Management for red spruce conservation in Québec: The importance of some physiological and ecological characteristics – A review by Daniel Dumais1,2 and Marcel Prévost1
ABSTRACT The physiological and ecological characteristics of red spruce (Picea rubens Sarg.) are reviewed and integrated into ecosys- tem management options. Red spruce is a shade-tolerant, late-successional conifer species found in the temperate forests of northeastern North America. Its wood, being of excellent quality, is prized by the forest industry. Unfortunately, this high-value species has been in decline throughout its entire range for the past 50 years. At high elevations, in northeast- ern United States, crown dieback caused by the combined effect of atmospheric pollution and climate is largely respon- sible of this decline. In other areas, such as Québec (Canada), the scarcity of red spruce is mainly caused by forest man- agement practices that are poorly adapted to the species’ ecophysiology. Many physiological studies have shown that the species is sensitive to full sunlight (at juvenile and advance growth stages), high temperatures and frost. It also has partic- ular microsite requirements for seed germination and early seedling establishment, such as the presence of large decaying woody debris. Hence, a management strategy adapted for red spruce should favour the use of partial cutting, maintain- ing some overstory and dead wood. This will emulate the natural dynamics of small canopy gaps and minimize the phys- iological stresses to regeneration. The ecophysiological aspects of natural and artificial regeneration of red spruce should be examined with respect to the increased use of partial cutting techniques.
Keys words: advance regeneration, balsam fir (Abies balsamea L.), ecophysiology, ecosystem management, frost suscepti- bility, light response, microenvironment, red spruce (Picea rubens Sarg.), seedling establishment, shade tolerance, silvicul- tural systems, thermosensitivity
RÉSUMÉ Les caractéristiques physiologiques et écologiques de l’épinette rouge (Picea rubens Sarg.) sont révisées et intégrées dans des options d’aménagement écosystémique. L’épinette rouge est un conifère tolérant à l’ombre et de fin de succession que l’on retrouve dans les forêts tempérées du nord-est de l’Amérique du Nord. Son bois, d’excellente qualité, est recherché par l’industrie forestière. Depuis 50 ans, cette essence de grande valeur a malheureusement connu un déclin sur l’ensemble de son aire de répartition. En haute altitude, dans le nord-est des États-Unis, le dépérissement de la cime causé par l’effet combiné de la pollution atmosphérique et du climat est en grande partie responsable de ce déclin. En d’autres endroits, comme au Québec (Canada), la raréfaction de l’épinette rouge est surtout attribuable à des pratiques d’aménagement forestier peu adaptées à son écophysiologie. Plusieurs études physiologiques ont montré que cette espèce est sensible à la pleine lumière (aux stades juvénile et de régénération préétablie), aux températures élevées et au gel. Elle requiert aussi des microsites particuliers, comme les gros débris ligneux en décomposition, pour la germination des graines et l’établissement initial des semis. Ainsi, un aménagement adapté pour l’épinette rouge devrait favoriser l’utilisation de coupes partielles qui maintiennent un certain couvert forestier et des attributs de bois mort, minimisent les stress à la régénération et s’apparentent à la dynamique naturelle des petites trouées. Les aspects écophysiologiques de la régénération naturelle et artificielle d’épinette rouge devraient être examinés en lien avec l’utilisation grandissante des diverses techniques de coupes partielles.
Mots-clés : aménagement écosystémique, microenvironnement, écophysiologie, épinette rouge (Picea rubens Sarg.), établissement des semis, régénération préétablie, réponse à la lumière, sapin baumier (Abies balsamea L.), sensibilité à la chaleur, susceptibilité au gel, systèmes sylvicoles, tolérance à l’ombre
1Ministère des Ressources naturelles et de la Faune, Forêt Québec, Direction de la recherche forestière, Service de la sylviculture et du ren- dement des forêts, 2700, rue Einstein, Québec (Québec) G1P 3W8. 2Corresponding author. E-mail: [email protected]
378 MAI/JUIN 2007, VOL. 83, No 3 — THE FORESTRY CHRONICLE species may also be reduced by exposure to ozone pollution (e.g., Cumming et al. 1988, Fincher et al. 1989, Amundson et al. 1991, Percy et al. 1992, Rebbeck et al. 1993). Although similar diebacks have periodically affected sugar maple (Acer saccharum Marsh.) and white birch (Betula papyrifera Marsh.) in eastern Canada during the last century (McLaughlin 1998, Loo and Ives 2003), red spruce has gener- ally been spared. However, the species is now in the same pre- carious situation in Canada as it is in some northeastern USA low-elevation forests. Crown dieback has not been a major problem in these stands, but restoration efforts, after more than a century of poorly adapted harvesting practices, are a Daniel Dumais Marcel Prévost failure (Pielke 1981, Seymour 1985, Peart et al. 1992). In Québec, red spruce decline has recently been cited as being important to the maintenance of biodiversity in temperate Introduction forests (Grondin and Cimon 2003). Red spruce (Picea rubens Sarg.) (Fig. 1) is a conifer species Many studies related to crown dieback have identified the native to the temperate forests of northeastern North stresses that hinder the physiological processes of this species America. Individuals of this species can live for 400 years and have revealed some of the ecophysiological characteristics (Cogbill 1996, Wu et al. 1999), attaining 26 m in height and that govern its establishment and growth. It was shown that 60 cm in diameter (Adams and Stephenson 1989, White and red spruce is poorly adapted to full sunlight at juvenile and Cogbill 1992, Farrar 1995). Its wood is light, straight-grained, advance growth stages, and that it is vulnerable to high tem- and resilient (Blum 1990) with physico-mechanical proper- peratures and frost, which may account for the widespread ties that are comparable or superior to those of other spruces post-harvest regeneration failures associated with the species. (Panshin and De Zeeuw 1980). The wood is denser than that Other studies revealed that red spruce also has specific of white spruce (Picea glauca [Moench] Voss) (Jessome microsite needs, which are critical for seedling establishment. 1977), and has a lower percentage of less desirable juvenile For example, large decaying woody debris provides microsites wood than does black spruce (Picea mariana [Mill.] BSP) conducive to the survival and growth of young seedlings (Fowler et al. 1988). For these reasons, red spruce is desirable (Perkins 1991, Seymour 1995). to both the forest industry and to specialized industries such This paper highlights some important physiological and as manufacturers of musical instruments. ecological characteristics that govern the natural population Unfortunately, this species has experienced a decline in dynamics of red spruce. We try to integrate these character- abundance and vigour throughout its entire range during the istics into ecosystem management options adapted to this last 50 years. At high elevations in the northeastern USA, valuable forest species. In an effort to assure the long-term crown dieback, growth decline and mortality are associated conservation of red spruce in Québec, we also suggest some with atmospheric pollution and a consistent loss of frost har- future research needs and provide some silvicultural recom- diness (Johnson 1992; McLaughlin and Kohut 1992; mendations. Sheppard et al. 1993a,b; DeHayes et al. 2001; Sheppard and Pfanz 2001). Sulphate and nitrate ions from acid precipitation cause a significant leaching of soil Ca (McLaughlin et al. 1993, Red Spruce in the Forest Landscape Friedland and Miller 1999, Fernandez et al. 2003, Hawley et Red spruce is predominantly a component of the temperate, al. 2006) and membrane-associated Ca (mCa) in the meso- late-successional forests of the Acadian Region (Rowe 1972) phyll cells of red spruce (Joslin et al. 1988, Thornton et al. and the New England States. The species is not found in 1993, Jiang and Jagels 1999, Schaberg et al. 2000a, Borer et al. boreal forest. Its range extends from the Appalachian 2005). In the frost-hardening process, mCa influences solu- Mountains of eastern Canada and USA in the north, to tion movement across membranes and the ability of cells to Pennsylvania in the south (Fowells 1965). This species is also resist dehydration and extracellular ice damage during frost sporadically distributed in the states of Virginia and North acclimation (DeHayes et al. 1997, 2001). Recently, Hawley et Carolina (Little 1971). Small populations of the species can al. (2006) showed the first evidence that losses in soil Ca, also be found in a narrow band that ranges from southeast- caused by factors such as acidic deposition, resulted in lower ern Ontario to the foothills of the Laurentian Mountains in foliar Ca and higher freezing injury in red spruce trees. Québec (Farrar 1995). Moreover, a 10°C reduction in the degree of frost tolerance Red spruce rarely forms pure stands, but associates with and associated increase in foliar injury was reported for both balsam fir (Abies balsamea [L.] Mill.), yellow birch (Betula mature trees and seedlings exposed to acid precipitation alleghaniensis Britt.), white birch, sugar maple, red maple (Vann et al. 1992, Sheppard et al. 1993b). Acid deposition has (Acer rubrum L.) and trembling aspen (Populus tremuloides been shown to inhibit root growth (e.g., Joslin and Wolfe Michx.) to form mixed conifer or conifer–hardwood stands 1992), desensitize stomatal response (e.g., Borer et al. 2005) (Blum 1990, Seymour 1992, Grondin et al. 1996). The red and increase water stress in red spruce (e.g., Eamus et al. spruce–balsam fir cover type is widely distributed in the 1989). Photosynthetic pigments, carbohydrate content, epi- Canadian provinces of New Brunswick and Nova Scotia, and cuticular wax production, frost hardiness and growth of this in the mountains of the coastal northeastern USA.
MAY/JUNE 2007, VOL. 83, No. 3 — THE FORESTRY CHRONICLE 379 Fig. 2. Distribution and relative abundance (%) of red spruce in different Québec bioclimatic domains (A: spruce, B: balsam fir–white birch, C: balsam fir–yellow birch, D: sugar maple–yellow birch, E: sugar maple–basswood, F: sugar maple–bitternut hick- ory). The relative abundance was calculated from the total basal area of red spruce trees in each ecological district using dendro- metric data from the three decennial forest inventory programs and the permanent plots belonging to le Ministère des Ressources naturelles et de la Faune du Québec (MRNFQ). Fig. 1. Left: Crown of mature red spruce (20 m height) in a yel- low birch–conifer stand (Portneuf County, Québec). Right: Trunk (40 cm dbh) of a mature red spruce in a yellow birch–conifer stand (L’Islet County, Québec). (e.g., shade tolerance) (Manley and Ledig 1979), and are under a high selection pressure (Manley 1972, Major et al. 2005). Hybrids mainly appear in disturbed areas (e.g., after In Québec, the balsam fir–red spruce association is mainly clearcutting) where the elimination of late-successional confined to mesic till soils with a high stone content, thin forests increases the juxtaposition of black spruce and red deposits and rocky outcrops (Grondin et al. 1996). Red spruce stands (Major et al. 2005). spruce is a common companion species in the hardwood for- est of southern Québec (sugar maple–basswood [Tilia amer- Important Physiological and Ecological Characteristics icana L.] and sugar maple–yellow birch bioclimatic domains) Light response (Fig. 2). It is also an important component of the mixed- Red spruce is a late-successional species that is adapted to wood forest (balsam fir–yellow birch bioclimatic domain) regeneration in the understory or in small canopy gaps. It is (Robitaille and Saucier 1998, Saucier et al. 2001), which cor- very shade tolerant (Fowells 1965), surpassing black spruce responds to its northern limit. In the mixedwood forest, red (Manley and Ledig 1979, Major et al. 2003a), white spruce spruce is principally a productive component of the yellow (Nienstaedt and Zasada 1990) and even balsam fir on some birch–conifer associations, which colonize mesic to subhydric sites (Boyce 1993). Seedlings only require 10% of full sunlight lower-slope tills (Grondin et al. 1996). for establishment (Place 1955, Pielke 1981), and only 50% of The genetic diversity of red spruce is low in relation to full sunlight for a good height growth (Blum 1990, Seymour commonly associated conifers and other companion species 1995). Shaded seedlings reach a maximum level of photosyn- (Fowler et al. 1988, Hawley and DeHayes 1994, Perron et al. thesis (Amax) at low light levels, compared to other conifers 1995, Rajora et al. 2000). This low diversity could have been (Manley and Ledig 1979, Fowler et al. 1988, Alexander et al. caused by shrinking populations of red spruce following gla- 1995). However, photosynthesis in red spruce is much less cier advances during the Pleistocene, 10 000 years ago efficient in full sunlight, where the risk of photoinhibition (Morgenstern and Farrar 1964), or by the recent arrival of the and photodamage is high (Alexander et al. 1995, Major et al. species, which is thought to have descended from a black 2003a). Hence, individuals have little capacity for photosyn- spruce population that was isolated during a glacial retreat thetic acclimation to a rapid increase in light intensity, which (Perron and Bousquet 2002). Although red spruce and black is typical of late-successional, shade-tolerant tree species spruce can hybridize within their broad zone of contact (Mohammed and Parker 1999). (Morgenstern and Farrar 1964, Manley 1972, Bobola et al. Red spruce is morphologically adapted to heavy shade. 1996, Perron and Bousquet 1997, Major et al. 2005), this phe- Young germinants have cotyledons that are larger and more nomenon is limited by natural barriers. The two species numerous than those of spruces that are adapted to full sun- inhabit different ecological niches (Fowells 1965, Gordon light (e.g., black spruce). This characteristic allows the species 1976, Major et al. 2003a), have ecophysiological differences to better capture weak light (Major et al. 2003b). Moreover,
380 MAI/JUIN 2007, VOL. 83, No 3 — THE FORESTRY CHRONICLE represents a major source of stress following clearcutting. In a natural forest setting, the intensity of light reaching the undergrowth is an overriding factor in red spruce develop- ment. Like eastern hemlock (Tsuga canadensis [L.] Carr.) (Mohammed and Parker 1999), physiological acclimation and growth of red spruce in the understory are favoured by moderate increases in irradiance (Davis 1989, Blum 1990) following the periodic formation of small canopy gaps (Busing and Wu 1990) by either windthrow or insect epi- demics (Fraver and White 2005).
Thermosensitivity Red spruce is highly sensitive to high temperature stress, which inhibits its ability to become established in exposed habitats. The species tends to colonize cool, sheltered sites (Blum 1990, Gordon 1994) in the forest understory, where temperatures are moderate and air humidity is usually high (Walter 1967, Johnson and Siccama 1983, Mosseler et al. 2003a), such as on north-facing slopes and in the periphery of lakes (Morgenstern and Farrar 1964). Soil surface temperatures between 20°C and 30°C favour germination of red spruce seeds but temperatures above 33°C can cause irreversible damage (Baldwin 1934). In seedbeds exposed to direct solar radiation, the temperature at the soil surface is often higher than this lethal threshold (Place 1955, Gray and Spies 1997). In the understory, seedling photosyn- thesis is strongly related to air temperature. Amax is attained between 15°C and 20°C, and diminished by 20% and 40% at 25°C and 30°C, respectively (Alexander et al. 1995). Above 32°C, a large drop in photosynthesis and stomatal conduc- tance is possible in seedlings and saplings (Day 2000). However, high temperature inhibition of photosynthesis is likely due to non-stomatal factors as well as an increase in res- Fig. 3. Red spruce advance regeneration under the canopy of a piration rate. In extreme cases, air temperatures that vary yellow birch–conifer stand (L’Islet County, Québec). Top: Live between 32°C and 40°C can result in irreversible foliar injury crown of a sapling (2.3 m height). Bottom: Small 40-year-old in saplings and mature trees (Fincher and Alscher 1992, Vann seedling (0.9 m height). et al. 1994). According to Johnson et al. (1988), abnormally warm like balsam fir, advance red spruce regeneration has a high summers can reduce radial growth of red spruce, although capacity to modify its crown architecture (Tucker et al. 1987, the mechanism of this growth inhibition is not understood. Parent and Messier 1995, Duchesneau et al. 2001) by favour- In northeastern USA, Hamburg and Cogbill (1988) attrib- ing lateral growth over height growth (Fig. 3). Lateral devel- uted the historical decline of red spruce populations to a opment increases light interception and reduces the quantity warming trend in mean summer temperatures. of resources required to maintain non-photosynthetic struc- tures (Waring 1991, McCarthy 2001). Frost susceptibility With its high physiological shade tolerance and capacity of The common occurrence of red spruce needles turning red morphological adjustment to low light, red spruce can sur- and dropping in winter and spring has typically been attrib- vive in the understory with limited growth for decades (Davis uted to foliage desiccation. Water lost from transpiring nee- 1989, Seymour 1995). It is therefore common to observe dles cannot be replaced when xylem cells and/or soil are advance regeneration that is more than 20 years old, but less frozen (Pomerleau 1962, Morgenstern 1969, Roche 1969). than 1 m in height, and saplings over 60 years old that are However, recent research has shown that the water content of shorter than 5 m (Davis 1989, Eagar and Adams 1992) (Fig. red spruce needles is sufficient to ensure the survival of 3). Over the long term, continued suppression may become foliage throughout the cold season (Boyce et al. 1992), and harmful to the species, although it is less problematic for red that needle desiccation is not correlated with the observed spruce than is for balsam fir (Westveld 1931). Red spruce can damage (Hadley and Amundson 1992). Several authors have undergo a succession of several suppression and growth peri- hypothesized that the symptoms of apparent drying of red ods before attaining the upper canopy (White et al. 1985, Wu spruce needles results from previous physiological and mor- et al. 1999). This type of scenario does not reduce its capacity phological frost damage (DeHayes et al. 1990b, Perkins et al. for release (Fowells 1965, Safford 1968, Seymour 1995). 1991, Strimbeck et al. 1991, Hadley et al. 1993a). Indeed, However, seedlings and advance growth red spruce have dif- many studies have shown that red spruce foliage is particu- ficulty acclimating to abrupt exposure to full sunlight. This larly sensitive to freezing temperatures. For example, the
MAY/JUNE 2007, VOL. 83, No. 3 — THE FORESTRY CHRONICLE 381 maximum frost tolerance of the current year’s foliage is gen- erally -30°C to -50°C (DeHayes et al. 1990b; Perkins et al. 1993; Strimbeck et al. 1995; Schaberg et al. 1996, 2000b), compared to -50°C to -90°C for balsam fir (DeHayes et al. 1990b, Strimbeck et al. 1995), and -80°C for black spruce and white spruce (DeHayes 1992, Major et al. 2003a). This com- paratively low frost tolerance would explain why the natural range of red spruce is generally confined to areas with an absolute minimum temperature above -40°C (Arris and Eagleson 1989). Ecophysiological research has shown that red spruce is slow to develop frost tolerance. In its natural environment, the maximum degree of hardening of saplings is reached in the middle of winter (Perkins et al. 1993), and frost tolerance may differ by up to 10°C between individuals in the same stand (Sheppard et al. 1989). A comparable range of harden- ing was also observed between previous- and current-year foliage of mature trees, with younger foliage being approxi- mately 10°C less tolerant than older foliage (Strimbeck and DeHayes 2000). This phenomenon indicates the reduced capacity of new leaves to complete their full acclimation to Fig. 4. A red spruce seedling growing on a partly decayed stump frost by mid-winter (DeHayes et al. 2001). Frost injury to red in a small canopy gap in a yellow birch–conifer stand (L’Islet spruce is thus a frequent phenomenon (Schaberg and County, Québec). DeHayes 2000) that has a negative impact on the growth and survival of trees (Wilkinson 1990, Johnson 1992, Tobi et al. during establishment (Place 1955, Page and Cameron 2006). 1995, Lazarus et al. 2004). Finally, unlike black spruce, red spruce rarely reproduces veg- Frost damage is also due to the fact that, in comparison etatively by layering (Stanek 1967, Gordon 1976). with other Nordic conifers, winter dormancy in red spruce is Like white spruce (Place 1955, Eis 1967), red spruce is par- not deep (Major et al. 2003a). For example, Schaberg et al. ticularly sensitive to water stress, desiccation and mortality (1996) noted that frost tolerance of seedlings strongly during seedling establishment. Red spruce seeds germinate decreased (-47°C to about -33°C) after four to eight days of best on sites that are cool, sheltered and moist (Gordon 1976). simulated winter thawing with temperatures above 0°C. Large decaying woody debris (Fig. 4), moist softwood litter, Similar reductions in frost tolerance were observed by and lightly scarified mineral soil are the best microsites for Strimbeck et al. (1995) in mature trees. This rapid deharden- germination and seedling establishment (Place 1955, Perkins ing, unique to red spruce, is less pronounced if the period of 1991, Seymour 1995). Because germinants produce shallow thawing is brief (DeHayes et al. 1990a, Perkins et al. 1993) and fibrous roots, (Place 1955, Blum 1990, Eagar and Adams if night-time temperature remains below 0°C (Hadley and 1992, Seymour 1995) they have difficulty penetrating hard- Amundson 1992, Schaberg et al. 1996, Lund and Livingston wood litter and organic matter to reach the moist mineral soil 1998). Therefore, periods of abnormally warm temperatures (Klein et al. 1991), and are rarely found on undisturbed for- during the winter (DeHayes et al. 1990a, DeHayes 1992, est floor (Perkins 1991, Battles and Fahey 2000). Hardwood Hadley et al. 1993b), followed by subfreezing temperatures litterfall tends to crush and smoother spruce seedlings (Strimbeck and DeHayes 2000), repeated freeze-thaw cycles (Koroleff 1954, Place 1955). According to many authors, (Lund and Livingston 1998) and direct sun exposure seedlings are considered to be established once they have (Lazarus et al. 2006) are frequent causes of frost damage. attained a height of about 15 cm (Westveld 1953, Frank and Bjorkbom 1973, Baldwin 1977, Blum 1990). Seedling establishment Biotic factors can also be harmful to red spruce regenera- Red spruce has some reproduction characteristics that can, in tion. The species is sensitive to the allelopathic leachable sub- part, explain why successful natural regeneration of this stances (e.g., phenolic compounds) produced by some ferns, species is difficult to obtain operationally. Seeds are only pro- which inhibit seed germination and seedling root develop- duced after a tree reaches 30 to 40 years of age (USDA Forest ment (Klein et al. 1991). Seed predation (pre- and post-dis- Service 1974). Good seed crops occur every three to eight persal) by small mammals is an additional factor that could years (Fowells 1965, White and Cogbill 1992). Mature cones explain the scarcity of red spruce regeneration (see Peters are not persistent and remain on the tree for only one year 2000 and Peters et al. 2003, 2004 for white spruce). after ripening (Morgenstern and Farrar 1964). Seed dispersal rarely surpasses 100 m from the parent tree (Randall 1974, Seedling response to competition Hughes and Bechtel 1997) and viability on the ground is less Once established, red spruce seedlings are sensitive to over- than 12 months (Frank and Safford 1970, Seymour 1995). head competition (Blum 1990). Fast-growing shrub species Similar to white spruce, Sitka spruce (Picea sitchensis [Bong.] tend to invade regeneration niches after large openings are Carr.) and Engelmann spruce (Picea engelmannii Parry), red made in the canopy (Seymour 1992, Smallidge and Leopold spruce seeds are very small and consequently, contain only a 1994). For example, red raspberry (Rubus idaeus L.) can delay small amount of internal reserves to support seedling growth red spruce development after cutting (Westveld 1931, Dibble
382 MAI/JUIN 2007, VOL. 83, No 3 — THE FORESTRY CHRONICLE et al. 1999). In the temperate mixedwood forest of Québec, red spruce than for either balsam fir or white spruce (Gordon intense competition by mountain maple (Acer spicatum 1976, Blum 1988, MacLean and MacKinnon 1997). This may Lam.) and pin cherry (Prunus pensylvanica L.) after clearcut- be because red spruce’s late budbreak (first week of June) ting is problematic for spruce regeneration (Archambault et (McLeod 1961, Blum 1988) does not coincide with the period al. 1998, DeGranpré et al. 2000, Laflèche et al. 2000, Hébert of budworm larval development (Greenbank 1963). 2003). Because red spruce is a late-successional species and Windthrow and spruce budworm damage create small- has a greater capacity for enduring suppression than its natu- scale canopy disturbances that favour the regeneration and ral rivals (Westveld 1931), it tends to reappear with canopy recruitment of late-successional species such as red spruce closure. However, the scarcity of red spruce seed sources can (Foster and Reiners 1983, White et al. 1985, Busing and Wu limit its potential for regeneration (Frank and Safford 1970, 1990, Fraver and White 2005). The understory in small Sullivan and Peterson 1994). Recruitment of new cohorts in canopy gaps is sheltered from full sunlight and temperature the stand depends on the presence of quality seed trees, a par- extremes (Barrett et al. 1962, Marquis 1965, Carlson and tially closed canopy and limited shrub competition. Groot 1997, Gray et al. 2002, Clinton 2003, Page and Because balsam fir possesses several reproductive and eco- Cameron 2006), and usually contains large woody debris physiological characteristics that make it a serious competi- (McCarthy 2001, Chen and Popadiouk 2002, Mitchell et al. tor, the species plays an important role in the natural regen- 2003) as well as light to moderate soil disturbance (Coates eration dynamics of red spruce (Westveld 1931, Seymour and Burton 1997). These are favourable conditions for red 1992, Fortin 2005). Balsam fir produces seeds at a younger age spruce regeneration. For example, old-growth stands in and more frequently than does red spruce (Seymour 1995). northern Maine are periodically opened by small natural gaps Fir’s larger seed reserves enable its germinants to develop rap- (< 50 m2) that represent 10% of the canopy area. These gaps idly. Fir seedlings quickly produce deep penetrating root sys- are sufficient in number and size to favour seedling establish- tems (Hopkins and Donahoe 1939, Westveld 1953, Place ment and the growth of advance regeneration, while main- 1955, Eis 1965, Roe et al. 1970, Knapp and Smith 1982) that taining the dominance of red spruce (Fraver and White increase their chances of first-year survival (Hannah 1988). A 2005). A similar micro-scale gap disturbance regime is known one-year-old fir is much more robust than any of the one- to maintain spruce (Picea engelmannii glauca hybrid) year-old spruces (Place 1952a,b). Furthermore, balsam fir is dominance in British Columbia sub-boreal stands (Kneeshaw better adapted to a broader variety of seedbeds (Simard et al. and Burton 1997). Balsam fir is an important gap-making 1998, Prévost and Pothier 2003). Therefore, fir advance species in Québec’s red spruce ecosystems. Fir not only has a growth is generally much more abundant than that of red short life span, but it is also vulnerable to both windthrow spruce (Ray 1941, Godin 1986, Perkins et al. 1992, Sullivan and spruce budworm attack. and Peterson 1994, Brissette 1996, Grondin et al. 1996). Given that balsam fir regeneration has better juvenile growth and Management Implications and Discussion rapidly adapts to increased light levels, it is better able to Over the last 200 years, forest harvesting has been prolific respond to canopy openings (Westveld 1931, Davis 1989, throughout the range of red spruce (Eagar and Adams 1992) Sullivan and Peterson 1994, Battles and Fahey 2000), which and very few virgin stands containing the species exist today make it a good gap-replacing species (Kneeshaw and (Busing and Wu 1990, Mosseler et al. 2003b, Villeneuve and Bergeron 1998, McCarthy 2001, Hill et al. 2005). Even after a Brisson 2003, Fraver and White 2005). In northeastern USA, major disturbance such as clearcutting, initial succession is diameter-limit cutting was widely applied to spruce–fir often dominated by the large balsam fir seedling bank stands to support the lumber industry during the 19th cen- (Westveld 1953, Foster and Reiners 1983, Battles et al. 1989, tury (Kelty and D’Amato 2006). In Québec’s red spruce Hughes and Bechtel 1997). Red spruce seedlings and saplings, ecosystems, diameter-limit cutting began at the end of 19th being much less numerous, are slower to respond to release century with light intensity harvesting of large, high quality (Battles and Fahey 2000) and are incapable of growing or trees (Ray 1941, Hatcher 1959). With the rapid expansion of even surviving under extreme environmental conditions. the pulp and paper industry, harvesting practices quickly Strategies for managing and conserving red spruce must take changed over the entire range of the species. Diameter-limits into account the different regeneration requirements and the were reduced and extensive clearcutting was practised from competitive interactions between these two companion the late 1800s through the mid-1900s. However, opposition to species (Sullivan and Peterson 1994). clearcutting resulted in a call by forestry professionals for a return to diameter-limit cutting in northeastern USA forests Response to natural disturbances in the 1940s (Heimburger 1941, Ray 1941, Ray 1956, Windthrow and spruce budworm (Choristoneura fumiferana Seymour 1992, Kelty and D’Amato 2006). In Québec, the Clem.) outbreaks are parts of the natural dynamics of north- mechanization of forest operations in the 1960s increased the eastern USA spruce–fir and spruce–fir–hardwood forests use of clearcutting (Fortin et al. 2003a). Clearcutting with (Seymour 1992, Fraver and White 2005) and Québec balsam protection for advance regeneration has been practiced in the fir–yellow birch mixedwood forests (Grondin et al. 1996). northeastern USA for about 60 years and in Québec since the Because it is shallow-rooted (McLintock 1957, Kelly and Mays 1980s (MERQ 1989). The practice was adopted in the USA 1989, Johnson et al. 1991), red spruce is prone to windthrow, following a re-evaluation of the silvicultural systems appro- although it is less vulnerable than balsam fir (Perkins et al. priate to spruce–fir forests (Kelty and D’Amato 2006). In 1992). Defoliation and mortality due to spruce budworm spruce–fir and spruce–fir–hardwood stands, this approach (Weiss and Millers 1988) are generally less problematic for has recently given way to partial cutting methods that are bet-
MAY/JUNE 2007, VOL. 83, No. 3 — THE FORESTRY CHRONICLE 383 ter-adapted to red spruce requirements, such as the shelter- regeneration, favoured shade-intolerant competitive species wood method (Seymour 1992, Fajvan and Seymour 1993). and removed seed sources. This resulted in a drastic reduction In Canada, selection cutting was recently recommended in in the amount of red spruce in its natural range (Frank and Ontario for coniferous and tolerant hardwood forests Blum 1978, Pielke 1981, Siccama et al. 1982, Scott et al. 1984, (OMNR 1998), and in Québec for balsam fir–yellow birch Gordon 1994, Smith and Nicholas 1999, Fortin et al. 2003a, and yellow birch–conifer types (MRNFPQ 2003). However, Loo and Ives 2003, Schwarz et al. 2003). Exceptionally, in the research is still necessary to develop silvicultural strategies eastern part of Nova Scotia, red spruce regenerates success- adapted to red spruce conservation. fully in clearcuts. This is most likely due to the common occurrence of fog, which diffuses the incident light and Diameter-limit cutting reduces the vapor pressure deficit. Foresters’ concerns about diameter-limit cutting date back to the early 1900s (Kenefic and Nyland 2006). The extensive use Logging with advance growth protection of this practice in the zone of red spruce distribution has Removing all of the overstory trees while protecting advance resulted in stand degradation over time, raising the question growth has the same type of negative impacts on red spruce of sustainability of structure and growth in stands with diam- physiology and regeneration as does clearcutting. In Québec eter-limit cutting. In the spruce–fir forests of Maine, much of balsam fir–red spruce stands, careful logging around advance the diameter-limit harvests in the late 1800s resulted in an growth (CLAAG) and related methods of protecting large increase in balsam fir in residual stands by the early 1900s advance regeneration (2 cm to 8 cm dbh) and small mer- (Kelty and D’Amato 2006). Moreover, repeated cutting to chantable stems (10 cm to 14 cm) during harvest are the successively lower diameter limits reduced the growth of red favoured silvicultural options (MRNFPQ 2003). This strategy spruce stands, because the trees with the poorest quality and is unsuitable for red spruce management because the rapid growth potential were retained on the site. Recent research in change in environmental conditions that occurs with over- the USA showed that the long-term use of diameter-limit cut- story removal could damage advance spruce regeneration ting reduced the growth of small residual overstory trees and prevent seed germination and seedling establishment. (Sokol et al. 2004). Because the large, healthy red spruce trees Where small merchantable stems are abundant and well which served as seed sources were, for the most part, elimi- developed, it is likely that they may protect red spruce regen- nated, the red spruce component of the spruce–fir–hardwood eration from the negative effects of high temperature and stands as well as the regeneration of red spruce, declined increased light intensity. However, it is known that CLAAG (Sendak et al. 2003; Kenefic et al. 2005, 2006). and related systems usually increase the proportion of balsam Diameter-limit cutting has been used in Québec’s temper- fir and black spruce in stands to the detriment of red spruce ate mixedwood forest. Ménard (1999) found that 50 years (Seymour 1992, Loo et al. 1997). Therefore, the use of partial after a diameter-limit cut (25 cm, at the stump, for balsam fir cutting methods that maintain some residual overstory are and 36 cm for red spruce) had been carried out, the propor- more appropriate for red spruce regeneration and conserva- tion of spruce in the stand had increased. In a similar study, tion. Archambault et al. (2003) discovered that not only had the basal area of red spruce increased, but also that the invasion Shelterwood cutting by competitive species was minimal in the 50-year period fol- Research in the northeastern USA indicates that partial cut- lowing a diameter-limit cut that removed 15% to 45% of the ting systems favour late-successional species such as red stand’s basal area. Cut diameters (stump) in this study were spruce over their competitors (Baldwin 1977, Seymour 1992, 20 cm for balsam fir and 40 cm for red spruce. However, the Fajvan and Seymour 1993). The presence of a residual canopy partial cutting also increased the density of balsam fir saplings reduces light intensity and limits competition from undesir- to the detriment of red spruce regeneration. Because harvest- able shade-intolerant species, while protecting advance regen- ing was carried out in winter, there was minimal soil pertur- eration from environmental extremes (Lundmark and bation and consequently, a lack of favourable microsites for Hallgren 1987, Groot et al. 1997, Orlander and Karlson 2000, spruce recruitment (Hannah 1988, Battles and Fahey 2000). Lazarus et al. 2004, Pritchard and Comeau 2004, Voicu and According to Fortin (2001) and Fortin et al. (2003a,b), over- Comeau 2006). However, the major impediment to manag- harvesting was a major cause of red spruce scarcity in diame- ing red spruce under any partial harvest system has long been ter-limit cutovers. These authors recommend harvesting bal- the risk of catastrophic windthrow, especially in dense stands sam fir (10 cm dbh) first and only removing large diameter (Westveld 1953, see Busing 2004). Better markets for small (≥ 30 cm dbh) red spruce trees over a long rotation period trees, improvement of logging technology and the practice of (≥ 30 years). This strategy respects the natural dynamics of light thinning from below have greatly lessened the problem spruce regeneration and should increase the proportion of (R. Seymour, personal communication). In the classic shelter- red spruce in the stand. wood system, the final cut (~ 10 years after seed cut) repre- sents a sudden change in microclimate that increases the risk Clearcutting of stress for seedlings and advance regeneration that are not Clearcutting is the least suitable practice for the conservation fully acclimated (Baldwin 1977). As suggested by Sendak et al. of red spruce. Because the species prefers shaded conditions (2003), maintaining a sheltering overstory for a longer period and is sensitive to extreme temperature fluctuations, it fails to and a slow-release strategy would be beneficial, allowing red regenerate in clearcut areas (Westveld 1931, Seymour 1992, spruce to acclimate and become established in several Dibble et al. 1999). Clearcutting of spruce–fir and mixed- cohorts, while limiting the growth of balsam fir and shade- wood stands during the last 200 years eliminated advance intolerant competitive species. This same result can be
384 MAI/JUIN 2007, VOL. 83, No 3 — THE FORESTRY CHRONICLE obtained by combining irregular shelterwood cuts that retain system is feasible and effective. In the context of the Québec spruce as seed trees with understory precommercial thinning mixedwood forest, where yellow birch and red spruce have to control species composition and favour red spruce advance similar longevity and regeneration dynamics, yellow birch is growth (Hannah 1988, Seymour 1995, Sendak et al. 2003). also likely to establish in openings. However, gap size is Like red spruce, balsam fir and tolerant hardwoods such as extremely important. Large gaps would favour yellow birch red maple grow well under shaded conditions (Seymour (Perala and Alm 1990, Fraver and White 2005), which is an 1995). This may be one of the reasons that foresters have intermediate shade-tolerant species, and may result in the problems establishing red spruce regeneration under classic invasion of shade-intolerant competitive species (McDonald shelterwood systems. In Maine, foresters recently experi- and Abbott 1994, Dale et al. 1995, Dibble et al. 1999, mented with an irregular group shelterwood with reserves sys- Raymond et al. 2003), high soil temperatures and frost tem. This expanding gap shelterwood approach inspired by injuries (Strong et al. 1997), whereas smaller gaps usually European silviculture, combines uniform shelterwood with favour the regeneration of shade-tolerant species like red group selection cutting, and is seen as a viable method to spruce and balsam fir (Kneeshaw and Bergeron 1998, restore natural age diversity and composition of Acadian McCarthy 2001). The intensity of thinning and the size of tree forests (Seymour 2005). In conifer and mixedwood stands in groups or small patches should therefore be adjusted accord- New Brunswick and Nova Scotia, research showed that both ingly. Furthermore, the timing between cutting and seed uniform and strip shelterwood cuttings were beneficial for crops is particularly important in successful red spruce regen- red spruce seedling establishment and did not induce wind- eration, owing to its scarcity in comparison with balsam fir throw problems (Baldwin 1977, NSDLF 1991, NSDNR 1994). and yellow birch. Preserving decomposing wood in canopy As expected, because of its shade tolerance and high ther- gaps also mimics the natural disturbance regime of spruce mosensitivity, the greatest stocking of red spruce regeneration regeneration (Mitchell et al. 2003) and provides favourable occurred in the most shaded areas. Unfortunately, to date, microsites for seed germination and seedling establishment. shelterwood systems have not been commonly used to regen- erate red spruce in temperate mixedwood forests in Québec. Artificial regeneration The restoration of red spruce on suitable sites can be accom- Selection cutting plished by artificial regeneration (Dibble et al. 1999, Rajora et According to Frank and Blum (1978), the concept of selection al. 2000). While uncommon in the northeastern USA system silviculture can be put into practice for red spruce (Seymour 1995), plantation establishment is frequent in New conservation and restoration. Recent research in USA showed Brunswick and Nova Scotia where the coastal climate is that this method can increase the proportion of red spruce in favourable to the ecophysiology of red spruce. In southeast- uneven-aged stands (Sendak et al. 2003; Kenefic et al. 2005, ern Ontario, where residual populations of red spruce are 2006). Selection cutting has an advantage over most shelter- very fragmented, planting was necessary to restore this wood methods because it maintains a continuous forest species. Recently, research into the genetics and reproductive cover, which is beneficial to advance regeneration (Westveld status of red spruce (Mosseler et al. 2000, Rajora et al. 2000) 1953, Trimble et al. 1974). However, the use of single-tree has included the establishment of seed orchards and manage- selection cutting is questionable in the absence of advance ment guidelines for artificial restoration. Red spruce is rarely, growth, because the sustained establishment of new red if ever, planted in Québec due to past establishment failures spruce seedlings may require openings larger than individual of this species in open areas (Doucet et al. 1996). However, tree crowns (Frank and Blum 1978). Indeed, when cutting direct seeding and planting in sheltered areas (e.g., small intensity is too light, it is possible that spruce regeneration is canopy gaps) are avenues that should be explored (Prévost et limited by insufficient soil disturbance (Battles and Fahey al. 2003). Recent restrictions concerning herbicide use on 2000, Prévost and Pothier 2003). Conversely, severe thinning Québec forest land may represent an additional challenge for may increase the proportion of balsam fir and other compet- foresters wishing to eliminate vegetative competition around itive species (Seymour 1992), modify stand structure red spruce regeneration. (Lorimer 1989) and cause windthrow (Blum 1990, Johnson et al. 1991, Seymour 1995, Busing 2004). The use of selection Research Needs and Concluding Remarks cutting in northeastern USA spruce–fir–hardwood stands is Failure of past forest practices to protect and encourage red limited for these reasons. spruce regeneration is partly attributable to the lack of knowl- According to Dibble et al. (1999), cutting individual trees, edge and consideration of species’ autecology, a key element groups of trees and creating small openings would be a better of ecosystem management and sustainable forestry (Kramer combined method for improving the regeneration of red 1986, Colombo and Parker 1999). Fortunately, research spruce. Group-tree selection cutting emulates the natural dis- efforts in recent years have filled in some of these gaps in our turbance regime (Coates and Burton 1997), ensuring suffi- knowledge. Numerous studies have shown the sensitivity of cient soil perturbation, a moderate microclimate and ade- red spruce to changes in its microenvironment. It has recently quate soil moisture (Gray et al. 2002, Page and Cameron been acknowledged that harvesting practices that remove 2006, Walters et al. 2006), which are all important factors for most of the forest overstory are inadequate for management red spruce regeneration and establishment. In the temperate of red spruce. Regenerating the species is a slow process that hardwood forest of eastern North America and in the British involves patience and timing, requires silvicultural systems, Columbia boreal mixedwood forest dominated by western such as some shelterwood variants and group-tree selection, hemlock (Tsuga heterophylla [Raf.] Sarg.), Lorimer (1989) and disturbance regimes that are adapted to the ecophysiol- and Coates and Burton (1997) found that a gap-based cutting ogy and regeneration requirements of red spruce.
MAY/JUNE 2007, VOL. 83, No. 3 — THE FORESTRY CHRONICLE 385 Foresters must integrate the most recent information ulation. In this perspective, ongoing and further ecophysio- about red spruce regeneration and conservation into their logical studies are essential to developing ecosystem manage- management strategies. Silvicultural trials must be conducted ment that is tailored to the particular physiological and eco- to determine the optimal gap size or partial cutting intensity logical characteristics of red spruce. needed to promote advance growth and seedling establish- ment. These studies must consider both ecophysiological fac- tors and the growth response of crop trees when establishing Acknowledgements the optimal conditions for red spruce regeneration. In stands The authors wish to thank Francine Bigras, Vincent Roy, with few seed trees, the extent and effect of inbreeding on Patricia Raymond, Robert Seymour and an anonymous seed production and viability are not well documented. reviewer for their helpful comments on a preliminary version However, it has been suggested that inbreeding in small pop- of this review paper. We also express our thanks to Debra ulations can be detrimental to traits such as fecundity Christiansen-Stowe for linguistic revision, Jean Noël for the (Mosseler et al. 2000). Since red spruce is often a companion ecological cartography (Fig. 2), and Lucie Jobin for her valu- species in Québec hardwood and mixedwood forests, the pos- able assistance during the bibliographic research. sibility of inbreeding should be considered. There is a lack of knowledge concerning the type of References microsites preferred by red spruce. For example, some studies Adams, H.S. and S.L. Stephenson. 1989. Old-growth red spruce have revealed the role of large decaying woody debris in the communities in the mid-Appalachians. Vegetatio 85: 45–56. establishment of many Picea species (e.g., Wagg 1964, Lees Alexander, J.D., J.R. Donnelly and J.B. Shane. 1995. Photosynthetic 1970, Knapp and Smith 1982, Côté and Bélanger 1991, and transpirational responses of red spruce understory trees to light Stewart et al. 1996, Timoney and Robinson 1996, DeLong et and temperature. Tree Physiol. 15: 393–398. al. 1997, Brang et al. 2003, Baier et al. 2006, Motta et al. 2006, Amundson, R.G., R.G. Alscher, S. Fellows, G. Rubin, J. Fincher, P. Peters et al. 2006). These microsites exhibit reduced vegetative Van Leuken and L.H. Weinstein. 1991. Seasonal changes in the pig- competition (Harmon and Franklin 1989) and higher mois- ments, carbohydrates and growth of red spruce as affected by expo- ture levels (Place 1955). However, not much is known about sure to ozone for two growing seasons. New Phytol. 118: 127–137. the regeneration ecology of red spruce on decaying wood. It Angers, A.A., C. Messier, M. Beaudet and A. Leduc. 2005. Comparing composition and structure in old-growth and harvested is also necessary to assess the preference of red spruce for (selection and diameter-limit cuts) northern hardwood stands in coarse woody debris (Jamie et al. 2006), especially since cur- Quebec. For. Ecol. Manage. 217: 275–293. rent forest management strategies result in reducing the vol- Archambault, L., J. Bégin, C. Delisle and M. Fortin. 2003. ume of this type of material (e.g., Clark et al. 1998, Hale et al. Dynamique forestière après coupe partielle dans la Forêt expérimen- 1999, Desponts et al. 2002, Roberge and Desrochers 2004, tale du Lac Édouard, Parc de la Mauricie, Québec. For. Chron. 79: Angers et al. 2005, Gibb et al. 2005, Ekbom et al. 2006). 672–684. Red spruce has the ability to respond positively to release Archambault, L., J. Morissette and M. 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