T E C H N I C A L R E P O R T 0 9 6

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1 T E C H N I C A L R E P O R T 9 6 Exmining the Potentil of Western Lrch, Sierin Lrch, nd Ponderos Pine s Regenertion Species in Crioo Region Ecosystems: 22-yer Results from EP

2 Exmining the Potentil of Western Lrch, Sierin Lrch, nd Ponderos Pine s Regenertion Species in Crioo Region Ecosystems: 22-yer Results from EP94.2 T.A. Newsome, J.L. Heinemn, nd A.F.L. Nemec

3 The use of trde, firm, or corportion nmes in this puliction is for the informtion nd convenience of the reder. Such use does not constitute n officil endorsement or pprovl y the Government of British Columi of ny product or service to the exclusion of ny others tht my lso e suitle. Contents of this report re presented for discussion purposes only. Funding ssistnce does not imply endorsement of ny sttements or informtion contined herein y the Government of British Columi. Uniform Resource Loctors (urls), ddresses, nd contct informtion contined in this document re current t the time of printing unless otherwise noted. ISBN Print version ISBN Digitl version Cittion Newsome, T.A., J.L. Heinemn, nd A.F.L. Nemec Exmining the potentil of western lrch, Sierin lrch, nd ponderos pine s regenertion species in Crioo Region ecosystems: 22-yer results from EP94.2. Prov. B.C., Victori, B.C. Tech. Rep Tr/Tr96.htm Prepred y Teres A. Newsome B.C. Ministry of Forests, Lnds nd Nturl Resource Opertions Crioo Region Willims Lke, B.C. Jen L. Heinemn J. Heinemn Forestry Consulting Vncouver, B.C. Amnd F. Linnell Nemec Interntionl Sttistics nd Reserch Corp. Brentwood By, B.C. Copies of this report my e otined, depending upon supply, from: Crown Pulictions, Queen s Printer 2nd Floor, 563 Superior Street Victori, BC v8w 9v For more informtion on other pulictions in this series, visit hfdctlog/index.sp 216 Province of British Columi When using informtion from this report, plese cite fully nd correctly.

4 Astrct Experimentl Project 94.2 (EP94.2) ws estlished in 1987 to exmine the performnce of western lrch (Lrix occidentlis), Sierin lrch (Lrix siiric), nd ponderos pine (Pinus ponderos) t ICHwk2, SBSmw, ESSFwk1, nd SBPSxc sites in the Crioo Region. Survivl nd growth of these non-ntive species over 22-yer period were compred with tht of lodgepole pine (Pinus contort vr. ltifoli), Dougls-fir (Pseudotsug menziesii), nd hyrid spruce (Pice engelmnnii gluc), which re common forest components in this region. Susequent to the instlltion of this experiment, climte chnge relted forest helth issues ecme n importnt mngement concern, nd the extension of species eyond their nturl rnge ws considered s mitigtion strtegy. Western lrch ws good cndidte for ssisted migrtion, nd interim guidelines re now in plce tht llow this species to e plnted in mny Crioo Region ecosystems. As result of these recent chnges, our 22-yer western lrch results re especilly relevnt to current forest mngement. Lodgepole pine, Dougls-fir, nd hyrid spruce survived nd grew ccording to generl expecttions for the individul iogeoclimtic units, except tht lodgepole pine sustined high (4%) first-yer mortlity t the ESSFwk1 site. At ll loctions, lodgepole pine grew more rpidly in erly yers thn Douglsfir or hyrid spruce, nd continued to e lrger t ge 22. Dougls-fir, once estlished, survived nd grew well t the ICHwk2 nd SBSmw sites ut did poorly t the SBPSxc instlltion. Hyrid spruce exhiited etter overll survivl thn ny other species ut grew considerly more slowly thn lodgepole pine or Dougls-fir throughout the 22-yer ssessment period. Western lrch hd higher survivl nd etter growth t the ESSFwk1 thn the ICHwk2 or SBSmw sites, possily ecuse the seedlot used t the ESSFwk1 instlltion ws etter dpted to moist conditions thn tht used elsewhere. After 22 yers, 79% of western lrch survived t the ESSFwk1 site, nd trees were pproximtely s tll s lodgepole pine. Within its current rnge, western lrch is not common in the ESSF zone, nd we speculte tht if more suitle seedlot hd een used t our ICHwk2 nd SBSmw sites, western lrch growth would hve t lest equlled tht t the ESSFwk1 loction. Western lrch hd very low survivl nd ws unle to increse in height t the SBPSxc site, pprently ecuse it ws s ill-suited s Douglsfir to the hrsh climtic conditions of the Chilcotin plteu. Sierin lrch survivl ws equl to or etter thn tht of western lrch t ll the experimentl loctions, ut fter 22 yers, it ws pproximtely 1 m shorter thn western lrch t the ESSFwk1 nd SBSmw sites. Both lrches sustined considerle forking dmge, ut from growth nd yield perspective, Sierin lrch hd the more serious defects. At the SBPSxc site, severe forking notwithstnding, Sierin lrch hd the second-est survivl fter lodgepole pine nd ws the only other species tht ws even mrginlly le to withstnd the climtic limittions of tht loction. Ponderos pine survived nd grew resonly well t the SBSmw nd ICHwk2 sites, ut tended to increse in dimeter t the expense of height nd ws sujectively oserved to hve lrge rnches nd wide crowns, trits tht do not suggest high economic vlue. Like ll other species except lodgepole pine, ponderos pine did poorly t the SBPSxc site. iii

5 ACKNOWLEDGEMENTS The uthors would like to thnk Crig Sutherlnd for co-ordinting nd overseeing the project, nd West Frser Mills Ltd. for cquiring the Sierin lrch seed nd providing ongoing technicl nd finncil support. We re grteful to Gerry Chpmn, George White, nd Doug Routledge for selecting sites nd estlishing individul experiments. We thnk Kirsteen Ling nd Lrry McCulloch for initil ssessments nd dt summries, nd Ev Fickell, Jnet nd Scott Zimonick, nd Industril Forestry Services employees for lter dt collection. We lso thnk Brry Jquish, De McKillop, Peter Ott, nd Alln Powelson for providing helpful review comments. Funding ssistnce ws received from the B.C. Ministry of Forests, the Forest Investment Account, nd West Frser Mills Ltd. Funding ssistnce does not imply endorsement of ny sttement or informtion in this report. iv

6 Contents Astrct... Acknowledgements Introduction Methods Study Ares Experimentl Design nd Tretments Mesurements Dt Clcultions nd Sttisticl Anlysis Results Survivl, Vigour, nd Overtopping Western lrch Sierin lrch Lodgepole pine Dougls-fir Hyrid spruce Ponderos pine Dmge nd Dmging Agents in Yers Hre nd frost dmge Leder dmge nd forking Species-specific dmging gents Growth Between-site comprison of size nd growth rtes for individul species Between-species size nd growth rte comprison t the ICHwk2, SBSmw, nd ESSFwk1 sites Stnd-level Responses Yer 22 tree condition ccording to growth nd yield dmge criteri Gross stnd volume per hectre Site index Discussion Lodgepole Pine, Dougls-fir, nd Hyrid Spruce Western Lrch Sierin Lrch Ponderos Pine Mngement Considertions Conclusion Literture Cited APpendices 1 Stocktype, seedlot, nd nursery informtion Assessment codes for vigour, overtopping, dmge, nd dmge cuse iii iv v

7 tles 1 ep94.2 site nd climtic chrcteristics Overlp of western lrch seed plnning zones lw1 nd lw2 with Crioo Region ICHwk, SBSmw, ESSFwk, nd SBPSxc suzones Site species comintions exmined in ep p-vlues indicting sttisticl significnce of interctions etween site nd species for size nd growth vriles etween yer 3 nd yer Site effects on men height of individul tree species in yer 3, yer 5, yer 1, yer 16, nd yer Site effects on men height nd dimeter growth of individul species for yer 1 16, 16 22, nd intervls Site effects on men ground-level dimeter in yer 3, yer 5, yer 1, nd yer 16, nd men dimeter t 1.3 m in yer Site effects on men stem volume for individul tree species in yers 16 nd 22, nd volume growth from yers Proportion of live trees ffected y specific dmge types nd dmging gents in the yer 22 growth nd yield ssessment t the ICHwk2, SBSmw, ESSFwk1, nd SBPSxc sites Men yer 22 site index estimted from yer 22 tree size for individul tree species t the ICHwk2, SBSmw1, ESSFwk1, nd SBPSxc sites Figures 1 ep94.2 study site loctions Survivl from plnting to yer 22 for western lrch, Sierin lrch, lodgepole pine, Dougls-fir, hyrid spruce, nd ponderos pine t individul sites Vigour nd overtopping of western lrch, Sierin lrch, lodgepole pine, Dougls-fir, hyrid spruce, nd ponderos pine in yers 3, 5, 16, nd 22 t individul sites Proportion of live trees of ll species ffected y hre dmge in yers 1 nd 3 t the ICHwk2 site only, nd live Sierin lrch nd Dougls-fir ffected y frost dmge in yer 1 t the SBSmw nd SBPSxc sites Survivl nd proportion of live western lrch, Sierin lrch, lodgepole pine, Dougls-fir, hyrid spruce, nd ponderos pine with leder or stem dmge in yers 1, 3, 5, 1, nd 16 t individul sites Yer 16 proportion of live lodgepole pine nd ponderos pine with western gll rust stem or rnch infections, lodgepole pine nd ponderos pine with pine needle cst, nd hyrid spruce with cooley gll delgid rnch glls t individul sites Comprison of yer 22 height, yer 1 16 height growth, yer height growth, nd yer height growth for individul species t the ICHwk2, SBSmw, nd ESSFwk1 sites Comprison of yer 22 dh nd yer dh growth for individul species t the ICHwk2, SBSmw, nd ESSFwk1 sites Comprison of yer 22 stem volume nd yer stem volume growth for individul species t the ICHwk2, SBSmw, nd ESSFwk1 sites Comprison of yer 22 gross stnd volume per hectre for individul species t the ICHwk2, SBSmw, ESSFwk1, nd SBPSxc sites Comprison of yer 22 gross stnd volume per hectre t ICHwk2, SBSmw, nd ESSFwk1 sites for western lrch, Sierin lrch, lodgepole pine, Dougls-fir, hyrid spruce, nd ponderos pine vi

8 1 INTRODUCTION Guidelines for selecting conifer species to regenerte hrvested sites in British Columi tke into considertion nturl distriutions, ecologicl suitility, nd fctors relted to forest helth nd productivity (B.C. Ministry of Forests 22). In the pst, the performnce of indigenous species outside their nturl rnges, nd of non-ntive species, ws lrgely specultive in British Columi ecuse our requirements were stisfied y the wide diversity of ntive species nd the generl good helth of our forests. In response to the recent mountin pine eetle epidemic (B.C. Ministry of Forests nd Rnge 27) nd projected climte-chnge effects on forest helth nd species distriution (Spittlehouse 28; Kliejuns et l. 29), there is now greter prcticl interest in expnding the rnge of cceptle species. One of the primry cndidtes for fcilitted migrtion is western lrch (Lrix occidentlis), which is the most productive interior conifer species in British Columi. This shde-intolernt species is est suited to medium to rich, modertely to slightly dry sites, nd under optimum conditions, it cn ttin site indices of up to 3 m (B.C. Ministry of Forests, Lnds nd Nturl Resource Opertions 213). It hs the enefit of hving reltively few serious helth prolems (Henigmn et l. 21), nd it produces high-qulity wood tht is suitle for rnge of products (Keegn et l. 1995; Prent et l. 28). At present, western lrch s nturl distriution in British Columi is restricted to the southestern portion of the province, ut recent in-depth modelling project y Rehfeldt nd Jquish (21) suggested tht its potentil niche, especilly s climte chnge is relized, extends considerly further north. On the sis of this work, seed plnning zones (LW1 nd LW2) were defined for western lrch in centrl interior British Columi, nd interim guidelines were implemented to llow the inclusion of up to 1% western lrch in mngement units within these zones (B.C. Ministry of Forests nd Rnge 21 1 ; B.C. Ministry of Forests, Lnds nd Nturl Resource Opertions 214). Sierin lrch (Lrix siiric), which is ntive to northestern prts of Europen Russi nd western Sieri, is lso of cdemic interest ecuse it hs high productivity in regions tht re climticlly similr to prts of north nd centrl British Columi. In prticulr, the Rivol provennce, which originted from stnd estlished in 1738 ner St. Petersurg to provide the Russin nvy with rw mteril for shipuilding, hs cught the interest of reserchers, since it is cple of ttining stnd volumes of up to 1 m 3 /h (Viherä-Arnio nd Nikknen 1995). Smith (1986) stted tht the most importnt fctor for successful introduction of exotic tree species is the similrity of climte nd soils to the ntive region. He points out tht drmtic movements of species cross mjor geogrphic rriers to similr hitts my e more successful thn seemingly modest extensions of nturl rnges. Possily ecuse we hve such welth of indigenous species, the introduction of exotic species hs never een seriously contemplted in British Columi. However, the process hs een successful in other prts of the world the most well-known eing the successful introduction of Dougls-fir (Pseudotsug menziesii) to the United Kingdom nd Europe, nd Monterey pine (Pinus rdit) to res in the southern hemisphere tht re 1 British Columi Ministry of Forests nd Rnge. 21. Interim mesures for the ssisted rnge nd popultion expnsion of western lrch for use s climte chnge dpttion strtegy in British Columi. Tree Improvement Br., Victori, B.C. Unpul. rep. 1

9 environmentlly similr to Cliforni. Sierin lrch is currently used to some extent in Scndinvin forest mngement; however, reserch to identify suitle provennces nd seed sources for tht region is ongoing (Mrtinsson 1995; Lrsson-Stern 23). Although long-term field experiments tht monitor species site reltionships re somewht limited y the smll numer of seedlots they involve (nd hence they involve little genetic vrition) nd y vrition tht cn e introduced when stock is sourced from more thn one nursery, they re nonetheless good source of informtion for considering the ssisted movement or introduction of tree species (Aitken et l. 28). In 1988, species study (Experimentl Project 94.2 [EP94.2]) ws undertken in the Crioo Region of south centrl British Columi to compre the performnce of western nd Sierin lrch with tht of lodgepole pine (Pinus contort vr. ltifoli), Dougls-fir, nd hyrid spruce (Pice engelmnnii gluc), which occur nturlly in tht geogrphic re. Out of interest out its ility to perform north of its nturl rnge, ponderos pine (Pinus ponderos) ws lso included in the experiment. Experimentl Project 94.2 ws designed to encompss the wide diversity of ecosystems tht occur in the Crioo Region (Steen nd Coupé 1997). It included single replictes on zonl sites in four iogeoclimtic suzones or vrints (Figure 1). Limittions to conifer seedling estlishment were nticipted to e most severe t the SBPSxc site, which hd high frost hzrd rting (Steen et l. 199). Limittions were lso expected to e somewht greter t ^ Study site! ^ Biogeoclimtic zone Bunchgrss Engelmnn Spruce Sulpine Fir ESSFwk1 site Wells! Interior Cedr Hemlock Interior Dougls-Fir Montne Spruce ^ Quesnel Su-Borel Pine-Spruce SBSmw site! ( Kersley Su-Borel Spruce! ( McLeese Lke Willims Lke Alexis Creek! ( Kilometres ep ^ Horsefly! ( ^ ICHwk2 site 15 Mile House! ( SBPSxc site 6 8 study site loctions. 2

10 TABLE 1 ep94.2 site nd climtic chrcteristics the ESSFwk1 site thn t the ICHwk2 or SBSmw sites, minly ecuse of the higher elevtion of the ESSFwk1 instlltion, nd ecuse tht iogeoclimtic vrint is suject to hevy snowlods (Steen nd Coupé 1997). Climtic chrcteristics of the reserch sites re provided in Tle 1. Experimentl Project 94.2 ws estlished long efore the interim guidelines for lrch use cme into eing (B.C. Ministry of Forests, Lnds nd Nturl Resource Opertions 214). Although none of our sites re precisely within the LW1 or LW2 seed plnning zones (Tle 2), the 22-yer western lrch responses re of interest ecuse ll the iogeoclimtic suzones we exmined hve some degree of overlp with the seed plnning zones. Site ICHwk2 SBSmw ESSFwk1 SBPSxc Site chrcteristics Opening numer 93A B 99(TFL 52) 93H 31(TFL 52) 93B Ltitude/longitude N ', W ' N ', W ' N ', W ' N52.42', W ' Site series Elevtion 114 m 85 m 134 m 125 m Reltive soil moisture regime Mesic to suhygric Mesic Mesic Mesic Actul soil moistue regime Fresh to moist Fresh Moist Modertely dry Slope position Mid lower slope Flt Mid slope Flt Topogrphy N/A Undulting N/A Flt Slope (%) 15 4% N/A 15 25% N/A Aspect NW N/A SW N/A Site history Clercut 1985 Brodcst urn 1986 Clercut 1985 Brodcst urn 1986 Clercut 1985/86 Brodcst urn 1986 Clercut 1983 Disc trenched 1986 Previous timer type CwFd 841 Unknown Unknown Pl 21 Recommended regenertion species 1 : Fd, Pl, Sx 2 : Bl, Cw, Hw 1 : Fd, Pl, Sx 2 : Bl 1 : Se, Bl 2 : Pl 1 : Pl 2 : none Climte c Men nnul temperture ( C) Men wrmest month temperture ( C) Men coldest month temperture ( C) Numer of frost-free dys Continuous frost-free period (dys) Men nnul precipittion (mm) Men summer precipittion (mm) Precipittion s snow (mm) Reltive soil moisture regime is moisture regime reltive to moisture vilility, while ctul soil moisture regime refers to moisture vilility without regrd to ecosystem (vlues from Steen nd Coupé 1997). Fd is Dougls-fir; Pl is lodgepole pine; Sx is hyrid spruce; Bl is sulpine fir; Cw is western redcedr; Hw is western hemlock. c From Climte BC, using normls (updted July 5, 215). 3

11 TABLE 2 Overlp of western lrch seed plnning zones lw1 nd lw2 with Crioo Region ICHwk, SBSmw, ESSFwk, nd SBPSxc suzones Suzone Totl re in Crioo Region (h) Are overlpping lw1 or lw2 (h) Proportion of suzone designted s lw1 or lw2 Loction of reserch site reltive to lw1 nd lw2 ICHwk % Less thn 1 km est of lw2 (northest of Willims Lke) SBSmw % Approximtely 2.5 km west nd 1 km est, respectively, of nrrow lw2 nd lw1 res tht occur to the est of Quesnel ESSFwk % 25 3 km est of lw2 (northest of Quesnel) SBPSxc % A minimum of 3 km from sustntive lw1 oundries tht surround the Chilcotin plteu Bsed on informtion provided y Geosptil Services, B.C. Ministry of Forests, Lnds nd Nturl Resource Opertions (Jn. 216). 2 METHODS 2.1 Study Ares 2.2 Experimentl Design nd Tretments The experiment includes single site in ech of four iogeoclimtic suzones or suzone vrints: (1) Quesnel vrint of the wet, cool Interior Cedr Hemlock zone (ICHwk2), (2) moist wrm Su-Borel Spruce zone (SBSmw), (3) Crioo vrint of the wet, cool Engelmnn Spruce Sulpine Fir zone (ESS- Fwk1), nd (4) very dry, cold Su-Borel Pine Spruce zone (SBPSxc). The SBPSxc site hd een clssified s IDFdk4 t the strt of the study (Newsome et l. 1995) ut ws lter reclssified when iogeoclimtic oundries were redrwn. Individul site nd climte chrcteristics re provided in Tle 1. The experiment consisted of four seprte rndomized lock designs, one t ech of the four iogeoclimtic suzones/vrints (sites). At ech site, plntings of up to six tree species were replicted in three locks (western lrch [Lw], Sierin lrch [Ls], hyrid spruce [Sx], Dougls-fir [Fd], lodgepole pine [Pl], nd ponderos pine [Py]). Site species comintions re shown in Tle 3. TABLE 3 Site (suzone/vrint) species comintions exmined in ep94.2 Site (suzone/vrint) Species ICHwk2 SBSmw ESSFwk1 SBPS c Western lrch Sierin lrch Lodgepole pine Dougls-fir Hyrid spruce Ponderos pine Dougls-fir nd ponderos pine were excluded from the ESSFwk1 site, which ws clerly ove the elevtionl rnge of those species, nd hyrid spruce ws excluded from the SBPSxc site due to the dryness of tht ecosystem (the site ws originlly thought to e IDFdk4). At ech site, ech species (six t the ICHwk2 nd SBSmw sites, four t the ESSFwk1 site, nd five t the SBPSxc site) ws rndomly ssigned to single plot mesuring m within ech of three locks. Blocks mesured 5 75 m t the ICHwk2, SBSmw, 4

12 nd SBPSxc sites, where five or six species were exmined, nd 25 1 m t the ESSFwk1 site, where only four species were exmined. Seedlings were plnted on grid within ech plot, t 2-m spcing, for totl of 144 trees per plot. In ech plot, the 44 perimeter trees were left s uffer, nd the inner 1 trees, which were rrnged on 1 1 grid tht ws pproximtely 2 2 m (4 m 2 ), were intended for eventul growth nd yield ssessment. Of these inner 1 trees, the first 5 (i.e., trees in the first five rows) were tgged t the time of plnting for regulr ssessment. In totl, 15 trees of ech species t ech site nd 315 trees in the experiment s whole were ssessed t regulr intervls. In yer 22, the remining 5 inner trees in ech plot were tgged to llow growth nd yield ssessment of ll 1 inner trees in ech plot. Stocktype, seedlot, nd nursery informtion is provided in Appendix 1. Seedlings were plnted in spring 1987, with the exception of western lrch t the ESSFwk1 site, which ws plnted in spring 1988 due to n insufficiency of stock in As result, western lrch plnted t the ESSFwk1 site ws grown from different seedlot thn stock used t the other sites. 2.3 Mesurements 2.4 Dt Clcultions nd Sttisticl Anlysis With the exception of western lrch t the ESSFwk1 site, the first 5 mesurement trees in ech plot were ssessed fter growth ws complete in yers 1 (1987), 2 (1988), 3 (1989), 5 (1991), 1 (1996), 16 (22), nd 22 (28). Assessments for western lrch t the ESSFwk1 site were delyed y 1 yer to compenste for the 1-yer plnting dely. Herefter, for simplicity of presenting results, western lrch results will e presented s if they followed the sme mesurement schedule s the other species. During ech ssessment, height (± 5 cm) nd dimeter (±.1 cm) were mesured. Dimeter ws mesured t ground level (GLD) up to yer 16, nd t rest height (1.3 m) (dh) in yers 16 nd 22. Survivl, vigour, type nd cuse of dmge, nd degree of overtopping of mesurement trees y competing vegettion were ssessed in yers 1, 2, 3, 5, 1, nd 16 ccording to Crioo Region Reserch Section codes tht were current for the ssessment yer (Appendix 2). In yer 22, survivl of the first 5 mesurement trees in ech plot ws ssessed, ut vigour, type nd cuse of dmge, nd overtopping were not ssessed s they hd een in previous yers. In the yer 22 ssessment, trees 51 1 (which hd not een ssessed previously) were mesured for height nd dh, so tht, together with dt for trees 1 5, gross stnd volume per hectre could e determined sed on volume in the known plot re of 4 m 2. In the yer 22 ssessment, dmge gent nd dmge severity dt were collected ccording to B.C. Ministry of Sustinle Resource Mngement (23) criteri for ll 1 mesured trees. Specific cuses of tree mortlity were not recorded in ny yers. In this report, for the first 5 mesurement trees, we present survivl for yers 1, 3, 5, 1, 16, nd 22, overtopping for yers 3, 5, 1, nd 16, nd vigour nd growth for yers 3, 5, 1, 16, nd 22. For the full set of 1 mesured trees, we present gross stnd volume, dmge gent, nd dmge severity results for yer 22. All dt were checked for outliers nd normlity prior to, nd s prt of, the nlysis. Individul tree volumes for yers 16 nd 22 were clculted using Kozk s (1988) tper equtions. Height growth ws clculted for the intervls corresponding to 1 16, 16 22, nd yers post plnting. Dimeter growth ws clculted for the intervl 1 16 yers post plnting sed on GLD nd for the intervl yers post plnting sed on dh. Volume growth ws clculted for the intervl yers. Ctegoricl vriles were 5

13 summrized s percent frequencies. Dmge summries included only trees with good, fir, or poor vigour; dmge to moriund trees ws excluded ecuse it ws inconsistently collected for trees tht were very close to deth. Site index ws estimted y dividing the re occupied y the first 5 mesurement trees (which hd een mesured repetedly since the strt of the study) into pproximtely.1-h suplots, selecting the lrgest dh plnted tree in ech suplot s site tree sed on yer 22 mesurements, nd pplying the following growth intercept models: western lrch nd Sierin lrch (Nigh et l. 1999), lodgepole pine (Nigh 1997), Dougls-fir (Nigh 1997), hyrid spruce (Nigh 1999), nd ponderos pine (Nigh 22). For the ICHwk2, SBSmw, nd ESSFwk1, six trees of ech species per site contriuted to the site index clcultions. In the SBPSxc, six trees contriuted to site index determintion for Sierin lrch nd lodgepole pine, nd five contriuted to the vlue for ponderos pine. Only two trees contriuted to site index determintion for western lrch ecuse so few chieved rest height. Similrly, site index could not e determined for Dougls-fir ecuse trees did not chieve rest height. Mixed-effects nlysis of vrince (ANOVA) ws used to ssess the interctive effects of site nd species on the height, dimeter, nd volume of trees (excluding those in moriund condition). All nlyses were sed on splitplot ANOVA model with (excluding the su-smpling [tree] error) fixed site (i.e., suzone/vrint) the grouping fctor for the min plots (i.e., locks), species minus the split-plot fctor, nd site species effects, nd rndom lock (nested in site) nd lock species effects. We tested for n overll site species interction (degrees of freedom = 12 rther thn the 15 tht would hve een used for design where ll 24 species site comintions were represented) nd constructed series of contrsts to test for site species interction t ech pir of sites (degrees of freedom for ech pir of sites = numer of species common to the two sites minus 1). To simplify interprettion of the interctions, seprte nlyses were lso performed y site nd y species. The Bonferroni method of multiple comprisons ws used to ssess the significnce of differences mong relevnt species or site mens. 2 3 RESULTS 3.1 Survivl, Vigour, nd Overtopping Western lrch Yer 22 survivl of western lrch t our four study sites decresed in the order of ESSFwk1 (79%) > ICHwk2 (5%) > SBSmw (43%) > SBPSxc (24%), trend tht ws estlished s erly s yer 1 (Figure 2). At the ESSFwk1 site, 5% of western lrch died within 1 yer of plnting, nd 13% hd died y yer 3, fter which survivl decresed very grdully over time. The proportion of the originl mesurement trees in good or fir vigour t the ESSFwk1 instlltion remined reltively constnt (67 74%) from yer 3 to yer 22 (Figure 3). At the ICHwk2, SBSmw, nd SBPSxc sites, survivl declined to 81%, 75%, nd 75%, respectively, during the first growing seson, fter which there were ongoing, reltively stedy losses to ge 22 (Figure 2). At the ICHwk2 site, western lrch vigour declined steeply etween yers 1 nd 16, nd there ws similr decline etween yers 5 nd 1 t the SBSmw 2 All dt nlyses for this report were generted using SAS/STAT softwre, Version 9.3 of the SAS System for Windows Copyright SAS Institute Inc. SAS nd ll other SAS Institute Inc. product or service nmes re registered trdemrks or trdemrks of SAS Institute Inc., Cry, NC, USA. 6

14 site (Figure 3). There ws some vigour recovery t oth loctions, resulting in out 4% of the originl plnted trees hving good or fir vigour in yer 22. At the SBPSxc site, lmost ll surviving western lrch hd poor vigour or were moriund from yer 3 onwrds (Figure 3). At the ICHwk2 loction, 33% nd 38% of surviving western lrch were overtopped in yers 3 nd 5, respectively, ut lmost none were overtopped in yers 1 nd 16 (Figure 3). Approximtely 2% of live western lrch were overtopped in yer 3 t the SBSmw site; the proportion of overtopped trees declined slightly y yer 5, nd in yers 1 nd 16, lmost none were overtopped. Overtopping of western lrch ws minor t the ESSFwk1 (mximum 12% in yer 5) nd the SBPSxc (mximum 6% in yer 16) sites Sierin lrch Yer 22 Sierin lrch survivl ws higher thn tht of western lrch t ll four sites, nd declined in the order of ESSFwk1 (81%) > ICHwk2 (73%) > SBSmw (66%) > SBPSxc (54%); gin, this trend ws estlished in yer 1 (Figure 2). There ws no first-yer mortlity of Sierin lrch t the ESSFwk1 instlltion, ut 16%, 24%, nd 32% of seedlings t the ICHwk2, SBSmw, nd SBPSxc sites, respectively, died within 1 yer of plnting. Declines in survivl fter yer 1 were firly grdul t ll loctions. At the SBSmw site, lmost ll Sierin lrch tht survived to yer 3 consistently hd good or fir vigour through yer 22 (Figure 3). In contrst, t the ESSFwk1 site where survivl ws highest, smll proportion of trees hd poor vigour; this ws lso true t the ICHwk2 loction, lthough y yer 22, nerly ll survivors t tht site were in good or fir vigour. At ge 22, the proportion of the originl mesurement trees tht hd good or fir vigour ws the sme (71%) t the ESSFwk1 nd ICHwk2 sites, nd ws slightly lower (66%) t the SBSmw instlltion. At the SBPSxc site, pproximtely one-hlf to two-thirds of survivors hd t lest fir vigour through yer 1, ut most were in poor condition in yers 16 nd 22 (Figure 3). Approximtely 2% of surviving Sierin lrch t the ICHwk2 site were overtopped in yer 3, nd this incresed to nerly 4% y yer 5 (Figure 3). In yers 1 nd 16, presumly s trees outgrew surrounding vegettion, there ws very little overtopping of Sierin lrch t this loction. Approximtely 2% of surviving Sierin lrch were overtopped in yer 3 t the ESSFwk1 site. The proportion decresed slightly y yer 5, nd lmost none of the ESSFwk1 Sierin lrch were overtopped in yers 1 or 16. Very few of the surviving Sierin lrch t the SBSmw or SBPSxc instlltions were overtopped in ny of the ssessment yers Lodgepole pine Lodgepole pine survivl t the ICHwk2 nd SBPSxc sites ws similrly high ( 89%) throughout the 22-yer ssessment period (Figure 2c), ut vigour trends differed etween these loctions (Figure 3c). At the ICHwk2 site, pproximtely one-third of surviving pine hd poor vigour in yer 3, fter which condition improved stedily until yer 1. A smll proportion of pine t this instlltion hd poor vigour gin in yers 16 nd 22; however, most (8%) were in good or fir condition. At the SBPSxc site, lthough more thn 9% of pine survived to yer 22, vigour declined fter yer 1; only 61% of the originl mesurement trees were in good or fir condition in yers 16 nd 22. At the SBSmw nd ESSFwk1 sites, lodgepole pine survivl declined to 85% nd 6%, respectively, within 1 yer of plnting (Figure 2c). 7

15 () Western lrch ESSFwk1 SBSmw ICHwk2 SBPSxc Survivl (%) P Yer () Sierin lrch Survivl (%) P Yer (c) Lodgepole pine Survivl (%) P Yer FIG 2 Survivl from plnting to yer 22 for () western lrch, () Sierin lrch, (c) lodgepole pine, (d) Dougls-fir, (e) hyrid spruce, nd (f) ponderos pine t individul sites. 8

16 (d) Dougls-fir ESSFwk1 SBSmw ICHwk2 SBPSxc Survivl (%) P Yer (e) Hyrid spruce Survivl (%) P Yer (f) Ponderos pine Survivl (%) P Yer FIG 2 Continued 9

17 () Western lrch 1 Good Fir Poor Moriund Overtoppedd (d) Dougls-fir Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 SBPSxc ICHwk2 SBSmw SBPSxc () Sierin lrch Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 SBPSxc (c) Lodgepole pine Proportion of trees (%) Proportion of trees (%) (e) Hyrid spruce Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 (f) Ponderos pine Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 SBPSxc ICHwk2 SBSmw SBPSxc FIG 3 Vigour (proportion of originl plnted trees) nd overtopping (proportion of surviving trees tht re overtopped) of () western lrch, () Sierin lrch, (c) lodgepole pine, (d) Dougls-fir, (e) hyrid spruce, nd (f) ponderos pine in yers 3, 5, 16, nd 22 t individul sites. Overtopping dt were not collected in yer 22. 1

18 After yer 1, survivl remined stle t the ESSFwk1 site, wheres t the SBSmw instlltion, it declined reltively steeply until yer 5 nd then stilized. In yer 22, lodgepole pine survivl ws similr (59 62%) t the two loctions. At the ESSFwk1 nd SBSmw sites, most pine tht survived to yer 3 hd good or fir vigour through to yer 22 (Figure 3c). At the ICHwk2 site, 34% nd 3% of surviving lodgepole pine were overtopped in yers 3 nd 5, respectively, ut nerly ll were tller thn the surrounding vegettion in yers 1 nd 16 (Figure 3c). At the ESSFwk1 instlltion, 25% of pine were overtopped in yer 3, ut lmost none were overtopped in yers 5, 1, or 16. There ws minor pine overtopping t the SBSmw instlltion in yers 3 nd 5 ( 6%) nd none in ny ssessment yer t the SBPSxc site Dougls-fir Dougls-fir t the SBSmw, ICHwk2, nd SBPSxc sites hd similr yer 1 survivl (91 93%) (Figure 2d). After yer 1, survivl t the SBSmw nd ICHwk2 loctions grdully diverged until it ws 73% nd 58%, respectively, in yer 22. At the SBSmw site, most surviving Dougls-fir were in good or fir vigour t ll ssessment dtes (Figure 3d). In contrst, more thn one-hlf of the surviving Dougls-fir t the ICHwk2 instlltion were in poor or moriund condition in yer 3, fter which vigour improved consistently until yer 1. Although Dougls-fir survivl declined etween yers 1 nd 16 t the ICHwk2 site, nerly ll live trees continued to hve good or fir vigour. At the SBPSxc instlltion, survivl declined stedily fter plnting to 18% in yer 22 (Figure 2d). Nerly ll surviving Dougls-fir t this site were in poor condition or moriund throughout the 22-yer ssessment period (Figure 3d). Approximtely one-hlf (49%) of the surviving Dougls-fir t the ICHwk2 instlltion were overtopped in yer 3, nd the proportion incresed to 66% y yer 5 (Figure 3d). Most survivors grew through surrounding vegettion fter yer 5, so tht 7% were overtopped in yers 1 nd 16. Overtopping of Dougls-fir ws less severe t the SBSmw site, ut gin, it incresed etween yers 3 nd 5 (from 26 to 34%) efore declining to less thn 6% in yers 1 nd 16. At the SBPSxc loction, none of the surviving Dougls-fir were overtopped in ny ssessment yer Hyrid spruce Throughout the 22-yer ssessment period, hyrid spruce survivl ws 91% t the ICHwk2 nd ESSFwk1 sites, nd 84% t the SBSmw site (Figure 2e). With the exception of short period of decline in condition in yer 16 t the SBSmw loction, most spruce consistently hd good or fir vigour (Figure 3e). Twelve percent of live spruce t the ICHwk2 site were overtopped in yer 3, nd this incresed to 3% y yer 5 (Figure 3e). In yers 1 nd 16, overtopping of spruce t this loction ws miniml (2 3%). At the SBSmw site, 11% of surviving spruce were overtopped in yer 3; the proportion incresed to 26% in yer 5, decresed to 3% in yer 1, nd then incresed gin to 15% in yer 16. At the ESSFwk1 instlltion, 46% of surviving spruce were overtopped in yer 3, ut the proportion decresed to 15% y yer 5 nd ws miniml in yers 1 nd 16 ( 4%) Ponderos pine Yer 1 ponderos pine survivl ws higher (94 99%) t the ICHwk2 nd SBPSxc sites thn t the SBSmw loction (8%) (Figure 2f). 11

19 Between yers 1 nd 5, survivl t the ICHwk2 site declined steeply to 59%, nd then grdully to 53% in yer 22. At the SBSmw site, ponderos pine survivl fter yer 1 declined grdully ut stedily until it ws 6% in yer 22. Ponderos pine survivl t the SBPSxc instlltion ws high until yer 1 ( 88%), fter which it declined precipitously to 34% in yer 16 nd 21% in yer 22. Ponderos pine vigour trends differed t the three loctions (Figure 3f). Most surviving ponderos pine t the ICHwk2 site hd poor vigour in yer 3, ut condition improved mrkedly y yer 1, likely ecuse survivors were outgrowing surrounding vegettion. Beyond ge 1, nerly ll survivors hd good or fir vigour. At the SBSmw instlltion, lmost ll surviving ponderos pine hd good or fir vigour t ll ssessment dtes, while t the SBPSxc site, vigour declined stedily through the 22-yer ssessment period (Figure 3f). At the ICHwk2 instlltion, 55% of surviving ponderos pine were overtopped in yer 3, nd the proportion incresed to 65% y yer 5 (Figure 3f). Ponderos pine grew through the vegettion etween yers 5 nd 1, nd there ws lmost no overtopping of tht species t the ICHwk2 site in yer 1 or yer 16. At the SBSmw loction, pproximtely 2% of ponderos pine were overtopped in yers 3 nd 5, ut none were overtopped in yers 1 or 16. There ws no overtopping of ponderos pine t the SBPSxc instlltion. 3.2 Dmge nd Dmging Agents in Yers Hre nd frost dmge Browsing nd clipping y hres ws common t the ICHwk2 site; in yer 1, hre dmge ffected 1 11% of surviving Sierin lrch, Dougls-fir, nd ponderos pine, nd 2 4% of lodgepole pine, western lrch, nd hyrid spruce (Figure 4). In yer 3 t this instlltion, hre dmge ffected 77% of surviving ponderos pine, 47% of western lrch, 4% of lodgepole pine, 36% of Dougls-fir, 16% of Sierin lrch, nd 15% of hyrid spruce. Hre dmge in yers 1 nd 3 ws minor t the ESSFwk1 nd SBPSxc sites, ffecting 5% of trees regrdless of species. No hre dmge ws recorded t the SBSmw instlltion in yer 1 or yer 3. At the end of the first growing seson, frost dmge ffected 2% of surviving Sierin lrch nd 1% of surviving Dougls-fir t the SBSmw site (Figure 4). At the SBPSxc site, frost dmge ffected 19% of surviving Sierin lrch nd 1% of surviving Dougls-fir t the end of the first growing seson. No yer 1 frost dmge ws oserved t other instlltions or mong other species. There ws lmost no recorded frost dmge to trees of ny species fter yer 1, lthough it my hve een present ut not recorded where more serious types of dmge ffected trees Leder dmge nd forking With the exception of yer 3 hre dmge t the ICHwk2 instlltion, leder injury nd stem forking were the most prevlent types of dmge tht ffected ll tree species t ll study sites. Western lrch nd Sierin lrch sustined the most extensive injury (Figure 5, ). Despite ongoing high survivl nd reltively good vigour t the ESSFwk1 site, 89% of surviving western lrch hd some form of leder dmge in yer 1. The proportion of trees with leder dmge declined to 38% y yer 16 t this loction, ut 69% of surviving lrch hd forked stems in yer 16. At the SBPSxc instlltion, ll western lrch tht survived to yer 16 hd leder dmge nd stem forks. Sierin lrch sustined similr mounts of leder nd forking dmge s western lrch t the ICHwk2 nd SBSmw sites ut ws less ffected thn western lrch t the ESSFwk1 loction. At the SBPSxc 12

20 () Hre dmge Proportion of trees (%) Yr1 Yr3 Yr1 Yr3 Yr1 Yr3 Yr1 Yr3 Yr1 Yr3 Yr1 Yr3 Western lrch () Frost dmge Sierin lrch Lodgepole pine ICHwk2 Douglsfir Hyrid spruce Ponderos pine Proportion of trees (%) SBSmw SBPSxc SBSmw SBPSxc Sierin lrch Douglsfir FIG 4 Proportion of () live trees of ll species ffected y hre dmge in yers 1 nd 3 (1989) t the ICHwk2 site only (occurrence of hre dmge ws minor t other sites), nd () live Sierin lrch nd Dougls-fir ffected y frost dmge in yer 1 t the SBSmw nd SBPSxc sites (these species did not sustin frost dmge t other sites, nor ws it recorded for other species t ny sites). Moriund trees re not included. instlltion, surviving Sierin lrch lso sustined slightly less forking dmge thn western lrch. Ponderos pine ws lso severely ffected y leder dmge nd/or forking t the SBPSxc site, where 73% of survivors were forked in yer 16 (Figure 5f). Yer 16 dmge ws not summrized for Dougls-fir t the SBPSxc site (Figure 5d), where nerly ll the trees were moriund; however, severe forking is ssumed to hve occurred sed on the verge yer 22 height of 22 cm. Dougls-fir nd ponderos pine sustined considerly less leder nd forking dmge t the ICHwk2 nd SBSmw sites thn t the SBPSxc instlltion, with mximum of 31% of Dougls-fir nd 22% of ponderos pine hving forks t ge 16. Lodgepole pine nd hyrid spruce hd more prevlent forking thn leder dmge (Figure 5c, e). In yer 16, 23% of lodgepole pine t the ICHwk2 site nd 35 41% t the other loctions hd forks. Hyrid spruce hd more extensive yer 16 forking dmge t the ICHwk2 (25%) nd SBSmw (41%) sites thn t the ESSFwk1 instlltion (13%) Species-specific dmging gents In yer 16, the proportion of live lodgepole pine tht hd western gll rust (Endocronrtium hrknessii) stem infection rnged from 6 to 1% t the ICHwk2, SBSmw, ESSFwk1, nd SBPSxc sites (Figure 6). The proportion of live lodgepole pine tht hd 13

21 Leder dmge Forked stem Survivl () Western lrch (d) Dougls-fir ICHwk2 SBSmw ESSFwk1 SBPSxc ICHwk2 SBSmw ESSFwk1 SBPSxc Proportion of trees (%) Proportion of trees (%) Proportion of trees (%) ICHwk2 SBSmw SBPSxc ICHwk2 SBSmw SBPSxc Proportion of trees (%) Proportion of trees (%) Proportion of trees (%) () Sierin lrch (e) Hyrid spruce (c) Lodgepole pine (f) Ponderos pine ICHwk2 SBSmw ESSFwk1 SBPSxc ICHwk2 SBSmw SBPSxc FIG 5 Survivl nd proportion of live () western lrch, () Sierin lrch, (c) lodgepole pine, (d) Dougls-fir, (e) hyrid spruce, nd (f) ponderos pine with leder or stem dmge in yers 1 (1987), 3 (1989), 5 (1991), 1 (1996), nd 16 (22) t individul sites. Moriund trees were not included. 14

22 western gll rust rnch infections ws more vrile, incresing in the order of ICHwk2 (1%) < SBPSxc (18%) < ESSFwk1 (21%) < SBSmw (4%). No ponderos pine hd western gll rust stem infection in yer 16, nd minor rnch infection (3% of live trees) ws oserved only t the SBSmw site. In yer 16, pine needle cst (Lophodermell concolor) infection ws recorded on 8% nd 1% of live lodgepole pine t the ESSFwk1 nd SBPSxc instlltions, respectively, ut ws not oserved on lodgepole pine t the ICHwk2 or SBSmw loctions (Figure 6). In contrst, pine needle cst ffected 35% nd 55% of ponderos pine t the ICHwk2 nd SBSmw sites, respectively, t ge 16 ut ws not oserved on trees t the SBPSxc instlltion. Cooley gll delgid rnch glls (Adelges cooleyi) were present on 92%, 95%, nd 68% of live hyrid spruce t the ICHwk2, SBSmw, nd ESSFwk1 sites, respectively, in yer 16 (Figure 6c), ut they did not sustntilly reduce tree vigour. () Western gll rust Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 Stem SBPSxc Brnch ICHwk2 SBSmw SBPSxc () Pine needle cst Lodgepole pine Ponderos pine Proportion of trees (%) ICHwk2 SBSmw ESSFwk1 SBPSxc ICHwk2 SBSmw SBPSxc (c) Cooley gll delgid Proportion of trees (%) ICHwk2 SBSmw Lodgepole pine ESSFwk1 Ponderos pine Hyrid spruce FIG 6 Yer 16 (22) proportion of live () lodgepole pine nd ponderos pine with western gll rust stem or rnch infections, () lodgepole pine nd ponderos pine with pine needle cst, nd (c) hyrid spruce with cooley gll delgid rnch glls t individul sites. Moriund trees re not included. 15

23 3.3 Growth There were significnt overll interctions etween site nd species for ll growth mesures tht were ssessed, in ll ssessment yers (Tle 4). When individul pirs of sites were compred, the site species interction for height ws most consistently significnt for ICHwk2 versus SBSmw, ICHwk2 versus SBPSxc, nd SBSmw versus ESSFwk1. For dimeter, the interction ws most consistently significnt for ICHwk2 versus SBSmw, ICHwk2 versus ESSFwk1, nd SBSmw versus ESSFwk1. Site species interctions for height growth during the yer 1 16 intervl were vrily significnt for the six pirwise comprisons. During the yer intervl, height growth interctions were significnt for ll pirwise comprisons, nd for the yer intervl, they were significnt for ll comprisons except ICHwk2 versus SBSmw. Dimeter growth interctions were vrile; they ecme significnt over time for some pirwise comprisons ut non-significnt for others. Stem volume nd volume growth interctions were significnt for ll pirwise comprisons except ICHwk2 versus SBPSxc. TABLE 4 p-vlues indicting sttisticl significnce of interctions etween site nd species for size nd growth vriles etween yer 3 (1989) nd yer 22 (28) Vrile Yer Overll ICHwk2 vs SBSmw ICHwk2 vs ESSFwk1 Site species interctions c ICHwk2 vs SBPSxc SBSmw vs ESSFwk1 SBSmw vs SBPSxc ESSFwk1 vs SBPSxc Height 3 <.1 <.1.2 <.1.7 < <.1 <.1.2 <.1.2 < <.1 <.1.68 <.1 <.1 < <.1 < < <.1 < <.1.26 <.1 Dimeter 3 <.1 <.1.11 <.1.2 < <.1 <.1 < < <.1 <.1 <.1.7 <.1.2 <.1 16 <.1 <.1 <.1.71 < <.1 <.1 < Volume 16 <.1 <.1 < <.1 <.1 22 <.1 <.1 < <.1 <.1 Height growth 1 16 <.1 < < <.1 <.1 <.1 <.1 <.1 <.1 < <.1.77 <.1 <.1 <.1 <.1 <.1 Dimeter growth 1 16 <.1 <.1 <.1.65 <.1.78 < <.1.27 <.1 <.1.24 <.1.61 Volume growth <.1.2 <.1.26 <.1.2 <.1 Interctions re considered significnt t p.5 (indicted y old font). Owing to the sence of some zone species comintions, the numer of degrees of freedom is not the sme for ech comprison presented in this tle. A significnt interction implies tht growth trends for conifer species re not prllel, while non-significnt interctions imply prllel growth trends. c For the overll site species interction, due to incompleteness the denomintor degrees of freedom = 12, rther thn 15 s would e the cse for design where ll 24 species site comintions re represented. For contrsts testing the site species interction for ech pir of sites, the denomintor degrees of freedom for ech pir of sites = numer of species common to the two sites minus 1. 16

24 3.3.1 Between-site comprison of size nd growth rtes for individul species Western lrch In ll ssessment yers, western lrch ws significntly tller t the ESSFwk1, SBSmw, nd ICHwk2 sites thn t the SBPSxc instlltion (Tle 5). Western lrch height ws sttisticlly similr t the ESSFwk1, SBSmw, nd ICHwk2 loctions in yers 3, 5, nd 1, ut in yers 16 nd 2, trees were significntly tller t the ESSFwk1 thn the ICHwk2 site. Western lrch height growth during the yer 1 16, 16 22, nd intervls ws lso significntly greter t the ESSFwk1, SBSmw, nd ICHwk2 sites thn t the SBPSxc instlltion (Tle 6). Western lrch t the ESSFwk1 site hd significntly greter height TABLE 5 Site effects on men height of individul tree species in yer 3 (1989), yer 5 (1991), yer 1 (1996), yer 16 (22), nd yer 22 (28) Height (cm) Species Yer ICHwk2 SBSmw ESSFwk1 SBPSxc p-vlue Western lrch 3 57 ± 6 78 ± 6 53 ± 6 1 ± ± ± 8 93 ± 8 22 ± ± ± ± 16 4 ± 2 < ± ± ± ± 38 c < ± ± ± ± 42 c <.1 Sierin lrch 3 54 ± 6 84 ± 6 63 ± 6 3 ± ± 1 c 138 ± ± 1 45 ± 11 c ± ± ± ± ± ± ± ± 28 c < ± ± ± ± 31 <.1 Lodgepole pine 3 36 ± 6 c 81 ± 6 59 ± 6 3 ± 6 c ± 11 c 162 ± ± ± 11 c ± ± ± ± 18 c < ± ± ± ± 23 c < ± ± ± ± 29 <.1 Dougls-fir c 3 29 ± 3 39 ± 3 14 ± ± 6 69 ± 6 2 ± 6.2 not ± ± 15 3 ± 17.4 included ± ± ± ± ± 6 <.1 Hyrid spruce 3 45 ± 3 5 ± 3 46 ± ± 6 8 ± 6 82 ± 6.48 not ± ± ± 1.6 included ± ± ± ± ± ± Ponderos pine 3 2 ± 2 56 ± 2 23 ± 2 < ± 3 15 ± 3 43 ± 2 <.1 not ± ± 7 95 ± 7 c <.1 included ± ± ± 31 c < ± ± ± 32 c <.1 Vlues re men ± 1 stndrd error. p-vlues re sed on ANOVA (p-vlues.5 re in old typefce). Mens ssigned the sme letter do not differ significntly ccording to the Bonferroni test. c Yer 16 vlues re not presented for Dougls-fir t the SBPSxc site ecuse ll trees were moriund (nd therefore were not included in the nlysis). 17

25 TABLE 6 Site effects on men height nd dimeter growth of individul species for yer 1 16, 16 22, nd intervls Height or dimeter growth (cm) Vrile Species Yer ICHwk2 SBSmw ESSFwk1 SBPSxc p-vlue Height growth Western lrch ± ± ± 9 23 ± 21 c < ± ± 9 36 ± 7 13 ± 13 c < ± 2 42 ± 2 53 ± 1 7 ± 3 c <.1 Sierin lrch ± ± ± ± 13 c < ± ± 1 25 ± 1 5 ± 1 < ± 2 53 ± 2 46 ± 2 1 ± 2 <.1 Lodgepole pine ± ± ± ± 6 < ± ± ± ± 8 < ± 2 47 ± 2 49 ± 2 25 ± 2 <.1 Dougls-fir ± ± not ± ± 1 9 ± 23 <.1 included ± 2 45 ± 2 2 ± 4 <.1 Hyrid spruce ± ± ± not ± ± ± 12.5 included ± 2 42 ± 2 43 ± 2.37 Ponderos pine ± 6 24 ± 6 47 ± 17 <.1 not ± ± 9 33 ± 15 <.1 included ± 3 4 ± 3 9 ± 3 <.1 Dimeter growth d Western lrch ± ± ±.3 c 2.6 ±.8 d < ± ± ±.2 <.1 Sierin lrch ± ± ± ±.3 c < ± ± ± ±.2.5 c Vlues re men ± 1 stndrd error. Lodgepole pine ± ± ± ±.6 c < ± ± ± ±.2.1 Dougls-fir e ±.4 9. ±.4 not ± ±.2 included.23 Hyrid spruce ± ± ±.5 not ±.2 3. ± ±.2 included.11 Ponderos pine ± ±.3 not 5.6 ± 1. c < ± ±.3 included 1.6 ±.5.1 p-vlues re sed on ANOVA (p-vlues.5 re in old typefce). Mens ssigned the sme letter do not differ significntly ccording to the Bonferroni test. c Indictes the Bonferroni test could not seprte mens lthough p.5. d Dimeter growth is sed on ground-level dimeter for the yer 1 16 intervl nd on dimeter t 1.3 m (dh) for the yer intervl. e Vlues re not presented for Dougls-fir t the SBPSxc site ecuse ll trees were moriund in yers 16 nd 22 (nd therefore were not included in the nlysis). growth thn those t the ICHwk2 site during the yer 1 16, 16 22, nd intervls, nd ESSFwk1 lrch lso hd significntly greter height growth thn SBSmw lrch during the yer nd intervls. In ll ssessment yers, western lrch ws significntly lrger in dimeter t the ESSFwk1, SBSmw, nd ICHwk2 sites thn t the SBPSxc instlltion (Tle 7). In yers 3 nd 5, western lrch t the SBSmw site hd significntly lrger dimeter thn t the ESSFwk1 or ICHwk2 sites. By yer 1, dimeter 18