Vertical gradients in photosynthetic light response within an old-growth Douglas-fir and western hemlock canopy

We examined needle-level light response of photosynthesis across a vertical light gradient within 45-55-m-tall western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees growing in a 400-500-year-old mixed species stand. We determined: (1) whether light-saturated photosynthetic rates, light compensation points, and respiration rates varied from the upper to the lower canopy, and (2) if light-saturated photosynthetic rates, light compensation points, and respiration rates varied between Douglas-fir and western hemlock. Over a 25-m gradient from the canopy top to the lower canopy, mean light-saturated photosynthetic rates, light compensation points, and respiration rates declined in overstory Douglas-fir and western hemlock needles, paralleling a 65% decline in the mean daily photosynthetic photon flux density (PPFD). At the canopy top, increasing light-saturated photosynthetic rates relative to lower canopy needles increased carbon uptake at high PPFD. In the lower canopy, reduced respiration rates relative to upper canopy needles increased carbon uptake at low PPFD by reducing the light compensation point. At all canopy positions, western hemlock had lower mean light-saturated photosynthetic rates, light compensation points and respiration rates than Douglas-fir. As a result, western hemlock had higher net photosynthetic rates at low PPFD, but lower net photosynthetic rates at high PPFD compared with Douglas-fir.     

Snag longevity of Douglas-fir,western hemlock,and western redcedar from permanent sample plots in coastal British Columbia

Snags are important both as structural components and as animal habitat in forests, but abundance is often low and their dynamics poorly understood in young, managed stands. Using a large data set of 19,622 snags from permanent plots in second-growth forests of coastal British Columbia, we modeled snag longevity (time from tree mortality to snag fall) for three species: Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western redcedar (Thuja plicata). Snag longevity was strongly related to species and snag size (diameter): the median snag longevity was 16 years for Douglas-fir, 11 years for hemlock and 5 years for redcedar. Western redcedar was predominantly in the subcanopy and its rapid fall rate was related to the small size of its snags. In addition to diameter, other attributes (height to diameter ratio, height, and live crown ratio before death) contributed significantly to models for one or two of the species. However, site level variables did not contribute significantly to any of the models. Snags greater than 50 cm diameter, especially Douglas-fir snags, have the potential for persistence well beyond 20 years in these second-growth forests, and could be important for wildlife.     

Annualized diameter and height growth equations for Pacific Northwest plantation-grown Douglas-fir, western hemlock, and red alder

Simulating the influence of intensive management and annual weather fluctuations on tree growth requires a shorter time step than currently employed by most regional growth models. High-quality data sets are available for several plantation species in the Pacific Northwest region of the United States, but the growth periods ranged from 2 to 12 years in length. Measurement periods of varying length complicate efforts to fit growth models because observed growth rates must be interpolated to a common length growth period or those growth periods longer or shorter than the desired model time step must be discarded. A variation of the iterative technique suggested by Cao [Cao, Q.V., 2000. Prediction of annual diameter growth and survival for individual trees from periodic measurements. Forest Sci. 46, 127–131] was applied to estimate annualized diameter and height growth equations for pure plantations of Douglas-fir, western hemlock, and red alder. Using this technique, fits were significantly improved for all three species by embedding a multi-level nonlinear mixed-effects framework (likelihood ratio test: p < 0.0001). The final models were consistent with expected biological behavior of diameter and height growth over tree, stand, and site variables. The random effects showed some correlation with key physiographic variables such as slope and aspect for Douglas-fir and red alder, but these relationships were not observed for western hemlock. Further, the random effects were more correlated with physiographic variables than actual climate or soils information. Long-term simulations (12–16 years) on an independent dataset using these annualized equations showed that the multi-level mixed effects models were more accurate and precise than those fitted without random effects as mean square error (MSE) was reduced by 13 and 21% for diameter and height growth prediction, respectively. The level of prediction error was also smaller than an existing similar growth model with a longer time step (ORGANON v8) as the annualized equations reduced MSE by 17 and 38% for diameter and height growth prediction, respectively. These models will prove to be quite useful for understanding the interaction of weather and silviculture in the Pacific Northwest and refining the precision of future growth model projections.     

Optimization of foliage photosynthetic capacity in tree canopies: towards identifying missing constraints

Plant canopies are characterized by huge within-canopy gradients in environmental and physiological characteristics, but distributions of foliage photosynthetic capacity in plant canopies do not follow simple optimality criteria. Various hypotheses have been put forward to explain the lack of "optimality", including missing physiological and structural constraints and/or missing fundamental relationships. In this issue, Dewar et al. (2012) highlight important optimality constraints due to finite minimum leaf dry mass per unit area the plants can achieve, while Peltoniemi et al. (2012) proposes that limited acclimation of hydraulic conductance to within-canopy light conditions is a key missing factor in canopy optimality. This commentary advocates that future large-scale model exercises need to introduce more realism in model predictions both by including hydraulic controls as well as consider additional constraints shaping foliage structure, chemistry and physiological activity.     

Douglas-fir and western larch: chemical and physical properties in relation to Douglas-fir bark beetle attack

Frequency of Douglas-fir bark beetle, Dendroctonus pseudotsugae Hopkins (Coleoptera: Scolytidae), attack on western larch (Lark occidentalis Nutt.) was negatively correlated with 3-carene content of the xylem oleoresin. Concentrations of all oleoresin volatiles from stem cores of standing trees were higher in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) than in western larch with the single exception of 3-carene. Successful brood production by the Douglas-fir bark beetle in standing trees of Douglas-fir but not western larch may therefore be explained, at least in part, by the high 3-carene content of western larch. Other differences between the species that might affect susceptibility to beetle attack include thinner phloem, higher phloem moisture content and larger diameter vertical resin ducts in western larch compared with Douglas-fir. Live standing western larch had no oleoresin exudation pressure, suggesting that this trait is not associated with resistance to attack by the Douglas-fir bark beetle in this species.     

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate

We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.     

Lygus hesperus as an agent of apical bud abortion in Douglas-fir nurseries in western Oregon

Causes of apical bud abortion in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were examined in two nurseries in western Oregon. Insect and damage surveys indicated that leafhoppers and Lygus hesperus Knight increased in abundance during August and September, coincident with damage accumulation. Caging studies confirmed that L. hesperus can cause bud abortion and indicated that nymphs of this insect could complete development to maturity on Douglas-fir seedlings.     

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting

Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).     

The impact of black cottonwood on soil fertility in coastal western hemlock forest

Black cottonwood (Populus trichocarpa Torr. and Gray) is a deciduous tree species that extends from Alaska through coastal regions of western Canada into the northwestern United States and as far south as Baja California. We examined the influence of black cottonwood on soil fertility within a forest dominated by Douglas-fir [Pseudotsuga menziessi (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg], and western red cedar (Thuja plicata Donn ex. D. Don.). Six circular 0.008 ha plots with a single cottonwood tree in the center of conifers were paired with six conifer plots (of the same size) without cottonwood. Litterfall, litter decomposition, properties of forest floor and mineral soil, and N mineralization were compared between plot types. Cottonwood litter had higher concentrations of almost all elements relative to conifer litter. Mass loss did not differ between cottonwood and fir/hemlock litter on cottonwood sites. Twice the amount of mull-like humus form (vermimull and mullmoder, 56%) was found in cottonwood plots compared to 28% in conifer plots. Higher pH (4.4) was found in the forest floor under cottonwood compared to conifer (3.9). Total N concentration (3.33 g/kg) and base saturation (68%) were higher in the mineral soil under cottonwood compared to conifers (2.98 g/kg total N and 50% base saturation). Net ammonification and net mineralization were both lower under cottonwood. These results suggest a variable effect of cottonwood on soil fertility within coastal western hemlock forests with some soil variables changed in a favourable direction and some in an unfavourable direction.     

Ants and birds reduce western spruce budworm feeding injury to small Douglas-fir and western larch in Montana

Western spruce budworm, Choristoneura occidentalis Freeman, larvae on small Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) and western larch (Larix occidentalis Nutt.) trees were protected from ant and bird predation during fourth instar to pupation. Significantly greater budworm densities were found on protected trees than on the controls at the end of the experiment, and significantly greater feeding injury was associated with increased budworm densities on fir. It is concluded that ants and birds remove budworm larvae feeding on small conifer trees and reduce injury up to 50%.     

Effects of shade on the morphology and physiology of amabilis fir and western hemlock seedlings

Successful regeneration of coastal montane sites harvested using alternative silvicultural systems may depend on the degree to which tree species can acclimate morphologically and physiologically to a variety of light environments. In a study to determine shade acclimation in montane conifers, one-year-old amabilis fir (Abies amabilis (Dougl.) Forbes) and western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings were grown in a nursery under four shade treatments: full sunlight (0% shade), 60% and 30% shade using shade cloth, and 30% shade using lath slats. Shading influenced shoot development, foliar physiology and morphological characteristics of both amabilis fir and western hemlock but in general, the effects were small. Shade levels of 60% were required to induce significant acclimation, and western hemlock appeared to respond more positively than amabilis fir and therefore was considered more shade tolerant than amabilis fir. Light quality had little influence on growth and development, as indicated by a lack of significant differences in physiology or morphology between seedlings grown under shade cloth or lath slats. There were indications that adequate nutrition levels may mitigate the effects of shade on seedling morphology and physiology.     

An in vitro method for screening Colletotrichum gloeosporioides as a biological control agent for western hemlock dwarf mistletoe

An in vitro system using detached western hemlock branches infected with dwarf mistletoe was developed to screen the virulence of five isolates of Colletotrichum gloeosporioides, a hyperparasite of dwarf mistletoe shoots and berries. Detached branches infected with dwarf mistletoe were placed in nutrient‐saturated rock‐wool blocks and mistletoe shoots were inoculated with a conidial suspension of C. gloeosporioides. One month after inoculation, lesions on mistletoe stems and berries were well developed. Infection levels for individual isolates varied from 40% to 60% of shoots and 60% to 80% of berries. Significant differences were found between the isolates and control (p = 0.02 and 0.001, respectively) while no differences were noted between the isolates for both the shoots and berries. Parallel inoculation of mistletoe shoots detached from hemlock branches on moist filter paper and in rock‐wool blocks failed because these shoots deteriorated rapidly, fragmenting into segments within a week. This in vitro test may provide an alternative method for rapid screening of potentially virulent C. gloeosporioides isolates.     

Potential effects of management practices on nitrogen nutrition and long-term productivity of western hemlock stands

The FORCYTE-10 computer model, developed by J.P. Kimmins and K. Scoullar for Douglas-fir forests in British Columbia, was modified to simulate growth and nutrient cycling of coastal western hemlock stands. Initial calibration indicated that predicted yield was extremely sensitive to the rate of mineralization of soil organic matter (SOM), variation in SOM C:N ratio with site quality, the soil extractable NO₃⁻:NH₄⁺ ratio, and the decomposition rate and N mineralization pattern of large and medium-size roots and woody debris. The predictions suggested that yield and SOM remain stable under a management system consisting of six successive 90-year rotations. More intensive utilization (e.g., shorter rotations, whole-tree harvesting and commercial thinning) causes depletion of soil and forest floor nitrogen and a small decline in site productivity in later rotations.     

Factors limiting early growth of western redcedar, western hemlock and Sitka spruce seedlings on ericaceous-dominated clearcut sites in coastal British Columbia

A 3 year field and pot study was conducted to determine the effects of several biotic and abiotic factors on the early growth of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western redcedar (Thuja plicata, Donn) and Sitka spruce (Picea sitchensis, Bong. Carr.) seedlings established on 2- and 8-year-old clearcut sites previously occupied by old-growth western hemlock and western redcedar forests (referred to as younger and older CH, respectively), and on adjacent 2-year-old clearcut sites previously occupied by second-growth western hemlock and amabilis fir (Amabilis amabilis (Dougl.) Forbes) forests (referred to as younger HA) in coastal British Columbia. The objective of the study was to determine which factors are associated with the poor growth characteristic of the salal (Gaultheria shallon, Pursh) dominated CH clearcut sites.

No soil moisture deficits were measured on any of the three types of clearcut sites at any time of year. The best seeding growth was on the younger HA sites followed by the younger CH sites and then the older CH sites. The better growth on the younger HA sites was associated with a higher availability of N and P in the first 20 cm depth of the forest floor. No differences in matric soil water potential and pH, and only small differences in soil temperature were measured between the three types of clearcut sites. Complete removal of the competing vegetation on the younger and older CH sites resulted in an increase in conifer seedling growth and in the availability of N (22–40%) and P (15–32%); however, it did not affect cellulose decomposition and matric soil water potential, and increased soil temperature only slightly. Both western hemlock and Sitka spruce seedlings were very responsive to differences in nutrient availability measured between types of clearcut sites and planting treatments. In contrast, western redcedar was not responsive. All three conifer species had very high mycorrhizal colonization on the younger CH sites, and this was not altered by the removal of the competing vegetation (mainly salal).

These results suggest that the nutritional stress and poor growth of conifers on salal-dominated CH clearcut sites in coastal British Columbia can be explained by:(1) inherently low forest floor nutrient availability; (2) competition between salal and conifer seedlings for scarce nutrients and nutrient immobilization in salal; (3) declining nutrient availability a few years after clearcutting and slashburning.     

Variation in water potential, hydraulic characteristics and water source use in montane Douglas-fir and lodgepole pine trees in southwestern Alberta and consequences for seasonal changes in photosynthetic capacity

Tree species response to climate change-induced shifts in the hydrological cycle depends on many physiological traits, particularly variation in water relations characteristics. We evaluated differences in shoot water potential, vulnerability of branches to reductions in hydraulic conductivity, and water source use between Pinus contorta Dougl. ex Loud. var. latifolia Engelm. (lodgepole pine) and Pseudotsuga menziesii (Mirb.) Franco (interior Douglas-fir), and determined the consequences for seasonal changes in photosynthetic capacity. The Douglas-fir site had soil with greater depth, finer texture and higher organic matter content than soil at the lodgepole pine site, all factors that increased the storage of soil moisture. While the measured xylem vulnerability curves were quite similar for the two species, Douglas-fir had lower average midday shoot water potentials than did lodgepole pine. This implied that lodgepole pine exhibited stronger stomatal control of transpiration than Douglas-fir, which helped to reduce the magnitude of the water potential gradient required to access water from drying soil. Stable hydrogen isotope measurements indicated that Douglas-fir increased the use of groundwater during mid-summer when precipitation inputs were low, while lodgepole pine did not. There was a greater reduction of photosynthetic carbon gain in lodgepole pine compared with Douglas-fir when the two tree species were exposed to seasonal declines in soil water content. The contrasting patterns of seasonal variation in photosynthetic capacity observed for the two species were a combined result of differences in soil characteristics at the separate sites and the inherent physiological differences between the species.     

Nutrition and bud removal affect biomass and nutrient allocation in Douglas-fir and western red cedar

Seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata J. Donn ex D. Don) were grown at high (250 mg l(-1)) and low (20 mg l(-1)) nitrogen (N) supply for a year. Before the second growing season, half of the seedlings in each nutrient treatment were allocated to the other treatment. Half of the seedlings in each nutrient treatment then had all growing points removed. Biomass and N, phosphorus (P) and potassium (K) concentrations of old and new shoots and roots were measured three times in the second year to test the interaction of current-year and previous-year nutrient supply on biomass and nutrient allocation in these two species with different growth habits. Pruned seedlings served as controls. Unpruned seedlings of both species increased in height throughout the second growing season, except for Douglas-fir in the N250 --> N20 treatment. Repeated pruning did not prevent new shoot growth, but resulted in a 12 to 52% reduction in biomass of new shoots and new and old roots. Seedlings receiving a low N supply in the first growing season were more severely affected by pruning than seedings receiving a high N supply. Growth was reduced more by pruning in western red cedar than in Douglas-fir. Concentrations of N, P and K were higher in pruned seedlings than in unpruned seedlings. Although dry weights of all plant parts in all treatments increased throughout the second growing season, some retranslocation of N, P and K was observed from old shoots of both species in the N250 --> N20 and N20 --> N20 treatments after August. Quantities of N, P and K retranslocated were greatest in seedlings grown the previous year in the high-N treatment.     

Underplanting western hemlock in a red alder thinning: early survival,growth, and damage

Underplanting conifers beneath thinned hardwood stands could shorten conversion to a more valuable crop species. Thinning the hardwoods to final crop-tree spacing could increase growth and marketable volume while releasing the site resources necessary to support conifers planted in the understory. In western Oregon, an experimental thinning of a 14-year-old red alder (Alnus rubra Bong.) stand provided an opportunity to test this concept. Initial efforts were directed toward testing survival and growth of wilding seedlings of western hemlock (Tsuga heterophylla [Raf.] Sarg.) planted in thinned and unthinned alder stands. Survival averaged 78% and 52% after the first and fifth growing seasons. Fifth-year height growth of surviving seedlings averaged from 38 to 49 cm under various thinning regimes but only 9 cm in the unthinned. Wildlife browsing and the pinning of seedlings by falling debris reduced growth and survival.     

Effects of mutual shading of tree crowns on prediction of photosynthetic light-use efficiency in a coastal Douglas-fir forest

Gross primary production (GPP) is often expressed as the product of absorbed photosynthetically active radiation and the efficiency (epsilon) with which a plant community uses absorbed radiation in biomass production. Light-use efficiency is affected by environmental stresses, and varies diurnally and seasonally. Uncertainty about epsilon can be a serious limitation when modeling GPP. An important determinant of epsilon is the amount and type of solar radiation incident on a canopy, because an abundance of light can trigger a photo-protective reaction, diminishing GPP. The radiation regime in a forest canopy is determined by the predominant sky conditions and by mutual shading of tree crowns. Shading effects, producing shifts in the amount of incident direct and diffuse solar radiation, have been largely ignored, however, because they depend on forest structure and are difficult to measure. We describe a new approach for estimating changes in mutual canopy shading throughout the day and year based on a canopy structure model derived from light detection and ranging (LiDAR). Proportions of canopy shading were then combined with eddy covariance data to assess the explanatory power for variance in epsilon by regression tree analysis over half-hourly, daily and weekly time scales. The approach explained between 75 and 97% of variance in epsilon, representing an increase of between 5 and 16% compared with models driven solely by meteorological variables.     

Temperature response of leaf photosynthetic capacity in seedlings from seven temperate tree species

Seedlings of seven temperate tree species (Acer pseudoplatanus L., Betula pendula Roth, Fagus sylvatica L., Fraxinus excelsior L., Juglans regia L., Quercus petraea Matt. Liebl. and Quercus robur L.) were grown in a nursery under neutral filters transmitting 45% of incident global irradiance. During the second or third year of growth, leaf photosynthetic capacity (i.e., maximal carboxylation rate, Vcmax, maximal photosynthetic electron transport rate, Jmax, and dark respiration, Rd) was estimated for five leaves from each species at five or six leaf temperatures (10, 18, 25, 32, 36 and 40 degrees C). Values of Vcmax and Jmax were obtained by fitting the equations of the Farquhar model on response curves of net CO2 assimilation (A) to sub-stomatal CO2 mole fraction (ci), at high irradiance. Primary parameters describing the kinetic properties of Rubisco (specificity factor, affinity for CO2 and for O2, and their temperature responses) were taken from published data obtained with spinach and tobacco, and were used for all species. The temperature responses of Vcmax and Jmax, which were fitted to a thermodynamic model, differed. Mean values of Vcmax and Jmax at a reference temperature of 25 degrees C were 77.3 and 139 micromol m(-2) s(-1), respectively. The activation energy was higher for Vcmax than for Jmax (mean values of 73.1 versus 57.9 kJ mol(-1)) resulting in a decrease in Jmax/Vcmax ratio with increasing temperature. The mean optimal temperature was higher for Vcmax than for Jmax (38.9 versus 35.9 degrees C). In addition, differences in these temperature responses were observed among species. Temperature optima ranged between 35.9 and above 45 degrees C for Vcmax and between 31.7 and 43.3 degrees C for Jmax, but because of data scatter and the limited range of temperatures tested (10 to 40 degrees C), there were few statistically significant differences among species. The optimal temperature for Jmax was highest in Q. robur, Q. petraea and J. regia, and lowest in A. pseudoplatanus and F. excelsior. Measurements of chlorophyll a fluorescence revealed that the critical temperature at which basal fluorescence begins to increase was close to 47 degrees C, with no difference among species. These results should improve the parameterization of photosynthesis models, and be of particular interest when adapted to heterogeneous forests comprising mixtures of species with diverse ecological requirements.     

Estimating cumulative defoliation of balsam fir from hemlock looper and balsam fir sawfly using aerial defoliation survey in western Newfoundland,Canada

Both hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) and balsam fir sawfly (Neodiprion abietis (Harris)) undergo periodic outbreaks in eastern Canada and cause significant growth and mortality losses to forests. Tree growth and mortality are closely related to cumulative defoliation estimates, which integrate annual defoliation over multiple years. Our objective was to determine a method to estimate cumulative defoliation of balsam fir (Abies balsamea [L.] Mill) due to these insects in western Newfoundland, using aerial defoliation survey data, as an essential input to modeling impacts for Decision Support Systems. Interpretation of aerial defoliation survey data for hemlock looper and balsam fir sawfly is problematic because both insects feed upon multiple age classes of foliage. Current-year (2008) aerial defoliation survey data were compared with ground estimates of defoliation by age class from 45 plots (450 trees and 395 mid-crown branch samples), representing a range of defoliation severity classes for each insect. Cumulative defoliation was calculated using defoliation per foliage age class, weighted by relative foliage mass for a given age of foliage. Three significantly different severity classes were defined based on cumulative defoliation values derived from aerial defoliation survey: (i) 1-year moderate (30–70%) defoliation, (ii) 1-year severe (71–100%) defoliation with calculated cumulative defoliation values of 19 and 39%, respectively, for balsam fir sawfly, 21 and 34% respectively for hemlock looper; and (iii) 2–3 years of moderate–severe defoliation, with cumulative defoliation ranging between 59 and 64% for balsam fir sawfly and 49% for hemlock looper. Defoliation severity from aerial defoliation survey alone hence can be misleading if defoliation measurements are not converted to cumulative defoliation values.