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Sources of Loblollg Pine - Managing Benefits and Risks

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Planting Nonlocal Seed Sources of Loblollg Pine - Managing Benefits and Risks Clem Lambeth. Soirtlzc)j-r~ TI-re I~~zpi-oi~er?lc~zt. Wc~!.r1-/incii.sci- Cor~iptr,l~.. Hot S17ririgs. AR 71 901; Steve McKeand, College (?f1Vlirili-(ii Resoicl-ces. lYor-tlz CL~I-olirzct Stcitc. L./2ii*e,-.sit~.. Raleigh. 'VC 2765.5; Randy Rousseau, Cefit~-ci/ FOI-est R~~S~LII-L.~ Slt~tiot?. l%fecidllie~ti~cim, U'icsX-1iJSr. KY 12007: ~r~d Ron Schrnidtling, Sout11e1-11 lrictitittr of Foresr Grrietics, L1.S. Foi-e.st Sei-rice, Sarrcilv-, MS 351574. ABSTRXCT: Seed rorrtr.c tesritlg of lohioll! p1tze (Pinu\ taeda), \t.lzich Degcifl 111 the IY20s, /?us aliolt.ed large recrlized prizetic pliitz.r fr-at?t ir.sir1~ ~~oizloc~cil seed so~i~-ces it? c~pet-crrioftiil pla~ztnrrot~s. Seed source testirzg ~~otlritl~tes, n~td deplo!.t?ie/lr g1ridelirte.r are stiil beiizg I-efi~led. Sotrze gette~~rl ej5ect.s of seed source nzo13einerzt c ~li~ be descr-ibed. k~ct riret-r are still gcips it7 (If ri~zde~-~tcr~ilit e.~~cti~ hat fa)- certuir? seed sorrrces cirtl be f?tol'e(J, ( 2 )112e degree of risk irli*ol\ ed. crlzd (3) Izoit. ct't-ruiit I~CI~IS SIICII IJS \tsood qllfllih i'a)? b!. seed sorrt-ce, erpecrtrll? \c.itii seed sorit-ce t7rat'el?reilt. Zrl Jorlze cases, seed soirrce tno\Tet?lerzt gaitrs can be acilie\*ed \t.irh little I-zsk: ,fir e.\-crnzple. pla~.rtit?g Lil.ilzgstotl Pnt-ish, Loltisic~izn t?zcirerial for rrrsr l-esistnt~ce in rnore eusterl~ GrrlfCoa.sru1 rrrerrJ. .-l!sct. r?zo~*r~?ze~zt of reed soltrces otte plant lzcrr-drrresszctrle 11orih can res~llt it7 irlcrensed grc~rt-111 ~t.itl1 little COIIC'~~-I~ for ~t.it?tet. rini~zage. Big gains it1 QI-oitrh, I?o\te~.el-, porn usitlg nor.rlocai seed solft-ces 171~1~ come cit srgnificnr~trisk. T~t-0 industr-iul escz17zpies of plantirzg trotzlocul seed sources atzd h o ~ . risks n el-e mntzcrged ore col el-ed: i I, Sorrth-to-north r?zrtr*enzelzr: Meadll/esr\.aco's tlse qf lob loll!^ pirze rot-tlz ofrlze ~rcrti\*e rcrirge in Kentuck?. u~zd r~lrroiiizdilzg ur-ecrs, and (21 Ensr-to-rr est tno\let?zet1r: We~.erl?aerr.ser's Lire of ,Yorth Ccu.olrrrri coastcri plcrir~fnl?tilies in .roritherrz il.rknt1sa.s a ~ t d soliri~ecrsr Oklnhot~zn. To deal ~tith the sigi1iJicntzt risks qf seed source 11rot.er71etzrs, otze t?ziist be nit are qf the risk factors, understui~d historic.ni clttizcrtic dcrtii ((71-e rlze risks hrgh ot- lolt. li.ithit1 a tl\ pica1 lzan~est I-oturion period), crtd ha1.e sill-icrilt~rral ci~zcl getzeric* stt-iirrgies to t?zirigtrte or i-educe I-isk. Po~siDleyettetic SII-arepies ir~clrtde rho/-o~tgh resting nrzd alloc.trtiorz of 01-cl1urd far?zilies qf tlze tzctizloc.al seed sct~ct-ce. de~.elopmetzt qf n "lnttd mce " (br-eeditzg und tesritzg for local ndaptariot1 of rile rloilloc~~l seed source), i111etpr-oi~er1ui.1ce h~ brids, aid irtterspec!fic. i~?.brid~. Erol7lples of rilest. czre dirciissed ii? t h i ~ article. Sorkriz. J. ,-lppl. For. 29(2 I: 96- 193. Key \Vords: Seed source. genetic gain. rtdaptabilit>. pro\.enance, hybrid. Importance of Nonlocal Seed Sources Reasons for numerous cases oi operational p~antingsoi nonlocal seed sourcec in the mutheastern Lnited State\ on the part of pri\ ate Iandou ners include: Increased fus~form rust iC~-oir~ir:r~ci?l tjircrc ~cunl. i E3erL.j Miy. Ex Shirai f. sp. Fir\ifr~i-,~lc~) resictance. For exam- ple. planting of Li\ ingcton Pari\h. Lou~tiana marer~al in high ru\t hazard areas 01' Flor~da. and southern Georgia, hlissiisipp~, and .llabama. Increa\ed grou th rate. For example. mo\ lng mater~al sIighti> north ~ i t h low ri\h of %Inter damage and making material best M rth IOU ri\h of drought damage. Increalled \ur\ i\ al. For example. planting drought-tol- ermt uestern source\ on the sand hrll\ of Georgia znd Alabama. * C\e of improxed famiires from source\ that are more geneticallq ad\ anced than the local seed jource. No trul_t local seed \ource. For example. pltintlngs just outside the lobloll! narural range. No Iooal seedlings for \ale ttn the open market. Occa- \ionalfy. a Iandouner nu) not be able to bu! seedlings from a truly local \eed wurce and must re\ort to the be\t alternative. NOTE. Clem Larnhrth can be reached at i 50 l )h2l-"; 'I 0, Fdx (50 1 634-S505; cIem.larnheth@ c\ieyerhaeu\rr corn 'L?anu\cripr rectsited Augti\t 23, 1003. accepted April 13, 3004 The old adage that "local is best" applies in the absence of information from well-conducted seed source trials but 96 SJAF 99(2 1 9005
Transcript
Page 1: Sources of Loblollg Pine - Managing Benefits and Risks

Planting Nonlocal Seed Sources of Loblollg Pine - Managing Benefits and Risks

Clem Lambeth. Soirtlzc)j-r~ TI-re I~~zpi-oi~er?lc~zt. Wc~!.r1-/incii.sci- Cor~iptr,l~.. Hot S17ririgs. AR 71 901; Steve McKeand, College (?f1Vlirili-(ii Resoicl-ces. lYor-tlz CL~I-olirzct Stcitc. L./2ii*e,-.sit~.. Raleigh. 'VC 2765.5; Randy Rousseau, Cefit~-ci/ FOI-est R ~ ~ S ~ L I I - L . ~ Slt~tiot?. l%fecidllie~ti~cim, U'icsX-1iJSr. KY 12007: ~ r ~ d Ron Schrnidtling, Sout11e1-11 lrictitittr of Foresr Grrietics, L1.S. Foi-e.st Sei-rice, Sarrcilv-, MS 351574.

ABSTRXCT: Seed rorrtr.c tesritlg of lohioll! p1tze (Pinu\ taeda), \t.lzich Degcifl 111 the IY20s, /?us aliolt.ed large recrlized prizetic pliitz.r fr-at?t ir.sir1~ ~~oizloc~cil seed so~i~-ces it? c~pet-crrioftiil pla~ztnrrot~s. Seed source testirzg ~~otlritl~tes, n ~ t d deplo!.t?ie/lr g1ridelirte.r are stiil beiizg I-efi~led. Sotrze get te~~rl ej5ect.s of seed source nzo13einerzt c ~ l i ~ be descr-ibed. k~ct riret-r are still gcips it7 ( I f r i ~ z d e ~ - ~ t c r ~ i l i t e . ~ ~ c t i ~ h a t fa)- certuir? seed sorrrces cirtl be f?tol'e(J, ( 2 ) 112e degree of risk irli*ol\ ed. crlzd ( 3 ) Izoit. ct't-ruiit I ~ C I ~ I S S I I C I I I J S \tsood qllfllih i'a)? b!. seed sorrt-ce, erpecrtrll? \c.itii seed sorit-ce t7rat'el?reilt. Zrl Jorlze cases, seed soirrce tno\Tet?lerzt gaitrs can be acilie\*ed \t.irh little I-zsk: , f i r e.\-crnzple. pla~.rtit?g Lil.ilzgstotl Pnt-ish, Loltisic~izn t?zcirerial for rrrsr l-esistnt~ce in rnore eusterl~ GrrlfCoa.sru1 rrrerrJ. .-l!sct. r?zo~*r~?ze~zt of reed soltrces otte plant lzcrr-drrress zctrle 11orih can res~llt it7 irlcrensed grc~rt-111 ~t.itl1 little C O I I C ' ~ ~ - I ~ for ~t.it?tet. rini~zage. Big gains it1 QI-oitrh, I?o\te~.el-, porn usitlg nor.rlocai seed solft-ces 1 7 1 ~ 1 ~ come cit srgnificnr~t risk. T ~ t - 0 industr-iul escz17zpies of plantirzg trotzlocul seed sources atzd h o ~ . risks n el-e mntzcrged ore col el-ed: i I , Sorrth-to-north r?zrtr*enzelzr: Meadll/esr\.aco's tlse qf lob loll!^ pirze rot-tlz ofrlze ~rcrti\*e rcrirge in Kentuck?. u~zd r~lrroiiizdilzg ur-ecrs, and (21 Ensr-to-rr est tno\let?zet1r: We~.erl?aerr.ser's Lire of ,Yorth Ccu.olrrrri coastcri plcrir~fnl?tilies in .roritherrz il.rknt1sa.s a ~ t d soliri~ecrsr Oklnhot~zn. To deal ~ t i t h the sigi1iJicntzt risks qf seed source 11rot.er71etzrs, otze t?ziist be nit are qf the risk factors, understui~d historic.ni clttizcrtic dcrtii ((71-e rlze risks hrgh ot- lolt. li.ithit1 a tl\ pica1 lzan~est I-oturion period), c r t d ha1.e sill-icrilt~rral ci~zcl getzeric* stt-iirrgies to t?zirigtrte or i-educe I-isk. Po~siDle yettetic SII-arepies ir~clrtde rho/-o~tgh resting nrzd alloc.trtiorz of 01-cl1urd far?zilies qf tlze tzctizloc.al seed sct~ct-ce. de~.elopmetzt qf n "lnttd mce " (br-eeditzg und tesritzg for local ndaptariot1 of rile rloilloc~~l seed source), i111etpr-oi~er1ui.1ce h~ brids, a id irtterspec!fic. i~?.brid~. Erol7lples of rilest. czre dirciissed ii? t h i ~ article. Sorkriz. J. ,-lppl. For. 29(2 I: 96- 193.

Key \Vords: Seed source. genetic gain. rtdaptabilit>. pro\.enance, hybrid.

Importance of Nonlocal Seed Sources R e a s o n s for numerous cases o i operational p~antings o i nonlocal seed sourcec in the mutheastern Lnited State\ on the part of pri\ ate Iandou ners include:

Increased fus~form rust iC~-oir~ir:r~ci?l tjircrc ~cunl. i E3erL.j Miy. Ex Shirai f. sp. Fir\ifr~i-,~lc~) resictance. For exam- ple. planting of Li\ ingcton Pari\h. Lou~tiana marer~al in high ru\t hazard areas 01' Flor~da. and southern Georgia, hlissiisipp~, and .llabama. Increa\ed grou t h rate. For example. mo\ lng mater~al sIighti> north ~ i t h low ri\h of %Inter damage and

making material b e s t M rth I O U ri\h of drought damage. Increalled \ur\ i \ al. For example. planting drought-tol- ermt uestern source\ on the sand hrll\ of Georgia znd Alabama.

* C\e of improxed famiires from source\ that are more geneticallq ad\ anced than the local seed jource. No trul_t local seed \ource. For example. pltintlngs just outside the lobloll! narural range. N o Iooal seedlings for \ale ttn the open market. Occa- \ionalfy. a Iandouner n u ) not be able to bu! seedlings from a truly local \eed wurce and must re\ort to the be\t alternative.

NOTE. Clem Larnhrth can be reached at i 50 l )h2l-"; 'I 0, Fdx ( 5 0 1 634-S505; cIem.larnheth@ c\ieyerhaeu\rr corn 'L?anu\cripr rectsited Augti\t 23, 1003. accepted April 13, 3004

The old adage that "local is best" applies in the absence of information from well-conducted seed source trials but

96 SJAF 99(2 1 9005

Page 2: Sources of Loblollg Pine - Managing Benefits and Risks

ma! not hold up once trial re\ult\ are akailable. Due to the potitical circumstances that often surround local terws non- local \eed source\. e\peciallj in the \ale of wedlings to

small priiate landouncr\. the Southern Forert Tree Im- proternent Co~nmittee (made up of uni~ers~tb . \[ate, federat. and in dust^ fcjre\t _renetlci\t\) iswed a SFTIC Porltion Statement in 1990 (unpubii\hed i . There \\ere \e\ era1 points in that po\ition \tatemen[ all of ~ h r r h are \ t i l l ialid and uieful toda!. Arnung them are:

" L't?irrfr~r~?rrcl l r 5 0 of !oc cri (oii/-c.r I \ uor 1 2 ~ 1 ti(r \cltfi!\. e~th~- ' r tilt. hert or rcrji~it citoiir. " ~E\limpIe\ itirnllar to those abote are en ). "Gcrrotir. cirfl>>,-citc.er eijtrorlg seed .roirrc*c.\ prct~.iil~ (it] oi~i7o~-r~i1llt~ for ~ t ~ f i ) r ~ t ~ ~ ~ d I ~ ~ ~ ? c J o \ ~ ' J z - cr-.s ro if~~.r.(~cl.s~ \ I C ' ) ~ ' , /?i j~ id i~io t i ( ~ ? ? u ~ c ' / z I I ~ ~ of I Z O ~ I / O C . I /

.rorrrceJ fct rpec.tfic pimt t r t ~ g et?\-i~nrlr?~~~~zr.r. "

Since \eed source moi ernent is a reality that i q not lil\dy to abate. the best course of action for the future ic to ensure that the forestrq cornmunit> is well ~nformed on the use of nonlocal seed source5 and the risks thereof.

Historical Seed Source Study Results The grandfather of all loblollj pine seed transfer studies

was Philip C. Wakele!'s Bosalusa. Louisiana. planting of 1927. The local Lixingston Parish. Lousiana loblolly pro- duced about twice the wood iolume through age 12 as did the loblolly from Arkansas. Georgia. and Texas. These differences persisted through age 35 (Wakeley and Bercaw 1965). reinforcing the widel> held belief that local seed sources are best. Differences in fusiform rust susceptibility uere also observed. uith the Texas and Arkansas sources being very resistant. the Liiringston Parish being moderately resistant. and the Georgia source being susceptible. Wake- ley's study had tho important shortcomings. The test was planted in only one location. and it uas not replicated.

The results of \hTakelel's pioneering study led to the establishment of the Southu ide Southern Pine Seed Source Study (SSPSSS). u hich u9as a cooperative effort initiated in 195 1 by the Southern Forest Tree Improl ement Committee. The early results of the SSPSSS gake the first indications that local sources ma! not: be the best for grou th and disease resistance (Wake19 196 1 . %'ells 1983. Wells and Wakeley 1966 j.

SimiIar types of seed source studies tn other forest tree species indicated that sources from harmer climates tend to grow faster than 10caI sources. ~f these sources are not moved to great11 differing climates. In lobIo1lj pine. thls is at least partly due to the \\arm-climate sources g r o ~ i n g longer in the fall than the sources from colder climates rJayawrckrama et al. 199th). Clirnatrc modeling of data from man) southern pine seed source ctudies has shown that the most important factor influencing yroueth and survival within their natural ranses is at erage ! etirlj minimu~n tem- perature at the seed source (Schmtdtling 2001). This cli- matic varlable has been used. not coincidentally, by horti- culturists to determine plant hardine.ss zones IUSDA 1990).

As far as seed transfers are concerned. the most impor- tant observation made in the SSPSSS and reinforced by a

study planted in \outhrrn Arlta~l\a\ tGrig\by 1973, Well\ and Lanlbeth 1983) &as that the \eed Tourcert from wect of the M I \ \ I \ \ I P ~ I R ~ i e r igere more d~wase and drought re\is- tant, but slouer grn\s rng than eastern \ource\. The one exceptton to t h ~ s Ea\u?;l'est character~zat~on 15 the ell- hnov, n t i \ rngsron Par~\h, L 4. iobiolt\. u hlch is located jurt to the Ea\t of the ,I\Il~\\l\\tppi Rl\er Thl\ \ource con~bines the fk,t gr(?%th. \inl~lar to eastern ~ u r c e s . and ru\t resis- tance \irnlIar to tse\tern \ource\ Thl\ situation undoubtedtj deriier from gene flou the JIil\issipp~ Riier Vallej In 3n eastiiard dlrectlon (Schmidtling et a1 1999).

Ob\er\at~ons from the\e tuo \tudies and cttherc haie re\ulted In large-xale n~otement of \eed \ources ea\t and ue\t acre\\ the hl i \ t~ \s tpp~ R i ~ e r In the eaitmard direction, loblollj ptne from Liiingston Par~\h. LA. and Ea\t Texas were planted In arear of hlgh rust hazard In blis\issippi. Alabama. Georgia. and Florida (Wells 1985 j. Thls was a kfery cuccessful interim 5olution before rust-reGstant strains of eactern seed \ource\ here deteloped. In the westward direction. large quantities of Carolina Coastal Plain seed sources uere pIanted rn southern .Arkansas (Lambeth et al. 1984). T h ~ s has resulted in large gains in gros th o\er local sources.

When grou n in ent rronments u here their performance is not dramatically altered bj maladaptatlon due to cold or drought. ieed sources hake the foilouing traits. in general (Wakeley 1961. Wells and Wakeleq 1966. Grigsby 1973. Wells and Lambeth 1983, Schmidtling 2001). Starting with northeast Flonda seed collectrons and gorng to more north- ern or western sources:

Growth rate: (as long as there are no maladaptarions): Decreases strong11 north and moderately west. Stem straightness: Increases strong11 north and moder- ately west. Fusiforrn rust resistance: Increases moderately north and strongly uest. Specific gratrity: Although there h a ~ e been a large number of reports of the effect of seography on specific g r a x ~ t ~ . there has been little uork on provenance dif- ferences in loblolb plne as compared M ith the plethora of results for grow'th and mct resistance (Zobel and van Buijtenen 1989). The more definitive studies suggest it increases moderatel! both north and uect ijayamick- rama et al. 1998a. Tauer and Loo-Dinkins 1990). This genetic trend IS opposite to the plantation (geographic) trend. Specific gratpity is often high in uarrn regions uith high summer ra~nhl l due to the longer growing season in those regions resulting in a higher proportion of sumerki~ood in the ring iZobe1 and \*an Buijtenen 1989). Therefore. plantations of Virginia seed source in Florida would have higher specific grai ity than the native plantations. Cold tolerance: Increases verj stronglj north, no strong effect west within the same plant hardiness zone. Drought tolerance: Increases moderately west, no known effect north.

Page 3: Sources of Loblollg Pine - Managing Benefits and Risks

These trend\ are not alktay\ 3 griidual cline with latitude or longitude becau\e the di\junct eaitern and we\tern pop- ulations are \ometime\ more different than the distance betueen them would ruggeit. Thi\ lnaj be due to a different evolutionary migratic,n of the tuo populations or a founder effect u hen the ~ L I o iiere \eparated eons ago. Trendi in \eed source trait\ appear t o be a\\ociated more uith changes in cl~mate. \pecificaiI temperature and rainfall. at the origin and seem to hake little to do LX lth \oil t! pe though the? are not a l ~ a y c complete1 independent (Schmidtling 3001 1.

The pre\iouil> mted \eed wurce difference5 are the rationale for attempt., to attain greater plantation yield or qualit) through the u ~ e of nonlocal \eed iources. However. there are known limit\ to \eed murce mokement, \ome of which can be disa\trous. il/Io\t notable of there is the fact that mocements e\en moderate distances north can result in poor plantation performance. Again. a quote from the S R I C Comn~ittee Poiition Statement:

". . . seed soz~rce dlfler-etzces c.un be cr 170tetztii11 pi-oble~?~ for urziizforr?ted lnt1~fui1'1ters '~11.110 ~ i i z k i ~ ~ ~ t ~ i i ~ g l j pfc~izt C ~ F I ill?- pi-ope!- seed sortl-ce. Ln\t.e~- yielcis c1rld e l m plcirztntioit fnilul-es ci1i7 I-esitlt. "

The conclusion thus far is that there are benefits to be had in using nonlocal seed sources. but there are definite risks. How does one go about mahng seed source decisions?

Considerations for Using Nonlocal Seed Sources

Considerations fall under tcx o broad categories: What are the potential benefits and risks? Benefits

Potential benefit5 must be evaluated in the context of the intended product u ith a thorough understanding of exactlq what trait benefits can be deriied for the intended product. Can the nonlocal seed source probide increased ~ r o u t h . quality, disease resistance. cold tolerance. or drought toler- ance? A landowner LI ho is pnmarii> interested in high-qual- ity lumber, for example. ma! do well to select sources with a high degree of straightness and fatorable uood quality traits if wood quality kill be ialued in the marketplace at hamest urns.

Cost and gain are tuo cntical componentc in financial decision-making. Hou et er. the differential cost of seedlings in toda! 's market u i l l rarel! be large enough alone to weigh heaicily In dec1sion4 of nvhether or not to plant a nonlocal seed source. Nonetheless, cost is a component in the overall equation and must be considered.

Financ~al benefits should be undentood ac clearly a\ possible to weigh risk appropriatelq before purchasing non- local seed sources. e\peciall! if there is a signitlcant risk involved. Landou ner\ in ho purchase geneticallj impro~ ed planting 5tock and do not haie their own ceed source trials should insist on useful information that can be converted to an estimate of financial benefit.

Risks and Silvicultural Practices to Mitigate Them

hlaladapration: What are the potential dot~nside\ in terms of cold or drought ditmage bawd on hi\torical seed \ource rntn entent trrali" Rotation: R I \ L o f lei\ due tct eniironmental extreme\ cannot be \eparated froin the harie\t cycle time. Loi\e\ near rotation age &re more \~gnlficant vr hen [he rotation 1s long.

* Sil\~culture: Sli\~cultural pritctlce\ can mitisate rome nrki\. For example. $011 preparttlon. planting deniity, and weed cctntroi can allel iate ntctirture \tre\s as\oci- ated tk~th maladaptation to drought. There is iome ek idence that proper nutrit~onal balance can reduce moi\ture stre\\ (Troth et al. 1986). Becauie tree vigor and couthern pine beetle tC)e~~d~-ocrorr~i.r 8-otztl-rlis Zirnm.) resii\tance are fa~orabl) correlated c Roberds et al. 2003). enhanced nutrient status ma) aid In overall tree health, whlch could improve insect resistance. Thus, fertilization can be a kaluabIe tool in ieed cource risk reduction. Historical climate perspecti\ e: Because forest crops are long-term in nature. it is neces5ar) to stud! historical patterns to full) understand risks ascociated with ex- treme climates. especiall! M hen mo\ing seed sources long distances. U'hat are the most extreme conditions that could occur during the rotation and h o ~ often do the) occur'? Fiftj >ears of cl~matic data are not too much. Test~ng information: Is there test information in the target planting area for the nonlocal seed source? HOM/ sound is i t and what do the results saj about risk? L4bilitj to absorb loss: Small priiate landouners obvi- ousl! cannot take the same risks that a large ~ndustrial landou ner can. The latter has se\eral thousands of hectares of stands of different ages and can absorb losses more easilq than some smaller prikrate landown- ers for u hom the loss of eLen a feu hundred trees could be a disaster. Other uncertainties: The old 5a>ing that "all of the information is net er in" holds in the area of seed source movement. Most testing s>ctems are not perfect. The prospects for and effects of global uanning are poorly understood. but come model\ predict that wme areas will become more drought! under \ome scenarios u htch could ha\ e ~mplicationcl for conirder~ng drought tolerance In cholce of proienance. .%ltzo. uhat other traitc ma) be of interert in the future that hake not \iet been characteri~ed In the iarrttus \eed wurces. For example. are rhere seed .tource d~fference$ In micro- tibrii angle or cellulo\e > ~ e l d per drj ton'

Guidelines and Industrial Examples of Seed Source ililovernent

Decades of seed source testing ha\ e produced results that can aid in deciding whether or not to plant nonlocal seed sources. Some simple guidelines are below. The recent USDA Forest Service Publication Sorithrt.1~ Pine Seed

98 SJAF 2912) 7005

Page 4: Sources of Loblollg Pine - Managing Benefits and Risks
Page 5: Sources of Loblollg Pine - Managing Benefits and Risks

rank changes. Although long-term field-test~ng is co\tly. it is the only method that insures \election of the be\[-adapted seed source.

West\ aco (no% Mead\+ e\t\ aco) ha\ been iucce\sful in its approach to the operational u\e of' lob[ol I! pine north of its natike range. The de\elopnlenr of cold-hard) loblollq pine wa\ preceded b) the de~el~tpnient of p~tch x lobloll! pine h>brid\. Thi\ uorh u a s begun a\ a cooperative agree- ment in 196-3 bettieen CSU.4 Fort.\t S e n ice Northea\t Forest Experiment Station ~ n d M'e\t\ act, t Little and T r e ~ t 1978). West\ aco e\tabl~\hed a number of pitch x, lobloll! pine hlbrid te\t\ In K e ~ i t u c k ~ . 'L'irginia, and U'e\t Virgtnia during the 1970\ and 1 Y X O i in conjunctlr>n uith lobloll\ pine trials to conipare the poi\ibtlit! of mo\ ing lobloll> northuard. Studle\ were establi\hed bq Lli'est\aco'\ Central Forest Research Center during the earl! 1970s tn Li\-ingston County. Kentucky. and MiNatr! Count). Tenne\see (Bar- bour 1972a. 197% and Barhour 1980). Prior to any \ignif- icant cold ex ent. lobIolly \ource\ from the 'Atlantic Coastal Plain and Louer Gulf Coagtal Plain outperforn~ed the more northern sources from the South Carolina piedmont and Virginia. H o u e ~ e r . the more eourherl! coastal sources cuf- fered extensiie foliage and cambial damage following sig- nificant cold e\ents. In man) caws. damage rewlted in mortalitq during that same e a r or a ueabenlng of the tree from which it ne\ er recovered. Foliage and terminal damage was more e\ ident in trees between ages one and eight. This type of damage is very \iiible in the late winter or early spring. U'ithin-source \anation among the more northern Virginia and P~edmont sources i 4 as significant. indicating that selections could be made for cold resimnce. Additional exploratorq tests indicated that cold-hard! selections could be made from all of the northern proxenances.

As is general1 the case. West\aco's operational plant- i n g ~ and research genetic trtalc uere establiched at the same time in the earl) 70's. Except for the most southerly or lower coastal plain material. almost an! native loblolly material could be grown effectil el!, in the Appomattox. Virginia area and the southuect Tennessee counties of Chester. Hardeman. Henderson. and Madison. Howecer. further extension of lobloll> into Tennes\ee and Kentucky u as needed to reduce the cost of traniportation to Vi'estk a- co's mill in ue\tern Kentucky. In addition. moiement of lobloll\ pine into the Parhersburg. M*e\t \iVirginla area to support the Luke. Marl land mill dlctated additional testing.

Intt~ally. cold reiiitance tiac, thought to be highl: corre- lated uith 40% grou th rate\. Man? of rhe initial cold-hard! selectioni exhtb~ted rather poor grou th performance M hen grown lthin the native range. Ho\\ e . ~ er. ~nclusion of higher census nuniberc ~ i t h ~ n the test popul:~tion hac led to a rethinking of this correlation. t'nf-c~rtunatel? . number\ of trees within ;1, select population are still not at the point where a large-scale breeding population can be fcjrmed. but an elite breeding population would certa~nl! fit this rather limited scale program. Specific mating de\ignc could be used to take advantage of the \light \artation that exists outside of the native range.

Continued \election efforts hace developed cold-hardy loblolly pine that suririie and grow in cold environrt~ents prerpiou\l> thought to be too inhospitable for loblollq pine. Deflned deplo) ment zone\ are based ctn the amount of cold re\istance needed fix \ur\ i t i a1 and rapid grotb th. Today. cold-hard! wurcei can he grou n in clme prctximit4 to the n~ills, thus lotiering tran\portation co\r. In add~tion. Ioblolly pine proved to be fii\ter SroLilng thktn either \hortleaf pine (PltrltLs c ~ ( - / ~ \ i t ~ i ~ ~ ~ ~ or p~tch ( P l i ~ ~ i s r \ y i d u ~ \ lobloll! hybrid. thus ailow ing for \hortsr rc.ttations. The prirnar! riik remain\ the uncenaint! of the en\ ironmcnt. Unprecedented mortal- it! from southcrn plne beetle occurred in 200 1 in northern plantation\ of lobloll! pine along the Cumberland Plateau in Tennei\ee. To date. u e hate not seen clouthern pine beetles in the more northern \tandi in Tenne\see and Kentuchy. but that ma) be becauw of re\tricted 111orement of the beetle. The size of the depio! ment population is also iorneu hat of a riih.. but thi\ \eem\ comparable to the lim~ted deploj ment of genotype' folloued in the natiie range of lob loll!^ pine.

East-\.Yest 3lok ernents In this section u e refer to mor ements east to uest or vice

cersa uithin the same plant hardiness zone. This type of mokement is more common and usuall) entails little cli- matic risk except in the case of moxement of seed courcec from high rainfall areas ro areas that experience frequent drought along the uestern fringe of the loblolly natural range. For example. planting South Georgia \eed cource in southeast Texas or North Carolina material in southuest Arkansas and Oklahoma should be viewed as a sisnificant risk. In one study in southeact Texas. no East Coast seed sources produced as much lolume per hectare as the local source in four 20->ear-old. unthinned plantings. two of which uere outcide the loblolly pine natural range (Long 1980). On the other hand. movement of matertal from South Georgia to \outheastern Louisiana or from central Louisiana to East Texas ma) not entail significant risk.

Some less risk! but beneficial moies in this category have alread! been mentioned - planting of Li\ ingston Par- ish. Louisiana material in high rust hazard areas of Florida. and southern Georgia. >lis\issippi. and Alabama (Wells 19851 and southhest Texaq material in the sandhills of Georgia and 'Alabama for drought re\ictance tJett and Guines\ 1992). Another common mokement i \ to plant more ea\ter-1) coastal sources moderate di\tances \t est. tx hich recult\ i n \~gnlficant gains in growth rate at little r ~ s k - for example. Georgia coastal source5 in coaital Miisis- \ippi and Alabama and Carolinas coastal material in central Georgta and .Alabama I Schmtdtling 2001. Sierra-Lucero et 21. 2ou2 1.

In a recent stud? (Sierra-Lucero er 31. 20021. Gulfcoa~ta l plain wurzes (GCP. i.r.. l o ~ i e r &li\ii\sippi and Alabama) gave une\pectedl> poor performance compared u ~ t h north Florida and Ioiier .4tlantlc coastal plain \ources (Agricul- tural Con\eri at ion Program ( ACP )). Until recent11 , most tree breeden hatie con\idered the GCP and ACP as com- parable provenance\ tor growth (e.g.. Lantz and Kraus 1987). In general. provenance grourh performance for

100 SJAF 29(7) 2005

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influenced by nur\ery culture. The local and nonlocal sources do not differ greatly in fir\t-year- \ur\ ival. Based on the 1980 drought result\. the dec~\ron \+a\ made in 1983 to extend the planting of XC materral onto deeper mountain \oils. wine of L% hrch are ju\t out\ide of the northern and western lintlts of the natural range o f lobloll> pine. This decision led to planting hOCr of' all contpanj landr in Ar- kantas and Oklahoma to the NC seed c,nurce.

Since 19:-3. approximatelj 1% N'eyerhaeurer trial\ comparing ea\tern and AR/OK wed source\ have been installed. including progenj te\t\ of otier 300 parent\. The grotvth rate of the NC \ource. almost ~ i t h o u t exception. significantlj exceeds that of the 1oi;iI \eed source. eten on the shalloi~.e\t and mo\t rock! \oils. During that period. there halie been six cignificant drought >ears that have provided the opportunitj to clarifq the r~ck of planting NC material. Sur\e!s of drought-related mortality are con- ducted in these progenj tests and in the thousandc of com- mercial plantations of nonlocal families that have accumu- lated ocer the !ears. Surveys indicate that drought mortality is higher in Oklahoma than southuest Arkansas in most drought >ears and is not related to soil uater holding ca- pacity. i.e.. there were no detectable trendc of seed source mortalit> differences being greater on shallower soils.

Each year for 10 years and nok during drought years only, 26 commercial plantations (20-80 hectares each) per seed source per each of two regions (fouthwest Arkansas and southeast Oklahoma) are surti e> ed b) helicopter, and a complete count of dead trees is made. The number of dead trees is dii8ided by the number of trees during the last inventor] to get percent mortalit). These surveys reveal that (1) even in a se\ ere drought year. such as 1998. mortality in the NC seed source is low and ( 2 ) men the local seed source is subject to drought mortalit> (Figure 2 ) . although at a lower rate.

There habe been worse drought periods than have been experienced in the past 20 )ears. Attempts ha\ e been made to model what might happen during such a period assuming a "worct case" approach (worse than what has been ob- serced) in terms of mortalit) as a function of soil moisture deficit for the NC seed source cersuc the local \eed source. Growth and ~ i e l d modeling of uorst case ccenarios hace indicated that ir is hi_chl\ unlikel! that the Ggnificant growth advantage of the NC seed source o.ier the local source uould be loct. eien in a uorst case \ltuatic>n.

Some additional action$ that are being taken b! Wsyer- haeuser to understand or reduce the ri\k are:

Genetic

Families are tested on a \ arietj of \ires and \traightness and growth are keq \election \;triables. Onlq nonlocal orchard farniliec, that ha\e been progeny tested in the area are planted commerciall_c, and they are planted in Familj blocks for ob\er\ation purposes. Families that do not perform uell during drought sur- veys are no longer deployed. but only a feu families have been eliminated on this bacir.

Figure 2. Mortality for Arkansas and North Carolina seed sources from aerial surveys in operational settings in southeast Oklahoma (a) and southwest Arkansas (b). Settings were 20 to 80 hectares each and 26 settings per seed source per year.

Individual trees u,ithin the best families have been selected on very tough sites and form the foundation of a "land race." These select trees ha\e been bred and their progeny are under test. Future orchards of nonlocal material (land-race de- rived) will be located in Arkansas so as to be contam- inated bq local rather than nonadapted. nonlocal pollen in Sorth Carolina. Interprovenance hq bridc are being testing for potential complementarity of grou th rate and drought resistance.

En.itironmentaI and Sill iculture

Mortality curveys during droughts continue. Climatic data are updated and each year so11 moisture deficits are calculated to undermnd the lerelc of ctress that produce mortality. Site preparation b~ mean\ of rlpprng a trench or tillage on the contour ic aimed at ensurln: adequ~ite moi\ture during the earl) establirhment year< bq proi~ding the seedling good sol1 structure for root penetration and capture of moiwrs I n the rlp. Fertilization throughout the rotation help5 keep trees nutr~tionally balanced and thu\ Iesr prone to ctress and/or beetle attacL. Stand densitie\ are lob b) comparl\on to manj \tands throughout the \outhea\tern Cn~ted States. whlch hope- fully reduces ctress as well. Herbaceou\ w e d s and hardwood control can reduce moisture \tress, especiallq in early years.

102 SJAF 29(1) 7005

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&litigating Seed Source Movement through Tree Improvement

There are rorne prom~\ing tree impro\ement ctrategie\ that can reduce the n\l\ of planting nonlrtcal reed source\: Tertinl famlliec for adaptation: Thorough te%ting for adap- tatlor1 under climatic extreme\ can help ~dentlf: potentiall! poor f a in~ l~e r that should be eliminated fr tm planting pro- gram\ and frrtrn wed orchard\. Land race de\elopnient: Breeding and te\tlng of the nonlo- cal reed wurce for adaptation tct the exotic en\ ironment can be a irer) effectlie vta: to increart adaptability and reduce rl\l\. The rerultant improked material i \ knovtn ar a "land race" tZobe1 and Talben 19841. Example5 of this were gt\en in the t u o indu\trial cake ctudiei aboie. Specie\ and iilterproienance hq bridi: Interllpecific hybrid- ization can be a berq effecti\e &a! to loi+er risk, of loss due to climat~c extremes. Interproienance hq brids have been less expjored. but a recent greenhouse stud) indicated that it might be pocsible to increase adaptability bithout losing growth rate uhen hlbridizing fast growins East coast ce- lections ~ i t h slo\ver grow ing but more drought-tolerant families. Figure 3 shou s growth and suritii a1 re\ults from a greenhouce stress tect. In the stress test. seedling5 were grown for approximatelj ceven months in cand boxes. and then water uas graduallq u ithheld until trees began to die (Burns et al. 1982). U'hen approuimatelj half of the seed- lings \+ ere dead. the test was reu atered and actual survival counts were made once seedling had a chance to recuperate.

Arkansas Hybrid North CaroIina

Arkansas Hybrid North CnmUnn

Figure 3. Survival (a) and growth (b) of Arkansas, North Caro- lina, and AR x NC hybrid seedlings of loblolly in a greenhouse moisture stress test. Means with the same letter do not differ significantly at P < 0.05.

Another study shot+ed that crossing fusiform rust-resis tant pro\ enance\ u rth fa\t-grow ing protenance\ can pro i ide coniplementar) combination\ of the t ~ o trait?. ant there was some e\idence of hererow\ 3% \$ell (Schm~drliri~ and Nelrcrn 19991. ?Id\ ante\ in controi-111a\c-pollinatior nou make the\e two option% mor-c: fea\ihle a\ commercia application\. Clon~ng: .4dapration differenc2\ an~ong f;ifnilitf\ uithin , nonlocal \eed wurce \ug_re\r diftPrence\ among clone* ulthin the bert hrnrlie\. an aienue that u ~ l l be elplotted ;tt

clone% become aidel! planted throughout the range o Ivhloll pine. Hoiseier. M ~ t h no genetic i anation M ithin , clone for bufkring againit cliinat~c extreme%. clone\ mu\ be ier! thoroughl: te\ted, more \o in the exotic eniiron nxnt than In itr natite area. Te\ting should be thorough botl In term\ of the number of s~tes and In term\ of a longer fielc a\ce\rtnent.

Conclusions

In man! repons of the South. use of nonlocal sources ot IoblolIy pine has become the rtandard rather than the ex- ception. For example. in the North Carolina State Unittersitb - Industrl, Cooperati\ e Tree Improiement Program. most companiec and state forectr) organizations in the Gulf coastal plain region plant Atlantic coastal plain sources of loblolly pine rather than the local source. The uell-docu- mented growth advantage of the ,4CP provenance (e.g.. Sierra-Lucero et al. 2002) and the lou risk from pests and climatic factors make the use of the nonlocal pro\ enance an economically attractiie decis~on in this region. Increased growth, reduction in fusiform rust disease. improied cold or drought tolerance, and better stem form and wood quality are all realized benefits from using well-tested. nonlocal provenances. Althoush the benefits of seed source move- ment are relatively easy to s h o ~ . Lte h a ~ e emphasized that there are substantial risks asrociated iirith the ure of some exotic sources. Long-term, uell-designed field trials are critical to understand risks.

Use of nonlocal sources i i frequent11 used in conjunction vt ith c i l i icultural systems declgned to reduce enitironmental ctrescee. uhich niaj include \pacing. fertilizatlon. intense site preparatior?. and increaied monitoring. L a n d o ~ ners not uiIIing or capable of implementing the\e measures will expertence increa\ed nik.

Literature Cited A "h~'.. S % \\I: 11 T 434kli 199- Spr:nf cold h ~ r d ~ n r \ \ ilnder \trtIng

fenetli ~ontroi In Oregon popuid:wn\ of Pic~iriior~iiyo t t r r t : : l r r t i I ur ~t tc , rr : ic . r i / Ccln J For Re\ 27 " 7 - 1-78

A L - R ~ B \ B %ti. 21 4 . A'--.[) C C 12 IL- i \ f i 7002 Populclt~on d\nclmlc\ of P I I I I I ~ i ~ i c d i i L bdwd on nu~!ear rni~ro\clteiltte% For Ecoi itlclnclge 163 263-27 1

B,R~OLK. H F l Y 7 7 d 1972 progi'r! irttm i1onc.k i n b~rg:n~ci clnd S C piedn~c\ni lohlitllj orchdrJ\ 'A e ~ ~ ~ ~ d i o Centrdi Fore\[ Re\rdrih Center E.ztclhli\hmr.nr Rept-irt Fi p

B 4 K R O : R. H F 1 9 7 3 1972 supplementdl piti h x IobIctlI! progen! tc\t \ite\tiaco Centrcll Fore\[ Re\eclrch Center E\tab~ish~neni Report 6 p

B A R B O L R . H F 19%) Lobiuli! wed wjurce for \ \ r \ t Kentuck) P 15-23 ln Proc 1980 Southern Yurier! Conference Lake Barhle!. Krntuch). Sept 2-3

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B L R ~ S . R., C C. LAMRETH. L.C. BCRRIF, AND C. TXUER. 1982. SurvlvabiI~ty ot container~zed ArkansasiOkiahom~. and North Caroltna seed sources under moisture stress. Weyerhaeuser Technical Report 050-1 204'8211. 9 P

BLRRIS, L.C . S D DLKE, A \ D C C LAUBETH 1982. Winter and drought damage suney of North Carolina. Louiwna, Texas. Pvlississipp~. Arkansas and Oht~hom,~ seed sources on XrhansaslOklahoma sites Ue)erhaeuser Technical Report 050-321lk82i 1 1-

FLORELCE. L X . I ~ D R R H I C ~ F I980 Further evidence tor lntrogression ot Prtlirl trledn btrrlr P rrlrindr~r Elecrrophoret~c cariab~lity and vdrlatlOn In resrstance to Crcirtclrtrrtm jii\if;jme S~l%ae Genetica 3-9 11-43

GIF~IORE. X R 1980 Extending the range of lobloll> plne In the XIisrissippi Riker Vallej Factor> relatins to grortth and iongekity. P 8- l4 it7 Proc I980 South Nurser) Conf Tech. Pub SATPI 7 icomp 1. Lantz. C t% fed I. Atlanta. GX USDl For Sen . State and Private forest^. Southeastern Area

G I L L ~ R E . '4 R . ALD D T F L ~ K 1976 Shctrtieaf ~ n d loblolly pine seed ongin trial\ In southern IIllnols 17-year rewits P 115-124 In Proc 10th Central States Forest Tree Improvement Conference September 22-23. 19'6. West Ldtayette. 13

GRIGSBY. H C 1973 South Carolina best of 36 loblolly plne reed sources tor southern Arkansas USDX For Sen Southern Forest Experiment Station. Research Paper SO-89 10 p

HODGE. G R . kc^ R J WEIR 1993 Freez~ny stress tolerance of hardy and tender families of loblolly pine Can. J For. Rei. 23 1592-1899.

J I\ 4% ICKR -\MA, K J S.. S E. ~ICKEAND. 4 V D J.B JETT. 199th. Phenological banation in helght Jnd dlameter grotcth in provenances and families of lobiolly pine Yew For I6 11-25

J-\'I IL'IICKRACt;\. K.J S . S E >ICKEA\ID. J B JETT. AND E .A. WHEELER. 1998b Date of earlyu-cmd-lateu-ood transition in pro\.enances and tamilies of loblollg. pine. and its relationship to growth phenologq and ju~enile wood specific gra\it! Can J For Res 27.1245-1353.

JETT. J B . W M. G L I ~ E S S I992 Growth and wood properties in a Carolina Sandhills pine seed source stud! South J Appl. For 16 164-169

JOLEF. E P. A\D 0 0 WELLS 1969 Ice damage in a Georg13 planting of lob loll^ ptne trom d~tterent seed sources USDA For Sen Research \ote SE-116 I p

KULB. T E., A L D K C STEILER 1985 Cold tolerance variation in lobloll> pine needle> from different branch type\. families. and environments P 358-367 in Proc ISth South For Tree Imp Conf. 1985. Long Beach. XIS

L-VBETH. C C . 4 L D L C B L R R ~ ~ l983 Ark ,'OA Seed source best tn 1981 field and nurser: survi~al te\ts Wejerhaeuwr Technical Note 050-1 204/8-32 5 p

L,\~HETH. C C . P Cf DO[ CHERT) L'L T GLAD>TOLE. R B ~ICCLLLOLGH. +LD 0 0 %ELLS 1981 Large-\cae plantin? ot hc~rth Carolina lob loll^ pine in lrk;in\as ~ n d Oklahorn~ 4 case ot gain cersus r~sh J For 321 13) 73h-7'JI

L-\TZ. C 'rZ . I ~ D J F KRILS 13Y7 A gu~de to southern plne seed .c,urces LSD4 For Srrt South For E'ip St Trchn Rep SE-43 7-i p

L E C, +\D I F TRFU 19-8 Poi\lbilitie\ ot pitch u lobloll> pine h\br:d\ rn rhe Interior South P :5 1-1SX In P ~ O L ot the Sqmpoblum tor the Cl.in.~gernent ot Pines n r the Interior South Knou~llle. Tenneisee \o\ernher'-4 pp 151-ISY

i t E lf l Y 4 0 Tcua4 and Lt)ui\i,in,t lohloll~ pine \tud> ~onfirrns , i~ l~or t~ ln ie r ) f I O C J I ited \ o~ r i c \ - South J 4ppl For - F r y ) i 37-1 -32

\ I \ ( n, ii LA 5 l'iiO Ettei: 1 1 t ~ C J iouri: ti,n he~ght grot~th ot pine -.cedlin;\ j Ft>r 4Y -&%)-A; t

* k d J 1 1 ) ~ 1 1 PI .L I I [ I~~ l o ~ l ( ~ l \ ~ . 7ir-< ~~LII~IC,. i t > nsiturci1 r age J For -4Y :-\-?-<I

RINK, C . -\ND 0.0. WELLS. 1988. Productivity cornpansons of 37-year-old loblolIq-shortleaf pine seed sources i n southern Illinors. North. J. Appl. For. 5 155-158.

ROBERDT. J H . B L. STROLI. F P H i IC , D P Gb AZE. S E. MCKEA~D, 2ND LH L o n 9003 Estimate\ of genetic parameten for oleoresin and orotvth t r ~ ~ t s In jukentIe lobioil! pine Can J Fur Res. 33(12): 2459 -2476

S C H ~ I I D T L I ~ G . R.C f994 Cse of procenance w t s to predict response to c![matli change LobiolI) ptne and ?jori%+ y ~ ~ c e Tree Phpstol. 14 805-8 17'

SCH~IIDTLILG. R C 1001 Southern plne seed sources USD.4 For. Serv Gen. Tech Rep SRS-4-4 15 p

Scw\rr~n:\c. R C . ~ L D C D N E L ~ C ~ 1996 Interpro\en~nce crosses in iobiolly plne usin? ~eiec;ed parents. For Gen 3 l i 5 5 6 6

SCH'LIIDTLI\G. R C . E C XRROLL. .?cGD T LIF ~ R C E 1999 Allozyme diversity of \elected and natur~l loblvily ptne populations. Siikae Genettca 48 35-43

SIERRA-LLCERO, \/, S E ?VICKE.ICD. D '4 HLBER. D L ROCK~~OOD, AND

T L WHITE. 20112 Performance ditkrences ~ n d penetlc parameters for four coastal prolenances ot lobiolly pine in the .outheastern United States. For Scl 4Y '32-742

SQLILLACE. X.E. ILD 0 0 WELLS. I981 Geo,or~ph~c vanation in monoterpenes In cort1ca1 oleoresin of lob loll^ pine Silvae Genetlea. 30.127-135

TALER. C.G., - \>D J 'A Loo-Dikkics 1990 Seed source vanatlon in speclfic gravity of iobIoIly pine grottn in a common envtronment In

Arkansas For. Scl 36(4) 1 133-1 145 TROTH. J L.. R.G C ~ L ~ P B E L L . -\>D H L. XLLE\~ L986 Chapter 1 Nutrients:

Cse of forest fertilization 2nd nutnent efficient genot>pe% to manage nutrient mess in son~fer stands P 91-12-i rn Proc of the Society of Amencan Foresters Natlonal Convention. 1985. Ph)siology Workln,o Group Stress phq.siology and forest productlkit? Hennessey, T.C., et al (eds) >lart~nus \ijhotf Publishers. The Hague. The Netherlands.

LSD-\. 1990 CSD;?t Plant hardiness zone map CSDA-Agricultural Research Sen ice hfisc. Pub 14'5

W + ~ E L E ' I . P C 1961 Rebuits of the southutde pine seed source study through 1960-61 P 10-21 In Proc 6th Southern Forest Tree Improbement Conference. June '-8. 1961. Gainesli~lle. FL

tk A ~ E L E ' ~ . P C . + \ D T E BERCIU 1965 Lobloil\ pne proienance test at 3ge 35 J For 63 168-1 7'1

W ~ L L S . O 0 1983 South~cide ptne zeed source ziudc-lob!olly pine dt 25 bears South J 4ppi For 7 ( 2) 62-7 1

%ELLS. 0 0 1985 LFS ot Li~ing~ton Par15h. Loui~ilind loblolly plne by forest products indu4rnes in the Southeait South J Appl. For. 9t3) 180-185

U E L L S 0 0 . ~ L D C C L iLreETw 1983 lob loll^ pine procenance test in \outhem -\rkania\ 35th bear re\ulti. South J App1 For 7( 1).71-75

W E L ~ ~ . 0 0 . + L O G R , \ k 1934 P!.int~ng lobioll~ pine north ~ n d west of its natural r;lnge P 26i-2hi ( 1 1 Proi Zrd Biennial Southern Sii\ iculturr: ContsrenLr 1984. 4t!anta. Gr\

% FLLS. 0 0 . I \ D P C V. , ~ E I E?I 1966 Ceograph:~ t dr1Lttion in survival. ~r tmth . and tuiitorv ru\r lnteccion ot planted :oololl) plne For Sct \lonograph I I 40 p

Lk ~\ . t \ . t H L ~ G F L . E C 1935 F~\e->r; ir re\ult\ ot I(~o1011> pine geographic ierd bourie :e\t\ P 16-25 i r : Proc 3rd 5r~thern Forest Tree Irnprovzment Cl)nt:rence. Jan 5-6 1955. \e.r Or!e~in\ L 4

Z %rtc ;i T G i 961 Ten-)e,lr re\ul:\ ) t a ~oopcrdt,. e i o n i o i l pine >eed \trurie te\t P 1S-q'i 111 Proc hii? 5t)uthern Fi'i~*t Tie:: Imprt7cement ConrerenLc June '- \ liih i Cia -e\\ lie FL

Z o b i B J : \ i , J T 7' ^ ' T /cjFf 4i-nI1cd :c re.[ -2t' rnprt)iernent John L i let. \ew P t r i \ I 505 13

70~3 B J %..i,! P . Ei ~ r i i t f O j c l !Lo< t ,r ,tion .~r:d i t \ irlu\es 5rrlnger \ crlds 'I orA \'I :b; ;I

Page 10: Sources of Loblollg Pine - Managing Benefits and Risks

Volume 29, Number 2 May 2005

Special Issue: Silviculture and Genetic Impacts on Productivity of Southern Pine Forests

Page 11: Sources of Loblollg Pine - Managing Benefits and Risks

PWDEW John A. Helms VICE-PRESIDEM" Marvin D. Brown

E,YECU~IVE V I C E - ~ ~ ~ D E ~ T iWLI CEO &lichaeI T. Goergen, Jr.

A D V ~ S I N G SALES William V . Brurnby

SOUTWERN JOLRVAL OF APPLIED FORESTRY E D ~ O R Ian Munn

Box 9581, Depattrnent of Forestry Mississippi State U., MS 39762-9681 (662) 325-4546; Fax: (662) 325-8726

[email protected]

GRVAL P R O D U ~ O N EDITOR Linda Edelman Cadmus ProfessionaI Communications

940 Elkridge Landing Road Linthicum, P/ID 2 1090 [email protected]

Management and Economics Daowei Zhang

Douglas R. Carter

Harvesting, Engineering, and L"tiifi3twn Robert B. Rummer

WiMli$e Management Darren A. &tiller

Forest Measurements Bernard Parresol

Forest Growth and Yield Quang V . Cao Mike R. Strub

Protection Henry Arnerson

J. Wayne Brewer

Soils. Hvdrology, and Watershed iManngernent Stephen H. Schoenholtz

Silvicrilture Thomas J. Dean

Emile S. Gardiner

Genetics and Tree Improvement Steven E. MlcKeand

~d address changes to Address Change De- trnent, Society of Arnencan Foresters, 5 4 0 Isbenor Lane, Bethesda, IUD 20814-2998. td .Clanuserip& (5 cop~es) and related corre- ndence to Ian >funn, Box 968 1. Department ot e\try. Missnsipp~ State U., MS 39762-9681 td other inquiries to SJAF, Society of Amer- 7 Foresterr, 5300 Grosvenor Lane. Bethesda, ) 20813-2 198. .mission to Reprint.-Individuals. and non- fit l~branes acting for them, may make fair of the mstenal in th~s journal; for example,

lying an article or portion of XI tnanicle for use teach~ng or research. Indivtduals may diso kte from the publlration if the customary dc- raledgernent of the source accompanies the ltatton. To reque\t permisston to make muI- e or sy\temattc reproductions or republish ern1 cancluding figures and art~cle excerpts) tdct the Copynght CIearance Center. Cus- ier Service Deputrnent, 222 Rolewood Dr. ivers. MA 0 1923 (978) 750-8400. Fax. (978) -470; www copynght.com. A fee for corn- cia1 use will be determined Jepend~ng on use.

SOUTHERN Volume 29, Number 2 May 2005

Summary of IEG-40 Meeting: Silviculture and Genetic Impacts on Productivity of Southern Pine Forests S. E. hlcKennd n n ~ i H. L. Allen

What is Ahead for Intensive Pine Plantation Silviculture in the South? H.L. Allen. T. R. For, and R.G. Campbell

Strategies and Case Studies for Incorporating Ecophpsiology into Southern Pine Tree Improvement Pro, ~ r a m s T.A. Martin, P.M. D o u g h e r ~ , k1.A. Topn, nrzd S.E. iClcKe~rnd

Risk Assessment with Current Deployment Strategies for Fusiform Rust-Resistant Loblolly and Slash Pines F. Bririytvnter, T. Kribisink, T. Byrnm, n n ~ i S. .tlcKrnnd

The Future of Tree Improvement in the Southeastern United States: Alternative Visions for the Next Decade T. D. B)rarn, T.J. itlzrllin, T. L. PVhite, and J.P. ran Brtijreizen

Planting Nonlocal Seed Sources of Loblolly Pine - Managing Benefits and Risks C. Lambetiz, 5. 1tfcKeni7cl. R. Ro~isseuii, nrzd I-i. Schrnidrliizg

A Review of the Biological, Social, and Regulatory Constraints ro Intensive Plantation Culture R. Ro~rsse~rti, D. K~rc:nztrrek, nir~l J. iClcrrriir

A -\Publication of the Society of American Foresters

The . S o ~ t f i ~ c x / - f i J o L ~ I - ~ I ~ L ~ o/ 4/>/>/ic~(/ t - < ) ~ - c ) i f r i

is p~~bl~\hect quarterl>-Fehruar! . >Id>. Xu;u\t, No~ernber-bj the S o ~ t e t j of Ainzr~c;in Forcsters. 5400 Grr ) \ L enor L ~ n e , Bethesda. ,LID 703 14-2 193 Cop;,r~ght G 7005 by the S o c ~ e t ; ~ ot Xmcrican Foreht- rr\ l S S N O 148-I4 I9 Sub\cr~ption ratcs are $52 tor ind~vlduul S A F members in the U.S and Canada ($72 ~n fore~gn countries), $26 tor \tudcnt EAF member\

in the L 5 and Candrt. and 57'5 tor 1n41- ~ i d i ~ a l non-EAF member\ t 595 ~n fore~gn ~ o ~ ~ n t r i e \ i. and 's 1 TO tor in\t~tutions ( 5 190 in toreign tnstlrutton\) All iubscr~pt~on\ start b ~ t h the tlr\t i\\ue ot the calendar year ,LIis\~ng I\sue\ w~l l be replaced tree if the cla~rn is made filthin I! months from U S. or Canadian ,tddrci\e\ or 6 month\ from other addresse\. Single issues Jre 520 for 1 n d 1 ~ lduals dnd $48 h r Inst1 tutioni.

Page 12: Sources of Loblollg Pine - Managing Benefits and Risks

Southern Jozrrnal of Applied Forestry

Editorial Policy

Soilthem fortmczl of Applied Forest? is a refereed profes- 3ublication sening practicing foresters and forest?-allied ionals. It incites contributions from a11 workers uho bq nce or research hake developed successful new practices or Jes. Subjects may be any of the goods, serxrce.;. and prob- ' forests and forestry in the South. first test of any manuscript is its usefulness to the field

- or land manager. hfanuscripts will be reciecked by the associate editorcs). and. normally, by experts in the subject being reported. Notification of the acceptabiltty of a manu- ,r review will be immediate. The formal retires process will inarily exceed 90 da>-s. Final acceptance or rejection of all ripts is the prerogati~e of the Editor. ddition to peer-renewed articles, the Salrrtrer~r Jorinzal of

The Society of American Foresters holds copyright to the Sotithem Jc~trnzal of rlppfied Forestn, and auhurs will be asked to assign their rights behre their contributions are published. A form will be provided for this purpose. Authors whose work is not subject to copyright. e.g.. federal government employees, should so state uhen they submit their manuscripts.

To request permission to reproduce material for course packets, conferences. and websites, and for republication or systematic or multiple reproduction of copyrighted material (including figures and article excerpts), see http://wuw.sdnetr.orgperiodicalslrepnntinfo. cfm.

Direct all correspondence pertaining to manuscript submission inctuding name, address. phone number. Fax number, and e-mail address of corresponding author. to

i Foresrn publishes edited but nonrefereed contributions I'echnical Notes, which describes useful ideas and findings. mally. only manuscripts not previously published will be Ian kfunn

:red. Pretiousl) published work includes that published in Editor. Solithem Journal ofl-lpplied Forestry

s and proceedings generally available in libraries. Box 968 1 Department of Forestry ~uscripts recised and returned to the Sortrizertz Joirt-tznf of bfississippi State C.. >IS 39762-968 1 Forest/? more than 6 months after review utll be treated as (662) 325-4546: Fax: (662 1 325-5726

snuscripts. imunn @ cfr.msstats.sdu

Guidelines for

and Form luscripts must be submitted in final form. The author is ,1b1e for accuracy of data. names. quotations. citations, and :a1 analysec. Engilsh or metrrc units may be used. as long as s consistent u i th~n each art~cle. Strict econom) of uords. formulas, and figures should be obsened and spec~alized avoided. 5lanuscnprs should be limited to S journal pages. :O t>-ped page, tncluding table5 and illustrations. nenciature and Termino1ogy.--Common names are used st plants and animals. Screnttfic name, are mcluded in less\ following the first use of' the common name The si of L'ruretf Srilrrr Tree5 i,t;7rr~ e LZHJ ,Lilrlll~i/i:edi bq E.L r i Xgric Handb 531 LSD.&. 1979) and the appendixe, ot Cn\ rr Tiyes of rile L nrrrtl Siclrr \ ( I I ~ L ~ C L I I I I I L ~ I ~ S 4 F 1980 ) ful reference\ tor plant name\. Technical u ~ s s in fore\tr; red fields to l l~i t k The Drc r10111117 of' F01.v t i t ' I S iF 1 qCj8 I

kescript.-'LIanuscrrpts must be double-.paced :hroughour i t r a table\. :llustr;ttion c,iptron\ and Iiieratiirrj cltation\l npie mdrg1n.l. drd printed on one \id? (-it irhitr p~iper \ \hould \ubmlt :he urig~nal ~ > p > dnd four ~cl i i : r : t j r . , i i copte, exr. in~luding i,r!;indi trgure\ and rabiz. t s ~ r h 'rlr;' i~ristndi ~d repn)Ju~:lon\ .t ~ t h rhrj duplicate i ( > v l t " k -,ho~iId he ed onl: irk [he final acceprcd ~n i f re; :\ed ,op> ot the 'rtpf Guideiii;e\ :or eiecironic preparat11)n MC -,ubmii\lon . tabIes. graphic\. and equat~on\ dril a i d i i ~ r ~ e at http I

afnet.orgipen~fdii~i~/n~)rtht'rn/regt~~n~~ici~n~~~ide L t rn and e Editor Thr titie. author\' names. atfiliation\. ~ c d cc~rnpietc e\. ,dung uith ;t~inc)i~ledgcmeilt\, and iootnort"\ ionceming , mind i.\penrnenr .ttatiitn pubitcarion nunllxr. .tkoulcf he on Page 2 \hould h a ~ e the i~;t~pietc: title. the abstr~ct. LC\ ckords. \ran c,t the bod? ot the manu4cnpt. Each page. ~nclud~ng

e ;itation\, table,. and figure caption\. 5hr)uld be numbered it~ieI> and an ~bbrec iated cenion of the title in the upper irner. Authors' name\ jhuiild appear onl) on page 1 ,raturc Cited.-Li,t all reterence., ~ lphabet ical l~ at the end ),iper kind cite them parenthetically tn the text b) the author-

Contributors

date system. e.g. (Beers 1978). L m t number of citations to three per set of parentheses. Theses and unpublished papers may be included sparingly. Smples of literature cited s ~ i e are ava~lable at http:,'l u cb cb . s a f n e t . o r ~ p e n o d i c A s / n o r i h e n v ' r e g I o and from the Editor.

Footnotes.-Authors are strongly encouraged to incorporate information in footnotes into the text. r\n> ~nfomation included for footnotes ~ b l l l be presented at the end ;tf the text as endnotes.

Illustrations and Tables Captions and tltle5 5hould contaln enough infomation so the

~llu,tration or table kill 5tmd done Edch illustration and table must be cited in the text. Tables khouid not duplicate data presented in figures md ice versa. Details dbout prepmnz table\ and figures are at luiable at http'iiebuis d n e : orgipenodiiti,, t'orrcienceitableguide. cfm and trom [he Editor

Illustrations.-411 photogr~ph.. line Jrdu ings. map\. and graph\ dre dtsign,:ed as t i p r e \ The> ih(i~iiil 5e bubmitted in the origlnal torm (no !;irser than h 5 i' 1 I in ;. ~nnoun ted . consecutieel~ nun~hrjrild. ;tnd identitied ~ t i r h sott p e ~ ~ : : on the reierse s ~ d e F!purc.\ ind) br ~ii~iudeci t,n di\i\. b ~ r or;_i:nal hard copie\ are ieql~irtd J\ ,ie!I Cdption\ \ h o i ~ i d be on .cprdts page at the end t i t the m ; i i ~ ~ ~ ~ r l p t . no t on the i'iriginiil-. o t ihtl tigures Labeling iilu\; be \Lirtlc~rntl> large to perm" redu~rion I b? '1s much as 50ci;i 2nd \houid be kept in red\ondble prop(,nioii to the figures Photograph, ,hould he b t , ~ ~ h ~ n d hire. 2!04\>. ir ith gocxi contrast, no \rndllc.r than 4 < 5 in nor larger thdn X K I0 in. Standard colurnn sidth is 7 3 in . d page consi\ts o f 2 columns

Illustr~tictns and ti?ure\ -,hould be prtiduced in a manner that ens~ire\ their re,idabillt; 2nd u\etulne\, uhen printed in black uhite If ioior reproduction i \ required. ~utl?or\ ecill be charged a publication ire b a w l on ac:ual cost

Tables.-T,lbles should be consihtent in torn1 i\ee recent i\we\ tor st)le) Each table should be tqped J iepctrate page. con\rcu- ttcely numbered and rit1t.d. at the end of the text ~xce \ \ i t e ly large tables or unnece\sar~l) deral1t.d. , t ~ t i \ t l i \ ,hould be &i\.olded


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