Assessing the relationships between stand development and understory vegetation using a 420-year chronosequence

The large-scale conversion of old forests to tree plantations has made it increasingly important to understand how understory vegetation responds to such landscape changes. For instance, in some forest types a reduction in understory richness and cover is thought to result from the development of canopy closure in plantations, although there is a paucity of empirical data demonstrating this relationship. We used a 420-year forest chronosequence as a case study to assess the relationship between stand age, tree canopy cover and understory vascular plant richness and composition in the Siskiyou Mountains of Oregon. The chronosequence consisted of six young managed (age 7–44) and nine older unmanaged (age 90–427) stands. All stands were similar in underlying geology, slope, elevation, and aspect. We found a non-linear relationship between stand age and richness, in which richness was highest in the youngest stands, reached a low in mid-aged stands (∼55 years), then increased in the oldest stands. We also found that percent tree canopy cover was correlated with total understory cover, richness, diversity, and species composition. In general, young stands were characterized by high shrub and graminoid cover and old stands were characterized by an abundant herb layer. Our work suggests that a major component of our study landscape is currently entering the forest stage (canopy closure) characterized by low levels of vascular plant species richness and cover. We use our results to discuss the potential effects of future forest management on understory plants.     

Analysis of the sensitivity of absorbed light and incident light profile to various canopy architecture and stand conditions

We analyzed the effect of simplifying assumptions in canopy representation of radiation transfer models, comparing modeled diffuse non-interceptance and photosynthetic photon flux density with measurements at different layers of complex pine-broadleaved canopy with large seasonal variation of leaf area index. The most detailed model included clumping of trees (i.e.,?stand density) and a vertical specification of leaf angle distribution and shoot clumping. A less detailed model replaced the vertically specified variables with their means. The most parsimonious model accounted for neither shoot clumping nor stand density. The vertical specification of shoot clumping and leaf angle distribution only slightly improved vertical and seasonal openness and light estimates over using mean values. Further simplification had little effect on total absorbed light but was more risky for estimates of the vertical distributions of openness and light absorbed by the canopy, which will affect photosynthesis estimates due to the non-linearity of photosynthetic light response. Including woody surfaces in winter, when leaf area was low, was essential for reproducing the measurements correctly. A sensitivity analysis showed that ignoring (i)?shoot clumping could result in a substantial overestimation of total absorbed light with errors increasing with decreasing leaf area and (ii) stand density in sparse stands could lead to substantial overestimation of total absorbed light, and the effect is largely independent of leaf area. Also, (iii) the effect of changing leaf angle distribution increased with decreasing leaf area, and was larger and more persistent along the leaf area range with increasing shoot clumping. Overall, accounting for the effect of tree clumping on absorbed light is most important in stands composed of species where leaves are not very clumped (e.g., broadleaved). However, even in forests with highly clumped shoots (i.e., coniferous), an accurate estimation of absorbed light distribution in stands requires incorporation of stand density in the model.     

Comparison of stand dynamics after dieback caused by pine wilt disease among pine forests with different management regimes in western Japan

In pine forests damaged by pine wilt disease, in western Japan, the effect of protection regimes of pine trees on the stand dynamics were examined in the following four stands: (1) lightly damaged stand (age 30–40 years) with no procedure in operation for protecting pine trees; (2) severely damaged stand (age 30–40 years) with no procedure in place for protecting pine trees; (3) severely damaged stand (age 50 years) with a selective cutting of infected trees; (4) severely damaged stand (age 30–40 years) with a selective cutting of infected trees. All the stands had been abandoned before the pine wilt disease damage. The understory structure of the severely damaged stand with no protection procedure was similar to that of the lightly damaged stand. Frequent invasion by tree species and acceleration in the growth of understory trees occurred after the dieback in the selective cutting stand. These results suggest that a deficiency in the canopy layer caused by the dieback resulted in low disturbance intensity in the early stages after the dieback, but the selective cutting increased the intensity by the reduction in the understory as well as the canopy layer. The intensity of the disturbance in the selective cutting stands was larger in the younger stand because it had a higher density of selectively cut pine trees. The different stand structure of pine forests occurred after the dieback because the intensity of the disturbance varied as a result of the selective cutting operation and the stand age.     

Three-dimensional light structure of an upland Quercus stand post-tornado disturbance

Light is the most common limiting factor in forest plant communities,influencing species composition,stand structure,and stand productivity in closed canopy stands.Stand vertical light structure is relatively simple under a closed canopy because most light is captured by overstory trees.However,wind disturbance events create canopy openings from local to landscape scales that increase understory light intensity and vertical light structural complexity.We studied the effects of an EF-1 tornado on horizontal and vertical(i.e.three-dimensional)light structure within a Quercus stand to determine how light structure changed with increasing disturbance severity.We used a two-tiered method to collect photosynthetic photon flux density at 4.67 m and 1.37 m above the forest floor to construct three-dimensional light structure across a canopy disturbance severity gradient to see if light intensity varied with increasing tornado damage.Results indicate that increased canopy disturbance closer to the tornado track increased light penetration and light structure heterogeneity at lower forest strata.Increased light intensity correlated with increased sapling density that was more randomly distributed across the plot and had shifted light capture higher in the stand structure.Light penetration through the overstory was most strongly correlated with decreased stem density in the two most important tree species(based on relative dominance and relative density)in the stand,Quercus alba L.(r=-0.31)and Ostrya virginiana(Mill.)K.Koch(r=-0.27,p<.01),and indicated that understory light penetration was most affected by these two species.As managers are increasingly interested in patterning silvicultural entries on natural disturbances,they must understand residual stand and light structures that occur after natural disturbance events.By providing spatial light data that quantifies light structure post-disturbance,managers can use these results to improve planning required for long-term management.The study also provides comparisons with anthropogenic disturbances to the midstory that may offer useful comparisons to natural analogs for future silvicultural consideration.     

杨树人工林冠层光合辐射分布的研究

通过对集约经营与粗放经营杨树人工林冠层上部,下部太阳总辐射和光合有效辐射（ＰＡＲ）的实测数据进行分析,建立了其相互转换的关系方程;研究了ＰＡＲ的日、季变化特征;根据Ｂｅｅｒ－Ｌａｍｂｅｒｔ方程,结合林分生长季内的叶面积动态及林冠各层次的叶面积指数（ＬＡＩ）,计算出生长季内各天的消光系数（Ｋ）及任一时刻冠层内不同深度的光分布,并对ＰＡＲ的透过率与林分消光系数Ｋ及累积叶面积指数之间的相互关系进行了研究     

Dependences of longleaf pine （Pinus palustris） natural reproduction on environments above ground

We studied relations between natural seedling reproduction and above ground environment in a longleaf pine ecosystem. Forty-eight 0.05 ha circular plots were sampled under single-tree selection, group-tree selection and control stands in three main longleaf pine areas in south Alabama, USA. We measured six above-ground environment factors, viz. canopy closure, stand density, basal area, average tree height, understory cover and PAR under canopy. We employed forward, back-ward and stepwise selection regression to produce one model. Three main variables：canopy closure, stand density and basal area, were left in the model; light, PAR and understory cover were not incorporated into the model at the 0.10 significance level. Basal area was a positive pa-rameter, while canopy closure and stand density were negative parame-ters. Canopy closure was the main parameter in the model. The model proved to be meaningful, and has potential to provide useful guidance for future work.     

Canopy development in young sugi (Cryptomeria japonica) stands in relation to changes with age in crown morphology and structure

In young sugi (Cryptomeria japonica D. Don) stands, crown shape (crown length/crown diameter) ratio, average branch inclination, and spatial density of foliage in the crown increased with stand age. Within crowns, foliage distribution increased from the apex downward and, until crown closure, reached a maximum near the crown base. After crown closure, the maximum occurred near the middle of the crown. In each stand, foliage distribution in the canopy showed almost the same vertical change over time as it did in individual crowns. The vertical distribution of foliage in the canopy moved upward with stand age, accompanied by an increase in canopy depth and leaf mass. The shape of the vertical distribution was almost symmetrical between the upper and lower halves in the closed stands, although slightly skewed downward. The logarithm of average spatial density decreased linearly as cumulative leaf mass increased with distance from the top of the canopy. The total cross-sectional area of the crowns exceeded the stand area from the middle of the canopy downward in the closed stands because of crown overlap. However, partly because of changes in crown morphology and structure, the increase in leaf mass with stand age did not always cause more severe crown competition.     

Effects of opening size and stand characteristics on light transmittance and temperature under young trembling aspen stands

While aspen (Populus tremuloides Michx.) cover provides protection from growing season radiative frosts, reduced light levels can limit growth of understory white spruce (Picea glauca (Moench) Voss). The creation of openings around individual spruce can provide more light, but the chance of radiative frost damage increases as gaps become larger. We examined the relationship between opening size and light and temperature levels in the center of circular gaps created in young aspen stands ranging from 1.8 to 8 m tall. Models for predicting transmittance as a function of sky view factor and transmittance under intact stands are presented. Results indicated a strong relationship between light levels and opening size after considering pre-treatment light levels. The relationship between opening size and temperature was weak with minimum temperatures being more affected by the height and density of the surrounding stand than opening size.     

Influence of fire intensity on structure and composition of jack pine stands in the boreal forest of Quebec: Live trees,understory vegetation and dead wood dynamics

North American jack pine (Pinus banksiana Lamb.) stands are generally characterized by an even-aged structure resulting from high intensity fires (HIF). However, non-lethal fires of moderate intensity (MIF), which leave behind surviving trees, have also been reported. The objectives of this study were two-fold: (1) assess the concurrent dynamics of live trees, understory vegetation and different types of coarse woody debris (CWD) during succession after HIF; and (2) document how MIF affects stand structure component dynamics compared to HIF. Stands affected by both HIF and MIF were selected. Tree characteristics and age structure, understory biomass, and CWD volume were assessed. Our results suggest that the structural succession of jack pine stands following HIF comprises three stages: young stands (<48 years), premature and mature stands (58–100 years) and old stands (>118 years). Canopy openness and jack pine density significantly decreased with time since HIF, while black spruce density and CWD volume significantly increased. The highest structural diversity was measured in the premature and mature stands. Compared to HIF, MIF increased mean jack pine basal area, decreased average stand density, delayed the replacement of jack pine by black spruce replacement in the canopy, decreased CWD volume, and significantly increased bryophytes mass. MIF increased the diversity of live trees and generally decreased CWD structural diversity. The study confirms the diversity of natural disturbance magnitude and successional processes thereby initiated. Thereafter, it appeared to be relevant for adjustment of disturbance emulating forest-management systems.     

Trade-offs between seedling growth,thinning and stand stability in Sitka spruce stands: a modelling analysis

The forest industry is increasingly adopting alternative silvicultural systems, involving regeneration beneath an existing forest canopy, rather than clear-felling and replanting. To apply these silvicultural systems in windy regions such as Britain and Ireland, it is essential that the interactions between thinning intensity, stand stability and seedling growth are properly understood. Here, we present a modelling analysis of the three key relationships between: (i) stand density and the proportion of incident radiation transmitted through a forest canopy as a stand is thinned; (ii) transmitted radiation and seedling growth, and (iii) stand density and stand stability. These relationships were examined using separate models of radiative transfer (MAESTRO), seedling growth, and stand stability/wind risk (ForestGALES). Output from the three models was synthesised to calculate whether a given stand thinned to a pre-defined stability limit would allow sufficient light to penetrate the canopy for seedling growth. A minimum transmittance of 20% was identified as a requirement for seedling growth, which corresponds to removing 45% of stand basal area. A thinning of this intensity left some stands susceptible to unacceptable wind damage, especially in old or previously thinned stands on soils where rooting is impeded. The results emphasised the fact that rooting conditions, thinning history and age of intervention are major constraints on the silvicultural options. In general, older stands are not suitable for conversion to continuous cover forestry (CCF) systems, and the transformation process should begin at pole stage, when heavy thinning does not leave the stand unstable. The analysis approach used here illustrates the potential for combining models to address complex forest management issues.     

Leaf area dynamics of a boreal black spruce fire chronosequence

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.     

Natural regeneration characteristics of Pinus sylvestris var. mongolica forests on sandy land in Honghuaerji, China

Natural regeneration in Mongolian pine, Pinus sylvesttis var. mongolica, forest at Honghuaerji of China （the original of the natural Mongolian pine, forest on sandy land） was studied in 2004. The total mean values of regeneration indexes were higher in mature stands （more than 80% individual stems were older than 50 years）, the maximum of regeneration index reached 29 seedlings, m^ 2, with lowest values in the younger stand, e.g., in 32-year old and 43-year old stands. The stand age was an important factor determining the natural regeneration, which was the best in the older stands in this investigation （e.g. about 80-year old）. The regeneration index seemed not to be closely in relation to canopy openness although Mongolian pine is a photophilic tree species. In each type of gaps, natural regeneration was very well. Regeneration indexes were satisfactory at the south and east edges in the circle gaps; and at the east edge of the narrow-square gaps. Results indicated that Mongolian pine, seedlings could endure shading understory, but it would not enter the canopy layer without gap or large disturbance, e.g., fire, wind/snow damage or clear cutting etc. These results may provide potentially references to the management and afforestation of Mongolian pine, plantations on sandy land in arid and semi-arid areas. Researches such as the comprehensive comparisons on regeneration, structure and ecological conditions and so on between natural Mongolian pine, forests and plantations should be conducted in the future.     

Instantaneous and potential radiation effect on underplanted European beech below Norway spruce canopy

In even-aged, 120-year-old Norway spruce stands with underplanted beech (in 1995) four permanent research plots were established, each 100 × 100 m in size. Twelve subplots were selected on each plot along a light gradient from complete canopy closure to open light conditions. On each plot, photon flux density (PFD) was measured continuously, and during same time interval (August 2008, 2009), color digital hemispherical photographs were taken. Cumulative PFD values for measuring points were compared with potential PFD radiation obtained from hemispherical photograph analysis for different angle of hemisphere and different parameters from hemispherical image analysis (gap fraction, total openness and direct, indirect and total amount of radiation). Cumulative and average daily values for the plots were compared; 120° hemispherical photograph angle, gap fraction and total openness were the variables that explained the largest proportion of variance in light transmittance. Determination coefficients between direct and total light component were highest for the total potential radiation and lowest for the diffuse light component. Comparison between potential and instantaneous light measurements for radial, height increment and SLA of young beech showed that instantaneous radiation measurements explained height increment best; 120° of hemisphere proved to be the best explaining angle.     

Understory succession in post-agricultural oak forests: Habitat fragmentation affects forest specialists and generalists differently

The herbaceous understory forms the richest stratum in temperate broadleaved forests in terms of plant diversity. Understanding the process of understory succession is thus of critical importance for the development of management guidelines for biodiversity restoration in post-agricultural plantation forests.We studied effects of stand age, forest fragmentation, and soil and canopy conditions on species richness and abundance of four species groups in the understory of post-arable oak plantations in southern Sweden: herbaceous forest specialists, habitat generalists and open-land species, and woody species.The group of forest specialists may approach the richness of continuously forested sites after 60-80 years in non-fragmented plantations, but many forest species were sensitive to habitat fragmentation. Open-land species richness decreased during succession, while the richness of woody species and of generalists remained stable, and was not affected by fragmentation. Abundance of generalists gradually decreased in non-fragmented plantations, probably due to competition from colonizing forest specialists. Soil pH in post-arable stands remained consistently higher than in continuously forested stands, which maintained differences in species composition. The development of a shrub layer seemed to imply a competitive advantage for forest specialists compared to generalist species.For successful recovery of a rich understory, we suggest that post-arable plantations should be established on loamy soils of intermediate to high pH proximate to older forest with source populations, and that a continuous overstory canopy cover of 70-80% is maintained by regular light thinnings and promotion of a shrub layer.     

Spatial patterns in mixed coniferous even-aged,uneven-aged and conversion stands

Different methodological approaches from the field of spatial statistics, the index of cluster size (ICS) and quadrat methods such as the two-term and three-term local quadrat variance (TTLQV and 3TLQV) and the new local variance (NLV) were tested to find a simple spatial measure to classify mixed coniferous uneven-aged, even-aged and conversion stands in the central Black Forest area of Germany. Altogether six stands were analysed with regularly distributed sample plots of 0.25 ha (50×50 m), each subdivided into 25 quadrats of 10×10 m. In each of the quadrats, diameter at breast height (dbh) for trees of the overstory (dbh>7 cm) was assessed and classified into three diameter classes. Height measurements were used to develop specific stand height curves for each stand and to calculate the standing volume per tree and per quadrat. The even-aged stands showed a regular distribution of the standing volume, while the conversion and uneven-aged stands were more clustered. This was detected using ICS, which proved to be a simple but very efficient measure for stand structure. The ICS also showed a highly random distribution of small and medium trees and a regular distribution of large trees of the overstory in the uneven-aged stand. Large and medium trees of one even-aged stand were also regularly distributed while conversion stands showed a regular, random or slightly clustered distribution of these trees. The more uneven the ages in the stands were, the larger were the phases detected by the NLV. The findings of the ICS were generally supported by the TTLQV and 3TLQV. The more uneven the ages in a stand were, the less clustered were the trees of different sizes of the understory. Clustering also decreased with increasing height of understory trees. The patterns detected in the investigated stands were related to the effect of different management regimes. Implications for the management of conversions stands based on the findings of the study are given.     

Root development across a chronosequence in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We investigated the biomass, vertical distribution, and specific root length (SRL) of fine and small roots in a chronosequence of Japanese cedar (Cryptomeria japonica D. Don) plantations in Nara Prefecture, central Japan. Roots were collected from soil blocks up to 50 cm in depth in five plantations of differing age: 4, 15, 30, 41, and 88 years old. Fine-root biomass reached a maximum (639 g m₋₂) in the 15-year-old stand before canopy closure, decreased in the 30-year-old stand (422 g m₋₂), and thereafter was stable. Except in the 30-year-old stand, fine-root biomass increased in deeper soil layers as stand age increased, and the depth at which the cumulative biomass of fine roots reached 90% exhibited a good allometric relationship with mean stem diameter. Both root-length density (root length per unit soil volume) and SRL decreased with soil depth in all stands, indicating that plants mainly acquire water and nutrients from shallow soils. The highest SRL was observed in the 4-year-old stand, but the relationship between SRL and stand age was unclear in older stands. The SRL in surface soils seemed to decrease with increases in root-length density, suggesting that branching of the fine-root system during development is related to density-dependent processes rather than age.     

Predicting understory maximum shrubs cover using altitude and overstory basal area in different Mediterranean forests

In some areas of the Mediterranean basin where the understory stratum represents a critical fire hazard, managing the canopy cover to control the understory shrubby vegetation is an ecological alternative to the current mechanical management techniques. In this study, we determine the relationship between the overstory basal area and the cover of the understory shrubby vegetation for different dominant canopy species (Pinaceae and Fagaceae species) along a wide altitudinal gradient in the province of Catalonia (Spain). Analyses were conducted using data from the Spanish National Forest Inventory. At the regional scale, when all stands are analysed together, a strong negative relationship between mean shrub cover and site elevation was found. Among the Pinaceae species, we found fairly good relationships between stand basal area and the maximum development of the shrub stratum for species located at intermediate elevations (Pinus nigra, Pinus sylvestris). However, at the extremes of the elevation-climatic gradient (Pinus halepensis and Pinus uncinata stands), stand basal area explained very little of the shrub cover variation probably because microsite and topographic factors override its effect. Among the Fagaceae species, a negative relationship between basal area and the maximum development of the shrub stratum was found in Quercus humilis and Fagus sylvatica dominated stands but not in Quercus ilex. This can be due to the particular canopy structure and management history of Q. ilex stands. In conclusion, our study revealed a marked effect of the tree layer composition and the environment on the relationship between the development of the understory and overstory tree structure. More fine-grained studies are needed to provide forest managers with more detailed information about the relationship between these two forest strata.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.     

A pre-harvest method for predicting understory light levels created by artificial canopy gaps

A new method for predicting understory light levels around an artificial canopy gap prior to the selective cutting is presented. In two even-aged coppice stands, hemispherical photographs were taken at a height of 1.3 m, and crowns of each canopy stem on the photographs were identified by field observations. To predict light levels after cutting, we used computerized images of the hemispherical photographs, in which we digitally erased all crowns to be cut (named ‘erasing method’). Light levels were calculated by an analyzing program; HEMIPHOT. Light levels predicted by the ‘erasing method’ were compared with that calculated by the ordinary photograph-analyzing method using photographs taken after the actual cutting. The correlations between the two methods were highly significant in the two stands. The ‘erasing method’ is eminently effective for the prediction of understory light levels in an artificial canopy gap.