Topographic and compositional influences on disturbance patterns in a northern Maine old-growth landscape

Disturbance patterns are strongly coupled with forest composition and structure, and patterns change through time in response to shifts in climate, anthropogenic impacts and other factors. Knowledge of the natural disturbance patterns for establishing baseline conditions for a forest type or ecosystem facilitates change detection for other elements of the biophysical system important to management and conservation. Dendrochronological reconstructions from old-growth forest remnants throughout northeastern North America document average decadal rates of disturbance of 5%–<10% over the last 150–300 years. Relatively frequent, low severity disturbance characterized by small gaps representing canopy openings made by 1–3 trees prevail in these forests dominated by varying mixtures of late-successional tree species. Few studies, though, have explicitly characterized differences related to composition or topographic setting in old-growth landscapes. We addressed this by comparing the temporal and spatial disturbance patterns reconstructed from tree rings at two spatial resolutions (0.5 ha and 200 m²). Sites were selected to assess the influence of topography (slope) and cover type in stands where red spruce (Picea rubens Sarg) and balsam fir (Abies balsamea L. Mill) were key components. Low rates of disturbance (average <10% per decade) and small gap sizes (≤30 m²) prevailed in all stands during the decades from 1850–1980. Episodic pulses of disturbance, of nearly moderate intensity in some stands, opened ca. 20–30% of the canopy area and were associated with wind events and/or insect outbreaks that differentially affected stands. We found no significant difference in the average temporal rates of disturbance related to cover type or topography in 0.5-ha plots. However, the influence of these factors was evident in comparisons of gap areas estimated for 200-m² plot sections. At this resolution, the largest canopy openings (≥100 m²) occurred most frequently in slope sites, enabling pulses of canopy accession for Betula alleghaniensis (Britton). Whereas the smallest canopy openings (≤30 m²) dominated softwood stands, favoring red spruce, balsam fir and other shade-tolerant species throughout the forest. The variable effects of common disturbance agents, regardless of topographic position and/or cover types, points to the important role of biological legacies determining stand structure and composition on subsequent disturbance events and long-term patterns.     

First decadal response to treatment in a disturbance-based silviculture experiment in Maine

Disturbance-based silvicultural systems generally seek to promote complex stand structures that are consistent with temporal and spatial patterns of natural disturbance while allowing for the sustainable harvest of timber. Gap-based harvesting systems are commonly used within this framework because they can be designed to approximate the frequencies and spatial patterns of a wide array of disturbance regimes. Patterns in stand-level growth, sapling recruitment and regeneration response were examined for one such gap-based system, the Acadian Forest Ecosystem Research Program (AFERP) in central Maine, that was designed to emulate the annual 1% disturbance frequency typical of the northeastern United States and Canada. A decade after treatment, stand-level differences in basal area growth and density between two gap-based treatments and an unharvested control were not statistically significant, largely due to low replication, but within-stand growth and regeneration responses differed strongly by spatial position relative to harvest gaps. Regeneration of shade-tolerant and intolerant species increased regardless of gap size, likely a response to increasing light availability from canopy openings due to harvesting and mortality. Further, there was evidence of gap size effects on sapling recruitment as large gaps (>1000 m²) favored the growth and survival of mid-successional species such as red maple and white pine, while small gaps (<1000 m²) favored late-succession species such as eastern hemlock and spruce. Overstory growth rates also differed by both species and position relative to harvest gaps with most species growing best in gaps and better along gap edges than in adjacent forest. Notably, overstory growth rates for white pine were not influenced by spatial position. These results suggest harvest gaps may have significant growth and regeneration impacts in adjacent, yet unharvested areas, which could lead to profound differences in forest development over the rotation. Obviously, longer-term studies of gap-based systems are needed to more clearly elucidate these responses.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

The role of gaps and tree regeneration in the transition from dense to open black spruce stands

Black spruce forests growing on clay soils in northwestern Quebec change structure from dense even-aged stands to open uneven-aged stands such that almost all forests older than 200 years have an open canopy. These forests become unproductive over time because they are prone to paludification. The main goal of our study was to document the transition between dense and open stands in terms of gap dynamics, with a focus on tree regeneration. Our objective was to determine whether forests remain open due to a lack of regeneration, a lack of growth or both. Nine stands along a 50–250-year-old time since fire gradient were sampled with the line intersect sampling method. Gap fraction increased with stand age and reached a maximum of 77% in the oldest site. In old-growth stands, gaps were interconnected due to the low density of these forests. Most of the gap makers were found with broken stems. Regeneration was dominated by black spruce layers and was relatively abundant (1.71 stems/m²). However, the majority of gap fillers were smaller than 1 m in height in stands of all ages. Instead of a lack of regeneration, the opening of the forests is due to a lack of growth associated with cold and wet organic deposits. Partial harvesting could be implemented on the most productive sites, while management techniques including soil disturbances will be required on low productivity sites to recreate good growth conditions.     

Regeneration of Norway spruce in canopy gaps in Sphagnum-Myrtillus old-growth forests

Gap-associated spruce (Picea abies (L.) Karst.) regeneration in Sphagnum-Myrtillus stands of south taiga forests (Central Forest Biosphere reserve, Tver region, Russia) was studied to evaluate the role of different disturbances in spruce dynamics. Sampled gaps (n=70) ranged from 40 m² to 1.7 ha in size, and from 1 to 70 years since disturbance moment. Formation of gaps lead to increase in the number of stems per ha in all gap size classes (small: 40–200 m², medium: 200–3000 m², and large: >3000 m² gaps). Spruce was the most important species in gap refilling, although its role was not the same in different gap classes. The highest values of relative abundance (compared to other species) were recorded in small gaps, and much lower values – in middle and large gaps. However, as refilling of gaps proceeded, spruce showed rather active regeneration in middle and large gaps and partly regained its abundance in middle-age disturbances. In general, all types of gaps studied supported spruce regeneration into the forest canopy. Almost perfect correlation between predicted outcome of spruce dynamics in gaps and its current role in the canopy of Sphagnum-Myrtillus stands suggests a good adaptation of this species to the current disturbance regime and a steady state of the these forests.     

Forest regeneration in artificial gaps twelve years after canopy opening in Acre State Western Amazon

The main objectives were to study the effect of gap size and canopy openness on the natural regeneration dynamics considering the parameters of sapling growth, recruitment, mortality, density, species composition and above-ground biomass accumulation. The study was carried out in 32 artificial gaps with sizes varying from 100 to 1200 m² and canopy openness from 10 to 45%, from the second to the twelfth year after gap creation. The gap size was measured using the vertical projection of the tree crowns on the ground (Brokaw's definition), and the canopy openness measurement by hemispherical photography. In the first five years, mean sapling growth (0.54 cm year⁻¹), mortality (3.9% year⁻¹) and AGB (26.2 Mg ha⁻¹ or 8.7 Mg ha⁻¹ year⁻¹) were significantly higher in the gaps than in the forest understorey (0.17 cm year⁻¹, 1.5% year⁻¹ and −0.59 Mg ha⁻¹ year⁻¹ respectively) and positively correlated with gap size and canopy openness. In the same period, recruitment was also significantly higher in the gaps (5.8% year⁻¹) than in the forest understorey (0.4% year⁻¹) but decreased with gap size and negatively correlated with canopy openness. In the first five years, the relative density of pioneer species was higher in the gaps but not significantly correlated with gap size or canopy openness. AGB increased linearly since canopy opening, and twelve years after gap creation it was still higher in larger (121.2 Mg ha⁻¹ or 10.1 Mg ha⁻¹ year⁻¹) rather than smaller (62.5 ha⁻¹ or 5.2 ha⁻¹ year⁻¹) gaps. Twelve years after gap creation there were no significant differences in the parameters of sapling growth, recruitment, and mortality which could be attributed to the original gap size and canopy openness.     

Effects of winter selective tree harvest on soil microclimate and surface CO2 flux of a northern hardwood forest

Soil surface CO₂ flux (S_flux) is the second largest terrestrial ecosystem carbon flux, and may be affected by forest harvest. The effects of clearcutting on S_flux have been studied, but little is known about the effect of alternative harvesting methods such as selective tree harvest on S_flux. We measured S_flux before and after (i) the creation of forest canopy gaps (simulating group tree selection harvests) and (ii) mechanized winter harvest but no tree removal (simulating ground disturbance associated with logging). The experiment was carried out in a sugar maple dominated forest in the Flambeau River State Forest, Wisconsin. Pre-treatment measurements of soil moisture, temperature and S_flux were measured throughout the growing season of 2006. In January–February 2007, a harvester created the canopy gaps (200–380 m²). The mechanization treatment consisted of the harvester traveling through the plots for a similar amount of time as the gap plots, but no trees were cut. Soil moisture and temperature and S_flux were measured throughout the growing season for 1 year prior to harvest and for 2 years after harvest. Soil moisture and temperature were significantly greater in the gap than mechanized and control treatments. Instantaneous S_flux was positively correlated to soil moisture and soil temperature at 2 and 10 cm, but temperature at 10 cm was the single best predictor. Annual S_flux was not significantly different among treatments prior to winter 2007 harvest, and was not significantly different among treatments after harvest. Annual (+1 std. err.) S_flux averaged 967 + 72, 1011 + 72, and 1012 + 72 g C m⁻² year⁻¹ in the control, mechanized and gap treatments, respectively, for the 2-year post-treatment period. The results from this study suggest selective group tree harvest significantly increases soil moisture and temperature but does not significantly influence S_flux.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Methods for studying treefall gaps: A review

As silvicultural objectives have changed over the last several decades, managers are increasingly designing cutting regimes that mimic natural disturbance with the hopes that such systems will restore forests to a more natural condition while optimizing harvest yield. Treefall gaps, canopy openings caused by the death of one or more trees, are the dominant form of disturbance in many forest systems worldwide. These gaps play an important role in forest ecology by helping to maintain bio- and pedo-diversity, influencing nutrient cycling, and preserving the uneven-age nature of late-successional forests. In gap literature, there are inconsistencies with regard to gap terminology, methods for identifying and studying gaps, and modeling gap disturbances. From the papers reviewed, the size of treefall gaps ranges widely from 10 to >5000 m²; we suggest that the maximum gap size should be set at 1000 m². Larger openings tend to have microclimates and return intervals significantly different than smaller treefall gaps. Two main definitions of treefall gaps exist: canopy gap: a ‘hole’ in the forest through all levels down to an average height of 2 m above ground and extended gap: canopy gap plus the area that extends to the bases of surrounding canopy trees. Although researchers have assumed a variety of gap shapes to simplify measuring gap size, gaps are often irregularly shaped and so we recommend that gap areas and shapes be determined from detailed field measurements. Gap age may be determined from tree ring analysis of released trees in or near the gap edge, the spacing of whorls on released saplings, or from decomposition of gap-making trees. Windthrow is the main cause of canopy gaps in a variety of ecosystems; other causes include insects, diseases, acidic deposition, drought, and climate change. Treefall-gap models have been developed to predict the following processes during gap making or infilling: (i) gap abundance, (ii) forest structure, (iii) spatial and temporal variations in light levels, (iv) canopy dynamics, and (v) soil nutrient and water regimes. We recommend a protocol for gap studies and identify future research topics.     

Regeneration of mahogany and Spanish cedar in gaps created by railroad tie extraction in Quintana Roo,Mexico

Seeds of mahogany and Spanish cedar were planted in gaps created by harvesting railroad ties at two locations in the Yucatan Peninsula. Gaps averaged 40 m² in area. Germination after 2 months was very low (2%) for Spanish cedar and moderately low (29%) for mahogany at the one location where it was measured, probably due to predation and fungal damage. Annual seedling mortality was 25–40%, and height growth was moderate to normal, with average height equal to 45 cm after 3 or 4 years, depending upon the location. We conclude that gaps from railroad tie harvesting are not adequate for the establishment of either species from seed. If, however, gaps created by harvesting could be concentrated into at least moderately open areas 400 m² in area or larger, they may be useful as sites for regenerating both species.     

Restoring longleaf pine (Pinus palustris Mill.) in loblolly pine (Pinus taeda L.) stands: Effects of restoration treatments on natural loblolly pine regeneration

Historical land use and management practices in the southeastern United States have resulted in the dominance of loblolly pine (Pinus taeda L.) on many upland sites that historically were occupied by longleaf pine (Pinus palustris Mill.). There is currently much interest in restoring high quality longleaf pine habitats to such areas, but managers may also desire the retention of some existing canopy trees to meet current conservation objectives. However, fast-growing natural loblolly pine regeneration may threaten the success of artificially regenerated longleaf pine seedlings. We evaluated the establishment and growth of natural loblolly pine regeneration following different levels of timber harvest using single-tree selection (Control (uncut, residual basal area ∼16 m²/ha), MedBA (residual basal area of ∼9 m²/ha), LowBA (residual basal area of ∼6 m²/ha), and Clearcut (complete canopy removal)) and to different positions within canopy gaps (approximately 2800 m²) created by patch cutting at two ecologically distinct sites within the longleaf pine range: Fort Benning, GA in the Middle Coastal Plain and Camp Lejeune, NC in the Lower Coastal Plain. The density of loblolly pine seedlings was much higher at Camp Lejeune than at Fort Benning at the end of the first growing season after harvesting. Following two growing seasons, there were no significant effects of canopy density or gap position on the density of loblolly pine seedlings at either site, but loblolly pine seedlings were taller on treatments with greater canopy removal. Prescribed fires applied following the second growing season killed 70.6% of loblolly pine seedlings at Fort Benning and 64.3% of seedlings at Camp Lejeune. Loblolly pine seedlings were generally less than 2 m tall, and completeness of the prescribed burns appeared more important for determining seedling survival than seedling size. Silvicultural treatments that include canopy removal, such as patch cutting or clearcuts, will increase loblolly pine seedling growth and shorten the window of opportunity for control with prescribed fire. Therefore, application of prescribed fire every 2-3 years will be critical for control of loblolly pine regeneration during restoration of longleaf pine in existing loblolly pine stands.     

Planting mahogany in canopy gaps created by commercial harvesting

Attempts at natural forest management of mahogany (Swietenia macrophylla King) have so far met with limited success, whilst many plantations are beset by the shoot borer Hypsipyla spp. In this paper we present preliminary results of an approach to enrichment planting that aims to balance economic returns (rapid growth and good silvicultural form) with intervention costs and changes to forest structure. Mahogany seedlings were planted in gaps created by selective timber harvesting and that ranged in vertical projected area from 91 to 542 m² (mean = 257 m²). Seedlings grew within the matrix of gap regrowth, with limited control of competing vegetation. Sixty-one percent of seedlings had survived by 4.4 years (equivalent to an annual mortality rate of 10.5% year⁻¹), and had reached a mean height of 4.5 m. Stocking levels of mahogany were similar to that of naturally regenerated commercial species in unplanted gaps of the same age, but mahogany seedlings were significantly taller. The incidence of shoot borer attack on mahogany stems was relatively low (54.7%), but, more importantly, most damaged stems (58%) responded by producing a single replacement leader. The cost of the proposed methodology (US$ 94 per gap over 4.4 years) was low compared to the high value of mahogany timber relative to other species in the forest. The implications of planting mahogany in gaps for forest management and the potential benefits to conservation of the species are considered.     

Calculation method for sunshine duration in canopy gaps and its application in analyzing gap light regimes

Understory light is essential to the growth and survival of plants, yet the light varies temporally and spatially within forest gaps. Measurement of understory light levels using light sensors is both time and labor intensive. Sunshine duration (SD) has a strong correlation with solar radiation and has been the variable most widely used for estimating solar radiation. The power of SD-based methods for estimating solar radiation lies in its ability to quickly estimate light levels. Although several calculation methods for SD within canopy gaps are available, all the methods oversimplify canopy gaps by classifying them as cylindrical or ellipsoid and thus have a relatively low level of accuracy. In this study we developed a calculation method for SD at any given point within natural canopy gaps and we used SD to estimate solar radiation within 12 canopy gaps on Mt. Taibai in China based on the Angström-Prescott model. We then evaluated the SD-based solar radiation by the total and direct solar radiation derived from a gap light index based on hemispherical photographs (HP). Finally, we analyzed the spatial-temporal characteristics of light levels within these 12 gaps by using the solar radiation derived from SD at hundreds of simulated points in each gap. Our results showed that (1) SD-based solar radiation was not statistically different from HP-based direct solar radiation and had a strong linear correlation with HP-based total solar radiation; (2) growing-season daily mean solar radiation within the 12 gaps varied from 0.08 to 13.28 MJ m⁻² day⁻¹ with an average of 4.13 MJ m⁻² day⁻¹; (3) solar radiation had a positive correlation with the ratio of the square root of the canopy gap area to the mean canopy height. This relationship was significant but solar radiation had no correlation with the canopy gap area; (4) among most gaps solar radiation was greatest in May while potential SD was longest in June 2008. From these results we can conclude that the SD-based method for estimating solar radiation developed in this study can quickly and accurately estimate light levels at any specified point within canopy gaps. SD-based solar radiation appears to be a good choice for studies on the spatial-temporal characteristics of gap light levels.     

Determinism versus chance in canopy gap herbaceous species assemblages in temperate Abies-Betula forests

It is not known if the species composition of herbaceous plant assemblies within tree-fall gaps is determined stochastically or whether it follows species-specific and environmentally-determined patterns. We applied three methods, fitting species abundance distribution models, comparing community similarities, and testing species-environmental variable association, to evaluate the relative importance of chance and deterministic rules in controlling herbaceous composition within canopy gaps in the Abies-Betula forests of Mt. Taibai, Central China. Herbaceous species abundance within canopy gaps was well fit by a neutral model, showing that relative abundance was qualitatively consistent with stochastic processes. Although species composition in gaps significantly differed from closed canopy sites (ANOSIM, R = 0.509, p = 0.001), there were no significant differences among gaps of different age groups (ANOSIM, R = 0.035, p = 0.191) or gaps of different size groups (ANOSIM, R = 0.089, p = 0.057). Similarly, gaps of similar age and size did not show significantly higher similarity (χ² = 2.30, df = 3, p = 0.513) in species composition than gaps of distinct age or size. Moreover, there were only eight species, including two light-demanding species, confined to larger gaps among the 69 species, and only an additional seven species found more commonly in larger gaps, whereas most herbaceous species were gap size generalists. Canonical correspondence analysis and random permutation tests suggested that only 27% of species with abundance ?5 were associated with environmental variables in gaps. In summary, the species composition in gaps was not constrained significantly by gap traits; rather species were distributed stochastically, likely through by random dispersal and recruitment limitation of species from the surrounding available species pool. Measures that introduce gap disturbance, such as selective harvesting, are still recommended when the maintenance of total biodiversity in forests is a concern.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Alternative silvicultural practices with variable retention to improve understory plant diversity conservation in southern Patagonian forests

Understory plants could can act as indicators of temperate forest sustainability, health and conservation status due to their importance in ecosystem function. Harvesting impacts on understory plant diversity depends on their intensity. Variable retention has been proposed to mitigate the harmful effects of timber harvesting, but its effectiveness remains unknown in southern Patagonian Nothofagus pumilio forests. The objectives of this study were to: (i) define a baseline of understory plant diversity in old-growth forests along a site quality gradient and under canopy gaps; (ii) evaluate stands with three different variable retention treatments compared to old-growth forests; and (iii) assess temporal changes during 4 years after harvesting (YAH). A 61 ha N. pumilio forest was selected. Understory plant (Dicotyledonae, Monocotyledonae and Pteridophyta) richness, cover (including woody debris and bare forest floor) and aboveground dry biomass were characterized in summer for 5 years. Before harvesting, baseline samples were conducted along a site quality gradient and outside/inside canopy gaps. Analyzed treatments include a control of old-growth forest (OGF) and three different harvesting treatments with variable retention: (i) dispersed retention (DR) of 30 m² ha⁻¹ (20-30% retention); (ii) aggregated retention (AR) with one aggregate per hectare and clear-cuts (28% retention); and (iii) combined dispersed and aggregated retention (DAR) with one aggregate per hectare and dispersed retention of 10-15 m² ha⁻¹ (40-50% retention). Data analyses included parametric and permutational ANOVAs, multivariate classification and ordinations.Before harvesting, 31 plant species were found, where richness, cover and biomass were directly related to site quality. The presence of canopy gaps did not have a significant impact on the measured variables. After harvesting, 20 new species appeared from adjacent associated environments (two from N. antarctica forests and 18 from grasslands and peatlands). At the stand level, understory values were higher in AR > DR > DAR > OGF. Most (81-95%) plant richness at baseline conditions was conserved in all treatments, where inside the aggregates understory remained similar to OGF. Combination of aggregated and dispersed retention (DAR) better limited exotic species introduction and protected sensitive species, improving conservation in harvested stands. Changes in understory variables were observed after the first YAH in all treatments; greater changes were observed in the harvested areas than in aggregates. Changes stabilized at the fourth YAH. As a conclusion, the location of retention aggregates should be selected to preserve species understory diversity of more speciose and diverse habitats or particularly uncommon stands. Implementation of different kinds (patterns and levels) of retention for improvement of biodiversity conservation in harvested forests should be included in timber and forest management planning.     

Sugar maple and yellow birch regeneration in response to canopy opening,liming and vegetation control in a temperate deciduous forest of Quebec

We examined how the density, growth and survival of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) regeneration are influenced by gap size, soil nutrient availability and understory vegetation. We used a factorial combination of (1) three gap sizes (small: <100 m²; medium: 100–300 m²; large: ∼1000 m²); (2) presence/absence of liming (92% CaCO₃ at 500 kg ha⁻¹, 1st year post-harvest); and (3) presence/absence of vegetation control (weeding twice a year; 1st to 3rd year post-harvest). We monitored height increment and survival of 1500 seedlings and saplings of both species from the 3rd to the 6th year post-harvest, and assessed density 6 years post-harvest. Both species exhibited a complex set of density, growth and survival responses across the combination of treatments. Compared to sugar maple, yellow birch had an overall lower density, greater growth, and similar survival rate; the two species attained maximum values in different gap size for density, and similar gap size for growth and survival. Liming had very little or no effect on the species. The growth of yellow birch was slightly but significantly greater when understory vegetation was controlled, particularly in medium and large gaps. These results suggest that a variety of canopy gap sizes can provide the right combination of understory conditions for regenerating these two functionally different tree species.     

Canopy gap characteristics and their implications for management in the temperate rainforests of southeast Alaska

In the coastal temperate rainforests of southeast Alaska, much progress has been made in describing landscape-level natural disturbances and formulating management systems that emulate those disturbances. Little is known, however, concerning canopy gaps, the dominant form of natural disturbance in the region. During June–August, 1991–1993, we characterized canopy gap patterns and dynamics at three sites in the western hemlock/blueberry/shield fern plant association in the northern portion of the Tongass National Forest.Forest area in canopy gaps ranged from 5.8 to 12.6% and averaged 8.7%. The proportion of forest area in expanded gaps ranged from 18.1 to 43.9% and averaged 27.4%. Gap and gapmaker (tree whose death or crown displacement results in the creation or expansion of a canopy gap) characteristics were generally similar among sites. The majority of canopy gaps were <50 m² in area, had a D/H ratio <0.50, were created from the death of one or two gapmakers, and had experienced gap expansion. The majority of expanded gap areas were <200 m². Gapmakers were usually snapped, had recently died (<20 years ago), and tended to be larger in diameter than surrounding overstory trees. Species composition of gapmakers was similar to surrounding overstory trees. A substantial amount of gap infilling takes place between 20 and 80 years following tree death, but gaps, or portions of gaps, can persist for >80 years. Forest turnover time was estimated to range from 230 to 920 years, and average 575 years. Canopy residence time was estimated to range between 210 and 840 years, and averaged 525 years.To emulate canopy gap dynamics in the plant association studied, forest managers should: (1) maintain a small proportion of a stand in openings within an otherwise undisturbed canopy; (2) use a combination of single tree selection and small group selection systems; (3) re-enter stands every 20–80 years; (4) select larger than average diameter crop trees in proportion to the species composition of the stand; (5) minimize soil disturbance and (6) select crop trees during re-entry so that the creation of new gaps and the expansion of old gaps is accomplished in approximately equal proportions.     

Slow responses of understory plants of maple-dominated forests to white-tailed deer experimental exclusion

We examined the response of understory plants in mature maple-dominated forests of southern Québec, Canada, following about 30 years of high deer densities, using a deer exclosure experiment. An exclosure and a paired control of 625 m² each were established on six sites in 1998. An exclosure and a paired control of 16 m² were added at each of the same sites in 2003 but under a recent canopy gap to determine if light could enhance plant responses. We measured plant richness and abundance, and aboveground biomass of different plant groups for 8 years in the understory plots and for 3 years in the canopy gaps. Four herbaceous species were also monitored individually in the same plots. No significant differences between treatments were found in plots under forest cover, except for lateral obstruction at 0–50 cm height which was higher in the exclosures. Under canopy gaps, however, tree seedling and total plant abundance were higher in deer exclosures than in control plots. Trillium erectum recovered partially as individuals were taller, had larger leaves and more frequently produced a flower or a fruit in the absence of deer browsing under forest cover. To a lesser extent, Erythronium americanum and Maianthemum canadense also exhibited signs of recovery but were still at the single-leaf stage after 8 years of recovery. In general, the different plant groups exhibited little recovery following deer exclusion, possibly because of the low light levels that prevailed in the understory of undisturbed maple-dominated forests. The higher latitude of the present study could also contribute to the slow recovery rates of the different groups of plants compared to studies conducted in northeastern USA. Variability among sites and years had an effect on detection of statistically significant differences. Trends are however appearing over time, suggesting that many understory plants are recovering very slowly following deer exclusion. Our results emphasize the importance of studying large herbivore–forest interactions on different groups of plants, but also on specific species, and under different latitudes to be fully understood.