The role of canopy gaps in the regeneration of coastal dune forest

In regenerating coastal dune forest, the canopy consists almost exclusively of a single species, Acacia karroo. When these trees die, they create large canopy gaps. If this promotes the persistence of pioneer species to the detriment of other forest species, then the end goal of a restored coastal dune forest may be unobtainable. We wished to ascertain whether tree species composition and richness differed significantly between canopy gaps and intact canopy, and across a gradient of gap sizes. In three known‐age regenerating coastal dune forest sites, we measured 146 gaps, the species responsible for gap creation, the species most likely to reach the canopy and the composition of adults, seedlings and saplings. We paired each gap with an adjacent plot of the same area that was entirely under intact canopy and sampled in the same way. Most species (15 of 23) had higher abundance in canopy gaps. The probability of self‐replacement was low for A. karroo even in the largest gaps. Despite this predominance of shade‐intolerant species, regenerating dune forest appears to be in the first phase of succession with ‘forest pioneers’ replacing the dominant canopy species. The nature of these species should lead to successful regeneration of dune forest.     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Comparison of old-field and forest revegetation dynamics in Provence

Abstract. This paper describes vegetation changes on cultivation terraces after abandonment, considered as secondary succession, and vegetation responses after clear-cutting, considered as regeneration. Species were grouped in life form and dynamic categories in order to infer dynamic patterns. By applying the Shannon diversity index to these characteristics, an index of functional diversity is obtained. Regenerative succession is considered here as a particular case of secondary succession, characterized by a fast vegetative regeneration of the dominant woody species, which controls the succession of the understorey species. Post-agricultural succession and forest regeneration succession show a similar relation between the diversity index and a forest index, indicating the relative amount of forest species. It appeared that the old-field dynamics resemble the deciduous oak-forest regeneration after the woody species have established. The species diversity at the end of the post-agricultural succession and in the forest is controlled by the nature of the dominant tree species.     

Disturbance and regeneration dynamics of a mixed Korean pine dominated forest on Changbai Mountain,North-Eastern China

We used dendrochronological methods to study disturbance history of a mixed Korean pine (Pinus koraiensis Siebold et Zuccarini) dominated forest on the northern slope of Changbai Mountain, North Eastern China, over 1770–2000. Frequent small-scale canopy gaps and infrequent medium-scale canopy disturbances dominated natural disturbance regime in the forest, which did not experience stand-replacing disturbances over the studied period. Percentages of growth releases in subcanopy trees were below 6% in most decades, suggesting that disturbances initiating these releases were of low intensity. Strong winds were likely cause of moderate disturbance events. Two episodes with increased disturbance rates (19% and 13%) were dated to the 1920s and 1980s, timing of the 1980s event was consistent with a hurricane occurred in 1986 on the western slope of the Changbai Mountain. Age structure and growth release analyses revealed species-specific regeneration strategies of canopy dominants. Shade-intolerant Olga bay larch (Larix olgensis Henry) recruited mainly before the 1860s. Recruitment of moderately shade-tolerant P. koraiensis occurred as several regeneration waves (1820s, 1850s, 1870–1880s, 1930s, and 1990–2000s) of moderate intensity. Shade-tolerant Jezo spruce (Picea jezoensis Carr. var. komarovii (V. Vassil.) Cheng et L.K.Fu) and Manchurian fir (Abies nephrolepis (Trautv.) Maxim.) regenerated continuously over the last 220 and 130 years, respectively. Enhanced recruitment of P. koraiensis, P. jezoensis, and A. nephrolepis was observed during the 1930s and 1990s, coinciding with increased growth release frequency in the 1920s and 1980s, and suggesting disturbance events of moderate intensity. Our results indicate that the current disturbance regime of the mixed Korean pine dominated forest maintains coexistence of light-demanding and shade-tolerant species and that change in wind climate may be particularly important for future forest composition.     

Spatial modelling of succession-disturbance dynamics in forest ecosystems: Concepts and examples

Over the last few decades it has become increasingly obvious that disturbance, whether natural or anthropogenic in origin, is ubiquitous in ecosystems. Disturbance-related processes are now considered to be important determinants of the composition, structure and function of ecological systems. However, because disturbance and succession processes occur across a wide range of spatio-temporal scales their empirical investigation is difficult. To counter these difficulties much use has been made of spatial modelling to explore the response of ecological systems to disturbance(s) occurring at spatial scales from the individual to the landscape and above, and temporal scales from minutes to centuries. Here we consider such models by contrasting two alternative motivations for their development and use: prediction and exploration, with a focus on forested ecosystems. We consider the two approaches to be complementary rather than competing. Predictive modelling aims to combine knowledge (understanding and data) with the goal of predicting system dynamics; conversely, exploratory models focus on developing understanding in systems where uncertainty is high. Examples of exploratory modelling include model-based explorations of generic issues of criticality in ecological systems, whereas predictive models tend to be more heavily data-driven (e.g. species distribution models). By considering predictive and exploratory modelling alongside each other, we aim to illustrate the range of methods used to model succession and disturbance dynamics and the challenges involved in the model-building and evaluation processes in this arena.     

Size distribution and expansion of canopy gaps in a northern Appalachian spruce-fir forest

Canopy gap area/age distributions and growth mechanisms were examined in a virgin subalpine forest in the White Mountains, New Hampshire, USA. The gap area distribution was negative exponential in form. Whithin gap tree ages varied widely in response to stepwise gap expansion caused by windthrow of peripheral trees or death of standing mature Picea rubens at gap edges. As a consequence, the density of small gaps may have been underestimated and the density of large gaps overestimated. The estimates of canopy turnover time, 303 yr, and of patch birth rate on an area basis, 3.3×10^-3 ha new patches/ha land area/yr, were not affected by the gap expansion phenomenon. However, any estimate of patch birth rate as numbers of new patches formed per year would have been too low. Because of increasingly widespread Picea death, the patch area/age distribution of this forest may not currently be in steady-state.     

Community dynamics of Ogawa Forest Reserve,a species rich deciduous forest,central Japan

Forest community dynamics were studied for 4 years in a 6 ha permanent plot of species rich, old-growth, temperate deciduous forest in Ogawa Forest Reserve, central Japan. The gap formation rate, recruitment, mortality, gain and loss rate in basal area during 4 years were 42 m2 ha^–1 yr^–1, 1.74% yr^–1, 1.19% yr^–1, 1.12% yr^–1 and 0.88% yr^–1, respectively. The turnover time calculated from them ranged from 58 to 240 years. Both the mortality and mortality factors were size dependent; trees in middle size class had smallest mortality, and the proportion of the trees killed by disturbances increased with size. Gap creations were concentrated in a particular year, suggesting a large heterogeneity in time. Spatial distribution of recruited trees were biassed to the old gaps (older than 4 years), especially that of the species with Bell-shaped dbh distribution (shade intolerant) strongly associated with the gaps. Recruitment in tree stems and the loss of basal area, thus had the larger variability than mortality of stems and this forest, and the species with L-shaped dbh distribution seemed to going to increase the importance in the future if the present trend continues to be held. The turnover time of population is positively correlated with the maximum dbh size of the species, indicating the slow change of the population of large sized species.     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.     

The role of the permanent soil seed bank in early stages of a post-agricultural succession in the Inland Pampa,Argentina

We studied the soil seed bank composition in four old fields of different ages, after abandonment from agriculture. Complete seed bank composition was assessed by direct seed separation from soil samples and identification to species. Most species found in the seed bank were not important in the present seral communities. Seed of the species that dominated the early succession were generally not found. Additionally, there were very few propagules rather than on the germination of in situ propagules. We suggest that pampean grasses evolved under that the course of post-agricultural succession will depend strongly on the pattern of arrival of exogenous propagules rather than the germination of in situ propagules. We suggest that pampean grasses evolved under disturbances of low intensity and/or a disturbance regime dominated by small gaps, in which open areas could be rapidly colonized from the edges and/or by remnant vegetative propagules. The changes produced by the introduction of agriculture triggered the invasion by exotic species adapted to the new disturbance regime.     

The role of anthropogenic influences in a mixed-conifer forest mortality episode

Abstract. Anthropogenic influences on recent tree mortality in mid-montane mixed conifer forests of southern California, USA, and northern Baja California, Mexico, were investigated. The Pinus jeffreyi-Abies concolor phase of the mixed-conifer montane community was sampled at three sites, characterized by: (1) low levels of air pollution and long-term fire suppression (Holcomb Valley in the San Bernardino Mountains, California); (2) chronic, severe air pollution and long-term fire suppression (Barton Flats in the San Bernardino Mountains, California); and (3) little air pollution and no fire suppression (La Corona Arriba in the Sierra San Pedro Martir in Baja California, Mexico). Similar percentages of dead standing trees, around 14 %, were found at both San Bernardino sites, but a comparatively low level, 4 %, was found in the Mexican forest. Anthropogenic effects, in particular fire suppression, may play an important role in enhancing the impact of natural stresses on the dynamics of mixed conifer forests.     

Spatial pattern of Quercus regeneration limitation and Acer rubrum invasion in a Piedmont forest

Abstract. Across eastern North America, there is a temporal trend from open Quercus forests to closed forests with increased Acer rubrum in the understory. We used a series of Ripley's K(d) analyses to examine changes in the spatial pattern of Quercus and Acer rubrum stems greater than 2.5 cm DBH over 45 yr in a 2‐ha mapped stand. Specifically, we asked whether changes over time were consistent with the hypothesis that Quercus is being competitively replaced by Acer rubrum. Both Acer rubrum and Quercus stems are spatially clumped, but have become less clumped over time. Stem mortality from Hurricane Fran (1996) was more clumped in all strata of the forest, at all spatial scales, than expected if damage had occurred to stems at random. Acer rubrum ingrowth occurred more often near established trees (all species) in the midstory, whereas Quercus ingrowth occurred less often near established trees in the midstory. The specific hypothesis that stems of Acer rubrum in the midstory of the forest are associated with a lack of Quercus regeneration was strongly supported. This effect occurred at all spatial scales tested, including scales larger than that at which direct competition for light can occur. Edaphic gradients in the plot are correlated with many of the observed trends at large spatial scales, and our results suggest that the presence of such gradients can generate complex spatial patterns over time.     

A note on the dynamics of montane virginPicea yezoensis forest in the Greater Khingan Mts,Northeastern China

The vertical growth ofLarix gmelini in the north of the Greater Khingan Mts., north-eastern China is very fast during the first 50 yr. ThoughPinus sylvestris var.mongolica shows a higher growth rate, it will be replaced by the larch population.Picea jezoensis is the most tolerant tree species and will replace all other trees.The fire cycle is 110–120 yr. Most of the montane areas from 800 to 1000 m a.s.l. are not occupied by spruce forest.There is a sapling bank with large numbers of saplings at ages below 20 yr in the understory in the spruce forest. The sapling bank is the key to maintain the steady state of the spruce forest. The persistent sapling bank in the life history of spruce governs the dynamics of the spruce forest controlled by the intensity and frequency of disturbances.The author is H. Q. Wu     

Community dynamics in relation to management of heathland vegetation in Scotland

The paper describes studies of post-fire succession in heathland vegetation in N.E. Scotland, dominated by Calluna vulgaris. A preliminary model (Legg, 1978) suggested good agreement between simulation of succession on the basis of a Markov chain and observations of stands at different stages of development after burning, at least in the earlier stages. Vegetation transitions are currently being recorded in permanent plots on burnt areas. First results confirm the view that (a) the post-fire succession has the properties of a Markov process, (b) this type of model remains valid when constructed from records of actual transitions, rather than data obtained by inference from evidence of transition. Comparing successional events in stands where, at the time of burning, the Calluna population was in pioneer-, building-, mature-and degenerate phases, shows that transition matrices generally agree with the Markov hypothesis, but not in the case of stands where Calluna was degenerate when burnt. The composition of establishing vegetation 1 year after fire is not confined to species normally associated with the early stages of succession, but reflects the composition of the stand before burning. Redevelopment after fire is described in terms of an initial floristic composition of species with strategies permitting early re-establishment, selected by the recurrence of the fire factor. Subsequent transitions represent changes in their relative abundance due to differing growth properties and competitive interactions. This interpretation applies only under conditions of recurrent incidence of fire (normally once in 10–15 yr). If fire does not recur, Calluna stands pass into the degenerate phase, where changes in the nature of relay floristics may come into play (e.g. with tree colonization).Nomenelature follows Clapham, Tutin & Warburg (1962) for vascular plants; Smith (1978) for bryophytes.     

Spatial and temporal dynamics of forest canopy gaps following selective logging in the eastern Amazon

Selective logging is a dominant form of land use in the Amazon basin and throughout the humid tropics, yet little is known about the spatial variability of forest canopy gap formation and closure following timber harvests. We established chronosequences of large‐area (14–158 ha) selective logging sites spanning a 3.5‐year period of forest regeneration and two distinct harvest methods: conventional logging (CL) and reduced‐impact logging (RIL). Our goals were to: (1) determine the spatial characteristics of canopy gap fraction immediately following selective logging in the eastern Amazon; (2) determine the degree and rate of canopy closure in early years following harvest among the major landscape features associated with logging – tree falls, roads, skid trails and log decks; and (3) quantify spatial and temporal differences in canopy opening and closure in high‐ and low‐damage harvests (CL vs. RIL). Across a wide range of harvest intensities (2.6–6.4 felled trees ha^?1), the majority of ground damage occurred as skid trails (4–12%), whereas log decks and roads were only a small contributor to the total ground damage (<2%). Despite similar timber harvest intensities, CL resulted in more ground damage than RIL. Neither the number of log decks nor their individual or total area was correlated with the number of trees removed or intensity of tree harvesting (trees ha^?1). The area of skids was well correlated with the ground area damaged (m²) per tree felled. In recently logged forest (0.5 years postharvest), gap fractions were highest in log decks (mean RIL=0.83, CL=0.99) and lowest in tree‐fall areas (RIL: 0.26, CL: 0.41). However, the small surface area of log decks made their contribution to the total area‐integrated forest gap fraction minor. In contrast, tree falls accounted for more than two‐thirds of the area disturbed, but the canopy gaps associated with felled trees were much smaller than for log decks, roads and skids. Canopy openings decreased in size with distance from each felled tree crown. At 0.5 years postharvest, the area initially affected by the felling of each tree was approximately 100 m in radius for CL and 50 m for RIL. Initial decreases in gap fraction during the first 1.5 years of regrowth diminished in subsequent years. Throughout the 3.5‐year period of forest recovery, tree‐fall gap fractions remained higher in CL than in RIL treatments, but canopy gap closure rates were higher in CL than in RIL areas. During the observed recovery period, the canopy gap area affected by harvesting decreased in radius around each felled tree from 100 to 40 m in CL, and from 50 to 10 m in RIL. The results suggest that the full spatial and temporal dynamics of canopy gap fraction must be understood and monitored to predict the effects of selective logging on regional energy balance and climate regimes, biogeochemical processes including carbon cycling, and plant and faunal population dynamics. This paper also shows that remote sensing of log decks alone will not provide an accurate assessment of total forest area impacted by selective logging, nor will it be closely correlated to damage levels and canopy gap closure rates.     

Uncertainty in detecting the disturbance history of forest ecosystems using dendrochronology

QuestionsUncertainty in detecting disturbance histories has long been ignored in dendrochronological studies in forest ecosystems. Our goal was to characterize this uncertainty in relation to the key parameters of forest ecosystems and sample size. In addition, we aimed to provide a method to define uncertainty bounds in specific forest ecosystems with known parameters, and to provide a required (conservative) minimal sample size to achieve a pre-defined level of uncertainty if no actual key forest parameters are known.LocationTraining data were collected from Žofínský Prales (48°40′N, 14°42′E, 735–830 m a.s.l., granite, Czech Republic).MethodsWe used probability theory and expressed uncertainty as the length (the difference between the upper and lower bounds) of the 95% confidence interval. We studied the uncertainty of (i) the initial growth of trees – if they originated under canopy or in a gap; and (ii) the responses to disturbance events during subsequent growth – on the basis of release detection in the radial growth of trees. These two variables provide different information, which together give a picture of the disturbance history. While initial growth date the existence of a gap in a given decade (recent as well as older gaps are included), release demonstrates the moment of a disturbance event.ResultsWith the help of general mathematical deduction, we have obtained results valid across vegetation types. The length of a confidence interval depends on the sample size, proportion of released trees in a population, as well as on the variability of tree layer features (e.g., crown area of suppressed and released trees).ConclusionsMost studies to date have evaluated the initial growth of trees with higher uncertainty than for canopy disturbed area. The length of the 95% confidence interval for detecting initial growth has been rarely shorter than 0.1 (error ± 5%) and has mostly been much longer. To reach 95% confidence interval length of 0.1 (error ± 5%) when detecting the canopy disturbed area, at least 485 tree cores should be evaluated in studied time period, while to reach a 0.05 interval length (error ± 2.5%) at least 1925 tree cores are required. Our approach can be used to find the required sample size in each specific forest ecosystem to achieve pre-defined levels of uncertainty while detecting disturbance history.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

吉林蛟河不同演替阶段针阔混交林木本植物幼苗空间分布与年际动态

幼苗的空间分布对群落更新具有重要意义。于2011年夏季在蛟河地区3个不同演替阶段针阔混交林样地内共设置了451个种子雨-幼苗观测样站,并于2012—2014年对幼苗样方内胸径1 cm的木本植物幼苗进行了连续3a的调查。对木本植物幼苗的数量分布以及水曲柳(Fraxinus mandshurica)、五角枫(Acer mono)、杉松(Abies holophylla)和东北枫(Acer mandshuricum)4个主要树种幼苗与大树的空间关系及其年际变化进行了分析,并用Syrjala检验分析了样站尺度上幼苗密度、丰富度空间分布在年际间的差异。结果显示:(1)木本植物幼苗的数量分布在不同群落和年际之间都表现出明显的差异,样站尺度上3个样地内的幼苗密度存在着较大的空间变异性,而幼苗丰富度的空间异质性较低。表明幼苗分布的空间异质性对幼苗密度有着重要影响。(2)样站尺度上幼苗数量和物种数的空间分布在不同年份之间是存在差异的。在一定程度上证实了种子产量、扩散方式和群落组成对幼苗空间分布的影响。(3))从死亡幼苗与大树空间关系来看,4个主要树种的死亡幼苗与成树表现出相似的分布格局,这既表明成体植株的空间分布特征也能够影响幼苗的空间分布格局,也从侧面说明了负密度制约效应对幼苗空间分布的影响。     

Influence of canopy opening on the environment and herb layer in a northern hardwoods forest

Single- (33–37 m²) and multi-tree (51–151 m²) gaps were created in an Allegheny Plateau northern hard-woods forest to investigate environmental and herb layer response to canopy opening. After gap creation, noon light on clear summer days was brightest north of opening center. At other times of the day, and when skies were overcast, there was no difference in the light quantity beneath opened and closed canopy. Nor was the distribution of soil moisture or of soil or air temperature greatly affected by gap creation. Species establishment tended to be higher near opening centers; otherwise, there was no pronounced effect of canopy opening on plant cover or species richness during the first four years after gap creation. Biotic responses were not significantly correlated with any environmental factor.     

吉林蛟河针阔混交林乔木幼苗组成与月际动态

以吉林省蛟河42 hm²针阔混交林动态监测样地为平台, 在样地内选取224个1 m × 1 m幼苗监测小样方, 基于2013年6至8月每月中旬的幼苗监测数据, 对乔木幼苗物种组成、数量特征、月际动态及幼苗与同种大树之间的关系进行分析。结果表明: (1)该群落共调查到乔木树种幼苗16种。物种组成在6月和7月间变化不大, 但与8月差异较大。物种组成在各样方间表现出极大的差异性, 不同种幼苗出现的样方数为1-159。(2) 6月至8月, 幼苗个体从1722株减少到1214株, 降幅达29.5%。不同种幼苗在个体数量上变异较大, 个体数超过100的幼苗有色木槭(Acer mono)、水曲柳(Fraxinus mandschurica)、紫椴(Tilia amurensis)和东北槭(Acer mandshuricum), 四者个体数之和占所有物种个体总数的84.26%; 不同种幼苗密度变异性较大, 只有色木槭、水曲柳、紫椴和东北槭密度大于1株·m^-2, 其他种幼苗密度均较低。(3)新生幼苗更新、死亡格局表现出明显的种间差异与月际间差异, 7月新生幼苗总体死亡率(48.9%)显著高于8月(28.3%); 整个调查季(6-8月), 新生幼苗总体死亡率(40.56%)明显高于多年生幼苗(7.34%)。就幼苗更新情况而言, 该研究群落一年中更新主要集中在6月。(4)色木槭、紫椴、杉松(Abies holophylla)幼苗的密度与同种成年个体的胸高断面积之和之间存在显著的正相关关系(p = 0.006、0.013、0.037), 表明三者幼苗密度随周围母树多度的增加而增加, 其他物种幼苗密度与同种个体数目或同种胸高断面积之和未表现出显著的相关关系。