Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Changes in spatial pattern of trees and snags during structural development in Picea mariana boreal forests

Questions : How do gap abundance and the spatial pattern of trees and snags change throughout stand development in Picea mariana forests? Does spatial pattern differ among site types and structural components of a forest? Location : Boreal forests dominated by Picea mariana, northern Quebec and Ontario, Canada. Methods : Data on the abundance, characteristics and spatial location of trees, snags and gaps were collected along 200 m transects at 91 sites along a chronosequence. Spatial analyses included 3TLQV, NLV and autocorrelation analysis. Non‐parametric analyses were used to analyse trends with time and differences among structural components and site types. Results : Gaps became more abundant, numerous and more evenly distributed with time. At distances of 1–4 m, tree cover, sapling density and snag density became more heterogeneous with time. Tree cover appeared to be more uniform for the 10–33 m interval, although this was not significant. Patch size and variance at 1 m were greater for overstorey than for understorey tree cover. Snags were less spatially variable than trees at 1 m, but more so at intermediate distances (4–8 m). Few significant differences were found among site types. Conclusions : During stand development in P. mariana forest, gaps formed by tree mortality are filled in slowly due to poor regeneration and growth, leading to greater gap abundance and clumping of trees and snags at fine scales. At broader scales, patchy regeneration is followed by homogenization of forest stands as trees become smaller with low productivity due to paludification.     

Differential leaf traits of a neotropical tree Cariniana legalis (Mart.) Kuntze (Lecythidaceae): comparing saplings and emergent trees

Cariniana legalis is an emergent tree that reaches the upper canopy in Brazilian Semideciduous Forest. Spatial contrasts in microclimatic conditions between the upper canopy and understorey in a forest may affect morpho-physiological leaf traits. In order to test the hypothesis that the upper canopy is more stressful to leaves than a gap environment we compared emergent trees of C. legalis, 28–29 m in height to gap saplings, 6–9 m in height, for the following parameters: leaf area, leaf mass area (LMA the dry weight:leaf area ratio), leaf thickness, leaf anatomical parameters, stomata conductance, and chlorophyll a fluorescence. Leaves from emergent trees had smaller leaf areas but greater LMA compared to saplings. Leaf thickness, palisade layer thickness, and stomatal density were higher for emergent trees than for saplings. The opposite pattern was observed for spongy layer thickness and spongy/palisade ratio. Stomatal conductance was also higher for emergent tree leaves than for sapling leaves, but the magnitude of depression on stomatal conductance near midday was more pronounced in emergent trees. The potential quantum yield of photosystem II, as determined by the F _v/F _m ratio was lower for leaves from saplings. The lower values of stomatal conductance, indicating restriction in CO₂ diffusion into the mesophyll can be related to higher photoinhibition observed in the saplings. Leaves from emergent trees and saplings exhibited similar values for apparent electron transport rates and non-photochemical quenching. Our results suggest that changes in leaf traits could be associated to dry conditions at the upper canopy as well as to the ontogenetic transition between sapling/emergent tree life stages.     

川西米亚罗亚高山暗针叶林岷江冷杉和糙皮桦空间格局及其关联性分析

基于2019年9月完成的川西米亚罗亚高山暗针叶林9.6 hm~2动态样地的调查数据,分析了主要优势种岷江冷杉(Abies fargesii var.faxoniana)与糙皮桦(Betula utilis)的径级结构,并运用O-ring统计中的成对相关函数g(r)和g₁₂(r),研究了两个优势种的空间格局及其种内、种间的关联性。结果表明：岷江冷杉与糙皮桦径级分布连续,均呈倒“J”型。岷江冷杉与糙皮桦的不同径级个体主要呈现由聚集分布向随机分布发展的趋势。岷江冷杉的小树对幼树、中树对幼树、中树对小树以及大树对中树的种内相关性在所有空间尺度上均为显著正相关(P<0.05),而相同径级糙皮桦的种内相关性则表现为由显著正相关(P<0.05)逐渐向无显著相关(P>0.05)变化。随着空间尺度的增大,岷江冷杉的大树对幼树、大树和小树由无显著相关(P>0.05)逐渐向显著正相关(P<0.05)变化,糙皮桦的大树对幼树由显著负相关(P<0.05)逐渐向无显著相关(P>0.05)变化。岷江冷杉大树对糙皮桦所有径级个体的种间相关性主要呈现由为显...     

Estimation of tree replacement patterns in an Appalachian Picea-Abies forest

Abstract. Patterns of tree species replacement in a Picea-Abies forest, determined by several different methods, are compared and the methods are assessed. Methods are grouped as either understory-based or gap-based estimates of replacement. The understory-based methods characterize canopy-understory interactions with spatial statistics, sapling density measurement, sapling frequency measurement, and successor sapling identification beneath live canopy trees. The gap-based methods include sapling density measurement, sapling frequency measurement, and successor sapling identification in tree-fall gaps. Methods except those based on frequency indicate a strong trend of replacement of all canopy species by Abies. Understory-based methods may underestimate canopy recruitment of intolerant trees, while gap-based methods relying on sapling density or frequency may overestimate recruitment of intolerant trees. Estimates based on the selection of successor saplings in the understory or in gaps are reliable. Gap successor estimates consider the process of gap capture and are useful in analyses of forest dynamics.     

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     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Successional trends 219 years after fire in an old Pinus sylvestris stand in northern Sweden

Abstract. We present results from repeated analyses (1962, 1993) of a permanent plot established in 1947, combined with retrospective stand age structure data, in an old Pinus sylvestris stand in Muddus National Park, northern Sweden. The study points towards a successional pathway governed by concurrent disturbance effects of climate variability, reindeer grazing and fire. This is intermediate to the two often advocated ideas on dynamics in boreal forests, that is, one of disturbance-related tree regeneration/mortality and one of continuous regeneration. When the plot was established in 1947 the tree layer (> 1.3 m) consisted of 300 individuals/ha of P. sylvestris and 62/ha of Betula pubescens. Subsequently the stand has become more dense and the species dominance has shifted. In 1993, 362 P. sylvestris and 62 Picea abies individuals were present per ha, while no Betula individuals were found. The number of dead trees increased from zero in 1947 to 200/ha (Pinus) in 1993. Pinus was also the most common species in the sapling layer (< 1.3 m) throughout the study period, though the number dropped from 8912/ha in 1947 to 51% in 1993. Dead saplings decreased from 2650/ha in 1947 to ca. 50% in 1962, and only 9% in 1993. Temporal variations in mortality and sapling mean height coincided with variations in snow depth, indicating a critical period in sapling development when saplings are exposed at the snow/atmosphere interface. The number of living Picea saplings increased slowly until 1993; no dead saplings were found. Most Pinus recruited shortly after the 1774-fire, and during the second half of the 1900s. The major part of the spruce regeneration took place during the later half of the 1900s. No successful Betula recruitment has occurred after the 1930s, and no live Betula were present in 1993, which might be explained as an effect of increased reindeer browsing – the reindeer stock has grown by 50% since 1961. Although subjected to high mortality, Pinus regenerated and maintained a seedling/sapling bank. In this way Pinus remained dominant in the tree layer after more than 200 post-fire years. The importance of the shade-tolerant Picea has slowly increased, while Betula has died off. Thus, even after 219 yr since fire there is an early successional trend in the stand. This suggests that an increased chronic disturbance (grazing/browsing by reindeer) has partly succeeded earlier discrete fire-disturbance events, and maintained a continuous seedbed favouring the shade-intolerant pine recruitment.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Scale‐dependent intraspecific competition of Taurus cedar (Cedrus libani A. Rich.) saplings in the Southern Turkey

Better understanding of the competitive interaction at the early development stages of the stand is crucial to help schedule silvicultural treatments for young stands and for the better management of the future stands. We used scale‐dependent analysis to improve our understanding of sapling dynamics in the pure Taurus cedar (Cedrus libani A. Rich.) stands in Southern Turkey. Using data from nine plots established at the western Taurus Mountains, diameter, height, and crown radii of saplings were compared, and spatial point pattern analyses were performed. We found significant differences for the mean diameter and height, and crown radii of saplings among the plots. Univariate pair correlation function showed that sapling pattern was regular only at small scales (r < 0.4 m) but was predominantly random. Bivariate pair correlation function revealed no evidence of spatial interaction between tall saplings and short saplings. Univariate mark correlation function revealed that strong intraspecific competition was detected at small scales (up to 0.55 m). This distance is reasonable for the juvenile age tending of Taurus cedar saplings and should be under consideration during silvicultural treatments to use the site productivity more efficiently.     

Spatial and temporal patterns of rowan (<Emphasis Type="Italic">Sorbus aucuparia</Emphasis> L.) regeneration in West Carpathian subalpine spruce forest

Non-random seed shadows are commonly seen in plant species whose seeds are dispersed by animals, in particular by birds. The behaviour of birds can influence the spatial pattern of seed dispersal and, consequently, the entire regeneration process of fleshy-fruited trees. This study examined regeneration patterns in a fleshy-fruited tree species, rowan (Sorbus aucuparia L.), growing in West Carpathian subalpine spruce forests, focussing on two problems: the temporal relationship between rowan regeneration and gap formation, and the spatial relationship between rowan regeneration and stand structure. It was found that rowan seedlings and saplings were recruited in advance of gap formation. Establishment of new rowan individuals in gaps was infrequent, but gaps enhanced their regeneration nearby under spruce canopy, where they occurred densely in a narrow belt about 15 m wide. Inside spruce stands, the highest density of young rowans was directly under crowns, especially near trunk bases. Few rowan saplings were found growing under mature rowan trees. The presence of a rowan seedling and sapling bank determines whether rowans fill spruce stand gaps. Dense rowan groves can develop mainly in extensive but slowly expanding gaps.     

天山云杉天然林不同林层的空间格局和空间关联性

天山云杉(Picea Schrenkiana)林是天山林区的重要针叶林。研究调查了分别代表天山云杉增长型种群和成熟型种群的两块标准地,应用Ripley's K系数法研究了天山云杉更新层、低林层、中林层和高林层的空间格局和不同林层的空间关联性,并应用Moran's I系数法研究了不同林层的林木高度的空间自相关性。结果表明:所有林层的天山云杉活立木都是显著聚集的,只是发生聚集格局的强度和聚集尺度不同。更新层天山云杉具有最大的聚集强度,从更新层到低林层,聚集强度急速降低。两块标准地中天山桦(Betula tianschanica Rupr.)、天山柳(Salix tianschanica Rgl.)和天山云杉活立木的空间关联性的表现不同。研究结果表明:增长型种群中天山桦、天山柳在所有尺度上都抑制天山云杉的生长;而成熟型种群中天山桦、天山柳在1-10 m尺度内对天山云杉种群的生长没有影响,10-40 m表现出正相关,40 m以上没有表现出明显的关联性;同时天山桦、天山柳基本上在所有尺度都抑制天山云杉的更新。在更新层和低林层间表现出正相关;而更新层和枯死木之间的空间关联性为正相关。     

Sapling communities in Amazonian white-water forests

Aim Structure and floristic composition of forest regeneration (trees between 1 and 10 cm diameter at breast height (d.b.h), ≥ 1 m growth height) was described in three forest types of Amazonian white‐water forests (várzea), in order to analyse whether floristic composition of saplings is related to the successional stage of the forests, whether it differs in comparision with the mature flora, and if there exists a zonation of sapling species along the gradients of flooding and irradiation. Location Mamirauá Sustainable Development Reserve, Western Brazilian Amazon. Methods The investigated forests were of the low and the high várzea type, on an annual average flooded around 4 and 1 months, respectively. The two low‐várzea forests belonged to the secondary and to the late‐successional stage, the high‐várzea forest also to the late‐successional stage. A total of 24 circular sample plots covering 1885 m² were installed. They were nested within three rectangular 1 ha permanent sample plots where individuals ≥ 10 cm d.b.h. were formerly inventoried. Average inundation and radiation [relative photosynthetically active radiation (rPAR)] at the forest floor was recorded in all inventoried plots. The Mean spatial Distribution Center (MDC; Ebdon, 1998 ) for all sapling species was calculated. Sapling species were grouped into associations with respect to their light‐demand and their location along the flood‐level gradient. Results Average flood height and average rPAR at the forest floor was highest in the secondary stage of the low várzea and lowest in the high‐várzea forest. Overall average density and basal area of the saplings averaged 2250 individuals and 2.3 m² ha^?1 in the secondary stage, 2330 individuals and 2.6 m² ha^?1 in the late‐successional stage of the low várzea and 5000 individuals and 4.8 m² ha^?1 in the high várzea. In all forest types, species richness of saplings was lower than species richness of trees ≥ 10 cm d.b.h., and amounted to 25 species in the secondary stage, to 35 species in the late‐successional stage of the low várzea and to 88 species in the high‐várzea forest. The amount of ‘immigrants’ ( Bazzaz, 1991 ) at the sapling level increased with proceeding forest succession and amounted to 24 and 29% in the low várzea and to 31% in the high várzea. Sapling species distribution, species richness and individual density was linked to both, the gradient of flooding and the gradient of irradiation. Species richness and individual density was highest in the high várzea and decreased with increasing influence of flooding. The high várzea was predominated by pronounced shade‐tolerant sapling species, whereas in the low várzea occured both, light demanding and shade‐tolerant species groups. Main conclusions Influence of flooding seems to be the main factor triggering species composition and structure of sapling communities in várzea forests. Proceeding forest succession reduces the impact of flooding because of the biogenical induced silting up of the forested sites. However, forest succession also alters forest architecture of the overstory and such the light conditions at the sapling level. Therefore, radiation is an important factor influencing species composition of sapling communities in várzea forests.     

青藏高原东缘林线杜鹃-岷江冷杉原始林的空间格局

为深入理解高山林线原始林中优势树种的空间结构特征和种间关系,以青藏高原东缘林线杜鹃-岷江冷杉原始林1 hm²样地调查数据为基础,采用成对相关函数g(r)函数,分析了优势种岷江冷杉(Abies faxoniana)和凝毛杜鹃(Rhododendron agglutinatum)各径级的空间分布格局和各径级间的空间关联性。结果表明:岷江冷杉和凝毛杜鹃径级分布连续,呈倒"J"型,均为进展种群。小尺度上,对于岷江冷杉活立木(小树+中树+大树)、小树、中树为显著聚集分布,大树为均匀分布,死树则为随机分布;凝毛杜鹃活立木(小树+中树+大树)、小树、中树、大树、死树均为显著聚集分布。小尺度上随径级增大岷江冷杉的分布格局从聚集分布趋于均匀分布。岷江冷杉大树与其小树的空间关联验证了Janzen-Connell假说中的距离制约效应。通过验证接受了随机死亡假说,发现种内的密度制约效应不明显。两个树种的枯立木对凝毛杜鹃小树均表现为排斥关系,而对岷江冷杉小树表现为相互独立的关系。两个优势种总体表现为相互独立,但种间不同径级间的空间关联性各异。     

Disturbance processes and spatial patterns of two emergent conifers in New Caledonia

Abstract Araucaria laubenfelsii Corbasson and Araucaria montana Brongn et Gris are emergent conifers in maquis and forest communities which are subjected to a combination of fire and cyclonic disturbances. Both species are able to survive fire once stems reach a critical size, but most seedlings and saplings are killed. Both species were found to be clumped at most spatial scales for both saplings and trees in maquis, probably resulting from a combination of patchy fire and a limited ability to disperse seeds. Comparisons of burned and unburned A. montana sites suggest that clumping increases only slightly after fire. The degree of clumping in the A. laubenfelsii sites, suggested that fire disturbances may occur relatively frequently in the region. Cyclonic disturbances are infrequent but may result in blow‐down of large individuals within both maquis and forest. All of the individuals blown‐down during the study had been previously fire‐scarred. Around tree blow‐downs, seedling and sapling densities can be high. This likely reflects both the low dispersability of Araucaria seeds and enhanced moisture from the shading of the adult, when it was alive. Disturbance by fire and wind play an important role in the regeneration dynamics and spatial pattern of these species in maquis in New Caledonia.     

Limits of pine forest distribution at the treeline: herbivory matters

Herbivores can affect future forest composition by feeding selectivity. At temperature-sensitive treelines, herbivory can exacerbate or constrain climate-driven distributional shifts in tree species. This study analyses the impact of herbivory in a Mediterranean treeline of widespread Pinus sylvestris and P. nigra pinewoods, testing whether herbivory damage reinforces or inhibits the climatic responses of these trees. We used naturally occurring sapling pairs of similar size and age of both species, thereby isolating plant characteristics from environmental effects in herbivore behaviour. Herbivory damage by ungulates proved higher than that caused by insects in saplings of both species. Low plant density and extreme abiotic conditions at the treeline could in part be responsible for the observed low incidence of insect herbivory. Ungulates preferred P. sylvestris over P. nigra, implying heavier browsing damage for a large number of P. sylvestris saplings, suffering reduced internode growth as a consequence. In addition, P. sylvestris could not compensate height-growth reductions due to browsing with higher growth rate than P. nigra. In fact, P. sylvestris showed similar or lower relative height growth with respect to P. nigra. Under a scenario of increasing aridity and maintenance of ungulate populations, the upward migration of P. sylvestris in its southern range could be restricted by higher drought vulnerability than P. nigra, a situation exacerbated by ungulate herbivory. Our results indicate that ungulate herbivory reinforces climatic response of coexisting P. sylvestris and P. nigra at treeline, favouring a potential change in community dominance towards Mediterranean P. nigra.     

Seed supply, drought, and grazing determine spatio-temporal patterns of recruitment for native and introduced invasive pines in grasslands

Aim The rate of grassland invasion by trees depends on the ability of the species to invade, i.e. their invasiveness, and on the susceptibility of the environments to invasion, i.e. their invasibility. Knowledge of the invasiveness of native and introduced tree species and of the environmental factors that contribute to invasibility is necessary to understand landscape evolution and assess required management measures. Our main aim was to explore this by estimating the separate effects of propagule pressure and environmental factors on the spatio‐temporal patterns of sapling recruitment, a key stage in the tree life cycle. Location Causse Mejean calcareous plateau (southern France). Methods The effects of seed supply and environmental variables (grazing, geological substrate, and duration or intensity of drought) on the spatio‐temporal patterns of sapling recruitment were assessed for the native Scots pine (Pinus sylvestris L.) and the introduced black pine (Pinus nigra Arn. ssp. nigra). Estimates were derived by inverse modelling with data of locations and ages of 4‐ to 20‐year‐old saplings and seed‐bearing trees in 32 sites. Yearly indices of drought were derived from a soil–water content model. Results For both species, seed supply was as important as the whole set of environmental factors in explaining sapling recruitment rates. Grazing and the duration of drought from July to August decreased sapling recruitment rates, which were also lower on hard limestone than on dolomite. Dispersal distances and effective fecundities were higher for the introduced P. nigra, which was less susceptible to drought but more affected by grazing than the native P. sylvestris. In grazed grasslands, shrubs facilitated sapling establishment of both species. Main conclusions This study shows how seed supply and environmental factors shape spatio‐temporal patterns of sapling recruitment for trees invading grasslands. Implications for landscape evolution and management, of the difference in invasiveness of the two pine species and of the hierarchy of environmental factors in determining invasibility, are discussed.     

Disturbance affects spatial patterning and stand structure of a tropical rainforest tree

The distribution and spatial patterns of plant populations in natural ecosystems have recently received much attention; yet the impacts of human‐induced disturbances on these patterns and underlying processes remain poorly understood. We used the sub‐canopy tree, Ryparosa kurrangii (Achariaceae), to explore the possible effects of such disturbances on stand structure and spatial patterning in an Australian tropical rainforest. We studied three populations that differed in their extent of habitat modification: anthropogenic disturbance (proximate settlement and roads) and internal damage by an invasive alien species, the feral pig (Sus scrofa). Populations were mapped, characterized, and three size cohorts (seedlings, saplings, trees) were analysed using a suite of spatial point pattern analyses (univariate: Diggle's G and F and Ripley's K; bivariate: Diggle's G and Ripley's K). Ryparosa kurrangii has a typical stand structure for a sub‐canopy tree species, but occurs at high densities locally (>400 stems ha^?1). At all sites, the tree cohort were randomly distributed and saplings were spatially aggregated at distances of up to 2–3 m. Between sites there were distinct differences in the size structure and spatial pattern of seedlings, the cohort most affected by recent habitat modification. That is, the least disturbed site had no aggregation among seedlings, the site with the greatest anthropogenic disturbance had many small, clustered seedlings that were spatially associated with trees, and the site with pig damage had clustered seedlings that had no spatial relationship with trees. We propose that habitat modification by anthropogenic and pig disturbance disrupts seed dispersal and establishment regimes, which leads to altered seedling spatial patterns. These disturbances could have long‐term implications for the population structure and health of R. Kurrangii.     

Disparate distribution patterns between canopy and subcanopy life-forms in two temperate North American forests

Quantitative vegetational data of canopy and woody subcanopy species (two life-forms adapted to occupy different strata at maturity) were compared with data collected in two temperate forest ecosystems to determine whether they exhibit a similar pattern of distribution. Tidal freshwater swamps (21 stands) and southern Appalachian forests (19 stands) were examined from data obtained using identical sampling methods. Separate structural analyses of the canopy, sapling, and subcanopy species were compared using the indirect ordination algorithm Detrended Correspondence Analysis. Environmental measurements collected in each stand were assessed for their relationship to the distribution of stands depicted by the ordination diagrams.Canopy trees and saplings showed a similar pattern of distribution, suggesting that the resource requirements of saplings and canopy-statured adults are similar. In contrast, the subcanopy species (species genetically adapted to an understory existence, i.e., shrubs and small understory trees) of neither ecosystem showed any discernable distributional relationship to the canopy or sapling layers (in tidal swamps, there was no clear way to even segregate subcanopy stands into communities). Environmental gradients associated with the subcanopy ordinations differed from those of the canopy and sapling strata in both forest systems, suggesting that subcanopy species partition different resources than do canopy species.If a lack of similarity in distribution patterns between canopy and subcanopy species is universal in temperate forests, then the common practice of combining sapling and subcanopy species in structural analyses may hinder our understanding of subcanopy structural patterns in forests.     

Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.