Manual of cultivated trees and shrubs (172) Nather, H. hardy in North America. Second ed,, 996 1958. IGermination of Swiss stone pine p. The Macmillan Co., New York. seed.l Cent. Ges. Forstwesen 75(1): 161- (191) Rietveld, W. J. 170. (In German.) Data filed 1969. USDA Forest Serv., Rocky (173) _ Mountain Forest and Range Exp. Sta., Correspondence, January 9, 1969. Institut Flagstaff, Ariz. fiir Waldbau, Austria. (192) Robbins, G. T. (174) Nederlandsche Boschbouw Vereeniging. Data filed 1945. USDA Forest Serv., South- 1946. Boomzaden: Handleiding inzake het west Forest and Range Exp. Stn., Inst., oogsten, behandelen, bewaren en uitzaaien Forest Genet., Placerville, Calif. van boomzaden. 171 p. Wageningen, (In (193) Roe, E. L Dutch.) Data filed 1939-1941. USDA Forest Serv.. (175) Newcomb, G. B. North Cent. Forest Exp. Stn., St. Paul, 1962, Geographic variation in Pinus at- Minn. tenuata Lem. PhD thesis. 190 o. Univ. (194) Rohmeder, E., and Loebel, M. Calif., Berkeley. 1940. Keimversuche mit Zirbelkiefer. Forst- (176) Nyman, B. wiss. Centralbl. 62: 25-36. (In German.) 1963. Studies on the germination in seeds of (195) Rossi, E. Scots pine. Stud. For. Suecica 2, 164 p. 1929. Sulla eerminabilita de seme di Pinus (177) Ohmasa, M. maritima ln rapporto alla temperatura. 1956. Tree planting practices in temperate Ist. Bot. R. Univ. Pavia Atti., Ser. IV, Asia: Japan. FAO For. Develop. Pap. 10, 1: 107-115. (In ltalian.) 156 o. (196) Rudolf, P. O. (1?8) O'Rourke, F. L. S. Data filed 1970. USDA Forest Serv., North 1961. The propagation of pines. Proc. Int. Cent. Forest Exp, Stn., St, Paul, Minn. Plant Propag. Soc. 11: 16-22, (197) Rudolph, T. D., Schoenike, R. 8., and Schantz- (179) Otter. F. L. Hansen, T, 1933. Idaho's record trees. Idaho For. 15: 1959. Results of one-parent progeny tests 37-39. relating to the inheritance of open and (180) Ouden, P. den, and Boom, B. K. closed cones in jack pine. Univ. Minnesota 1965. Manual of cultivated conifers. 526 p. For. Note 78, 2 p. Martinus Nijhotr, The Hague. (198) Sargent, C. S. (181) Pearson, G. A. 1905. Manual of the trees of North America, 1931. Forest types in the southwest as de- 826 p. Houehton, Miffiin and Co., Boston termined by climate and soil. U.S. Dep. and New York. Agric. Tech. Bull. 247,144 p. (199) (182) Pennsylvania Department of Forest and Water. 1965. Manual of trees of North America Data filed (n. d.). Mont Alto, Penn. (exclusive of Mexico). Ed. 2, 934 p. Dover (183) Posey, C. E., and McCullough, R. B. - Pub., Inc., New York. 1969. Tenth year results of a shortleaf pine (200) Savory, B. M. seed source study in Oklahoma. Okla. 1962. The taxonomy of Pinus khasya ( Royle) Agric. Exp. Stn. Bull. 8-668, 14 p. and. Pinus insularis (Endlicher). Empire (184) Poynton, R. J. Forest. Rev. 41 (1): 6?-80. 1961. A guide to the characteristics and (201) Schmitt, D., and Namkoong, G. uses of trees and shrubs quoted in the 1965. Pine species in the Harrison Experi- price list of the Forest Department. Re- mental Forest Arboretum. USDA Forest pub. S. Afr. Bull. 39, 50 p.- Serv. Res. Pap. SO-18, 18 p.

636 PINUS

(202) Schubert, G. H. (221) Stoeckeler, J. H., and Jones, G. W. 1,952. Germination of various coniferous 1957. Forest nursery practice in the Lake seeds after cold storage. USDA Forest States. U.S. Dep. Agric., Agric. Handb. Serv., Calif. Forest and Range Exp. Stn. 170, I24 p. Res. Note 83, 7 p. (222) and Rudolf, P. O. (203) 1956. Winter coloration and gron'th of jack -_ 1955. Effect of ripeness on the viability of pine in the nursery as affected by seed sugar, Jeffrey, and ponderosa pine seed. - source. Z. tr'orstgenet. Forstpflanzenzuecht. Soc. Am. For. Proc. 55: 67-69. D: tol-loD, (204) _ (223) and Slabaugh, P. E. Data filed 1969. USDA Forest Serv., Rocky 1965. Conifer nursery practice in the prairie- Mountain Forest and Range Exp. Stn., plains. U.S. Dep. Agric., Agric. Handb. Flagstaff, Ariz. - 279. 93 p. (205) -.- Heidmann, L. J., and Larson, M. M. (224) Straun, W. H. 1970. Artificial reforestation practices for Correspondence, 1969. N. C. State Forest the southwest. U.S. Dep Agric., Agric. Serv., Morganton, N. C. Handb. 370,25 p. (225) Sudworth, G. B. (206)- and Rietveld, W. J. 1908. Forest trees of the Pacific slope. 441 p. Data filed 19?0. USDA Forest Serv., Rocky USDA Forest Serv., Wash., D.C. Mountain Forest and Range Exp. Stn., (226) Swingle, C. F. (compiler). Flagstaff, Ariz. 1939. Seed propagation of trees, shrubs, and (20?) Sen Gupta, J. N. forbs for conservation planting. SCS-TP- 1936. Seed weights, plant percents, etc., for 27, USDA Soil Conserv. Serv., 198 p. forest piants in India. Indian Forest Rec. Wash., D.C. (n. s.) Silvic. 2(5):175,22I. (22?) Swofford. T. F. (208) Shafiq, Y., and Omer, M. 1958. Stratification harmful to some lob- 1969. The effect of stratification on Eermina- lolly and slash pine seed. Tree Plant, tion of Pinus brutiu, seed. Mesopot. Agric. Notes No. 32, p. 5-6. 4: 96-99. (228) Takayama, Y. (209) Shaw. G. R. 1966. [Studies on the seed orchard of Jap- 1914. The genus Pil'as. Arnold Arbor. Publ. anese red pine (Prlnus densiflora Sieb. & Zucc.). I. On the 1000-seed weight of the 5, 96 p. grafted (210) Sherry, S. P. crops from the clones of Jap- 1947. The potentialities genetic anese red pine.l J. Jap. For, Soc. 48(5): of research 199-208. (In Japanese.) in South African forestry. Pretoria, Dep. For., 11 p. (229) Thompson, N. S. (211) Shoulders, E. 1968. The response of pine cone scales to 1961. Effect nurserybed changes in moisture content. Holzfors- of density on lob- chung 22(4): 124-125. lolly and slash pine seedlings. J. For. b9: 576-579. (230) Thulin, I. J., and Faulds, T. 1968. The use of cuttings in (212) Simak, M., Ohba, K., and Suszka, B. the breeding and 1961. Effect afforestation of Pittus radiata. N. Z. For. of X-irradiation on seeds of dif- 13(1): 66*77. ferent weight from individual trees of Scots pine (Pittus syluestris L.). Bot. No- (231) Ticknor, R. L. tiser 114(3): 300-312. 1969. Review of the rooting of pines. Proc. Plant Propag. 132-737. (213) Slayton, S. Int. Soc.19: Data filed 1969. USDA Forest Serv., J. W. (232t Tkachenko. M. E. Toumey Nursery, Watersmeet, Mich. 1939. [General forestry.] ?45 p. Goslestek- (214) Smouse, P. E. hizdat, Leningrad. (In Russian.) 1970. Population studies in the Eenus Pizzs (233) Troup, R. S. L. PhD thesis, 126 p. N. C. State Univ., 1921. The silvicuiture of Indian trees. Vol. Raleigh. 3, p. 1013-1095. Clarendon Press, Oxford, (215) Snorv, A. G., Jr. (234) Turner. E. E. 1960. Silvical characteristics of Vireinia Correspondence, 1968. Louisiana State For. pine. USDA Forest Serv., Northeast. For- Comm., Woodworth, La. est Exp. Stn., Stn. Pap. 131, 22 p. (235) USDA Forest Service. (216) Squillace, A. E. 1948. Woody-plant seed manual. U.S, Dep. 1966. Geographic variation in slash pine. Agric. Misc. Publ. 654, 416 p. Forest Sci. Monogr. 10, 56 p. (236) _ (217) and Bingham, R. T. Data filed (n.d.). South. Forest Exp. Stn., -- 1958. Localized ecotypic variation in r.estern New Orleans, La. white pine. Forest Sci. 4(1): 20-33. (237 ) (218) _ and Silen, R. R. -- Data filed 1928-193?. Intermt. Forest and 1962. Racial variation in ponderosa pine. Range Exp. Stn., Moscow, Idaho. Forest Sci. Monogr. 2, 27 p. (238) __ (219) Steinhoff, R. J. Data filed 1952. North Cent. Forest Exn. 1964. Taxonomy, nomenclature, and varia- Stn., St. Paul, Minn. tion rvithin the Pinus fl,enil.is complex. ( 239 ) PhD thesis. 81 p. Mich. State Univ. --- Data filed 1957. 1959. 1966-68. Eastern Tree (220) Steven, H. M.. and Carlisle, A. Seed Lab., Macon, Ga. 1959. The native pinervoods of Scotland. (210) _ 368 p. Oliver and Boyd, Edinburgh and Data filed 1966-19?0. Intermt. Forest and London. Range Exp. Stn., Moscow, Idaho.

637 PINAS

(24r\ (258) Wellner, C. A. Data filed 196?. Project 2302 fot developrnent 1962. Silvics of western white pine. USDA of blister rust resistant rvestern rvhite Forest Serv., Intermt. Foresf and Range - pine. Intermt. Forest and Range Exp. Exp. Stn. Misc. Publ. 26,24p. Stn., nfoscorv, Idaho. (259) Wells, O. O. (242\ __ 1964. Geographic variation in ponderosa Data filed 1969. Bend Nursery, Bend, Oreg. pine. 1. The ecotypes and their distribu- (243\ tion. Silvae Genct. 13(4): 89-103. ( ___ -- Data filed 1969. Coeur d'Alene Nursery, 260 ) 1969. Results of the southwide pine seed Coeur d'Alene, Idaho. source study through 1968-69. Tenth (244\ South" Conf. Forest Tree Improv. Proc.: --- Data filed 1969. Lucky Peak Nursery, Boise, tt1-129. Idaho. (261) and Wakeley, P. C. (245\ 1966. Geographic variation in survival, Data filed 1969. Mt. Sopris Tree Nursery. growth and fusiform-rust infection of Carbondale, Col. - planted lobloliy pine. Forest Sci. Monogr. (246) -_ 11. 40 p. Data filed 1969. Placerville Nursery, Placer- (262) and Wakeley, P. C. ville, Calif. 1970. Variation in shortleaf pine from sev- (247\ -- eral geographic sources. Forest Sci. 16(4): Data filed 1969. Div. Timber Manage., Re- 415-423. gion 4, Ogden, Utah. (263) Western Forest Tree Seed Council. (247a)- Uyeki, Homiki. 1966. Sampling and service testing rvestern conifer seeds. 36 p. 1927. The seeds of the genus Pirtus as an (264) aid to the identification of species. Suigen Wrieht, J. W. p. (Korea.) Coruespondence, 1969-19'/0. Sch. For., Mich. Agric. For. Coll., Bull. 2, 129 State Univ., East Lansing. (248) Vandemillen, E. (265) Data filed 1969. USDA Forest Serv., Eveleth 1970. Genetics of eastern white pine. USDA Nursery, Eveleth, Minn. Forest Serv. Res. Pap. WO-9, 16 p. (249) VanDeusen, J. H. ( 2ri6 ) - Data filed 1969. USDA l'orest Serv., Ilocky 1962. Genetics of forest tree improvement. . Mountain Forest and Range Exp. Stn., FAO For. and Forest Prod. Stud. 16, Rapid City, S. Dak. 399 P. (250) Veracion, V. P. 1267 ) Lemmien, W. L.. and Bright, J. 1964. Correiation of cone size and rveight -- 1963. Geographic variation in eastern white with the numbers, size and weight of seeds pine-6 year results. Mich. Agric. Exp, of Benguet pine (Pizas in.sularis, Endl.). Stn. Q. Bull. 45(4): 691-697. Philippines Bur. For. Occas. Pap. 16, 11 p. (268) - p2,lsy, S. S., Polk, R.8., Jokela, J. J. (25t\ and Read, R. A. 1966. Correlation of the size of seeds to the - 1966. Performance of Scotch pine varieties in germination and early growth of Benguet the North Central Region. Siivae Genet. - pine (Pinus insularis, Endl.). Philippines 15(4): 101-110. Bnr. For. Occas. Pap.26,7 p. (269) Yanagisawa, T. (252) Vines, R. A. 1965. Effect of cone maturity on the viability 1960. Trees, shrubs, and'rvoody vines of the and longevity of coniferous seed. Gov. Southwest. 1,104 p. Univ. Texas Press, Forest Exp. Stn., Meguro (Japan) Bull. Austin. 172:45-94. (253 Wahlenberg, W. G. (270) York, H. H., and Littlefield, E. W. 1946. Longleaf pine. 429 p. Charles Lathrop 1942. The naturalization of Scotch pine, Park For. Found.. Wash.. D.C. northeastern Oneida County. J. For. 40(7\: 552-559. (254) Wakeley, P. C. pines. (277) Zarger,T. G. 1954. Planting the southern U.S. Dep. Tenn. Agric., Agric. Monogr. 18, 233 p. Correspondence, October 25, 1968. Valley Auth., Norris, Tenn. (255) and Barnett, J. P. -- pine (272) Zavarin, E., Critchfield, W. 8., and Snajberk, K. 1968. Viability of slash and shortleaf con- seed years. For. 840- 1969. Turpentine composition of Pinus stored for 35 J. 66: torta X Pinus banksiana hybrids and hy- 841. brid derivatives. Can. J. Bot. 47(9\: 1443- (256) Wappes, L. 1932. Wald und Holz ein Nachschlagebuch (273\ Zobel. B. fiir die Praxis der Forstrvirte, Holzhindler 1953. Geographic range and intraspecific und Holzindustriellen. Vol. 1, 872 p. J. variation of Coulter pine. Madrofro 12: Neumann, Berlin. t-1. (257) Weidman. R. H. (274\ Zobel, D. B. 1939. Evidence of racial influences in a 25- 1969. Factors affecting the distribution of year test of ponderosa pine. J. Agric. Res. Pinus pungerls, an Appalachian endemic. 59:855-888. Ecol. Monogr. 39(3): 303-333.

638