Recruitment Processes and Species Coexistence in a Sub-boreal Forest in Northern Japan

We investigated the recruitment of saplings (across the 2 m-heightthreshold) of six species,Picea jezoensis, Abies sachalinensis,Betula ermanii, Picea glehnii, Acer ukurunduense andSorbus commixta,in a sub-boreal forest, northern Japan. Data were collectedin a 2.48-ha plot over six growing seasons (1989–1994).We used path analysis to analyse the relationships between therecruitment rates of saplings and the stand structural attributessuch as mother tree abundance, stand crowdedness, stand stratification,Sasabamboo density on the forest floor, and fallen log abundance.The combination of stand structural attributes affecting recruitmentrates of the six sub-boreal forest tree species differed markedlyamong the species and corresponded to species composition. Itis suggested that the size-structure dynamics of adult treesof the sub-boreal forest are regulated largely by differentregeneration processes among the species and only slightly byinterspecific competition between adult trees because interspecificcompetition between adult trees was not evident. The dynamicsof species coexistence of the sub-boreal forest should be describedas a process combining the diversity of recruitment processesof saplings of the component species and the diversity of interspecificcompetition between adult trees. We propose the boundary conditionhypothesis for species coexistence in the sub-boreal forest,that the persistence of each component species is ascribed largelyto the different recruitment processes of saplings (boundaryconditions for adult tree growth dynamics) and only a littleto interspecific adult tree competition. Climax forest; safe site; regeneration niche; mode of competition; species diversity     

北京地区油松人工林不同演替类型空间结构对林下植被及土壤的影响

以北京地区油松(Pinus tabuliformis)人工林不同演替类型林分为研究对象,研究油松纯林、油松-栓皮栎(Quercus variabilis)混交林和栓皮栎纯林三种不同演替类型林分的空间结构、林下植被和土壤水分的变化规律及其相互作用关系。结果表明：(1)林分水平及垂直空间结构、草本层物种多样性、更新幼树生长、土壤持水和透气性能等指标在三种不同演替类型林分间差异显著(P<0.05),林分空间结构参数中的角尺度、林层指数和开敞度显著影响了各类型林分的灌草多样性,混交度、林层指数和大小比数显著影响了更新幼树的生长,混交度和林层指数显著影响了土壤水分的变化(P<0.05)。(2)松栎混交林灌草生物量、天然更新幼树的生长以及土壤水分物理状况均好于纯林,并主要受林分混交度和林层指数的共同作用。(3)各演替类型林分内均存在栓皮栎更新幼树,混交林栓皮栎更新幼树数量最多、长势最好,对林地资源的竞争最为激烈。因此,可以通过调整林分空间结构实现种间关系及林地资源的调控,以充分发挥森林生态系统的各项功能与价值。     

藏东南川滇高山栎种群不同生长阶段的空间格局与关联性

运用点格局分析方法中Ripley L函数,对藏东南川滇高山栎种群不同生长阶段的空间格局及其关联性进行分析.结果表明： 藏东南川滇高山栎种群径级结构呈单峰型,小树与中树的数量在整个种群中占绝对优势,整个种群自然更新状况良好,属增长型种群.川滇高山栎种群在从小树发育到大树的过程中,小树、中树在小尺度上均呈明显的聚集分布,而大树在整个尺度上基本上呈随机分布.小树与中树、大树间在小尺度上有显著的相关性,而中树与大树不相关.川滇高山栎种群在从小树、中树发育到大树的过程中,其空间格局由聚集分布到随机分布.川滇高山栎种群的自然更新受种间竞争的影响,也受种内竞争的影响.在自然环境条件相似下,影响川滇高山栎种群分布格局的最主要因素是其自身的生物、生态学特性.     

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.     

The interspecific relationships of seaweeds and their role in the formation of communities of Ahnfeltia tobuchiensis (Kanno et Matsubara, 1932) Makienko, 1970 (Rhodophyta)

The character and intensity of interspecific relationships between common species of an Ahnfeltia bed community (Ahnfeltia tobuchiensis, Ahnfeltiopsis flabelliformis, Chondrus armatus, and Ptilota filicina) were investigated in relation to light levels and concentrations of dissolved inorganic nitrogen and phosphorus compounds. Growth and photosynthetic rates of seaweeds were measured in mono and co-cultures, as well as in a medium containing the metabolites of attendant species. Under low light conditions, A. tobuchiensis showed the highest competitive ability among the species we studied. The experimental results indicated both negative (exploitative competition, allelopathic inhibition of production processes of a competitor) and positive (stimulation of growth and photosynthesis of accompanying species) relationships between seaweeds. Increased nutrient concentrations and light levels resulted in a substantial reduction in the intensity of interspecific relationships. The dynamic system of positive and negative relationships may regulate the A. tobuchiensis community structure; the intensity of interspecific relationships depends on the availability of resources such as light and nutrients.     

The relative importance of above- versus belowground competition for tree growth during early succession of a tropical moist forest

Competition between neighboring plants plays a major role in the population dynamics of tree species in the early phases of humid tropical forest succession. We evaluated the relative importance of above- versus below-ground competition during the first years of old-field succession on soil with low fertility in Southern Mexico, using the premise that competition for light is size-asymmetric, unlike competition for nutrients. Plant growth is thus expected to be disproportionally impeded by larger neighbors. We studied how growth and survival of 3.5–5.5 m tall saplings of Cecropia peltata and Trichospermum mexicanum, two pioneer species that dominate the secondary forests in the study region, varied with the abundance and size of neighboring trees in 1–2 year old secondary vegetation. We found that local neighborhood basal area varied 10-fold (3 to 30 cm² m^-2) and explained most of the variation in diameter and height growth of the target saplings. Most growth variables were strongly affected by the neighbors bigger than the focal trees with no significant additive effect of the smaller neighbors, indicating asymmetric competition. Smaller neighbors did have a small but significant additive effect on the diameter growth of Cecropia saplings and stem slenderness of Trichospermum saplings. We conclude that competition for light was more important than belowground competition in this initial phase of moist tropical forest successional, despite the low soil fertility.     

湖北木林子保护区山矾属植物空间分布格局及其关联性

物种共存机制是生态学研究的重要问题。以同属物种山矾、光叶山矾为研究对象,分析其空间分布格局和种内、种间关联性特征。将山矾、光叶山矾分为4个径级,分别归属幼树、小树、中树和成年树4个生长阶段。采用单变量成对相关函数分析空间分布格局特征;双变量成对相关函数分析种内、种间关联性。结果显示：山矾和光叶山矾径级结构基本一致,均属增长型种群。在完全空间随机模型下,山矾和光叶山矾主要呈聚集分布,逐渐过渡为均匀分布;在异质泊松模型下,只在小尺度上聚集,较大尺度范围呈随机分布。种内关联上,山矾各生长阶段主要呈正关联,逐渐向无关联过渡,光叶山矾除小树与成年树外,其余均呈正关联;排除生境异质性影响后,小尺度呈正关联,其余尺度无关联。种间关联上,在幼树、小树物种对中主要呈负关联,中树、成年树物种对中多为无关联。研究表明,同属物种山矾和光叶山矾呈种内聚集、种间分离的空间构型,在竞争排除作用下实现同属共存,有利于亚热带常绿落叶阔叶混交林群落结构稳定和物种多样性维持。     

沙地樟子松天然林南缘分布区林木竞争、空间格局及其更新特征

以呼伦贝尔沙地樟子松地理分布南缘天然林为研究对象,采用Hegyi竞争指数与点格局等分析方法,研究了沙地樟子松天然林的种群结构、种内竞争、空间格局及幼树更新等特征。结果表明:1)所调查区域天然沙地樟子松纯林处于中幼龄阶段,属增长型种群;2)其竞争指数与对象木胸径服从幂函数关系CI=242.24D~(-1.12)(R~2=0.91);3)幼树在小尺度上呈聚集分布,中树与大树在中大尺度上呈随机分布;幼树与中树在小尺度上呈正相关性,中树与大树在小尺度呈负相关性;4)竞争指数与更新幼树和存活更新幼树的密度均呈显著正相关性。在林分管理中需要充分考虑林木竞争、空间格局以及种群更新的关系,本研究可为沙地樟子松天然林的经营管理与保护提供重要的科学依据。     

Unusual Fine Root Distributions of Two Deciduous Tree Species in Southern France: What Consequences for Modelling of Tree Root Dynamics?

The spatial distribution of fine roots of two deciduous tree species was investigated in contrasting growing conditions in southern France. Hybrid walnut trees (Juglans regia×nigra cv. NG23) and hybrid poplars (Populus euramericana cv. I214) were both cultivated with or without annual winter intercrops for 10 years on deep alluvial soils. Soil samples for measuring the fine root distribution of both trees and crops were obtained by soil coring down to 3-m depth at several distances and orientations from the tree trunk. The distribution of live fine roots from walnut and poplar trees was patchy and sometimes unexpected. In the tree-only stands, fine root profiles followed the expected pattern, as fine root density decreased with increasing depth and distance from the tree trunk. However, many fine root profiles under intercropped trees were uniform with depth, and some inverse profiles were observed. These distributions may result from a high degree of plasticity of tree root systems to sense and adapt to fluctuating and heterogeneous soil conditions. The distortion of the tree root system was more pronounced for the walnut trees that only partially explored the soil volume: in the tree-only stand, the walnut rooting pattern was very superficial, but in the intercropped stand walnut trees developed a deep and dense fine root network below the crop rooting zone. The larger poplars explored the whole available soil volume, but the intercrop significantly displaced the root density from the topsoil to layers below 1 m depth. Most tree root growth models assume a decreasing fine root density with depth and distance from the tree stem. These models would not predict correctly tree–tree and tree–understorey competition for water and nutrients in 3D heterogeneous soil conditions that prevail under low-density tree stands. To account for the integrated response of tree root systems to such transient gradients in soils, we need a dynamic model that would allow for both genotypic plasticity and transient environmental local soil conditions.     

Spatial pattern formation and relative importance of intra- and interspecific competition in codominant tree species, <Emphasis Type="Italic">Podocarpus nagi</Emphasis> and <Emphasis Type="Italic">Neolitsea aciculata</Emphasis>

Spatial patterns, their changes due to mortality, and intra- and interspecific competition of two codominant tree species, Podocarpus nagi and Neolitsea aciculata, were analyzed at Mt. Mikasa, Nara City, Japan. Podocarpus nagi has a higher shade tolerance but a narrower seed dispersal range than N. aciculata. We inferred the mechanisms of spatial pattern formation and coexistence of the two species. Podocarpus nagi and N. aciculata trees were clumped and showed a spatial repulsion from each other. Patches dominated by either P. nagi or N. aciculata were formed. Podocarpus nagi trees were less clumped with increasing tree size, although no significant change in spatial patterns due to mortality was detected. A patch formation of the P. nagi population seemed to be induced by the narrow seed dispersal range. On the other hand, N. aciculata trees were more clumped and more repulsive from P. nagi trees with increasing tree size. The distribution of N. aciculata trees shifted to more clumped than expected from the random mortality over the research period. Post-dispersal mortality due to competitive exclusion by P. nagi affected the patch formation of the N. aciculata population. The relative importance of intraspecific competition to interspecific competition on the relative growth rate increased with increasing tree size. The shift corresponded to an increasing spatial repulsion between the two species. The patch formation of P. nagi population may create the spatial refuge of N. aciculata from P. nagi and may enable avoidance of interspecific competition and the coexistence of the two species.     

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。     

Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

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).     

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...     

乌拉山自然保护区白桦种群的年龄结构和点格局分析

白桦群落是乌拉山森林植被的主要类型之一,在高海拔阴坡、半阴坡以纯林形式分布.根据乌拉山自然保护区白桦林不同林龄结构设置3个典型样地,采用种群径级结构代替年龄结构、点格局分析(Ripley's K-Function)方法探讨了乌拉山白桦种群年龄结构、空间分布规律和种群动态.结果表明:(1)乌拉山自然保护区白桦种群径级结构呈典型的“金字塔”型,种群自然更新良好,属增长型种群;(2)由于种内不同个体间为争夺空间和资源,种群在第Ⅲ、Ⅳ径级死亡率较高,自疏作用明显;(3)白桦种群的存活曲线接近于Deevey Ⅰ型曲线;(4)在研究尺度内白桦种群以幼树、中龄树为主时呈聚集分布,而成龄树或老龄树占多数时呈随机分布,即随着种群年龄的增加,其分布格局逐渐由集群分布向随机分布转变.乌拉山白桦种群在小于1.5m的尺度呈聚集分布,即具有2株以上个体“丛生”现象.在环境条件相似的情况下,白桦种群自身的生物、生态学特性是影响其分布格局的最主要因素.     

Intercropping Competition between Apple Trees and Crops in Agroforestry Systems on the Loess Plateau of China

Agroforestry has been widely practiced in the Loess Plateau region of China because of its prominent effects in reducing soil and water losses, improving land-use efficiency and increasing economic returns. However, the agroforestry practices may lead to competition between crops and trees for underground soil moisture and nutrients, and the trees on the canopy layer may also lead to shortage of light for crops. In order to minimize interspecific competition and maximize the benefits of tree-based intercropping systems, we studied photosynthesis, growth and yield of soybean (Glycine max L. Merr.) and peanut (Arachis hypogaea L.) by measuring photosynthetically active radiation, net photosynthetic rate, soil moisture and soil nutrients in a plantation of apple (Malus pumila M.) at a spacing of 4 m × 5 m on the Loess Plateau of China. The results showed that for both intercropping systems in the study region, soil moisture was the primary factor affecting the crop yields followed by light. Deficiency of the soil nutrients also had a significant impact on crop yields. Compared with soybean, peanut was more suitable for intercropping with apple trees to obtain economic benefits in the region. We concluded that apple-soybean and apple-peanut intercropping systems can be practical and beneficial in the region. However, the distance between crops and tree rows should be adjusted to minimize interspecies competition. Agronomic measures such as regular canopy pruning, root barriers, additional irrigation and fertilization also should be applied in the intercropping systems.     

The spatial relationship between post‐crop remaining trees and the establishment of saplings in Pinus sylvestris stands in Spain

Question: What is the spatial relationship between remaining trees and the establishment and development of recruited saplings? Location: The Pinus sylvestris forest Pinar de Valsaín, in the Sistema mountain range (central Spain). Methods: Three 0.5 ha plots have been analysed. The saplings were located in a 2 m x 2 m grid, characterizing their spatial pattern through a nested ANO VA. The spatial pattern of stems was analysed using the L(d) function. To analyse the spatial relationship between stems belonging to different cohorts, the intertype L_rs(d) function was used. Finally a new function K_rx(d) is presented as a method to analyse the relationship between the spatial distribution of stems and the sapling density (a sampled continuous variable). Results: The mother trees show cluster pattern at scales of ca. 12 m ‐ 22 m, leading to a spatial pattern at 14 m–16 m for the saplings during the regeneration period. At the beginning of the shelter phase, saplings less than 1.30 m in height show spatial repulsion from the old crop at distances above 10 m, whereas taller saplings show repulsion at shorter distances, due to the suppression of sapling development near the mother trees. At the end of the regeneration period, saplings < 1.30 m appear under the last remaining mother tree canopies. Conclusions: In the stands analysed, located at the southern limit of Pinus sylvestris distribution, this species behaves as half‐shade tolerant. This study shows that the K_rx(d) function might be widely applied to analyse the relationship between patterns that occur at different scales or between a point pattern and a continuous variable, being a useful tool for analysing some forest processes.     

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.     

Spatial segregation of pines and oaks under different fire regimes in the Sierra Madre Occidental

Surface fire can modify spatial patterns and self-thinning in pine-oak ecosystems. Spatial pattern analyses were used to compare pattern development and interspecific spatial interactions in trees and seedlings in five Madrean pine-oak stands with different recent fire histories. Interspecific and intraspecific patterns were compared in small (< 15 cm dbh) and large (< 15 cm dbh) diameter classes of the pines (Pinus durangensis, P. teocote, and P. leiophylla) and oaks (Quercus sideroxylla, Q. crassifolia, and Q. laeta) that collectively dominated the five stands. Numbers of juvenile trees in 2.5 × 2.5 m subplots were correlated with cumulative distances to adult trees. Small pine and oak trees were intraspecifically clustered at all scales, irrespective of fire regime. Large pines were strongly clustered only in stands with longer fire-free intervals, and patterns of large versus small pine trees were regular or random in frequent fire stands. These patterns were consistent with fire-induced mortality of maturing trees under frequent fire. Large and small pines were segregated from small oaks at short and long distances in one stand with a 32-year fire-free interval, implying that two or more dynamic factors had produced regular patterns at different scales. Such regular spatial patterns at short distances were not seen in other stands. Therefore, there was little evidence for direct competition between oaks and pines. The results reported here are consistent with studies from other pine-oak ecosystems showing that different fire regime and site factors interact to influence stand development processes and relative dominance of pines and oaks. In some stands, the continued absence of fire could foster increasing tree densities and an intensification of local neighborhood effects, producing segregation of pine and oak species at longer distances. This revised version was published online in August 2006 with corrections to the Cover Date.     

Woody vegetation spatial patterns in a semi-arid savanna of Burkina Faso,West Africa

Spatial patterns of woody individuals were studied in a semi-arid savanna of West Africa located in Burkina Faso at and around 14° 12 N and 2° 27 W. The study was based upon a 10.24 ha plot within which individuals were mapped. Spatial pattern analysis was carried out using second order characteristics of point processes as K functions and pair correlations. The overall density amounted to 298 individuals ha^-1. The most abundant species were Combretum micranthum G. Don., Grewia bicolor Juss. and Pterocarpus lucens Lepr. Anogeissus leiocarpus (D.C.) G. et Perr. was also an important constituant of this vegetation type, owing to its taller stature. Clumped spatial distributions were identified for all species except for two, for which complete spatial randomness (CSR) was found (including P. lucens, a dominant woody plant). No regular pattern was found even when tall individuals were considered alone. Aggregation dominates interspecific relationships, resulting in multispecific clumps and patches. The overall aggregation pattern was constituted by two different structures. A coarse-grain pattern of ca. 30–40 m was based on edaphic features, and expresses the contrast between sparse stands on petroferric outcrops and denser patches on less shallow soils. A finer-grain pattern made of clumps ca. 5–10 m wide, with no obvious relation to pre-existing soil heterogeneity. There was no overall pattern for saplings (between 0.5 m and 1.5 m in height) irrespective of species, and thus no obvious common facilitation factor. For species with a high recruitment level there was no significant relationship between mature adult and saplings. The only case of clumped saplings with randomly distributed adults was found in P. lucens. However, this cannot be unequivocally interpreted as density dependent regulation since the existence of such a process was not consistent with the spatial distribution of dead P. lucens individuals (victims of the last drought). The mean density around dead P. lucens was lower than around surviving ones, indicating that the last drought tended to reinforce clumping rather than promote a regular pattern of trees. Spatial pattern analysis yielded no evidence supporting a hypothesis of stand density regulation through competition between individuals. Other processes, as surface sealing of bare soils or insufficient recruitment, may play a more important role in preventing a savanna-like vegetation from turning into denser woodlands or thickets.