Regeneration development under shelterwoods in a lowland red spruce balsam fir stand

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1 Regenertion development under shelterwoods in lowlnd red spruce lsm fir stnd 31 Dvid Pothier nd Mrcel Prévost Astrct: The shelterwood system cn e used to estlish regenertion nd to improve the growing conditions of seedlings nd, therey, increse their proility of survivl fter finl hrvest. To determine the cutting intensity tht est promotes the development of red spruce (Pice ruens Srg.) regenertion, n experiment, including four repetitions of five tretments (control; low, moderte, nd hevy prtil cuttings; nd cler-cutting), ws estlished in lowlnd stnd in Queec, Cnd, dominted y red spruce nd lsm fir (Aies lsme (L.) Mill.). Regenertion development responses to tretments vried ccording to cutting intensity, species, nd position reltive to the skidding trils in the residul stnds. Ten yers fter tretment, red spruce seedlings were well estlished ut were smller thn those of lsm fir nd deciduous species. Red spruce seedling height ws generlly greter in prtil-cut skidding trils, s ws light vilility. However, in clercuts, the size of red spruce seedlings estlished in skidding trils seemed to e negtively ffected y the considerle quntity of woody deris strewn over them. Among the prtil-cut tretments, tree regenertion lef iomss ws the highest where 6% of the initil sl re ws removed. Thus, this tretment is n ttrctive lterntive to cler-cutting in such lowlnd stnds where wtering-up is nticipted fter finl hrvest. Résumé : L coupe progressive d ensemencement peut être utilisée pour étlir l régénértion et pour méliorer les conditions de croissnce des semis, ce qui ugmenterit leur proilité de survie près l coupe finle. Dns le ut de déterminer l intensité de coupe mximisnt le développement de l régénértion d épinette rouge (Pice ruens Srg.), un dispositif expérimentl, comprennt qutre répétitions de cinq tritements (témoin; coupe prtielle d intensités file, modérée et forte; et coupe totle), été étli dns un peuplement ml driné et dominé pr l épinette rouge et le spin umier (Aies lsme (L.) Mill.) de l province de Quéec, Cnd. L réction du développement de l régénértion ux tritements vrié en fonction de l intensité de coupe, de l espèce et de l position des semis pr rpport ux sentiers de dérdge dns les peuplements résiduels. Dix ns près l ppliction des tritements, les semis d épinette rouge étient ien étlis, mis étient plus petits que les semis de spin umier et des espèces cducifoliées. Les semis d épinette rouge étient générlement plus huts dns les sentiers de dérdge des coupes prtielles tout comme l étit l disponiilité de l lumière. Cependnt, dns les coupes totles, l tille des semis d épinette rouge étlis dns les sentiers de dérdge semlit négtivement ffectée pr l quntité considérle de déris ligneux répndus sur et utour d eux. Prmi les tritements de coupe prtielle, l plus forte iomsse foliire de l régénértion d espèces rorescentes été oservée dns l intensité de coupe qui enlevé 6 % de l surfce terrière initile du peuplement. Ce tritement constitue donc un choix intéressnt pour remplcer l coupe totle dns ces peuplements ml drinésoùune remontée de l nppe phrétique est nticipée près l coupe finle. Introduction The shelterwood method is generlly intended to estlish nturl regenertion y grdully removing the overstory through one or more prtil cuttings followed y finl hrvest. This silviculturl system is normlly used where dense cnopy hinders seedling estlishment nd when stnds composed of intermedite to shde-tolernt species re desired. In mixed conifer stnds dominted y red spruce (Pice ruens Srg.) nd lsm fir (Aies lsme (L.) Mill.), the shelterwood method often hs the secondry ojective of promoting regenertion of red spruce t the expense of less desired lsm fir, which is more vulnerle to spruce udworm (Choristoneur fumifern (Clem.)) defolition (McLen nd McKinnon 1997; Solomon et l. 23) nd hs lower commercil vlue thn spruce (Sendk et l. 23). Red spruce seedling estlishment cn e improved y prtil cutting, especilly if, through logging opertions, the humus lyer cn e distured loclly to expose receptive minerl soil, s is the cse for other spruce species (Wurtz nd Zsd 21; Nilsson et l. 22; Prévost nd Pothier 23). Becuse red spruce nd lsm fir re oth very shde tolernt, they cn e present in the regenertion strtum elow dense cnopy, where seedlings survive ut, generlly, sty smll. These smll seedlings re commonly rooted in the upper prt of the orgnic lyer, which dries out fter cler-cutting or the finl hrvest of the shelterwood method, sitution tht often results in extensive seedling mortlity (Jlnczy 1969; Örlnder nd Krlsson 2; Nilsson et l. 22). Red spruce is lso prticulrly sensitive to full light Received 13 Decemer 26. Accepted 11 July 27. Pulished on the NRC Reserch Press We site t cjfr.nrc.c on 17 Jnury 28. D. Pothier. 1 Déprtement des sciences du ois et de l forêt, Université Lvl, Quéec, QC G1K 7P4, Cnd. M. Prévost. Ministère des Ressources nturelles et de l Fune, Forêt Quéec, Direction de l recherche forestière, 27, rue Einstein, Quéec, QC G1P 3W8, Cnd. 1 Corresponding uthor (e-mil: dvid.pothier@sf.ulvl.c). Cn. J. For. Res. 38: (28) doi:1.1139/x7-135 # 28 NRC Cnd

2 32 Cn. J. For. Res. Vol. 38, 28 Tle 1. Men merchntle sl re of smple plots immeditely efore nd fter tretment ppliction. Cutting intensity Control Low Moderte Hevy Totl Bsl re efore tretment (m 2 /h) Red spruce 2.2± ± ± ± ±14.5 Blsm fir 1.8± ± ± ± ±1.4 Other conifer species 6.± ± ± ± ±13. Deciduous species 5.4± ± ± ± ±4.9 Totl 42.4± ± ± ± ±7.1 Bsl re fter tretment (m 2 /h) Red spruce 2.2± ± ± ±2.5.±. Blsm fir 1.8± ± ±.9 2.3±2.8.±. Other conifer species 6.± ±3.2.7±1.4.8±1.5.±. Deciduous species 5.4±4.7.2±.6.±..4±.5.±. Totl 42.4± ± ± ±1.4.±. Note: Bsl re per hectre ws clculted for trees tht hd dimeter t rest height >9 cm. Ech men nd stndrd devition ws clculted for four 2 m 2 m plots. Cutting intensity trgets for removl of merchntle sl re re s follows: control, %; low, 15% ; moderte, 3%; hevy, 4%; totl (clercut), 1%. nd to temperture extremes t the juvenile stge (Dumis nd Prévost 27). Moreover, seedlings of oth these species tend to die from the rise in the wter tle tht cn occur on cler-cut wetlnds (Roy et l. 2). Hence, the shelterwood method cn lso hve the ojective of developing tree seedlings until they hve reched minimum height under the moderte microclimte of the sheltering overstory to ensure tht they cn survive the sudden chnges in environmentl conditions cused y finl hrvest. This minimum height ws clculted to e t 2 cm y Frnk nd Bjorkom (1973) nd Bldwin (1977) for red, white (Pice gluc (Moench) Voss), nd lck (Pice mrin (Mill.) BSP) spruces nd lsm fir in northestern North Americ, wheres Sikström nd Glöde (2) considered seedling height etween 1 nd 2 m necessry to minimize Norwy spruce (Pice ies (L.) Krst.) mortlity in Sweden. Thus, on mny sites, the shelterwood method is used not only for estlishing new seedlings of the desired species, ut lso for cclimting dvnce regenertion to incresed light intensity nd for stimulting their growth so s to minimize mortlity fter the finl hrvest. Moreover, the development of this regenertion should uild up sustntil lef iomss tht would decrese wter tle rise fter the finl cut y incresing site evpotrnspirtion (Pothier et l. 23). Accordingly, the ojectives of this study re to determine the seed cut intensity tht (i) promotes estlishment of red spruce seedlings, (ii) reduces development of rodleved species, (iii) stimultes dimeter growth of residul stnding trees, nd (iv) mitigtes wter tle rise fter finl hrvest. These ojectives will e chieved y nlysing the results of n experiment tht includes four repetitions of five prtil cutting intensities. The results on wter tle rise during the first 5 yers following prtil cutting re lredy ville (Pothier et l. 23). This pper dels with the 1 yer survey of regenertion estlishment nd development nd of shelter tree growth response. Mterils nd methods Study re The 3 h stnd is locted in the estern prt of the sugr mple (Acer scchrum Mrsh.) sswood (Tili mericn L.) ioclimtic domin known s St. Lwrence Lowlnd ecologicl region 2 (Sucier et l. 1998), ner Villeroy (4683 N, W), Queec, Cnd. The study re is chrcterized y flt topogrphy, men nnul precipittion of 183 mm (837 mm of which flls s rin), nd men nnul temperture of 4.5 8C. The growing seson typiclly extends from the eginning of My to the end of Septemer, during which rinfll verges 51 mm nd men ir temperture reches C. The soil is gleyed, humo-ferric podzol chrcterized y the presence of compcted lyer t depth of 8 1 cm, which is responsile for the poor dringe of the site. The orgnic lyer is out 1 cm thick, nd roots of dominnt trees re found minly to depth of 25 cm. The experiment ws estlished in 1994 in stnd dominted y red spruce nd lsm fir. The reminder of the stnd ws composed of estern hemlock (Tsug cndensis (L.) Crr.), estern white cedr (Thuj occidentlis L.), red mple (Acer rurum L.), nd yellow irch (Betul lleghniensis Britt.). The merchntle sl re (BA) of this 8 yer old stnd ws pproximtely 4 m 2 /h (Tle 1), nd its dominnt height ws ner 18 m. The understory ws minly composed of numerous, smll (<3 cm in height) lsm fir seedlings tht were estlished on n lmost continuous crpet of Sphgnum spp. nd fethermoss (mostly Pleurozium schreeri (Brid.) Mitt. nd Hylocomium splendens (Hedw.) BSG). Other species were sprse in the understory nd included red mple, Coptis groenlndic (Oeder) Fern., Oxlis montn Rf., nd Dennstedti punctiloul (Michx.) Moore. In 1994, complete, rndomized lock design with four repetitions (locks) ws estlished in the stnd (Fig. 1). In ech lock, five 5 m 5 m experimentl units (EUs) were delimited, nd ech received one of the following tretments: n uncut control; three prtil-cutting levels tht imed to remove 15% (low), 3% (moderte), or 45% (hevy) of the merchntle BA; nd clercut (removl of 1% of the merchntle BA). Becuse we tried to control the composition, the BA, nd the men tree size mong EUs of ech lock, it ws not possile to use EUs >.25 h, even # 28 NRC Cnd

3 Pothier nd Prévost 33 Fig. 1. Lyout of experimentl units (EUs) in the complete experimentl lock design. For ech EU, the uppercse letter refers to the cutting intensity (C, control; L, low prtil cutting; M, moderte prtil cutting; H, hevy prtil cutting; nd T, totl cutting), nd the numer is the repetition (lock). C1 L2 T2 C2 T1 5 m x 5 m experimentl unit 2 m x 2 m centrl plot M1 H2 H1 L1 M2 H3 M3 L3 N min rod T3 C3 M4 H4 C4 L4 T4 4 m 2 regenertion plot skidding trils though opertionl hrvested locks ner the study re covered pproximtely 1 h. This cn limit the scope of the results, especilly for cler-cut EUs, which were expected to produce higher wtering-up thn ws mesured (Pothier et l. 23). In the prtil-cut tretments, trees to e removed were mrked ccording to low thinning prescription (i.e., the lower crown clsses were mrked first). These tretments were pplied in Mrch 1995 to tke dvntge of the presence of 1 m thick snow cover tht protected the soil nd dvnce regenertion from mchine trffic nd other potentil hrvest-relted dmge. The hrvesting system ws composed of hrvester nd forwrder tht were restricted to 4.6 m wide trils (Fig. 1), which corresponded to pproximtely 25% of the stnd re (the strips etween skidding trils were 13.6 m wide). Trees were delimed on site, nd logging deris ws plced on the skidding trils for the most prt. Becuse prescried prtil cuttings were defined without tking into ccount the nrrow, cler-cut skidding trils, the 15%, 3%, nd 45% prtil cuts resulted in the ctul removl of out 4%, 5%, nd 6% of the BA, respectively (Tle 1). Tree inventory A forest inventory ws conducted immeditely fter hrvesting during spring 1995, t which time ech tree ws numered. This inventory ws repeted during fll 1999 nd gin in fll 24. This inventory consisted in mesuring the dimeter t rest height (DBH) (±1 mm) of ll trees (>1.1 cm) of ech species in 2 m 2 m plots locted in the centrl prt of ech of the 2 EUs (Fig. 1). Thus, these 2 m 2 m centrl plots were surrounded y 15 m uffer strips tht isolted these plots from the influence of other tretments pplied in djcent EUs. Moreover, five dominnt trees per plot were selected to determine their totl height. From the repeted mesurements of ech numered tree, we were le to clculte their dimeter increment for 5 nd 1 yer periods s well s to determine tree mortlity. Regenertion inventory A regenertion inventory ws performed t the end of every growing seson from 1995 to 1999 nd then in 21 nd 24 (yer 1). This inventory only took into ccount smll regenertion, i.e., <1.3 m in 1995, composed of commercil species nd ws crried out y counting the seedlings per species nd height clsses in 16 permnent, circulr, 4 m 2 plots systemticlly estlished in ech centrl plot of the 2 EUs (5 tretments 4 locks, 32 plots totl; Fig. 1). Twelve of these sixteen 4 m 2 plots were locted elow the shelter trees, nd the four others were plced in skidding trils. This regenertion inventory excluded smll trees tht were>1.3 m in 1995, ecuse they were lredy mesured in the tree inventory. However, the regenertion inventory took into ccount seedlings tht were <1.3 m in 1995 ut tht hd reched this height in the following yers. Seedling height clsses were s follows: (i) 3 cm; (ii) 31 1 cm; nd (iii) 11 4 cm. The medin of these height clsses ws used to clculte the dimeter of the seedlings, with the liner regressions pulished y Roussopoulos nd Loomis (1979). Then, this clculted dimeter ws used to clculte regenertion lef iomss ccording to the species-specific power equtions developed y Roussopoulos nd Loomis (1979). For lsm fir, red spruce, nd red mple, these equtions ll hd R 2 vlues >.9. Moreover, within ech plot, we clculted the men height nd the percentge of ground covered y woody deris left fter logging s well s the regenertion stocking, which ws de- # 28 NRC Cnd

4 34 Cn. J. For. Res. Vol. 38, 28 Tle 2. Proility vlues for red spruce, lsm fir, nd deciduous species of different height clsses ssocited with fctors nd interctions sumitted to mixed model of covrince nlysis tht took into ccount irregulr time intervls etween repeted mesurements of regenertion stocking. Red spruce Blsm fir Deciduous species Source of vrition > cm >3 cm >1 cm > cm >3 cm >1 cm > cm >3 cm >1 cm Covrile < < Cutting intensity (CI) Position (P) Yer (Y).1 <.1 <.1.91 <.1 <.1.41 <.1 <.1 CI P < CI Y.2694 <.1 <.1 < < <.1 P Y CI P Y < Note: The tree height clsses of >, >3 nd >1 correspond to stocking of seedlings 3, 31 1, nd 11 4 cm, respectively. Position refers to loction of regenertion plots (etween or in skidding trils). fined s the percentge of the 4 m 2 plots with t lest one live seedlings of the considered species. Light vilility In July 1997, the proportion of incident solr irrdince tht reched the seedlings ws clculted using hemisphericl photogrphy. A self-levelled Nikon cmer mounted on monopod nd equipped with Sigm 8 mm fish-eye lens ws plced just ove the seedlings in ech of the 32 regenertion plots, nd photogrphs were tken using color slide film. The processed slides were scnned with Nikon LS-2 35 slide scnner, nd the digitized imges were nlyzed using the gp light nlyser softwre developed y Frzer et l. (1999). This softwre clculted the mount of direct plus diffuse rdition elow the cnopy for growing seson tht ws specified to extend from mid-my to mid- Septemer. This index, expressed s percentge of the rdition found ove the cnopy, ws individully clculted for the 32 digitized imges. For ech EU, percentge of rdition ws verged for regenertion plots locted under the shelter trees (12 plots) nd those locted in the skidding trils (4 plots). Sttisticl nlyses Regenertion dt (stocking nd numer of seedlings per hectre) were sumitted to mixed model of covrince pplied to completely rndomized lock design with repeted mesurements using the MIXED procedure of the SAS system (SAS Institute Inc., Cry, N.C.). The fixed effects used in these nlyses were the yers of inventory, the cutting tretments, the positions of the regenertion plots reltive to the skidding trils, nd the interctions mong these fctors. The locks nd the interction etween the locks nd the cutting tretments were used s rndom effects. Covriles sumitted to these nlyses corresponded to the stocking or the numer of seedlings per hectre mesured efore the ppliction of the cutting tretments when the independent vriles were stocking nd numer of seedlings fter tretment, respectively. Moreover, power covrince structure, with time intervl etween two consecutive inventories s the power, ws used in the model to tke into ccount the irregulr time intervls etween repeted mesurements of ech regenertion plot (Moser 24). For tree dt, the 1 yer dimeter growth ws sumitted to n nlysis of covrince for completely rndomized lock design using the generl liner models (GLM) procedure of the SAS system (SAS Institute Inc., Cry, N.C.). The covrile used in this nlysis ws tree dimeter mesured t yer, nd the clssifiction vriles were the locks nd the cutting tretments. This nlysis llowed us to detect significnt effect of the interction etween cutting tretments nd tree dimeter t the eginning of the experiment. To help understnd this significnt interction nd to fcilitte presenttion of the results, we grouped tree dimeter t the eginning of the experiment into four clsses: (1) DBH < 1 cm, (2) 1 cm DBH < 2 cm nd (3) 2 cm DBH < 3 cm, nd (4) DBH 3 cm. These DBH clsses, together with the cutting tretments, were then sumitted to n nlysis of vrince for completely rndomized lock design nd Tukey multiple comprison test ws performed to detect differences mong the levels of the significnt interction etween DBH clsses nd cutting tretments. Solr rdition elow the cnopy nd percentge of ground covered y woody deris were oth sumitted to n nlysis of vrince for completely rndomized lock design tht ws followed y Tukey multiple comprison test to determine sttisticl differences mong the levels of the interction etween cutting tretments nd position reltive to the skidding trils. Results Dmge to dvnce regenertion cused y winter hrvesting ws very limited in ll cutting tretments, except in skidding trils of totl cler-cutting. Immeditely fter cutting (1995), this dmge decresed stocking of red spruce nd lsm fir (the two min species tht composed dvnce regenertion) y pproximtely 15% in totl cler-cuttings (Fig. 2). Anlyses of vrince indicte tht some fctors significntly explined the vrition in regenertion stocking of red spruce, lsm fir, nd deciduous species (Tle 2). However, these significnt effects differed mong species depending on the regenertion height clss considered. Hence, when red spruce stocking of ll height clsses ws considered, no differences were oserved mong cutting intensities (Tle 2, height clss >), which were ll ssocited with stocking vlues >7% for ech mesurement period (Fig. 2). The significnt interction etween yer nd regenertion plot position clculted for red spruce stocking # 28 NRC Cnd

5 Pothier nd Prévost 35 Fig. 2. Chnges in men regenertion stocking over time (n =4 locks) for red spruce, lsm fir, nd deciduous species fter ppliction of five cutting tretments: untreted control (C); low (L), moderte (M), nd hevy (H) prtil cuttings; nd totl cler-cutting (T). Regenertion stocking ws clculted for the three seedling height clsses defined in Tle 1. Stock ing (%) Stock ing (%) Stock ing (%) seedling height > cm seedling height > 3 cm seedling height > 1 cm Red spruce C L M H T C L M H T C L M H T Blsm fir C L M H T C L M H T C L M H T Deciduous species C L M H T C L M H T C L M H T of ll height clsses cn e explined y the grdul increse over time in the skidding trils, wheres stocking etween the trils remined constnt ner 1% during the entire study period (dt not shown). For red spruce seedlings >3 cm, the pttern of chnge in stocking over time differed mong tretments s indicted y the significnt interction etween cutting intensity nd yer (Tle 2). In 1999 (i.e., 5 yers fter tretment), red spruce regenertion stocking ws proportionl to cutting intensity, wheres the control ws the only tretment tht significntly differed from the others fter 1 yers (24) (Fig. 2). In the cse of red spruce regenertion >1 cm, tretment effects were only pprent fter 1 yers, t which time stocking ws proportionl to cutting intensity (Fig. 2). Ten yers fter tretment, the results on the numer of red spruce seedlings per hectre (Tle 3) followed the sme proportionl response to tretment s tht found for stocking (Fig. 2). For lsm fir nd deciduous species, cutting tretment effects were similr to those oserved with red spruce, except tht height development ws much fster s indicted y stocking vlues of lsm fir nd rodleved seedlings >1 cm (Fig. 2). The threefold interction occurring mong cutting intensity, position, nd time ws significnt for the stocking of red spruce seedlings >1 cm (Tle 2). This mens tht red spruce stocking vlues etween nd in skidding trils followed different trends for t lest one tretment during 1 yer. Such result is well illustrted y red spruce regenertion estlished 1 yers fter cutting etween nd in skidding trils (Fig. 3). In this figure, stocking of red spruce seedlings >1 cm for the three prtil cuttings ws higher in skidding trils thn etween them, wheres the opposite ws true in the cse of cler-cutting. Seedling environment differed ccording to the cutting tretments nd to their position reltive to the skidding trils. Hence, the vilility of solr rdition, s clculted through nlyses of hemisphericl photogrphs tken over ech regenertion plot, ws greter in skidding trils thn etween them for the three prtil cut tretments, wheres no differences were oserved in totl cler-cuttings (Tle 4). However, the undnce of woody deris, s estimted y percentge of ground coverge, ws greter in skidding trils of ll cutting tretments (Tle 5). The difference etween the two positions reltive to the skidding trils (etween nd in skidding trils) ws prticulrly importnt in totl cler-cuttings (Tle 5), where the hrvested volume ws more importnt thn in prtil cuts. An nlysis of vrince concerning the growth dt of residul trees indictes tht the interction etween cutting tretment nd dimeter of residul trees efore tretment ws significnt 1 yers fter cutting. This result mens tht the effect of t lest one cutting tretment on dimeter growth differed mong initil tree dimeters. Multiple-comprison testing indictes tht it is mostly smll-dimeter trees tht enefit from prtil cuttings, s cn e seen from the effects of these tretments reltive to the controls (Fig. 4). Moreover, the highest prtil-cutting intensity tended to produce the est improvement in dimeter growth for ll dimeter clsses (Fig. 4). The numer nd size of tree seedlings llowed us to clculte seedling lef iomss estlished fter tretment. For ll tretments, regenertion lef iomss ws for the most prt composed of lsm fir (*7%), red spruce (*2%), nd red mple (*1%). In every tretment except the control, regenertion lef iomss ccumulted slowly during the first 5 yers following cutting nd then shrply incresed until the end of the 1 yer survey period (Fig. 5). At this time, regenertion lef iomss ws proportionl to cutting intensity nd rnged from out 55 kg/h in the control tretment to 7 kg/h in the clercuts (Fig. 5). Discussion Red spruce lsm fir stnd regenertion is lrgely dependent on dvnce growth. Compred with red spruce, l- # 28 NRC Cnd

6 36 Cn. J. For. Res. Vol. 38, 28 Tle 3. Men numer of seedlings per hectre 1 yers fter tretment pplictions for red spruce, lsm fir, nd deciduous species of three height clsses. Red spruce seedlings/h Blsm fir seedlings/h Deciduous species seedlings/h Cutting intensity > cm >3 cm >1 cm > cm >3 cm >1 cm > cm >3 cm >1 cm Control Low Moderte Hevy Totl Note: Vlues re mens of the four locks for ech tretment. See Tle 1 for seedling height clsses. Fig. 3. Men red spruce regenertion stocking (n = 4 locks) 1 yers fter ppliction of five cutting intensities for plots estlished etween skidding trils (B) nd in skidding trils (I). Regenertion stocking ws clculted for the three seedling height clsses defined in Tle 1. There were no skidding trils in the controls; hence only one vlue is shown. Stocking (%) seedling height > cm seedling height > 3 cm seedling height > 1 cm B I B I B I B I B I Control Low Moderte Hevy Totl sm fir produces good seed crops more frequently (Seymour 1995), is etter dpted to vriety of seededs (Simrd et l. 1998), nd hs more extensive root development (Plce 1955); consequently, it often domintes the regenertion strtum (Frnk nd Bjorkom 1973). This sitution is generlly not welcomed y forest mngers, ecuse lsm fir is shorter lived, is less resistnt to spruce udworm nd decy, nd hs lower commercil vlue thn red spruce (Sendk et l. 23). Hence, silviculturl tretments, such s shelterwood cutting nd scrifiction, hve een proposed to improve seeded conditions nd growing environment of red spruce regenertion (Frnk nd Bjorkom 1973, Bldwin 1977, Sendk et l. 23). However, in this study, the estlishment of red spruce seedlings ws not prolem, ecuse stocking of this species ws present t levels >7% in ll tretments s well s the untreted control t the eginning of the experiment (Fig. 2). Thus, the principl gol of the shelterwood method ws to improve the growing conditions of these smll red spruce seedlings to increse the proility of survivl fter finl hrvest. Relevnt informtion out red spruce seedling development cn e drwn from the pttern of chnge in stocking over time for regenertion of different height clsses (Fig. 2; Tle 3). Five yers fter cutting, stocking of red spruce seedlings >3 cm ws proportionl to cutting intensity, with 5% vlue for the heviest prtil cutting. Ten yers fter tretments, ll prtil-cutting intensities hve reched the generlly ccepted 6% stocking vlue for red spruce seedlings >3 cm nd, thus, could e considered s hving well regenerted. However, oth deciduous species nd lsm fir were lso well estlished, with seedlings eing generlly tller thn those of red spruce (Fig. 2; Tle 3). Vigorous red mple sprouts, in prticulr, contriuted to the greter stocking vlues of deciduous species >1 cm, 1 yers fter cutting. Therefore, if forest mngers wnt to mintin the spruce proportion s it ws in the preceding mture stnd, precommercil thinning should e plnned over the medium term to djust future stnd composition. However, to mke precommercil thinning successful on these lowlnd sites the lef iomss of the remining trees must e sufficient to void wtering-up, which could e detrimentl to tree survivl. The present shelterwood experiment ws performed following oservtions of high seedling mortlity in djcent 1 h cler-cut locks. This regenertion mortlity ws presumed to hve resulted from wtering-up, which forced forest mngers to prescrie costly tretments such s dringe nd plnting. However, in our experiment s.25-h clercut EUs, seedlings of ll species hd low mortlity rtes nd were le to use ville resources efficiently, such tht their numers nd growth levels were higher thn those of the other tretments (Fig. 2; Tle 3). The low mortlity rte oserved in these.25 h cler-cut EUs most likely stemmed from restricted wtering-up cused y low rinfll during the first yer fter tretment, from the smll size of the cler-cut EUs, nd from the presence of uncut forest round the experimentl re (Pothier et l. 23). Becuse of their smll size, cler-cut EUs could inccurtely represent opertionl cler-cut res, which re much lrger nd, thus, more prone to wtering-up nd relted dmge to regenertion. Height development of red spruce seedlings ws different etween the two positions reltive to the skidding trils, s is demonstrted y the stocking vlues for different height clsses 1 yers fter tretment (Fig. 3). In the three prtilcut tretments, red spruce seedlings estlished in the skidding trils tended to e tller thn those under the shelter trees (Fig. 3). This result cn e explined y the difference in the vilility of solr rdition etween these two environments. Indeed, the percentge of solr rdition, clculted using hemisphericl photogrphs, ws systemticlly 4% higher in the skidding trils of ech prtil-cut tretment (Tle 4) thn etween the trils. # 28 NRC Cnd

7 Pothier nd Prévost 37 Tle 4. Amount of direct nd diffuse solr rdition elow the cnopy, expressed s percentge of rdition received ove the cnopy, clculted using nlyses of hemisphericl photogrphs tken ove 16 regenertion plots in ech experimentl unit (12 etween skidding trils nd 4 in skidding trils except in the control tretment, where ll 16 plots were locted etween skidding trils). Cutting intensity Position Control Low Moderte Hevy Totl Rdition etween skidding trils (%) 27.±6.3f 42.2±6.9e 47.2±9.4d 5.6±1.8c 88.±6.8 Rdition in skidding trils (%) 46.5±3.5d 51.1±5.c 54.3± ±2.6 Note: Vlues re mens ± SDs clculted from the mens of the four locks. Vlues followed y different letters indicte significnt differences ( =.5) ccording to Tukey multiple comprison test. Tle 5. Percentge of ground covered y woody deris during the summer following cutting for ech cutting tretment nd ech position reltive to the skidding trils. Cutting intensity Position Control Low Moderte Hevy Totl Woody deris etween skidding trils (%).±.c 4.±15.7c 8.5±2.c 2.8±11.1c 6.8±2.7c Woody deris in skidding trils (%) 51.9± ± ± ±34.1 Note: For ech experimentl unit, the percentge of ground covered y woody deris ws clculted for sixteen 4 m 2 plots (12 locted etween skidding trils nd 4 locted in skidding trils). Vlues re mens ± SDs clculted from the mens of the four locks. Vlues followed y different letters indicte significnt differences ( =.5) ccording to Tukey multiple comprison test. According to Vlkonen et l. (22) nd Strnd et l. (26), the greter growth of Scots pine (Pinus sylvestris L.), Norwy spruce, nd lodgepole pine (Pinus contort Dougl. vr. ltifoli Engelm.) seedlings locted fr from the shelter trees is etter explined y decresing nutrient competition thn y incresing light level. However, if nutrient vilility ws more importnt thn light vilility, different tll seedling stocking vlues would hve een oserved for seedlings exposed to the sme light level ut tht hd een estlished t vrious distnces from shelter trees. This cn e verified y compring, on the one hnd, the stocking levels of tll seedlings estlished in the skidding trils ssocited with low prtil cutting nd, thus, tht re reltively fr from shelter trees nd, on the other hnd, the stocking levels of tll seedlings estlished elow the shelter trees of the moderte prtil cutting nd, thus, tht re reltively close to the shelter trees; oth sorts of seedlings were exposed to out 47% of solr rdition (Tle 4). However, red spruce seedling stocking levels were the sme for oth situtions, i.e., pproximtely 6% for seedlings tller thn 3 cm nd 6% for seedlings >1 cm (Fig. 3). Slightly different results cn e oserved in comprison of the stocking levels otined, on the one hnd, for tll seedlings in the skidding trils ssocited with moderte prtil cutting nd, on the other hnd, for tll seedlings under the shelter trees ssocited with hevy prtil cutting; in oth instnces, the seedlings were exposed to out 51% of solr rdition (Tle 4). However, lthough the red spruce stocking level ws similr for seedlings >3 cm (pproximtely 75%), it differs for seedlings >1 cm: 25% for seedlings in the skidding trils of the moderte prtil cuttings nd 8% for seedlings etween the skidding trils of the hevy prtil cuttings (Fig. 2). Consequently, it is thus possile tht light nd nutrients interct to explin differences in height growth of red spruce seedlings estlished in different loctions reltive to the skidding trils. A more detiled study, including ccurte mesurements of light nd nutrient vilility, is needed to quntify the impct of ech fctor in these lowlnd environments. In the cse of the cler-cut tretment, red spruce stocking results show trend opposite to those oserved in prtil cuttings, ecuse tll seedlings were more frequent etween the skidding trils thn in the skidding trils (Fig. 3). This cn e relted to the percentge of ground covered y logging slsh, which ws much greter in the skidding trils of cler-cut EUs thn in those of prtil cuttings (Tle 5). Furthermore, we oserved tht logging slsh tended to e thicker nd more compcted in totl cuttings thn in prtil cuttings. Hence, it seems tht logging slsh in skidding trils of cler-cut EUs ws undnt enough to deteriorte seeded conditions, to dely red spruce estlishment, nd (or) to shde smll seedlings with the consequence of reducing their survivl nd their height growth. The prtil-cut tretments not only ffect seedling growth ut lso the dimeter growth of residul trees, prticulrly smll-dimeter trees (Fig. 4). This result supports the oservtion tht smll-dimeter trees respond to vrious silviculturl tretments fster nd with greter mgnitudes thn lrge-dimeter trees, (e.g., Shen et l. 2; Del nd Tppeiner 22; Mäkinen nd Isomäki 24; Prévost et l. 25). The prtil cuts were pplied very crefully y experienced hrvester nd forwrder opertors, so tht very few injuries were oserved on residul trees. This certinly contriutes to the slightly etter dimeter growth of prtil cut tretments over controls for lrger dimeter clsses, lthough most of these differences were not sttisticlly significnt. In this regrd, the hevy prtil cutting, which removed 6% of the initil sl re, ws ssocited with the lrger dimeter increment mong ll initil dimeter clsses (Fig. 4), which suggests tht this cutting intensity must e considered y forest mngers on low windthrow hzrd res. The residul trees left on res suject to prtil cutting were responsile for mitigting wter tle rise during the first 5 yers fter tretment (Pothier et l. 23). During # 28 NRC Cnd

8 38 Cn. J. For. Res. Vol. 38, 28 Fig. 4. Ten yer dimeter growth increment of trees (ll species) remining fter four cutting intensities: untreted control (C) nd low (L), moderte (M), nd hevy (H) prtil cuttings. Trees were grouped in four dimeter t rest height (DBH) clsses. Within ech DBH clss, the different letters indicte significnt differences ( =.5) mong tretment mens. Tretment mens (rs) nd SEs (error rs) were clculted on the sis of the men vlues otined from the four repetitions (locks). 1 yer dimeter increment (cm) C L M H C L M H C L M H C L M H DBH<1 1 DBH<2 2 DBH<3 DBH 3 Fig. 5. Chnges in men lef iomss over time for regenertion of ll tree species fter ppliction of five cutting tretments. Lef iomss ws clculted ccording to the species-specific equtions developed y Roussopoulos nd Loomis (1979). Lef iomss (kg/h) totl cutting hevy cutting moderte cutting low cutting control Yer this period, the lef re supported y these trees ws the lrgest source of evpotrnspirtion in the prtil-cut tretments nd the control, wheres cler-cutting, in which very few smll trees were left stnding, resulted in the lrgest wtering-up (Pothier et l. 23). However, ecuse the finl step in shelterwood method is creful cler-cutting round dvnce growth, vigorous nd undnt regenertion strtum must e estlished to mitigte, in turn, the nticipted wtering-up following logging. Therefore, in the sence of significnt windthrow, finl hrvest ws delyed until sufficient lef iomss ccumulted in the regenertion strtum. This lef iomss ccumultion ws prticulrly sustntil during the second 5 yer period fter tretment (Fig. 5), which justifies the decision to dely the finl hrvest commonly pplied s soon s the desired regenertion is well estlished, i.e., 5 yers fter tretment (Fig. 2). Ongoing monitoring of the wter tle following finl hrvest will help quntify the impct of this 5 yer dely on the nticipted wter tle rise. Ten yers fter tretment, tree regenertion lef iomss ws proportionl to cutting intensity, nd logiclly enough, the hevy prtil cutting tht removed out 6% of the initil sl re produced the highest regenertion lef iomss mong prtil-cut tretments (Fig. 5). Thus, s result, hevy prtil cutting presents the potentil dvntge of helping to mitigte wtering-up following finl hrvest. Moreover, ecuse the trees remining fter hevy prtil cutting hd sufficient lef re to control the wter tle rise immeditely fter tretment (Pothier et l. 23), the shelterwood method using this first-cut intensity is n ttrctive lterntive to cler-cutting to chieve forest-mngement ojectives in this type of lowlnd conifer stnd. Acknowledgements We thnk Michel Beulieu, Alin Lnglois, Gilles Audy, Louis Fucher, Dniel Plourde, Jolène Lemieux, nd mny summer students for their help in the field. We lso thnk Sylvin Turis for his work nlyzing hemisphericl photogrphs nd Donld Kellough nd three nonymous reviewers for helpful comments on the mnuscript. References Bldwin, V.C Regenertion following shelterwood cutting in New Brunswick softwood stnd. Cn. For. Serv. Mritimes For. Res. Cent. Inf. Rep. M-X-76. Del, R.L., nd Tppeiner, J.C. 22. The effects of prtil cutting on stnd structure nd growth of western hemlock Sitk spruce stnds in southest Alsk. For. Ecol. Mnge. 159: doi:1.116/s () Dumis, D., nd Prévost, M. 27. Red spruce conservtion in mnged forests of Queec: importnce of some physiologicl nd ecologicl chrcteristics review. For. Chron. 83: Frnk, R.M., nd Bjorkom, J.C A silviculturl guide for spruce fir in the Northest. USDA For. Serv. Gen. Tech. Rep. NE-6. Frzer, G.W., Cnhm, C.D., nd Lertzmn, K.P Gp nlyser (GLA): imging softwre to extrct cnopy structure nd gp light trnsmission indices from true-colour fisheye photogrphs, users mnul nd progrm documenttion. Simon Frser University, Burny, British Columi, nd the Institute of Ecosystem Studies, Millrook, N.Y. Jlnczy, A Estlishment of white spruce nd lsm fir in shelterwood fellings of field spruce in Nov Scoti. Deprtment of Fisheries nd Forestry, Cndin Forest Service, Ottw, Ont. Int. Rep. M-48. McLen, D.A., nd McKinnon, W.E Effects of stnd nd site chrcteristics on susceptiility nd vulnerility of lsm fir nd spruce to spruce udworm in New Brunswick. Cn. J. For. Res. 27: doi:1.1139/cjfr Mäkinen, H., nd Isomäki, A. 24. Thinning intensity nd longterm chnges in increment nd stem form of Norwy spruce trees. For. Ecol. Mnge. 21: doi:1.116/j.foreco Moser, E.B. 24. Repeted mesures modeling with PROC MIXED. In Proceedings of the 29th SAS Users Group Intern- # 28 NRC Cnd

9 Pothier nd Prévost 39 tionl Conference, 9 12 My 24, Montrel, Que. SAS Institute Inc., Cry, N.C. Pp Nilsson, U., Gemmel, P., Johnsson, U., Krlsson, M., nd Welnder, T. 22. Nturl regenertion of Norwy spruce, Scots pine nd irch under Norwy spruce shelterwoods of vrying densities on mesic-dry site in southern Sweden. For. Ecol. Mnge. 161: doi:1.116/s (1) Örlnder, G., nd Krlsson, C. 2. Influence of shelterwood density on survivl nd height increment of Pice ies dvnce growth. Scnd. J. For. Res. 15: Plce, I.C.M The influence of seed-ed conditions on the regenertion of spruce nd lsm fir. Cndin Deprtment of Northen Affirs nd Nturl Resources, Forestry Brnch, Ottw, Ont. For. Res. Div. Bull Pothier, D., Prévost, M., nd Auger, I. 23. Using the shelterwood method to mitigte wter tle rise fter forest hrvesting. For. Ecol. Mnge. 179: doi:1.116/s (2) Prévost, M., nd Pothier, D. 23. Prtil cuts in tremling spen conifer stnd: effects on microenvironmentl conditions nd regenertion dynmics. Cn. J. For. Res. 33: doi: /x Prévost, M., Dumis, D., nd Plmondon, A.P. 25. Effets du dringe sur l croissnce et le sttut nutritif d un peuplement d épinette noire de structure inéquienne: résultts de 1 ns. For. Chron. 81: Roussopoulos, P.J., nd Loomis, R.M Weights nd dimensionl properties of shrus nd smll trees of the Gret Lkes conifer forest. USDA For. Serv. North Cent. For. Exp. Stn. Res. Pp. NC-178. Roy, V., Ruel, J.-C., nd Plmondon, A.P. 2. Estlishment, growth nd survivl of nturl regenertion fter clercutting nd dringe on forested wetlnds. For. Ecol. Mnge. 129: doi:1.116/s (99)17-x. Sucier, J.-P., Bergeron, J.-F., Grondin, P., nd Roitille, A Les régions écologiques du Quéec méridionl (3e version) : un des elements du système hiérrchique de clssifiction écologique du territoire mis u point pr le ministère des Ressources nturelles du Quéec. L Ordre des Ingénieurs forestiers du Quéec, Quéec, Que. Auelle 124(Suppl.) Sendk, P.E., Brissette, J.C., nd Frnk, R.M. 23. Silviculture ffects composition, growth, nd yield in mixed northern conifers: 4-yer results from the Penoscot Experimentl Forest. Cn. J. For. Res. 33: doi:1.1139/x3-14. Seymour, R.S The Northestern region. In Regionl silviculture of the United Sttes. 3rd ed. Edited y J.W. Brrett. John Wiley & Sons, New York. pp Shen, G., Moore, J.A., nd Htch, C.R. 2. The effect of hitt type nd rock type on individul tree sl re growth response to nitrogen fertiliztion. Cn. J. For. Res. 3: doi: /cjfr Sikström, U., nd Glöde, D. 2. Dmge to Pice ies regenertion fter finl cutting of shelterwood with single- nd doule-grip hrvester systems. Scnd. J. For. Res. 15: Simrd, M.-J., Bergeron, Y., nd Sirois, L Conifer seedling recruitment in southestern Cndin orel forest: the importnce of sustrte. J. Veg. Sci. 9: doi:1.237/ Solomon, D.S., Zhng, L., Brnn, T.B., nd Lrrick, D.S. 23. Mortlity ptterns following spruce udworm infesttion in unprotected spruce fir forests in Mine. North. J. Appl. For. 2: Strnd, M., Löfvenius, M.O., Bergsten, U., Lundmrk, T., nd Rosvll, O. 26. Height growth of plnted conifer seedlings in reltion to solr rdition nd position in Scots pine shelterwood. For. Ecol. Mnge. 224: doi:1.116/j.foreco Vlkonen, S., Ruusk, J., nd Siipilehto, J. 22. Effect of retined trees on the development of young Scots pine stnds in southern Finlnd. For. Ecol. Mnge. 166: doi:1.116/s (1) Wurtz, T.L., nd Zsd, J.C. 21. An lterntive to cler-cutting in the orel forest of Alsk: 27-yer study of regenertion fter shelterwood hrvesting. Cn. J. For. Res. 31: doi:1.1139/cjfr # 28 NRC Cnd

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