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PINAS
Pinaceae-Pine family
Growth habit, occurrence, and use.-The ge-nus Pinus, one of the largest and most importantof the coniferous genera, comprises about 95species and numerous varieties and hybrids.Pines are widely distributed, mostly in theNorthern Hemisphere from sea level (Pi'nuscontorta var. contorta) to timberline (P. albi-cantl;i,s). They range from Alaska to Nicaragua,from Scandinavia to North Africa. and fromSiberia to Sumatra. Some species, such as P.syluestris, are widely distributed-from Scot-land to Siberia-while other species have re-stricted natural ranges. Pinus canariensis, forexample, is found naturally only on the CanaryIslands, and P. torreyana numbers only a fewthousand individuals in two California localities(table 1) (4e).
Forty-one species of pines are native to theUnited States. These species are also the mostwidely planted in the United States. Plantinghas extended the range of a number of them,including P. strobus, P. banksinna, P. rad'inta,P. pond,eroso var. scopulorum, P. resi,nosa, and,P. oirginiana (71, 266). Many of the nativeeastern pines, especially those from the southernStates. do not do well in the western States. Thesame appears to be true for many western pineswhen they are planted in the eastern States(120,201).
Many introduced pines have been planted andgrown successfully in the United States. Fourof these-P. sglaestris, P. thunbergi,ana, P.densiflora, and P. nigra-have become natural-ized in parts of New England and the LakeStates (261r, 270).
Many other pine species have been suce.ess-fully planted outside their native range. Theseinclude P. patuln in South Africa; P. radinta inSouth Africa, New Zealand, Australia, andSouth America ; P. insularzs in East Africa; P.etliottii var. elliottii, P. taedn, P. pinaster, and,P. palustris in South Africa, New Zealand, andAustralia; P. ponderoso in Australia and New
. 'Timber Management Research, USDA Forest Serv-rce.
'Pacific Southwest Forest and Range Exp. Stn.
598
PINUS L. Pineby Stanley L. Krugman 1 and James L. Jenkinson 2
Zealand; P. canariensis in North Africa andSouth Africa; P. cari.bea in South Africa andAustralia; P. halepereszs in South America; P.muricata in New Zealand and Australia; P.sgluestris, P, strobus, P. contorta, and P. ni'grain Europe; P. merkusii in Borneo and Java 128,152, 169, 266).
The pines are evergreen trees of variousheights,-often very tall but occasionally shrubby(table 3). Some species, such as P.lnmbertionn,P. monticola, P. ponderosa, antd. P. strobtr's, growto more than 200 feet tall, while others, as P.cembroides and P. Ttumila, may not exceed 30feet at maturity.
Pines provide some of the most valuable tim-ber and are also widely used to protect water-sheds, to provide habitats for wildlife, and toconstruct shelterbelts. The bulk of naval storesstill comes from pines, and the seeds of somespecies are a valuable food source. Increasingly,pines are planted to improve man's environ-ment.
Sixty-five species and varieties which are nowbeing planted or which have a potential in theUnited States are listed in table 1; included are40 species and varieties native to the UnitedStates, one to Mexico, one to the Caribbeanregion, 11 to Europe, Africa and the near east,and 12 to Asia.
Geographic races and hybrids.-The impor-tance of planting seeds or seedlings from theproper source cannot be stressed to strongly.Seed origin is extremely important in determin-ing the ability of a species to grow and succeedin a given environment. Many pines with anextensive range, as well as some of limitednatural range, have developed geographic racesthat are morphologically and physiologicallydistinct (gz). These differences make each racebest suited for growing in certain environments.As a general rule, seeds from sources in moistregions are smaller and produce faster growingand less deeply rooted seedlings than seeds fromsources in drier regions. Southern seed sourcescommonly differ from northern sources by beingfaster growing, capable of growing longer in aseason, more susceptible to damage by winter
freezes, less susceptible to late spring and earlyautumn frosts; and having seed dormancy thatcan often de overcome with a shorter presowingtreatment (122, 218, 260, 266).
For many of the pines, detailed data aboutgeographic rdces are not available. In somecases our understanding of races of pines nativeto the United States comes from plantings inother countries. When appropriate, this infor-mation is included in the following species sum-maries:
Pinus attenuata-From morphological andnursery studies, knobcone pine in California canbe separated into two major groups, one northand the other south of the Monterey-San LuisObispo County line. Seed weight tends to in-crease from north to south. and seeds of north-
PINUS
ern origins require a longer stratification period(3 wee[s or more) than seeds of southern origin(less than 3 weeks). Seedlings from northernsources tend to be more frost resistant thanthose from southern sources. These differencesamong sources appear to be clinal and reflectthe latitudinal disliibution of the species (175).
Pinu,s balfouriana-Seeds of northern origin(Lake Mountain, California) are longer thanthose of southern origin (Mineral King, Cali-fornia) and have persistent seed wings com-pared to the detachable wings of southernsources (154.).
Pinus banksiana-Sources differ in seed size,growth, form, and susceptibility to insect anddisease damage. Seeds tend to be larger fromthe warmer parts of the range (71) . In Minne-
TAsLp l.-Pinus: nomenclature, occutrence, and uses; data comTtilers
Scientific names and synonyms Common names Occurrence IJses' Data compilersfor the species
P. albicaulic Engelm.
P. arietata Engelm. ...P. balfouriana var.
aristata (Engelm.) Engelm.
P. armandii Franeh.
P. artcnua'a Lemm.P. tuberculatc Gord.
P. balfouriana Grev. & Balf. --------
P. bankeiana Lamb.P. diaaricata (Ait.) Sudw.
P. brutia Ten.P. halepensis vat. brutia
(Ten.) Elwes & HenryP. canariendt C. Smith
P. caribaea MoreletP. bahamensis Griseb.P, hondurensis Loock
whitebark pine Subalpine in the northernRocky Mountains andCoast Mountains ofBritish Columbia throughthe Cascade Range tothe southern SierraNevada.
bristlecone pine, Small, scattered subalpinefoxtail pine, areas in California,hickory pine. Nevada, Utah, Arizona,
and New Mexico.Armandpine Moderate-highelevations
from central and south-western China to north-ern Burma and south-eastern Tibet; Taiwanand Hainan.
knobcone pine Rocky slopes and ridgesfrom southwesternOregon and Californiato northwestern BajaCalifornia.
foxtail pine, Subalpine in KlamathBalfour pine. Mountains and southern
Sierra Nevada,California.
jack pine, Canada, from the Mackenziescrub nine. River and Alberta tobanksiana Nova Scotia and southpine, black into the Great Lakespine, gray region and New Englandpine. States.
Calabrian pine Crete, Cyprus, and Lebanonnorth through Turkey.
Canary Islandprne,Canary pine.
Caribbean pine
Dry, exposed slopes of thecentral and westernCanary Islands.
Bahama Islands, westernCuba, Isle of Pines, andCentral America fromBritish Honduras toNicaragua.
T,W,E - R. J. Steinhoff.
w,E S. L. Krugman.
H, W Do.
T, H, W, D. H. Dawson.S, E.
H,E
T,E
S,E
T,W,E
T,E
G. H, Schubert.
P. O. Rudolf.
S. L. Krugman.
P. O. Rudolf.
S. L. Krugman.
599
PINAS
Tnnln l.-Pi,nus: nomenclature, occu,rrence, and uses; data compilers-Continued
Scientific names and synonyms Common names Occurrence Uses' Data compilersfor the species
P. cembra L.- - ---P, montana Lam.
P. cembroidet Zucc.-
P. clauta (Chapm.) Vasey
P. con,orta Dougl. var.con,orta
P. contorta vat. latiloliaEngelm.
P. contorla yat. nrurrayana(Grev. & Balf.) Engelm.
P. couhcri D. Don..
P. densiflora Sieb. & Zucc.---------------
P. ecfrinnra Mill.
P. edulis Engelm.P. cembroides var. edulis
(Engelm.) Voss
P. elliottii var. densa Little& Dorman.
P. elliottii Engelm. var. elliottii-P. caribaea Morelet
P. engelntannii Carr.P.Iatifolia Sarg.P. apacheca Lemm.
P. /e.tifis James
P. gerardiana Wall.P. aucklandii Lodd.P. chilghoza Ehh.
P. glnhra Walt.
Swiss stonepine, cembranpine, arollaprne.
Mexican pinyon,nut pine,plnyon.
sand pine,spruce pine.
shore pine,lodgepolepine, beachpine, coastprne.
Rocky Mountainlodgepole pine.
Sierra Nevadalodgepole pine.
Coulter pine,nut pine,big-cone pine.
Japanesered pine.
shortleaf pine,southernyellow pine.
pinyon,Coloradopinyon pine,nrr+ nina
South Floridaslash pine.
slash pine,swamp pine,pitch pine,yellow slashpine.
Apache pine,Arizonalongleaf pine.
limber pine,RockyMountainwhite pine.
chilgoza pine,Gerard pine.
spruce pine,cedar pine.
High elevations in the Alpsand Carpathian Moun-tains of central Europe.
Semiarid mountain regionsin Mexico. southeasternArizona. southwesternNew Mexico. and south-western Texas.
Sandy plains of central andcoastal Florida and thesouthern tip of Alabama.
Low elevations on thePacific Coast fromsoutheastern Alaska toCalifornia.
Rocky Mountains, fromCanada to Utah andColorado, and the BlackHills.
High elevations in theCascade Range-SierraNevada chain fromsouthwestern Washing-ton to Baja California.
Coastal mountains fromcentral California southto northern BajaCalifornia.
Midelevation in the moun-tains of Japan andKorea north to easternManchuria and adjacentu.s.s.R.
Coastal plain of south-eastern United States toeastern Oklahoma andeastern Texas.
Semiarid regions in Utah,Colorado, Arizona, andNew Mexico.
Southern and centralFlorida, and LowerFlorida Keys.
Coastal Plains from SouthCarolina to eentralFlorida and southeasternLouisiana.
Mountains of southeasternArizona and south-western New Mexicosouth through the SierraMadre Occidental, Mexico.
Subalpine in the RockyMountains and southernSierra Nevada.
Mountains of easternAfEhanistan, northernPakistan. Kashmir. andnorthern India.
Coastal Plains from SouthCarolina to northernFlorida and west tosoutheastern Louisiana.
P. O. Rudolf.
S, L. Krugman.
L. Jones.
S. L. Krugman.
J. E. Lotan.
S. L. Krugman.
Do.
S. L. Krugmanand P. O.Rudolf,
J. M. McGilvray.
G. H. Schubert.
R. L. Barnes.
Do.
G. H. Schubert.
R. J. Steinhoff.
P. O. Rudolf.
J. P. Barnett,
T,W,E
w, E___-,,,.
T,H
w,E
T,H,W,S
T,H,W
H,E
E
T,H,W
T,H,E
T,E
T,H,E
T,E
T,W,S,E
T,H,E
T.H
600
PINAS
and, uses; data compilers-Conlinued
Scientific names and synonyms Common names OccurrenceData compilersfor the speciesIJses t
P. halependc Mill, - .---.
P . alepensis Poir.
P. helilreichii ChristP. heldreiehii' var. leucodermis
(Ant.) Markgr.P.leucodermis Ant,P. nigra'rar, Ieucodermis ( Ant. )
Rehd.
P. insularis Endl...-.--- .
P. khasaa Royle
Aleppo pine,Jerusalempine.
Heldreich pine,Balkan pine,Bosnian pine,graybark pine.
Khasi pine,-,
----- Jefrrey Pine .. --
Korean pine,eedar Pine.
sugar pine,pino real.
Chihuahua pine,yellow pine,prno real,
Merkus pine,Tenasserimpine.
singleleafpinyon, nutpine, pinyon.
western whitepine, Idahowhite pine,silver pine.
Swiss mountainpine, Mughopine, dwarfinountain pine.
bishop pine,prickle-conepine, SantaCruz Islandpine.
Austrian pine,black pine,Europeanblack pine.
Mediterranean region, from T, W' ESpain and Morocco toTurkey and Jordan.
High elevations from cen- W' Etral Yugoslavia tonorthern Greece and insouthern Italy.
High elevations from east- S' Eern India to SouthVietnam. and mountainsof northern Luzon,Philippines.
Mountains of southwestern T, H, W' E'-Oregon, California,western Nevada andnorthern Baja California.
Mountains from southeast- Tern Siberia to SouthKorea; central Honshu,Japan.
Mountains from western T' WOregon through Cali-fornia to western Nevadaand northern BajaCalifornia.
Mountains of central T, EArizona and southwest-ern New Mexico souththroush the Sierra MadreOccidental in northernMexico.
Mountains of southeastern T, E . -
Burma to North andSouth Vietnam; rvesternSumatra: Luzon andMindoro in the PhiliP-pines,
Semiarid mountains from H, W' Esoutheastern Idahotbrough the Great Basinrangei to central andsoufhern Calif ornia andnorthern Baja California.
Southern British Columbia T' Wto western Montana,northern Idaho andnortheastern Oregon;south in the CascadeRanse. Klamath Moun-tainJ and Sierra Nevadato central California.
Subalpine in the Pyrenees, W' EAlns. Carpathian andBaikan Mbuntains ofcentral and southernEurope.
Coastal California and H' Enorthern Ba.ja California ;
Santa Rosa, Santa Cruz'and Cedros Islands.
Southern Europe, northern T' W' S, EMorocco, SicilY, andCyprus.
P. O. Rudolf.
Do.
P. ietrregi Grev. & B?lf_. .
P. ponderoso' var. jefiregi
S. L. Krugman.
Do,
P. O. Rudolf.
S. L. Krugman.
W. J. Rietveld.
S. L. Krugman.
A. L. Roe.
R. T. Bingham.
P. O. Rudolf.
S. L. Krugman.
P. O. Rudolf.
(Grev. & Balf.) Engelm.
P. koraiensis Sieb. & Zttcc'---------------
P.lantbertiano Dougl.
P. leiophylla var. chihuahuana(Engelrn) Shaw
P, chihuahuana Engelm.
P. merkusii Jungh. & de Vriese -----
P. monophvlla Torr. & Fr6m. ------P. cembroides var,
monophglia (Torr. & Fr6m.)Voss
P. rrronticolo Dougl.
P. ntugo Turra- --P. montana Mill.
P. muricaaa D. DonP. remorata Mason
P. nigra Arnold-,P.laricio Poir.
601
PINAS
T.qslr l.-Pinus: nomenelature, occztrrence, and uses; data compilers-Continued
P. palu*ric Mill, ____ longleaf pine, Coastal plains from south- T. H. EP. australh Michx. f. southein pine, eastern Virginia to cen-
longstraw pine tral Florida and west toeastern Texas.
P. paruiflora Sieb. & Zucc.-----___---____ Japanese Mountains throughout EP_. pe.ntaphyllc Mayr ivhite pine. .fapan from iofittre"ttF. htnekomatsu Kyushu north to south_
Miyabe and Kudo ern Hokkaido; KoreanIsland of Utsuryo.
P. patula Schiede and Deppe Mexican Eastern Mexico from T, W-weeping pine. Tamaulipas south to
Oaxaca.
Scientific names and synonyms Common names Occunence
High mountains of westernEulgaria, southern Yugo-slavia, eastern Albania,and northern Greece.
Coastal areas in Morocco.Portugal, Spain, southernFrance, western Italy,Corsica, Sardinia, andnortheastern Algeria.
Portugal, Spain, and northshores of the Mediter-ranean Sea to Lebanon:northeastern Turkev:Ibiza,-S;;di;i;;si.iiJ,Crete, and Cyprus.
Extreme southwesternNew Mexico and south-eastern Arizona throughthe Sierra Madre Occi-dental to Durango.
Southern British Columbiasouth to central ldaho,and through the CascadeRange, Coast Ranges,and Sierra Nevada tosouthern California.
Rocky Mountains fromMontana south to NewMexico, Trans-PecosTexas and northernMexico: west to easternNevada and east tocentral Nebraska.
Eastern Siberia south tc,northern Mongolia, Man-churia. Korea. andthrough the Kuril Islandsto central Honshu, Japan.
Appalachian Mountainregion from Pennsylvaniato northeastern leorgia.
Semiarid mountains ofextreme southern Cali-fornia and northern BajaCalifornia.
Coastal central Californiaand northern GuadaluoeIsland, Mexico.
Great Lakes region fromsoutheastern Manitobaand Minnesota east toNewfoundland, NovaScotia and New Jersey.
Uses t
T, E.
T,W,S
T,H,W,E
W,E,H
T,S,E
T, H, W,E, S.
J. M. McGilvray.
-. P. O. Rudolf.
S. L. Krugman.
P. O. Rudolf.
Do.
S. L. Krugman.
S. Little.
S. L. Krugman.
Do.
D. H. Dawson.
P. ponderotavat, arizonica Arizona pine,(Engelm.) Shaw Arizona pon-
P. arizonica Engelm. derosa pine,Arizonayellow pine.
P. pon.derosa Laws. var. ponderota ponderosa pine,bull pine,rock pine,westernyellow pine,blackjack pine.
P. ponilerota var. rcopulorunt Rocky MountainEngelm. ponderosa
prne, westernyellow pine,blackjack pine.
P, peuce Griseb...- -.,P. encelsa ltan". peuce
(Griseb.) Beissn.
P. pinaster Ait.-P, maritima Poir.
P. pinea L.
P. purnita RegelP. cembra iat. pum;ta pitl.
P. pungent Lamb. -
P. quodrilolia Parl.P. cembroides yar.
parryana Yoss
P. radiata D. DonP. insignis Dougl.
P. reeinosa Ait.
Balkan pine,Macedonianpine, Greekstone pine.
- maritime pine,cluster pine,pinaster pine.
,- Italian stonepine, umbrellapine, stoneprne.
Japanesestone pine,dwarf Siberianpine.
Table Mountainpine, hickorypine, mountainpine.
Parry pinyon,nut pine,prnyon.
Monterey pine,rad.iata pine,rnsrgnrs prne.
red pine,Norway pine,hard pine,pitch pine.
T,H,S,E Do.
T,H.W W. J. Rietveld.
T,S,E R. J. Boyd.
T, H, W,s, E.
J. L. VanDeusen.
602
PINAS
TASLp L-Pinus: yromenclature, occurrence, and uses; data compilers-Continued
Scientific names and synonyms Common names Occurrence Uses' Data compilersfor the species
P. risida Mill.
P. roxburghii Sarg..P.Iongifolia Roxb.
P. rabiniana Dougl.
P. eerotina Michx.P . rigida var. serotina
(Michx.) Loud.
P, sibirica Du TourP. cembra var. sibirica
Loud.
P. urobilormis Engelm. . . .
P. flerilis var. refl'era Engelm.P. reflera (Engelm.) Engelm.
P. ilrobus L.
P, ryhtestria L.
P. taeda L.
P. thunbergiana FrancoP. tlrunbergiiParl.
P. toneyana Parry-- - -..
P. drginianaMill.
P. wallichiana A. B. Jacks.P. excelsaWall.P. g rifr.tlLii McClellandP. nepalensis de Chambr.
Northeastern United States T, H, W' Eand the AppalachianMountain region, fromLake Ontario and Maineto western KentuckY andnorthern Georgia.
Monsoon belt of the outer T, S, EHimalayas, from north-eastern West Pakistanto Bhutan.
Low-elevation dry sites T' Hin the Coast Rangesand Sierra Nevada,California.
Coastal Plains from south- Tern New JerseY to centraiFlorida and west tocentral Alabama.
Ural Mountains east T, Ethrough western andcentral Siberia tonorthern Mongolia.
Mountains of southwestern T, H, WColorado, Arizona, andNew Mexico south in theSierra Madre Occidentaland Oriental to centralMexico,
T, H, W,S, E.
S. Little.
P. O. Rudolf.
S. L. Krugman.
B. Benson.
P. O. Rudolf.
W. J. Rietveld.
R. E. Graber.
T,S,E R. S. Walters.
T,H,W B. F. Mclemore.
pitch pine,hard pine,bull pine.
Chir pine,longleafIndian pine,
Digger pine,gray prne.
pond pine,marsh pine.
Siberianstone pine.
southwesternwhite pine,border limberpine, Mexicanwhite pine.
eastern whitepine, northernwhite pine,white pine,soft pine,Weyinouthpine.
Scotch pine,Scots pine.
loblolly pine,oldfield pine.
Japaneseblack pine.
Torrey pine,Soledad pine,Del Mar pine.
Virginia pine,scrub pine,Jersey pine.
blue pine,Bhutan pine,Himalayanpine.
Newfoundland and NovaScotia west to south-eastern Manitoba, souththrouEh the GreatLakes and AppalachianMountain regions toIowa, western KentuckY,and northern Georgia.
Eurasia, from Scotland andSpain to Finland andTirrkev. across U. S.S.R.to the Sea of Okhotskand Manchuria.
Coastal Plains and Pied-mont from Delaware tocentral Florida to east-ern Texas, southernArkansas, and southernTennessee.
Maritime in South Koreaand in Japan from north-ern Honshu south tonorthern Ryukyu Islands.
Santa Rosa Island andlow coastal bluffs in SanDiego Co., California.
Aooalachian Mountain and'Piedmont regions, fromLonE Island south andwesf to central Alabama,v'estern Tennessee, andsouthern Indiana.
Mid- and hiEh-elevationsin the Himalayas, u'estto eastern Afghanistanand east to southernChina and northernBurma.
P. O. Rudolf.
S. L. Krugman.
S. Little.
P. O. Rudolf.
T,W,S,E
E
T,W
T,E
1 T: timber production, H:
603
habitat or food for wildlife, W: watershed, S: shelterbelt, E: environmental forestry'
PINAS
sota there is a change in cone serotiny fromclosed cones in the north to predominantiy opencones in the south (19f). Winter colorafion ofneedles is less in seedlings from lower latitudestlran in those from higher latitudes (ZeZ). InCanada, height growth was greater for sourcesfrom areas with longer growing seasons; selec-tions moved north made bettei treigtrt growththan selections moved south (i04). -
Pi,tr,us brutia-several varieties have beendescribed in the Black Sea region. The var.pitltyusa is found along the nortliern and north-eastern shores of the Black Sea. and var. eI-darica is found in the central Transcaucasus(152).In northern California, an Afghanistansource related to var. eldarica is out-growingvar. pithyusa and appears to be both fiost anddrought hardy and of good form (90, 1 ZZ) .
Pinus canariensis-This pine is found natu-rally only in the Canary Islands where it rangesfrom-.2,100 to 7,200 feet above sea level (15-Z).Seedlings of several sources from the variousislands and elevations, when exposed to lowwinter temperatures in the Institute of ForestGenetics' nursery at Placerville, California,showed marked differences in cold hardiness.Seedlings from 4,000 feet showed more colddamage than seedlings from 6,200 feet onTenerife Island. Seedlings from 6,200 feet onPalma Island were badly damaged, suggestingthat sources from different islands rn'iy als6differ in susceptibility to low temperature"(122).
Pintts caribaea-Three geographic variantsare recognized. The var. caribaeaiative to Cubaand the Isle of Pines has persistent seed wings,while the other two varie[ies do not. In testslnSouth Africa, var. caribaea showed bettergrowth than var. hondut'ensis from the mainlandof Central America. The var. hondur.ensts hasthe largest seeds; var. bahamensis from theBahama Islands has the smallest (150).
Pinus cembro-A number of cultivars of thisspe_cies have been reported (51, 1g0).
Pitttts cembroides-Two varieties have beendescribed. The var. remota found on theEdwards Plateau in southwestern Texas hasvery thin seedcoats. The var. bicolor- is found insouthwestern New Mexico and southeasternArizona (137\.
Phtus clattsa-ln central Florida, this smallpine has closed cones. But in western Florida ithas an open-cone variety, vat. immuoinata-Choctawhatchee sand pine, whose cones"ripen inSeptember and shed seeds during October (es,138).
Pintts cotttorta-Four varieties are recog-Lized -thouSh only three are listed in the table"s.'r'he fourth, var. bolanderi, has a restrictedrange on the northern California coast-the
604
Mendocino White Plains-and has serotinouscones like those of the Rocky Mountain variety,var. Iatifolia. "Ihe other varieties have coneswhich open at or soon after maturity; the opencones of var. contorfo persist indefinitely whilethose of var. lnumo,yana do not (lL\.In tests innorthern Europe, seedlings of var. contortogrew faster, and were more branchy and lesswinter hardy than those of var. lntifolia frominterior British Columbia and the Rocky Moun-tains (6y'). The var. mul.raAarLa has the largestseeds. At temperatures of 50' to 68" F. the var.Iatifolta germinates twice as fast as coastalsources (44).
Differences between sources also are foundwithin a given variety. Seeds from high eleva-tions in central British Columbia germinatedfaster at 68' F. than at other temperatures, andseed from low elevations germinated faster attemperatures above 68'F. (8/). Commonly, thesouthern sources grow faster than the northernsources of a given variety.
Pintts cot.tlteri-This pine is endemic to Cali-fornia and Baja California. No races have beendescribed, even though it grows in isolatedstands from 500 to 7,000 feet and on fertile tovery poor soil types. Mount Diablo sources areconsidered to have the poorest form, with.qreater branching than any other source (Z7g\.
Pinus densiflora-A number of cultivars havebeen described (150) . This pine is widelyplanted and hardy in the Lake States, NewEngland, and southern Ontario (Z6t+).
Pinus echin,o,ta-Ten-year results in a range-wide test suggest that in the southern UniiedStates the southernmost sources from east ofthe Mississippi River are superior to northernsources in both survival and growth (260).Seeds from northern sources should be used inthe northern extremities of the range (1llr,260, 262). In an Oklahoma test, an Arkansasseed source showed the best performance, evensurpassing a local source (189\.
Pinus edulis-A single leaf variety, var.fallar, is found in the mountains at 6,0b0 feetin central and eastern Arizona, and in the GrandCanyon and parts of New Mexico. Its seeds tendto be larger and have a thicker seedcoat thanthe more common variety, var. edulis. In con-trast to var. edulis, var. fallar seldom producesseeds in quantity (1Sf).
Pinus elliottii-Two varieties are recognized.The var. densa is found only in south Florida.Compared to typical slash pine, var. elliottii, thevar., dettsa germinates faster, has a grasslikeseedling stage with crowded needles. and hasheavy wood with very wide summer rings (lgg,110, 216). Sources of var. elliottii in north-eastern Florida appear to be the poorest, as they
are susceptible to ice damage and are lessdrought resistant than northern and westernsources (216,260).
Pinus flerilis-seedlingq from--- southernstands are'faster growinglhan seedlings fromnorthern stands (219).
Pi,nus halepensis-In Israel at least two ele-vational ecotypes are recognized, and others canbe expected in other parts of the range (1I8,152).
Pinus heldreichii-Pinus tt'eldreichid' var.leucodermis, considered by sorne to be a timber-line tree, commonly is found on drier sites andoften on soils formed on limestones (51). Thevar. heldreichiiforms open forests in the moun-tains at elevations between 3,000 and 5,000 feet(180).'
Pi,nus insulnris-In tests in northern Rho-desia, trees from seed of Philippine origin weremore vigorous and had better form than thosefrom India, Burma, or Vietnam origins (152,2oo).
Pinu.s ieffreui-Evidence for distinct ceqgraphic rices ii lacking, but differences in seedorigin are still important. Generally, trees fromseed collected east of the Sierra Nevada areslower growing, more drought resistant, andless susceptible to cold damage (85). Also treesfrom high-elevation sources tend to be slowergrowing than those from lower elevations (93) '
Pinus koraiensis-The Siberia, China, andKorea sources most likely should be considereda geographic race distinct from that in Japan.
Pinus leiophylla var. chihunhunnn-Two dis-tinct forms have been found' One has poor formwith a short, crooked bole and many branches;the other has good form and grows to heightsof 80 feet (152),
Pinu.s merkusii-At least two distinct racesare recognized, one on the Asian mainland andthe other on the island of Sumatra. Seeds ofmainland origin are larger than those fromSumatra. Trees from mainland origins passthrough a grasslike stage and tend to be uni-nodal with a straight, cylindrical bole, but thgydo not grow as tall as those from Sumatra. TheSumatra trees may reach 200 feet in height andtend to be sinuous in gron'th (40)' Some con-sider the two races as separate species; P. mer'kusii on Sumatra, and P. merkusiana, on theAsian mainland (41).
Pinus monophglla-No varieties are recog-nized, but there are variants which have partlyor mostly two needles in a fascicle, rather thanjust one (137).
P inus m o ntic ol,o-Local vari ati ons associatedwith elevation and site are recognized. Pro-genies from high-elevation origins grow fasterat high elevation than those from low-elevation
PINUS
origins (217). Seeds from northern Idaho areil;tt"" ihan'those from washington and cali-fornia (295).---Pir*; mugo-A number of horticulturalvarieties hav6 been described. They vary from asnrawlins shrub, vat' pumilio, to a small tree'v-ai. iostFata Q8:D. Varieties also diffe-r in seedsize and germination capacity (1p7\.-Seedlingsof io*-"t6uation origins-are not hardy at highelevations (256).
Pinus muricata-Trees from origins northand south of Fort Ross, California, differ ing"o*ttt form, foliage color, and cone shape' I1triito*i", tiees fiom northern sources tendto grow larger with fuller and more com-pact
""oiuns (a-r )l In tests in Australia, the northern
souices maintained better growth rate and formthan the southern sources (69).
ptnus nigru-Several distinct varieties andnumerous cultivars are recognized. The var'iiia*iin" from the Crimea, Turkey, andCyprus tends to have the largest seeds (17-,500
to'ZO,SOO per pound) ; the var. c,ors,i9a.na frornCorsica tras ttr6 smallest seeds (28,000 to 36'000per pound) (152, 187). Wood in the Corsicanirari6tv is' considerably better than -that intypicai Austrian pine, var. austrinca, !.rom theddtt
"n peninsula- and eastern Alps. Stands of
iir. iitiarfco planted in Belgium are consideredone of the more cold hardy varieties. Othervarieties are var. cebennensis from the Pyreneesin southern France, vat' hispanica ftom Spain,and var. mauritanica from Morocco and Algeria.
Some general statements can be made forvarious geographic sources (152):
Western Groups: Sources in France andSpain are often proven to be 4rought re-siitant and are indifferent to soil type.
Central Groups: Sources in Corsica and Italygrow well
-and have good form. But-they
need a high humidity and grow poorly onlimestone soils.
Eastern Group;: Sources in the Balkan andCrimea are6s appear to do well on thepoorer limestone soils.
Pinus palustris-Seedlings of different geo-graphic origin differ in height, growth, coldiesi-stance, and survival (71, 260'). Southeasternand central Louisiana sources did poorly; south-ern Florida sources failed outside of their areaof origin. Central Gulf Coast sources grow wellthroughout the Gulf Coast (260).
Piius p ara ifl or a-Several horticultural f ormsare cultivated, (123) . Pinu,s paraifl'ora is thoughtto consist of two geographical varieties whichintergrade in central Honshu, Japan (49).
Piitrs patula-Because of its rapid growth,this species has become one of the most im-nortant sources of wood in the summer rainfall
605
PINUS
areas of South Africa (148).It also grows wellin East Africa, New Zealand, and Australia(128, 152). A variety, var. longepeduneulata,from the States of Oaxaca and Chiapas, Mexico,has been reported. The cones of this varietyopen quickly at maturity, thus differing fromvar. patuln which has closed cones. The seeds ofvar. Iongepedunculata are black with brownmarks, while those of var. patuln are pure black(1 t8) .
Pinus peuce-Il Europe seeds from the Rilaand Pirin Mountains in Bulgaria are consideredto produce the best trees (77I).
Pinus pinaster-This species is highly vari-able, and at Ieast four main races are recognized(152). The French or Atlantic race, which isitself quite variable, is commonly found on thecoastal sands. The Portuguese race is also foundon coastal sands but is superior to the Frenchrace in growth, form, and drought resistance;it has done well in tests in South Africa andwestern Australia, and appears to have someseed dormancy (105, 266). The Corsican race ismost commonly found in the mountains. TheMoroccan race shows differences between moun-tain and near coastal origins, in that sources ofmountain origin fail when planted on the coast.The mountain sources are thought to be frostresistant (152).
Pittus ponderoso-The three main varietiesare var. ponderosa, var. arizoniea, and var.seopulort+m. These varieties differ in seed andcone size, needle length and number per fascicle,speed of germination, rate of growth, form, re-sistance to drought and low temperatures, andsurvival (31, 71, 218, 257, 259). Within thevarieties there are also source differences insome of these characteristics. Speed of germina-tion at different temperatures was found tovary for seeds from the Pacific Northwest,Rocky Mountains, and the Southwest. South-west sources reached maximum germination atthe lowest temperature (91).In tests in Oregonand Washington, growth rate generally in-creased with seed origins from east to west, andfrom south to north in the eastern part of therange; slowest grou'th was shown by sourcesfrom eastern and southeastern parts of therange (218\. In tests in California, growth rategenerally increased as seed source elevationdecreased (33). In tests in northern Arizona,eastern and southeastern sources did well, butnorthern and western sources failed-as didthe southernmost source (195\. Northernsources of var. scopttlorum are characterizedby a relatively good growth rate and frost re-sistance (257). Southern sources of this varietyare slower growing but frost resistant. In Cali-fornia, sources of var. ponderosa west of the
606
Sierra Nevada are faster growing but less frostresistant than sources east of the crest (122).
Closely related to P. ponderosa in appearanceis P. toasltoensfs. This rare pine, which is oftenwrongly identified as P. ponderosa var. ponder-oso, is known to be in only three areas: eastslope of Mount Rose, Nevada; southern WarnerMountains, northeast California; and BaldMountain Range in the eastern Sierra Nevada.Both its male and developing, second-year fe-male cones are purple to purplish black, unlikeponderosa pine. Washoe pine flowers in July;its cones mature in August and September andopen in September. Cones and seeds are handledin the same manner as var. ponderosa, and itsseeds germinate promptly after a 60-day coldstratification (47, 122).
Pinus ptmgens-Seed weight, cone length andwidth appear to decrease as elevation increasesand as latitude decreases. Cone serotiny is mostcommon in the southern part of the range. Nodistinct varieties are recognized (27[t).
Pinus radinta-Monterey pine occurs natu-rally in four relatively small and well separatedpopulations. The var. binata occurs on SanGuadalupe Island and is slower growing thanthe three mainland populations (Monterey,Cambria, and Afro Nuevo Point). The Cambriapopulation has the largest corr€s orrd seeds, andthe Monterey population has the smallest (70).In tests in Australia, trees of Cambria origindid not do as well as those from the Afro NuevoPoint and Monterey populations (69).
P'intts 1'psi11ssa-lhis species is consideredone of the least variable of the pines, and nosubspecies or varieties are recognized, (74,266).However, some height growth, form, and woodquality differences may exist among populationsin the Lake States, New England States, andWest Virginia. In northern sources, seeds areoften smaller, lammas frequency is generallyless, and frost resistance is higher than insouthern sources (74).
Pinus rigida-No distinct geographic sourcesare known, but there are variations in form anddevelopment between populations. Throughoutmost of the range, trees consistently open conesand shed seeds soon after maturity; in southernNew Jersey the vast majority bear cones thatremain closed at maturity and only open atirregular intervals (6,71). The latter trees arecharacteristic of areas with a history of wildfires (71).
Pittus sabin,iana-Seeds from populations inmild climate areas germinated more quicklyafter stratification than those from the colderareas. Seedlings of southern origin grew longerthan those of northern origin. Larger cones weremore frequent in the northwest part of therange than in the Sierra Nevada (82).
Pinus serotina-Cone serotiny is greater insouthern and coastal populations than in north-ern and Piedmont populations (211t)'
P inus si,b i.r i c a-Di sti n ct diff erence s in gr owthrate, branching habit and fat content of seedsexist between certain populations. No varietiesare recognized, but several forms have beendescribed. Pinus sibiri,cu f. humistrata is adwarf form that grows on mountain summitsand ridges. P'itttts si'biricaf. coronans has a wideand dense crown, is reasonably drought resist-ant, and is found from sea level to 6,600 feet'Pimts si,birica f . turf osa grows on peat (185).
Pinus syluestris-Scotch pine, the mostwidely plan-ted introduced species i1 tlre UnitedStatei, js probably the most intensively studiedof all pin-es. The first cornparative pine seedsource trials involved this species. It is the pinewith the greatest natural range' and it growsin many different ecological situations (266,268). Numerous varieties, forms, and ecotypeshave been described. A conservative estimate ofthe number of geographic varieties ranges from21 to 52 (268). There is also considerable varia-tion within named varieties. Sources differ inmany characteristics including seed size, ger-mination, dormancy, seed and cone color, treeform, growth rate, structure of root system,flowering characteristics, needle color, and sus-ceptibility to heat, cold, or drought (28, 18-6,gZ-0, ZOO, 263). Seed size decreases from thesouth (44,200 per pound in Spain) to the north(127,000 per pbund in Lapland) (101). Growthrate tends to
-decrease from southern to north-ern sources. In tests in Sweden, southernsources grew faster and later in the autumnthan did northern sources (117) . Similar resultswere found in Michigan tests, where certainFrench sources grew three times as tall asnorthern Finnish and northern Siberian sources(266, 268). The more southern sources, however,were more susceptible to low temperatures thannorthern sources. Under Canadian prairie condi-tions, Russian and Finnish sources survivedbetter than more southerly sources (tnZ). tnMichigan, needles of Spanish, southern France,Balkans, and Asia Minor origins remainedgreen during the winter, while those of Siberianand Scandinavian origins turned yellow (266,268).In tests conducted in Sweden, seed origininfluenced germination rate at a given tempera-ture. Sources from northern latitudes or highelevations germinated well over a wider rangeof temperatures than those of southern latitudesor low elevations (117). Sources in the extremenorthern parts of the range and certain sourcesfrom Turkey and Greece show the greatest seeddormancy (101). Lack of a fully developed em-bryo accounts for part of the dormancy ofnorthern sources (116). In some European
PINUS
localities, introduced sources have producedbetter trees than local sources'
Pi,nus strobus-Seeds from the western partof the range are lighter than seeds from theeastern paft, and seeds of southern origin re-qui." u iottg"" stratification period than thoseof northern origin (72, 162, 264). Field testsfru"" A"-onstrafed that southern sources, suchu* tftot" from the southern Appalachians, tendio gro* faster and to continue shoot elongationlon"ger in the fall than do northern sources (73,iosi. ln artificial freezing studies and fieldobservations in the northern Lake States, south-ern sources are more sensitive to low tempera-tures (162, 196). In fall, seedlings of easternoiigin 'had'blueigreen foliage compared to theyellow-green foliage of northwestern sources(267).' Piitu,s foeda-Numerous field studies havedemonstrated definite geographic variations ingrowth rate, drought and cold hardiness, dis-ease
iesistance, and survival. Seedlings of Marylandorigin tend to grorv less than those of otheroriEin when pianted in different locations(260,261). Soulhern sources outgrow northern.ou"".. in South Africa (210).In many of thetests, local seed sources were best. Sources westof the Mississippi River are more drought anddisease resistant than most of the easternsources. And southern sources tend to be moresusceptible to low temperature damage thannorthern sources (260).
Pintts thunbergiana-The better formed treesare from inland sources (1/+5).
Pi,nus torreyan,a-In a common planting onthe California mainland, trees of mainlandorigin grew taller and had a single trunk, whiletrees from a Santa Rosa Island source wereslower growing, bushy, and freely branched.The islald souice produced larger cones (86).
Numerous pine hybrids have been described.As a conservative estimate, at least 200 first-generation and second-generation hybrids aswell as backcrosses, crosses between varietiesof the same species, and crosses involving threeor more diffeient species either occur naturallyor have been produced artificially (122,1/t1, 169,266). No attempt is made here to provide yieldstatistics for the numerous hybrids, since suchdata are highlv variable. The number of soundseeds produced depends on the species and indi-viduaitrees involved, as well as on the environ-mental conditions under which the cross is made.
Some natural hybrids are relatively common;e.g., P. palustris I taeda, known as Sonder-egger pine, which occurs in Louisiana and else-where in the South (36), and P' contorta ybartksiann, in central Alberta and southwesternMackenzie (272). Most other hybrids occasion-
607
PINAS
ally occur where the two parent species arenaturally associated. Examples of this in Cali-fornia are P. ponderosa y jeffreEi, P. jeffreyi7 cotilteri, P. radiata )( attenuata (48, 169).In the South are P. taeda X echinata in eastTexas, and P. taeda y seroti,na throughout theircommon range (tnS, 27L, 266). The hybridbetween P. densiflora and P. thmtberginnaoccurs naturally in Japan but has also beenproduced spontaneously in plantations of thesespecies in Michigan (26.1). In Europe, P. syl-aestri.s occasionally crosses naturally with bothP. nigra and P. mltgo where these species areplanted near one another (266). These are buta few of the many hybrids which have beenreported to occur naturally.
Several pine hybrids have been produced inrelatively large numbers by controlled pollina-tion methods. These include P. rigida \ taeda,which is an important hybrid in the reforesta-tion program in Korea, and P. attenuata yrad'iata, which is being planted in California andOregon (83, 106). Many other hybrids are beingproduced in smaller numbers and tested fortheir suitability in various parts of the UnitedStates.
Flowering and fruiting.-In certain species,reproductive structures are first formed whenthe trees are only 5 to 10 years old-e.g., P.atten.uata, P. banksiana, P. clausa, and P. con-
torta; in others they do not form until the treesare much olCer-e.g., P. lnmbertiana and. P,resinosa (table 3). Pines are monoecious withmale (microsporangiate) and female (mega-sporangiate) strobili borne separately on thesame tree. Male strobili predominate on thebasal part of new shoots, mostly on older lateralbranches in the lower crown. Female strobiliare found most often in the upper crown, pri-marily at the apical end of the main branches inthe position of subterminal or lateral buds. Butfrequent exceptions will be found to this generalscheme. For example, P. banksiana, P. clausa,P. rad'iata, or P. attenttata are multinodal in thebud, and female strobili are found occasionallyat a secondary whorl position (71, 122). Pintnattenuata, P. radiata, and P. airginiana fre-quently produce female strobili in all parts ofthe crown (71,122). In temperate climates, theearliest stages of male and female strobili canbe detected in the developing buds during thesummer or fall; the male develops l" to severalweeks before the female strobilus (76, 122, 169) .
Male and female strobili of the southern andtropical pines emerge from buds in late winter;e.g. P. elliottii var. densa, P. elli,ottii var. el-Iiottii, P. glabra, and P. palustris. Strobili ofother pines emerge from the bud in early spring,or late spring and early summer (table 2). Themale strobili are arranged in indistinct spirals
Tasl,n 2.-Pinus: phenology of flotuering andfruiting
Species Location Floweringdates
Cone ripeningdates
Seed dispersaldates
Datasource
P. albicaulis--. ,,,- CaliforniaP. aristata ArizonaP. armandii--, .---.-.. CaliforniaP. attenuata-- ----, doP. balfouriana-, doP. banksiana ,- , Lake StatesP.bru,tia ., ., CaliforniaP.canariensis-,.--,-. doP. caribaea Cuba
P. cembra -. GermanvP, cembroides . CaliforriiaP. clausa Florida
- JulYJuly-August
, April-May- ,--AprilJuly-AugustMay-JuneMarch-MayApril-May
- January-Feb-ruary.
MayMay-JuneSeptember-De-
cember.
August-SeptemberSeptember-OctoberAugustJanuary-February---September-OctoberSeptemberJanuary-MarchSeptemberJuly-August
August-OctoberNovember-DecemberSeptember
September-OctoberAugust-September
September-OctoberAugust-September-
doOctober-NovemberSeptember
August-September
not shedSeptember-OctoberAugust-September -closed cone', - .- ,September-OctoberSeptemberclosed coneSeptember-OctoberSeptdmber
not shedNovember-December-September "
Fall'September-October "
doOctober n
September-OctoberOctober-NovemberSeptember-October
September-November
P. contortavat. contortavar. Iatifolia
YAT, mUrrA,AAnd,P. coulteriP. densifloraP, ecltinata.,P. edulisP. elliottii
vat. densa
var. elliottii
California May-JuneRocky Moun- June-July
tains.Caliiornia MayJune
do dodo Anril
South Carolina M-arch-AprilArizona June
SouthernFlorida.
Florida
January-April
lqQ qt.n181, 2061001001 00 00E
t oo100150
19412229, 235
122,235oary
l qo
122,235122
132,133,13/+
30
P.engelmannii, ArizonaP. flerilis California,
Montana.
508
January-Feb- September-October- October 59ruary.
May November-December November-February- - 62,235June-July August-September . September-October, 122
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611
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PINUS
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612
in clusters 0.5 to 2 inches long (57, 787, 209,225). Before ripening they can be green oryellow to reddish purple, but are light brownto brown at the time of pollen shed; in mostspecies they fall soon after ripening. Femalestrobili emerge from the winter bud shortlyafter the male strobili and are green or red topurple (51, 71, 181, 225). At the time of pollina-tion they are nearly erect, and 0.4 to 1.5 incheslong and sometimes longer. After pollination,scales of the female strobili close, and thestrobili begin a slow development. At the end ofthe first growing season they are about one-eishth to one-fifth the lengbh of mature cones.Where temperatures are favorable, developmentcontinues through the winter as in P. elliottiivar. elliottii in Florida, and in P. attenuato andP. ponderoso at low elevations in the SierraNevada (122,25I). Fertilization takes place inspring or early summer about 13 months afterpollination, and the cones begin to grow rapidly.Growth of a new shoot leaves the developingcone in a lateral position. As the cones maturethey gradually turn from green, purple, orviolet purple to yellow, light brown, reddishbrown, or dark brou'n (table 3).
Cones and seeds of most snecies maturerapidly during late summer and fall of thesecond year (table 2). Cones of a few speciesmature during late winter of the second year orearly spring of the third year; e.g., P. attenuata,P. brutia, and P. merkusii (25, 10,722). Seedsof P. attemnta and P. bruti,a are mature duringthe fall, about 16 to 18 months after pollination,but the cones are not fully developed until latewinter (25, 122). Pittus leiophAlla, P.leiopltUllavar. ehihtnhuan.a, and P. pinea require a full 3years for their seeds and cones to ripen (5-l ,136). The requirements of P. torreyana are stillunclear. This species has been described as need-ing 3 years to mature its seed, but there isevidence that the seeds mature in 2 vears. withthe cones requiring 3 years to open tiZZ).
The interval between large cone crops isvariable and depends on the species and environ-mental factors. Some species consistently pro-duce a large crop every year, while others showa cyclic pattern of 2 to 10 years between Iargecone crops (table 3).
Mature cones (fig. 1) vary widely in size andweight. Those of P. rnugo are 1to 2 inches longand weigh about 0.06 ounce, while those of P.lambertiana may be 12 to 25 inches long andweigh about 1 to 2 pounds. Those of P. sabinianaand P. cotilteri often weigh more than 2 pounds(51,180,225).
The mature cone consists of overlappingwoody scales, each of which bears two seeds atthe base on the upper surface. The cones of mostspecies open on the tree shortly after ripening,
PINUS
and the seeds are rapidly dispersed (table 2).Drying causes the cone scales to separate owingto differential contraction of two tissue systems:woody strands of short, thick-walled, tracheid-like cells extending from the cone axis to thetip of the cone scales, and thick-walled scleron-chyma cells in the abaxial zone of the scale. (2,229). In a few species with massive cones, thescales separate slowly, and seeds are shed overperiods of several months; e.g., P. coulteri, P.rorbtrt'ghii, P. sabiniana, and P. tomeEana(225, 233) .
In some species part or all of the mature conesremain closed from several to many years oropen on the tree only at irregular intervals; e.g.,P. attenuata, P. banksiana, P. brut'ia, P. elausa,P. contorta var. latifolia, P. ltalepensis, P.mtrlicata, P. pi,nastet', P. ptmgens, P. radiata,and P. t'igida (51 , 71, 225) . In addition to theirclosed-cone habit (serotinous cones), P. bank-siana, P. clausa, P. r'igida, and P. contortahaveforms whose cones open prornptly at maturity(29,71, 196, 197). The closed-cone habit is theresult of three factors: (a) extremely strongadhesion bet'*'een adjacent, overlapping conescales beyond the ends of the winged seeds (738,126); (b) cone structure; and (c) the natureof the two tissue systems in the scales describedabove. The scales apparently are held togetherby a resinous substance. The melting point ofthe resin seal for P. contorta var. latifolia isbetween 113" and 722" F. (ltl). Heat, especiallythat from fire, melts the resin and permits thecones to open. Stiil other species shed partlyopened or unopened cones, and the seeds aredispersed only when the cones have disinte-grated on the ground; e.g., P. albicuulis, P.eembra, P. pu,mila, and P. sibirica (51, 169, 185,225) .
Commonly, cones that opened on the tree areshed rvithin a few months to a year after theseeds are dispersed. In some species, however,opened cones may remain on the trees for up to5 years or indefinitely; e.g., P. attenuata, P.eontorta, var.Iatifolia, and P. rigida (71,225).
Mature seeds vary widely in size, shape, andcolor (2t+7u) (fig. 2). They range in length fromone-sixteenth to one-tenth inch (2 Lo 3 mm.) forP. banltsi,anlc, to more than three-fourths inch(19 mm.) for P. sabin,iana. They are ellipsoid( P. radiata ), p ear -shap ed ( P. pu,mila ), cylindric(P. gerardian,a), more or less triangular (P,'pttngens), ovoid (P. petrce), or convex on theinner and flattened on the outer side (P. pinea)(51, 180). The seedcoat, which may be reddish,purplish, greyish, brown, or black, and is oftenmottled, can be rather thin and papery to hardand even stony (5/, 209,225).
In most species a membranous wing is at-tached to the seed, but in some species the wings
613
PINAS
P. albicauliswhitebark pine
P. banksianajack pine
P. engelmaniiApache pine
P. flexilislimber pine
P. attenuataknobcone pine
P. clausasand pine
P. leiophylla var. chihuahuanachihuahua pine
dispersal, /2 y.
P. cembroidesMexican pinyon
..& "t.rt'
t\
tt rli"\\4'
614
Frcunn l.-P'inus: mature cones collected before seed
P. monophyllasingleleaf pinyon
P. rigidapitch pine
P. strobiformissouthwestern white pine
P. ponderosa var. arizonicaArizona pine
PINAS
P. ponderosa var. scopulorumRocky Mountain ponderosa pine
P. serotinapond pine
P. virginianaVirginia pine
% X-Continued.
-il
{,'t'l
.*
*
P. pungensTable Mountain pine
P. sylvestrisScotch pine
iw
ffirF
FIcuRE l.-Pinus: mature cones collected before seed dispersal,
615
pP. edulispinyon
&;-*
P. glabraspruce pine
P. koraiensisKorean pine
(*"** .$F
P. balfourianafoxtail pine
P, gerardianachilgoza pine
PINUS
&nP. albicauliswhitebark pine
','J
P. banksianajack pine
P, contorta var. murrayanaSierra Nevada lodgepole pine
,r,'r,.4b
P. aristatabristlecone pine
P. brutiaCalabrian pine
P. elliottii var. elliottiislash pine
P. halepensisAleppo pine
P. lambertianasugar pine
q-pP. attenuataknobcone pine
P. cembroidesMexican oine
P. engelmanniiApache pine
P. armandiiArmand pine
P. bungeanalacebark pine
P. coulteriCoulter pine
t&"p
P. flexilislimber pine
'
&ffiP. merkusii P. monophyllaMerkus pine singleleaf pinyon
flpP. clausasand pine
P. densiflora P. echinataJapanese red pine shortleaf pine
wP. heldreichiiHeldreich pine
tP. leiophylla var. chihuahuanaChihuahua pine
P. insularisKhasi pine
P. jeffreyiJeffrey pine
616
FrcuRo Z.-Pinus: seeds as shed naturally from their cones, 1 X: some are wingless when shed.
PINUS
are absent or rudimentary; e.g., P. rnlbi,cauli,s,P. armandii, P. cembro,, P. flerilis, P. gerar-d'iana, P. koraietrcis, P. pumila, P. sibirica, andP. strobiformis (209, 225, 233, 2rt7a) (fiq. 2).In others the wing or modified "wings" mayremain attached to the cone scales when theseeds are shed; e.g., P. cembroides, P. edulis, P.gerardiana, P. monophulla, and P. qzutdrifolia(225, 233). The seed wings are easily detachablefrom the seed of most hard pines except P. pinea,P. ronbttrghii, and P. can,ariensis,' those of thesoft pines are firmly attached except for P.aristata and certain sources of P. balfouriana(169,209,233).
The mature seed consists of a seedcoat whichencloses an embryo imbedded in a food-storagetissue, the endosperm (female gametophyte).Attached at the micropylar end of the whitishendosperm is a brown papery cap, the remnantof the nucellus. The endosperm and papery capare covered by a thin, brown, membranous ma-terial-the remnant of the inner layer of theovules integument (fig. 3).
seed coa t
endos perm
cotyl edons
hypocotyl
radicle
Ripe cones can be collected from standingtrees, from newly felled trees, and from animalcaches. To avoid large yields of immature seed,collection from animal caches should not beginuntil late fall, when the seeds are definitelymature (205) . For most species, cone collectionsfrom standing trees should start as soon as thecones are ripe and just cracking, since mostseeds are shed promptly from opening cones.For closed-cone species, collections can be de-layed without loss of seed, and frequently delayis even desirable. Although seeds may be ma-ture in the fall, the closed cones are very difficultto open at that time; additional maturation onthe tree facilitates both cone opening and seedextraction (25,122).
To avoid extensive collections of immatureor empty seeds, it is advisable to first checkripeness of seeds in small samples of cones fromindividual trees. A mature seed has a firm whiteto yellow, or cream-colored "endosperm" and awhite to yellow embryo which nearly fills theendosperm cavity. With experience this visualcheck is very useful, and with some species itmay be essential.
Ripeness for some species can be estimatedby changes in cone color. Colors of green andripe cones for most species are listed in table3. For example, P. Ttond.erosa var. ponderosacones are mature when the color changes fromgreen or yellow green to brownish green, yellowbrown, or russet brown; P. resinoscl. cones turnfrom green to purplish with reddish brown onthe scale tips; P. strobrts cones turn from greento yellow green with brown on the scale tips orto light brown. For some species such colorchange comes too late to be a useful index; e.g.,in P. palttsti'is the ripe cones are still green incolor and may have already started to openbefore turning brown (25It).
For species in which changes in cone colormay not be useful, flotation tests of cone specificgravity may be. These tests are based on thefresh weight of the cone. Species for which conespecific gravity has been related to seed maturityand their appropriate tests are shown in table 4.The easiest way to determine if cones havereached a desired specific gravity range is tosee if samples of freshly picked cones will sinkor float in liquids with known specific gravities.Pitttts Ttonderosa seeds are ripe when the coneswill float in kerosene, P. strobu.s is ripe whenthe cones just float in linseed oil, and P. glabrais ripe when the cones just float in SAE 20motor oil. Cones from standing or felled treesshould be tested only when fresh, since excessivedrying will lead to erroneous conclusions as tomaturity.
Collecting from felled trees should only takeplace after the seeds mature. Otherwise there
L.
FIGURE 3.-Pinus ponderosa, ponderosa pine: longitudi-nai section through a seed, 6 X.
Collection of cones.-Cones should be collectedfrom trees superior in growth and form char-acteristics. Larger cones generally contain mores-eeds, but normaily all cones are collected exceptthose with obvious disease and insect damage.Widely spaced, dominant trees with full crownsproduce the most seeds per cone, provided ade-quate pollen from other trees is available. Whentrees are isolated and pollen from other trees isIimited, seed yield tends to be low. In dense,young stands most species usually produce litileseed. Those species which form fire thickets areexceptions i e.g., P. attenuata, P. banksinno,, P.clausa,, P. rigida, and P. serotina (71).
618
PINAS
TAsLp .-Pitttts: speci,fi,c grauity of ri.pe cones und liquids used for testing ri.peness ba flotati,on
SpeciesSpecific gravity
of ripe conesFlotation
test liquid'Data
source
P. aristataP. contorta var. latifolia
0.59-0.800.43-0.89
1.1-00.88
0.80-0.86
<0.89<0.90
0.880.81-0.860.70-0.800.80-0.89
0.88-0.970.80-0.86
<0.85<1
0.80-0.940.88
0.85-0.950.92-0.970.88-1.00
0.88<1.00
kerosene -. p061l,g
Seff zo ,r"t"i"if, oil-part kerosene to 4 parts lin;;ed ;lt 36s1,kerosene z0l,
P, densiflora,P. echinataP. edulisP. elliotti.i
vat. denso,vzr. elliottii
P. glabra,P. jeffreuiP.Iambertianrt,P. palustrisP. ponderosa
valf. arlzonrcavat. ponderosavat. scopulorum
P. rad.iataP, resinosaP. serotinq,P. strobif ormisP. strobusP. sylaestrisP. taedaP. airginiana
SAE 20 motor oilSAE 20 motor oilSAE 20 motor oil
!t4251,
90320325 ltSAE 20 motor oil
kerosenekerosenewater_ketosene "
19171
1307111tr,251,191oa<66, 131
68
sic" eih"iotlinseed oiI
SAE 20 motor oil, or 1 part keroie". io a pa.ts ti"seea olt
' Test should be made as soon after picking as possible to prevent excessive drying; the liquids have the followingspecific gravities: kerosene 0.80, 95 percent ethanol 0.82, SAE 20 motor oil 0.88, linseed oil 0.93. Five or morefreshly 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 somespecies, e.g., P. taeda and P. echinata, nearlymature cones can ripen in the crown on felledtrees, but in other species they may not (251).With some species, slightly immature seeds canbe successfully ripened in the cones after re-moval from the tree-(P. elliottii var. elliottii)(23, 25L); in cold moist storage-(P. lam-bertiatta) (121) ; or in proionged cold dry stor-age in closed gsntainsrs-(P. ui,rginiana) (37).Such methods should only be attempted ifcompletely mature seeds cannot be collected.
Cones most often are hand-picked. eitherfrom ladders or by climbing the trees. Forsome species, hand cutters oi a cutting hookmust be used to detach the cones, and hooks maybe needed to bring the cone-laden branches tothe picker. With certain species, mechanicaltree shakers have been used for the rapid har-vesting of cones; e.g., P. elliottii var. ell;iottii,P. palustris, and P. taeda (119). With a fewothers; e.g., P. edulis, P. monophylla, and P.quadrifolia, large amounts of seeds are col-lected by shaking the tree or beating the crownto extract the seeds, and then gathering themfrom the ground (204,235).
Cone processing and seed extraction.-Conesshould be dried immediately after collection toavoid mold development and excessive internalheating, which lead to rapid seed deterioration.Drying can be accomplished in 2 to 60 days by
immediately spreading the fresh cones in thinlayers on a dry surface in the sun, or on traysin a well-ventilated building, or by placingthem in sacks hung from an overhead rackprotected from rain (205, 221, 223, 25/+). Thecones should dry slowly to prevent "case hard-ening." After initial drying, the cones can bestored temporarily in well-ventilated bags ortrays. For many species, ripe cones open satis-factorily under the above conditions, but conesof some species may require additional heat ineither a cone drying kiln or a heated shed. Prop-erly air-dried cones of a few species may opensatisfactorily after a few hours in a kiln, butthose of other species may require several days.Cones of most species can be opened at kilntemperatures not exceeding 130' F. and ahumidity of about 20 percent; but cones of afew specieS, €.g., P. banksiana, P. clattsa, P.ponderoso" var. scopulorum, require highertemperatures (205, 221,223, 235) (table 5).
Cones stored long enough in containers tohave dried without opening or cones dried undercool conditions may not open properly duringkiln drying. In such cases, the cones must besoaked in water for 72 to 24 hours, and thenkiln dried before they will open satisfactorily(223\.
Serotinous cones have beerr opened by dippingthem in boiling water for 10 to 120 seconds.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
Kiln dryingViable period for seeds
in cold storage 2
Species Time inboilingwater
Air-dryingtime 1 Time
Meanuemper-ature
Datasource Time Data source
Seconds000
15-3000
10*30000
010-30
00
" 30-6000
0-1200
0-3000
Days15-30
2-815
1-32-8
3-103-20
z-to
2-8'|
2-20
2-30
2-20tJ--t- D
z-a.)
A
izlb-to
15
3-10
D_I')l-2
4-io
4-ri4-t2
30
Hours000
480
2-400
100
206
12212612678,12278,122I q.)
liz11 4, 138
63, 72963, 12214s23s63, 722100100196,264159, 25[204
oaqr)
25425t,oq<10q
118,122100122,1467qolaa
100
122, 2.13
122,171,63
7AO
254700
178,17/,119 16)6)
23526,722,23526, 122, 235o/100
"F. YearsI 122,166I 166,204
16 isz, pog16 122,20210 122,213
s iip, soe
is iip, sos3 1501+ 1/e
iis
P. albicaulis" -,P. aristata- ..
P, armandii -P. attenuata-P. balfourianaP. banksiana
r20
16ii
r20
t40
t20
105
100130
t20
120
120 3.D
P. brutia
P, canariensisP. caribaea,-. -P. cembra"----.-P. cembroides "P, clausaP. contorta
var. contorta
var. Iatifolia
vat. muffaaand,P. coulteri
P. densifloraP. echinataP. edulis'P. elkottii
vat, densa
var. elliottii
P, engelmanniiP. fleri,IisP, gerardianaP. glabraP. halepensis
P. h.eldreichiiP. insularisP. jeffreyi
P.Iambertiana
P. merkusii. .
P. monophylla n
P, monticola
P. mugoP. muricataP. nigra
P. palustrisP. paruifloraP. patula.P. pinaster
P. pineaP. ponderosa
.val', arxzon1.cavat. ponderosa
var. scopulorum
P. purnila"P. pungens
620
130
02-4
096
06-8
00
720
480
60
0.)
0
720
t45
1nttD
2-535
77
7+
122,202
202, 242
iip, zop122,766
s8, lrr,159, 255
1Ir, 157
122, 202112,166
soe
00000000000000000000000000
15-3000
00000
00
8-100
8-100
6000
4810000
oA
0qA
000
1A
0484824
048
048
0
3-105-205-20
5-7
o-lo-l2-3
120
rto
1io
35 113,296
5 122,219
1+ ii, to10 78,112
18 166,122
2L 202,2/t6
2+ ia
20 ioz, st o
5 112,174
io+ s8
110
rto120-T ID
100
I.LDIID
110120
16i
5-10
2l11
18
18 26,202
15+ 6,
PINAS
T.q.sln 5.-Pittus: cone processing schedules and uiuble periods lor seeds in cold storage-Continued
Cone processing schedule
Kiln dryingViable period for seeds
in cold storage'Species Time in
boilingwater
Air-dryingtime' Time
Meantemper-ature
Datasource Time Data source
P. quadrif olia n
P. radiata
P, resinosa,P. rigida u
P. rosburghiiP. sabinianaP. serotinn,P. sibirica" -P. strobif ormisP, strobusP. sylaestris
P. taeda- -, ,P. tlrunbergianaP. torreyanaP. airginiana
712,202
98, 213 , 24852, 9856, 112
238-so,
ss98,115
159,235202122, 166112,235
Seconds0
60-12060-120
0
000
00000
0-3000
Days2-8
0o-t
Hours0
48-720I
104
9lr62l|6221, 238
1qo
101
176, 2127 0o159, 25.110a 1ry1
Years
it30114J-5
ql
1015
9+11
65+
L20
130
ii
^;d-t
5-205-20
4848
04-t2
10-160
4800,.
L,O105
130120
105
170
'Air-drying times are for a temperature range of 60" to 90' F. When kiln-drying is used, it should be p_recededby an air-drying period that was not reported for most species. A period of 3 to ? days is recomntended rvhere notimes are listed.
'Period after which at least 50 percent of the seeds germinated. Storage temperature ranges for most specieswere either 0' to 5' F. or 33' to 41' F. The iower range is preferred. Seed moisture contents betrveen 5 and 10percent 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 cleaningcones (122). This procedure, by melting the processes, empty seeds of many spec,ies canresins between the ione scales and by wetting be separated from the sound seeds by flotationthe 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-tracted by placing the cones in a large mechan-icai tumbler or shaker, or in a small manualshaker for small lots. Seeds are then dewingedby machines of various types, by being flailedin a sack, or by rubbing. Dewinging of a fewspecies can be simplified by first wetting theseeds, then ietting them dry; wings are loosenedby this method and can then be fanned oat (223,25/t). Care must be exercised in the use ofmechanical dewingers, since they may injurethe seed. Seeds of three species-.P. aristata(206), P. palustris (25tt), and P. syluestris(115)
-are especially susceptible to mechanical
damage and must be dewinged very carefully.The seeds are cleaned by using mechanicaiclipper cleaners, fanning mills, screens, or Viability may be reduced after seeds havegravity separators to remove the mixture of been immersed in an organic liquid such asbroken seed wings, pieces of cone scale, and ethanol, pentane, or petroieum ether. The re-other impurities. duction, however, can be minimized by using
Flot.atiort. IiqtLicl.f or separating Data
Species e trtpt 11 seetis source
P. brutiaP. coulteriP. ech.inataP. echinataP. elliottiivar.
eiliottii.P. glabraP. halepensisF. nigraP. palustrisP. pinasterP. pineaP. sabinian.aP. strobusP. syluestri.sP. taecla
rvater 118water 129959f ethanol 159-water )5 ^lrvater 2 5lr
951 ethanoi 15959/. ethanol 11895/.ethanoi 118,17/tpentane 15895!f ethanol 118,17t+rvater 77,118rvater .122i00!'. ethanol 221petroleum ether .127rvater 159,25I
627
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a short immersion time and by evaporating alltraces of the liquid from the seeds before theyare placed in storage (-25).
When water is used for floating off the emptyseeds, the remaining sound seeds should bedried to a moisture content between 5 and 10percent before the seeds are placed in storage.
Numbers of cleaned seeds per pound andsome data for computing yields of cones andseeds are given for OS species and varietiesin table 6.
Seed storage.-For most pines, high seed via-bility can be maintained for long periods oftime with the proper storage methods. Pinusresinosa seed stored 30 years still producedvigorous seedlings in the nursery (96), as didthose of P. echinata and P. elliottii var. elliottiistored for 35 years (255). Seeds ofmany speciesare now routinely stored for periods of 5 to10 years. Storage temperature and seed mois-ture content are the two most important factorsaffecting the success of seed storage. As ageneral rule, seeds should be dried to a moisturecontent between 5 and 10 percent. Tempera-tures of 0o to 5" F. are preferred for mostspecies for long-term storage (122, 255), btrta range of 33' to 41" F. also has been used.The viable periods for seeds stored under theseconditions are listed in table 5. Seeds of a fewspecies such as P. insularis and P. utalliclianahave remained viable for several years at ordi-nary room temperature (38, 56), but suchstorage is not recommended. Some seed lotsdeteriorate rapidly following removal from coldstorage if they are held at room air tempera-ture before sowing. Seeds should not be re-moved from storage more than a week beforestratifying the seeds at low temperatures,sowing, or testing (251t).
Pregermination treatments.-Most pines oftemperate climates shed their seeds in the fall,and the seeds germinate promptly during thefirst spring. For some species, such as P. cembraot P. peuce germination may take place duringthe second or even third year after dispersal.Pine seeds display highly variable germinationbehavior when sown following extraction orstorage. The type and degree of dormancy varyamong species, geographic sources of the samespecies, and lots from the same source. Seeddormancy may result from prolonged extractionat too high temperatures, and dormancy mayincrease with prolonged storage (97, 122).Seeds of many species ordinarily germinatesatisfactorily without pretreatment, but germi-nation is greatly improved and hastened byfirst subjecting the seeds to cold stratification,especially if the seeds have been stored.
Stratification is accomplished by first soaking
624
the seeds in water for 1 or 2 days and thenplacing them in a moist medium or in a plasticbag and holding them at a temperature between33o and 41' F. for a specified period of time(Chapter 6). Recommended periods for bothfresh and stored seeds of Pi,nus are listed intable 7 for 64 species and varieties.
TArLU'7.-Pinus : recommended cold strati.fica-tion periods (33'-41" F. in a moist medi,um)and other pr e g ermination treatments
Species
Recommendedcold strati-
fication period Datasources
Freshseed
Storedseed
P. albicaulis - ,,..-. -P. aristata-, ,
P. armandiiP. attenuataP, balfouriana ,,..P. banksianaP. brutiaP, canariensis -.-.P, caribaea - -,,.P. cembrat'P. cembroidesP. clausa- -,--P. contorta
vat. contortavar. latifoliavat. murraaana
P. coulteri
P. densifloraP. ech,inataP, edulisP. elliottii
vat. densavar. elliottii
P. engelmanniiP. flenilisP, gerardianaP. glabraP. halepensisP. heldreichiiP. insularisP. ieffreuiP. koraiensis 'P.Iambertiana -
P. merkusiiP. monophylla.P. monticola
P. mugoP. muricataP. nigra. ,...P. palustrisP. paraiflora2. -- ,-P. patulaP. peuce
P. pinasterP, pinea" .-P. ponderosa
.vat. arxzonxc&,, ,
vat. ponderosa , .
vat, scopulorum
Days Days90-120 90-120
0 0-3090 9060 6090 90
0-7 0-70 0-450 0-2000
90-270 90-2700 0-30
2t 2l
0 20-300 30*560280 21-90
0 0-200-15 15-60
0 0-60
30 300 15-6000
2r-90 2r-900 0-30
28 2800
30-42 30-42000 0-60
90 9060-90 60-90
0028-90 28-90
30-120 30-120
000 20-300 0*600 0-30
90 9060 600-60 60-180
100112,20t,o" 100
10q193, 2rr878, 208152, 19615099,722,19r,78,108,722qo
1002t 3, 24463, 12263,108,122,
24697,99,122122,159204
239753 , 22r, 23528599, 2397AA
15,1607 <O
94, 99, 108102,15263, 1088,9,22663,97,108,
1 d),)
102,152
97 , 122, 235 ,237, 246
122,19663, 122,235122, 796, 235235, 25tro70099,108,122,
238105, 152152
15263 , 122, 2t r122,249
00
000
600
030-6020-60
Tlsr,u 1.-Pinus : recommended cold strati'f'ca.--'i;; Tteriod's (33"-41' F. in a mo.'i'st medi'um)
and,' oth'er ' pregermination treatments-
Continued
PINAS
prolonged cold stratification for 6 to 9 months isiiruch irore effective (100, 121). Acid scarifica-tion is not recommended for seeds of pines(122).'
Seeds of P. cembra, P.ltorainsi,s, P. paruifl'ora,and P. si,birica are suspected of having imma-ture embryos at the fime of collection. Ger-mination has been increased by placing theseeds first in a warm moist environment forseveral months and then in cold stratificationfor several more months (table 7, footnote 2)(8,9,173,238).
Germination tests.-For reliable tests of seedviability the seed is allowed to germinate undernear-optimum conditions of aeration, moisture,tempeiature, and light' On the basis of ex-tensive experience and experimentation, stand-ardized sebd tests for a number of pine specieshave been established by the Association ofOfficial Seed Analysts (70), the InternationalSeed Testing Association (108), and certain re-gional organizations such as the Western Foresttree Seed Council (263). Recommendations andprocedures given by these orgariz-ations andbthers who test seedi are summarized in table 8.In addition, the results of seed tests conductedunder known conditions are given for 61 speciesand varieties.
The germination of pine seeds can be effec-tively tested in any medium or container thatprovides good aeration and holds adequatemoisture. For a number of species, light, com-monly supplied by a cool white fluorescent l-a-p'is required for reliable tests. When light isnecess-ary, the usual exposure is 8 hours in each24-hour period. Different temperatures areemployed-in seed testing; constant 68' F. andalternating 86'/68" F. regimes are most com-mon. When alternating temperatures are used,the higher temperature ordinarily is for 8 hoursand tfie lower-is for 16. The duration of mosttests is from 3 to 4 weeks. Usually 400 to 1'000seeds per test are adequate. Germination isepigeal (fig. a).
Cutting tests are commonly uesd for roughdeterminations of seed quality. Such tests canalso provide information on soundness and canbe used as an emergency guide in fall sowing offresh seeds with embryo dormancy. X-raymethods too supply good information on sound-ness. Estimates of viability from the abovetests are the most subject to error, since theseeds are not actually germinated (221 , 223,2sL) (Chapter 7).
Biochemical methods employing a rapid vi-ability indicator such as one of the tetrazoliumcompounds can also be used, but are not gen-erally recommended. The results are highly cle-pendent on the analyst's experience, and the
Species
Recommendedcold strati-
fication period Datasources
Fresh Storedseed seed
P. pu,mila_P.pungens, -
P. quadrif oliaP. radiataP. resinosaP. rigida ,P. ronburghiiP. sabiniana o
P. serotinaP. sibirica "P. strobif ormisP. strobusP. sylaestrisP. taedoP. thunbergianaP. toneyanaP. rirginianaP. wallichiana
Days Days120*150 120-150
000 0-30
0-7 7-200600 0-3000
60-120 60-1200 0-30
60-90 60-9060-120 60-120
60 600 15-90
30-60 30-600 30-60
30-90 30-900-30 300-15 15-90
8,97,722
108, 12297,122,19352,2351q.)122,165,235tt4238/, 191122,193,23591, 101, 23516118,99122, 23597, 23956, 99
'P. cembra: cold stratification period has. been re-duced to 90 days by first soaking the seeds rn concen-iit't"a ii ti""i" "t"ia"tot 3 to 5 hours, or by mechanically.^."iiivits Tt " ieeds,(729, 1g/*), b:ut acid treatment isnot now recommended,'*; i;: ;";b;;',"p. koiaiensis, P. paruiflora, P. sibirica:In some lots embryos may be immature and requre a;;"* ;.i.t stratification period (2 months-at 70" tog0; ir.ip;uceding the cold-stratification period (9, 99,oary\'"iij. edulis (204), P. pinaster (1-52). and ^P'
pineausii,
-Cooa ELrmlnation'has been obtain-ed after soak-
i"i-J""aii" E"ta iao' F.) water for 24 hours with nosu-bsequent stratifi cation.-'; p.-iiiii"t a cold stratification pe,riod.of 60- days wassufficient when seeds were first soaked in sulturtc acroi""'lO-*i"nies (9.q, 122, 23s), but acid treatment isnot now recommended''*; i;: ;'"ii;;iio :- germination is speede.d bv cracking.the*rlcf<' seedcoats, t-etore stratification (122, 165, 235\'
Seeds of some species may exhibit extremedormancy; e.g., those that requile. more than6O duy.'of str'atincation (table 7-) ' The dor-tnu".v may be due to physiological or physicalfactois. A pretreatment may be, needed to over-come a phisiological block in the embryo; e'g',p. i"mUnitiana* (121), or effect a phvsicalchange in the seedcoat to make it permeable towatei; e.g., P. sabini'ana (82, 121). The dor-mancy can also be more complex; -an anatom-ically immature embryo with a physiologi-calblock may be coupled with an impermeableseedcoat as in P. cimbra. Acid scarification ofseedcoats has been used with several species;e.g., P. cerlbra', P. peuce, and P. sabiniana,b:ut
625
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age of the seed. The viability estimates oftenare much higher than the germination capaci-ties obtained from germination tests (Zpg')(Chapter 7).
Nursery and field practice.-Pines are suc-cessfully nursery grown in most parts of theUnited States and in virtually all soil types.The soil should be fertile, and of good drainageand aeration. In most large nurseries the soilis fumigated in the fall or spring before sowingto control soil-borne diseases, insects, nema-todes, and weed seeds. Nursery practices aresummarized for 35 species and varieties intable 9.
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I Lt)ll !iFIcURE A,-Pinus res,inosa, red pine: seedling develop-
ment at 1, 7, and 30 days after germination.
In temperate regions, pine seeds can be sownin the fall or spring. It is now common practiceto spring-sow nondormant seeds. Dormant seedstoo may be spring-sown, but they must be pre-treated. Some nurseries pretreat dormant seedsof all species in the same manner; however,this is not to be recommended. The pregermina-tion treatment used on each specles seed lotshould be that which achieves best germinationfor that lot. Seeds with embryo dormancy canbe sown in fall without a pretreatment. Com-pared to seedlings produced by spring-sownseeds, those from fall-sown seeds are commonlyIarger and better developed after one season.With fall-sown seeds, however, sowing must beIate enough to avoid fall germination, so thatseedlings are not subject to winter freeze dam-age and mortality. Fall-sown beds also are moresubject to losses from rodent damage.
PINUS
The seeds can be either drill-sown or broad-cast by hand or machine, but most largenurseries drill-sow in beds because it is moreeconomical. The amount of seed sown per unitarea and the sowing density vary with thespecies, seed size and germination capacity, andthe desired density. The stock density influencesthe vigor and size of the seedlings and trans-plants produced. The stock density will dependon the species, the length of time seedlings willrerrain in the nursery bed, and whether theywill be transplanted.
Seeds are sown at densities selected to pro-duce from 75 to 75 seedlings per square fbot.Higher tree survival factors are obtained whenmedium-to-low sowing densities are used. Mostnurseries sow seeds at a slightly higher densityif the-seedlings are to be placed in transplanlbeds for one or more years. A lesser densityis desired for 2-0 than 1-0 seedlings. Depend-ing on the species, seed lot, and nursery, sowingdensities range from 2 to 20 ounces of seed per100 square feet of bed. One northwestern nurs-ery drill-sows P. monticola at 35 seeds perlineal foot in rows spaced 3.5 inches aparf toobtain a density of 120 seedling per squarefoot for 2-0 seedlinSs (Zttg\. A second wesrernnursery drill-sows P. monticola at 18 seeds perlineal foot with 6 inches between rows to obt-aina density of 35 seedlings per square foot forits 2-0 seedlings (916). Ultimately, then, ex-perience determines the correct sowing densityfor agiven species and nursery for a particularplanting situation. Average nursery-germina-tion has ranged from 20 to 85 perient of thegermin_ation capacities found ln laboratorytests. Of the seeds that germinate, as little as19 and as much as 90 percent produce useableseedlings; the average has been about 5b per-cent.
At the time of sowing, seeds are drilled orpressed firmlv into the soil. then uniformlvcovered withlA 16 ri inch soil, sand. or mulch.Fall-sown seeds should be placed slighily deeperthan, spring-sown seeds t-o protecfthem frbmwind erosion and frost action. Large-seededspecies, as P. albieaulis, P. lambertinna, and p.mon,oph11lla, are covered to a depth s1 r/2 inch.Smaller seeds require the least coverlng. fhesouthern pines, P. echinata, P. elliottii, p. palu,s-tris, P. taed.a, and P. airgintatta, are pressedinto the soil surface and covered with burlapor chopped pine straw. Such materials protectthe seeds from birds or displacement by rainand help maintain soil moisture. Pinus corttorta(var. cdntorta and. var. latif olia), p. densifl,ora,and P. tlttmbergiana are sown /s inch deep; p.banksiana, P. eanatiensis, P. edulis, and p.
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nxonticola are commonly sown /a inch deep.Care should be taken not to sow the seed toodeep.
Germination is complete for most speciesfrom 10 to 50 days after spring sowing. Butsome lots of dormant seed species, even afterpregermination treatment, may continue togerminate several months to a year after sow-ing; e.g., P. albicau,lis, P. cembt'a, P. peuce, andP. strobif ormis (122).
In most nurseries, fungicides are used to con-trol damping-off, and sprays are used duringthe season to control insects and other diseases.fn southern nurseries the fusiform rusts on P.taeda and P . elliottii and brown spot on P . palus-tris, and, in Lake States nurseries the sweetfernblister rust on P. banksiana, P. sylaesf.r'is, andP. Ttonder"oso are controlled by repeated applica-tions of fungicides.
Generally, transplants or older age classes arerecommended for difficult sites. In the LakeStates and Prairie Plains, for example, P.banksiana is grown as 1-0 or 11/z-0 stock foreasy to average sites, 1-1 or 2-0 for difficultsites, and 2-1, l-2, or 2-2 for windbreaks. Mostof the white pines are grown as 2-0 and 3-0or as transplants 2-1 and 2-2. Tn specializednurseries, pines are routinely grown as con-tainer plants. Either young seedlings or seedsare planted directly in containers with a soilmix. Depending on the nursery, partial shadeis provided during the germination and seedlingestablishment phases. Normally, container-grown pines are cultured 1 to 5 years beforeoutplanting. Care must be taken not to growpines in too small a container for too long atime since they will become container-bound orrootbound.
Methods of mulching. watering, shading, pestcontrol, lifting, and other nursery operationshave been described in several regional hand-books (95, 107, 221, 223, 25/t). Each one con-tains much useful information on nurseryproduction of pine seedlings.
Ali pine species can be vegetatively prop-agated either by rooting or grafting (122, 178,231). However, rooting success for most speciesdecreases rapidly when scions are taken fromtrees older than 5 years. PitttLs radiata, P. at-tenuata, P. den.siflora, and P. thunbergianaare relatively easy to root. But only P. radiatais extensively propagated by rooting cuttingsunder nursery and greenhouse conditions (230).Selected trees of many other species are clonedby rooting of cuttings. Grafting also is rou-tinely used to propagate rare material or toclone individual plants, as in seed orchard pro-grams designed to produce genetically improvedforest tree seed (122).
PINAS
Literature and Other f)ataSources Cited
(1) Albert, R.Data filed 19?0. USDA Forest Serv., Region
3, Coronado Natl. Forest.(2) Allen, R., and Wardrop, A. B.
1964. The opening and shedding mecha,rismof the female cones of Pin.us racliata. Aust.J. Bot. 12: L25-134.
(3) Allison, A. C.Correspondence, 1969. Virginia Dep. Nat.
Resour.(4) Altman, P. L., and Dittmer, D. S.
1962. Biological handbook of growth. 608 p.Fed. Am. Soc. Exp. Biol., Wash., D. C.
(5) Andreev, V. N.1925. Pintrs peuce Griseb., 1n [Dendrology,
vol. I, Gymnosperms p. 96.1 Ukraine StatePrinting 0ffice. (In Russian.)
(6) Andresen, J. W.1963. Germination characteristics of Pintts
rigida seed borne in serotinous cones.Broteria 32 (3-4) : 151-178.
(7) -----1965. Stratification of seed of Pinus strobi-
fornis. Tree Plant. Notes no. 72, p. 5-7,(8) Asakawa, S.
1957. IStudies on hastening the germinationof the seeds of the five-leaved pines.] Gov.Forest Exp. Stn., Meguro (Tokyo) Bull.100: 41-54. (In Japanese.)
(e)
-Correspondence, June 17, 1969. Minist. Agric.
For., Meguro, (Tokyo) Japan.(10) Association of Official Seed Analysts.
1965. Rules for testing seeds. Proc. -t-ssoc.Off. Seed Anal.54(2): 1-112, illus.
(11) Baker. L. A.Correspondence, 1969. Oreg. State Forest.
Dep., Dwight L. Phipps Forest Nursery,Elkton, Oreg.
(12) Railey, L. H.1939. The standard cyclopedia of horticul-
ture. 3 vol., 3,639 p. The Macmillan Co.,New York.
(13) Barnett, J. P.Data filed 1968. USDA Forest Serv., South.
Forest Exp. Stn., Nerv Orleans, La.(14)
1969. Long-term storage of longleaf pineseeds. Tree Plant. Notes 20(2\: 22-25.
trDt -- -1970. Flotation in ethanol affects storabilityof spruce pine seeds. Tree Plant. Notes21 (4) : 18-19.
(16) ---- and Mclemore, B. F.1967. Improving storage of spruce pine seed.
Tree Plant. Notes 18(2): 16.(17) -- s1l Mclemore, B. F.
1967. Study of spruce pine cone maturity andseed yield. Tree Plant. Notes 18(2): 18
(18) Barton, L. V.1930. Hastening the germination of some
coniferous seeds. Am. J. Bot. 17: 88-1.15.(19) Bates, C. G.
1930. The production, extraction. and germi-nation of lodgepole pine seed. U.S. Dep.Agric. Tech. Bull. 191, 92 p.
(20) Bennett, M.Communication, 1969. Arizona Cypress Gar-
dens, Sedona, Ariz.
631
PINUS
(21) Benson, D. A.Data filed 1969. USDA Forest Serv.. Eastern
Tree Seed Lab., Macon, Ga.(22) _
Data filed 1970, USDA Forest Serv., EasternTree Seed Lab., Macon, Ga,
(23) Bevege, D. I.1965. An investigation of cone and seed
maturity of slash pine in southern Queens-land. Aust. For.29(3): 135-148.
(24) Bonninghausen, R. A.Communication, 1968. Florida Forest Serv.,
Tallahassee.(25) Boskok, T. E.
1970. ISeed maturation period in Pinusbrutia, Picea orientolis and Abies born-muelleriana.l Orm. Arast. Enst. Tek.Butll. 42: 64 p. (In Turkish.)
(26) Boyd, R. J., Jr.Data filed 1970. USDA Forest Serv., Intermt.
Forest and Range Exp. Stn., Moscow,Idaho.
(27) Britton, N. L., and Shafer, J. A.1908. North American trees. 849 p. Henry
Holt and Co., Nev'York.(28) Brown, J. H.
1969. Variation in roots of greenhouse grownseedlings of difrerent Scotch pine prove-nances. Silvae Genet. 4(4\: 111-112,(29) Burns. R. M.
Data filed 1968. USDA Forest Serv., South-east. Forest Exp. Sta., Marianna, Fla.
(30) Byrd, R. E.Communication, 1968. Labelle. Florida.(31) Callaham, R. Z.1962. Geographic variability in growth of
forest trees. In Tree Erowth. T, T. Koz-lowski (ed.), p. 311-325. Ronald Press Co.,New York.
(32) --- 1963. Provenance research: investigation of
genetic diversity associated with geog-raphy. Unasylva 18(2-B) :78-74.
(33)
-
and Liddicoei, A. R.1961. Altitudinal variation at 20
ponderosa and Jeffrey pines.59(11): 814-820.
(40) Cooling, E. N. G.1968. Fast growing timber trees of the low-
land tropics. Pinus merkusii. No. 4. L69 p.Commonw. For. Inst., Dep. For., OxfordUniv.
(41) -- and Gaussen, H.
1970. In Indo China: Pinus m,erkusiana sn.nov. et non P. merkuszli Jungh. et de Vriese.Trav. Lab. For. Toulouse Tome 1 Vol. 8.Art. T, 8 p.
(42) Cnm, W. H., and Brack, C. G. E.1953. Performance of Scotch pine races under
prairie conditions. For. Chron. 29G\:aqi rAo
(43) Critchfield, H. M.Correspondence, October 9,
Mountain Tree Farm andHelena, Calif.
1969. GlassNursery, St.
(34) Carlisle, H., and Brown. A. H. F.1968. Biological flora of the British Isles.
Pinus sylaestris L. J. Ecol. b6(1): 269_307.
(35) Cerepnin, V. L.1964. [The importance of Pinus syluestris
seed origin, weight, and colour in selec-tion. I Sel. Drev. Porod er Vost. Sibiri. n.58-68. (In Russian.)
(36) Chapman, H. H.1922. A new hybrid pine (Pinus palustris X
Pinus taeda). J. For. 20t 729_784.(37) Church, T. W., Jr., and Sucoff. E. I.1960. Virginia pine seed viable two months
before natural cone opening. USDA ForestServ., Northeast. Forest Exp. Stn. Res.Note 102, 4 p.
(38) Claveria, J. R.1953. Growing Benguet pine (pinus insu-
laris) in Cebu Province.-philippine J. For.9:57-76.
(39) Cocozza, M. A.1961. Osservazioni sul circlo riproduttivo di
Pin.us heldreicftii Christ. var. leucorlermisAnt. del monte pollino. Accad. Ital. Sci.For. 10: 9T-110. (In Italian.)
632
(44) Critchfield, W. B195 ?. ileographic variatio n in P inus contor ta.
Maria Moors Cabot Found. Publ. 3, 118 p.(45) __
1963. Hybridization of the southern pines inCalifornia. South. Forest Tree Imnrov.Comm. Publ. 22: 40-48.
(46) __1966. Phenological notes on Latin American
Pinus and. Abies, J. Arnold Arbor. 47(4\:313-318.
(47)
-
and Allenbaugh, G. L.1965. Washoe pine on the Bald Mountain
Range, California. Madroiio 18(2): 68-64.(48)
-
and Krugman, S. L.1967. Crossing the western pines at Placer-
ville, California. Univ. Wash. Arbor. Bull.30(4): 78*81,92.
(49)
-
and Little, E. L.. Jr.196-6. Geographic distribution of the pines of
the world. U.S. Dep. Agric. Misc. Publ.991, 97 p.
(50) Curtis. J. D.1955. Effects of origin and storage method
on the germinative capacity of ponderosapine seed. USDA Forest Serv.f Intermt.Forest and Range Exp. Stn. Res. Note26,5 p.
(51) Dallimore, W., and Jackson, A. B.1967. A handbook of Coniferae and GinkEo-
aceae. Ed. 4, rev. by S. G. Harrison, 229p. St. Martin's Press, New York.
(52) Dansbury, C.Correspondence, 1968. Washington Crossing
State Forest Nursery, Titusville, NewJersey.
(53) Day, R. J.196?. Whitebark pine in the Rocky Moun-
tains of Alberta. For. Chron. 4B(B) | Z7g_282.
(54) Debazac, E. F.1964. Manuel des conifdres. 172 p. Ecole Nat.
Eaux For6ts, Nancy.(55) Delevoy, G.
1935. Note preliminaire sur I'influence deI'origine des graines chez le pin maritime.Bull. Soc. Cent. For. Belg. 42(g): 9?_105.
(56) Dent, T. V.1.947. Seed storage with particular reference
to the storage of seed of Indian forestplants. Indian Forest Rec. (N.S.) Silvic.7(1): 1-134.
years inJ. For.
(74)
(75)
(52; Derr, H. J.1955. Seedbed density affects longleaf pine
survival and grorvth. Tree Plant. Notesno. 20, p.28-29.
Dimitroff, I.1926. [Study of the seed material of Pinus
peuce.l Ann. Univ. Sofia. Fac. Agric. 4:259-306. (In Bulgarian.)
Dorman, K. W., and Barber, J. C.1956. Time of flowering and seed ripening in
southern pines. USDA Forest Serv.,Southeast. Forest Exp. Stn. Pap,72,75 p.
Duff, C. E.1928. The varieties and geographical forms
of Pinus pinaster Soland., in Europe andSouth Africa, with notes on the silvi-culture of the species. S. Afr. Dep. For.Bull. 22-a, 55 p.
Duffield, J. W.1951. Interrelationships of the California
closed-cone pines with reference to Pinusmuricata D. Don. Ph.D. Thesis Univ. Calif.Berkeley. 114 p.
1953. Pine pollen collections dates-annualand geographic variation. USDA ForestServ., Calif. Forest and Range Exp. Stn.
cone ripe-Notes no.
R,es. Note 85, I p.Eden. J.
Correspondence, 1969. Calif. Div. Forest.,Davis State Forest Nursery, Davis, Calif. (g2)
Edwards, M. U.
PINUS
and Lester, D. 'f.1970. Genetics of red pine. USDA Forest
Serv. Res. Pap. WO-8, 13 p.Fritts. H. C.
1969. Bristlecone pine in the White Moun-tains of California, Growth and ring-widthcharacteristics. Tree-Ring Pap. 4, 44 p.Univ, Ariz. Press, Tucson.
Gifford, E. M., and Mirov, N. T.1960. Initiation and ontogeny of the ovulate
strobilus in ponderosa pine. Forest Sci.6(1); 19-25.
Goor, A. Y.1955. Tree planting practices for arid areas.
FAO Forest. Dev, Pap. No. 6, 126 p.and Barney, C.'W.
1968. Forest tree planting in arid zones. 409p. Ronald Press Co., New York.
Graber, R. E.1965. Germination of eastern white pine seed
as influenced by stratification. USDAForest Serv. Res. Pap. NE-36, 9 p.
Graber, R. E.Data fiIed 1967. USDA Forest Serv., North-
east. Forest Exp. Stn., Upper Darby,Penn.
Data fi]ed 1970. USDA Forest Serv., North-east. Forest Exp. Sta., Upper Darby,Penn.
Griffin, J. R.1962. Intraspecific variation in Pirzas sobi-
niano Dotgl PhD thesis, 274 p. Univ.Ca1if., Berkeley.and Conkle, M. T.
1967. Early performance of knobcone andMonterey pine hybrids on marginal timbersites. USDA Forest Serv. Res. Note PSW-156, 10 p.
Haasis, F. W., and Thrupp, A. C.1931. Temperature relations of lodgepole
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and Czabator, F. J.1961. Length of stratification and Eermina-
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( 143)
635
PINAS
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637
PINAS
(24r\
-Data filed 196?. Project 2302 fot developrnent
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(242\ __Data filed 1969. Bend Nursery, Bend, Oreg.
(243\ -- Data filed 1969. Coeur d'Alene Nursery,
Coeur d'Alene, Idaho.(244\
--- Data filed 1969. Lucky Peak Nursery, Boise,Idaho.
(245\
-Data filed 1969. Mt. Sopris Tree Nursery.
Carbondale, Col.(246) _
Data filed 1969. Placerville Nursery, Placer-ville, Calif.
(247\
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