长白山次生林乔木树种空间分布格局

应用RipleyK函数,分析了长白山一个5．2hm^2次生林样地内乔木树种的空间分布以及主要乔木树种的幼树、小树、大树的空间分布和空间关系。结果表明：共调查到34种乔木,其中常见种27种,少见种7种,共计15688株;在27个常见种中,有23个种在空间上主要呈聚集分布,另外4个种主要表现为随机分布;在7个少见种中,均主要表现为随机分布;簇毛槭、假色槭、水曲柳、臭松和红松随着种群发育（幼树→小树→大树）聚集强度逐渐减弱,直至趋向于随机分布,其他树种的空间分布随种群发育未表现出明显的变化规律;在10个主要树种组成的45个种对中,有11个种对表现出显著正相关,1个种对为显著负相关。     

Spatial and temporal patterns of seed attack and germination in a large-seeded neotropical tree species

It has long been argued that seed dispersal enhances recruitment in tropical trees by allowing offspring to `escape' strong density/distance-dependent attack by insects, pathogens and rodents. Here we examined the effects of canopy openness and parent-offspring distance upon the frequency and timing of Chlorocardium rodiei seed attack and germination within a 15-ha plot of Guyanan tropical rain forest. Seeds were artificially dispersed beneath parent trees, in the understorey away from trees and in gaps. Analysing our data from an 85-week period of regular monitoring, we found that the main spatial gradients, canopy openness and distance to nearest adult conspecific, do not lead to differences in the final number of seeds attacked by infesting scolytid beetles or rodents. The timing of beetle attack, however, varied along the distance gradient and this difference affords seeds at further distances a `window' in which to germinate and produce a seedling before attack. Canopy openness was not a good predictor of rooting success, but distance was strongly associated with root and shoot formation success and the mean time to shoot formation. There was a strong negative effect of distance on the likelihood of a seed being colonised by scolytid beetles prior to removal by rodents and shoot failure was strongly associated with prior infestation. We believe these results bring a key point to bear on the well-established notion of distance-dependent attack on seeds in tropical rainforests, viz. that seed characteristics (size, germination syndrome) and the timing of attack may be more important in explaining patterns of early seedling recruitment than distance. Our studies suggest that advantages accrued through dispersal in species like Chlorocardium will depend heavily on the `race' between seed germination and attack. In the case of Chlorocardium, the `race' can be lost at considerable distances due to its prolonged dormancy and the temporal fluctuations in fruitfall and rainfall which influence attack and germination. The results presented here suggest that the lag between seed attack and germination in tropical trees can regulate the influence of parent-offspring distance on cohort recruitment at this life history stage. Received: 5 March 1998 / Accepted: 6 December 1998     

Spatial patterns of mycorrhizal infectiveness of soils long a successional chronosequence

 This study quantified intersite variation and spatial pattern in arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) infectivity of soils among six sites constituting a successional chronosequence in southwestern Ohio, USA. The study sites included an active agricultural field (chronic disturbance), a site which had been stripped of its surface soil (pseudo-stripmine, acute disturbance), 5- and 10-year-old fields, a 25- to 30-year-old prairie restoration, and an undisturbed, mature forest. AM infectivity was lower in the agricultural field, successional fields, and prairie than in the mature forest, but there was no clear correlation between time since disturbance and the overall level of AM infectivity. Spatial structure in AM infectivity decreased with time since disturbance. In the pseudo-stripmine site and active soybean field, semivariance analysis attributed 44–50% of the total variance in AM infectivity among samples to spatial structure, whereas spatial dependancy accounted for only 18% of total variance in the mature forest. Kriging of AM infectiveness demonstrated small, isolated areas in the disturbed plots that were devoid of AM infectiveness, whereas the kriged AM maps of the other four sites showed AM infectiveness to become progressively more homogeneous. ECM infectiveness was lacking from 35–50% of the samples from the disturbed sites, and both overall ECM infectiveness and ECM diversity increased with time since disturbance. Approximately 44% of the variance in ECM infectiveness was related to spatial structure in the two disturbed sites, and large areas entirely devoid of ECM infectivity were present on the kriged ECM maps for these sites. There was less spatial structure in ECM in the old fields and prairie and very little in the mature forest. The results of this study emphasize the need to explicitly evaluate spatial heterogeneity in mycorrhizal infectivity in studies of the role of mycorrhizae in succession. Accepted: 4 August 1995     

不同区域典型树木的空间分布格局及关联性

以温带长白山(CBS)、暖温带关帝山(GDS)、亚热带黑石顶(HSD)样地监测数据为基础,采用成对相关函数g(r)分析了3个样地共有的3科(松科、壳斗科、蔷薇科)树木的空间分布格局及其关联性.结果表明:不同样地3科树木分布数量及结构特征存在差异.松科在GDS分布数量较多,呈双峰型径级结构,在CBS和HSD分布数量少,径级结构近似正态分布;壳斗科在CBS和GDS分布数量少,分别呈双峰型和偏正态径级结构,在HSD分布数量多,呈倒"J"型径级分布;蔷薇科在GDS分布数量多,呈"L"型径级分布,在CBS和HSD分布相对较少,径级结构分别呈倒"J"型和"L"型分布.3科树木空间分布格局在不同样地表现也不同,松科的大径级个体在CBS和GDS小尺度上呈均匀分布,在HSD呈聚集分布,中、小径级在3个样地均呈聚集分布;壳斗科在CBS以大径级个体为主,呈近似随机分布,在GDS和HSD以中、小径级为主,呈聚集性分布格局;蔷薇科在3个样地均为聚集分布.3科树木的聚集程度均随尺度增加而降低.大径级的壳斗科个体在CBS和HSD与松科呈不相关或小尺度负相关关系,中、小径级的壳斗科在CBS和GDS与松科呈负相关关系,但在HSD与松科表现为正相关;松科与蔷薇科在3个样地均表现为负相关;中、小径级的壳斗科与蔷薇科在CBS和GDS呈正相关关系,但在HSD呈负相关关系.总之,3科树木空间分布格局及关联性随径级、尺度而变化且在不同样地内有不同表现.     

Spatial and temporal patterns of seed predation on three tree species in an oak-pine forest

We experimentally examined spatial and temporal patterns of seed predation on three tree species in an oak-pine forest in southern Maine. USA, Rodents were the principal seed predators. Rates of seed loss varied with tree species and exposure to three different suites of potential seed predators (all vertebrate seed predators, medium-sized rodents [primarily squirrels] and small rodents [primarily mice and voles]), Acer rubrum seeds were removed more slowly (mean of 17.7% within 2 d across all habitats and treatments in 1991) than seeds of either Quereus rubra or Pinus strobus (e,g., > 99% removed within 2 d across all habitats and treatments for each species in 1991), Levels of final removal (cumulative removal at final census) varied with year but not with microhabitat; i.e., seed predation in four types of forest gap was not significantly different than in intact forest. Both field data and experimental feeding trials with captive Peromyscus leucopus and Clethrionomys gapperi suggest that a significant proportion of while pine seeds is eaten at time of detection, while red oak acorns are more likely to be cached. Captive Peromyscus leueopus cached and ate significantly more red oak acorns than Clethrionomys gapperi. These results suggest that red oak mast may be more important in overwintering success in Feromyscus than in Clethrionomys. Nevertheless, both species did consume red oak acorns, suggesting that in southern Maine these species are not substantially inhibited by high tannin levels in red oak acorns, as has been suggested by researchers elsewhere.     

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence

We measured the response of dark respiration (R_d) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communites in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R_d at a reference temperature (R₁₀) and the extent to which R_d increased with temperature (described by E_o, a parameter related to the energy of activation, or the change in R_d over a 10°C range, Q₁₀). Site averaged E_o ranged from 44.4 kJ mol^–1 K^–1 at the 60-year-old site to 26.0 kJ mol^–1 K^–1 at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E_o or Q₁₀) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A_max/R_d) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.     

Spatial patterns of soil pathogens in declining Mediterranean forests: implications for tree species regeneration

Soil-borne pathogens are a key component of the belowground community because of the significance of their ecological and socio-economic impacts. However, very little is known about the complexity of their distribution patterns in natural systems. Here, we explored the patterns, causes and ecological consequences of spatial variability in pathogen abundance in Mediterranean forests affected by oak decline. We used spatially explicit neighborhood models to predict the abundance of soil-borne pathogen species (Phytophthora cinnamomi, Pythium spiculum and Pythium spp.) as a function of local abiotic conditions (soil texture) and the characteristics of the tree and shrub neighborhoods (species composition, size and health status). The implications of pathogen abundance for tree seedling performance were explored by conducting a sowing experiment in the same locations in which pathogen abundance was quantified. Pathogen abundance in the forest soil was not randomly distributed, but exhibited spatially predictable patterns influenced by both abiotic and, particularly, biotic factors (tree and shrub species). Pathogen abundance reduced seedling emergence and survival, but not in all sites or tree species. Our findings suggest that heterogeneous spatial patterns of pathogen abundance at fine spatial scale can be important for the dynamics and restoration of declining Mediterranean forests.     

川西亚高山红桦-岷江冷杉林优势种群的空间格局分析

以红桦-岷江冷杉林4 hm²样地调查数据为基础,采用点格局分析法O-ring函数,分析了红桦和岷江冷杉2个优势种群不同龄级的空间分布格局,以及各龄级间的种内和种间关联性.结果表明：红桦种群径级分布呈单峰型,为衰退种群;岷江冷杉种群径级分布呈倒“J”型,为进展种群.岷江冷杉与红桦大树在所有尺度上均为随机分布,其余龄级在小尺度上为聚集分布,随尺度增大趋于随机分布和均匀分布,最大聚集强度随龄级增大而减弱.空间关联主要发生在小尺度上.岷江冷杉种内关联以小尺度正关联为主,红桦种内关联以小尺度负关联为主.对于2个种群的相同龄级,小龄级、中龄级和大龄级间,分别以空间无关联、负关联和无关联为主.2个种群间不同龄级间以负关联为主,龄级相差越大,其负关联越强.     

浙江省典型天然次生林主要树种空间分布格局及其关联性

松阔混交林和常绿阔叶林是亚热带地区天然次生林代表性的森林类型,通过研究主要树种分布变动趋势,可以有效地指导森林经营措施。在该地区4个1hm~2典型样地,在0—30m尺度范围内综合分析了主要树种的空间分布格局及种间关联性,并对同一树种分发育阶段在不同森林群落中的种群空间分布格局进行了比较,以探讨亚热带地区天然次生林群落空间格局形成和种群维持机制。样地1和2为松阔混交林,其中样地1的马尾松密度较低,样地3和4为常绿阔叶林。研究结果表明:(1)以完全随机模型为零假设时,样地1的主要种群在小尺度(10 m)呈聚集分布,随尺度增加呈随机分布;样地2—4的主要种群在所有尺度呈聚集分布,随尺度增加聚集强度逐渐减弱;以异质泊松模型为零假设时,4个样地的主要种群在大部分尺度呈随机分布;(2)青冈和苦槠的小树(5.0 cm≤DBH10.0 cm)在4样地的所有尺度以聚集分布为主,大树(DBH≥10.0 cm)在松阔混交林样地呈随机分布趋势,但在常绿阔叶林样地青冈在0—20 m尺度、苦槠在所有尺度呈聚集分布;(3)松阔混交林中建群种马尾松和其他树种的种间关联性,在样地1的小尺度为负相关,随尺度增加为不相关,在样地2的所有尺度为负相关;常绿阔叶林中建群种青冈和其他树种的种间关联性在在样地3的小尺度为负相关,随尺度增加为不相关,在样地4的所有尺度为负相关;所有伴生树种的种间关联性以负相关为主。结果说明,种群空间分布格局及其关联性随群落结构和空间尺度的不同而出现变化,在松阔混交林和常绿阔叶林群落格局形成中除了扩散限制和生境异质性以外,密度制约机制在松阔混交林中发挥了重要作用,而在常绿阔叶林中其作用随着树木生活史阶段的提高而减弱。     

Patterns of species diversity and soil nutrients along a chronosequence of vegetation recovery in Hainan Island, South China

Understanding the pattern of species diversity and soil factors can enhance our knowledge of the mechanism of vegetation recovery, however, there is still a gap in the knowledge of succession rate and trend for species diversity in relation to soil nutrients during the vegetation recovery process. Patterns of species diversity and soil nutrients during the tropical vegetation recovery as well as the correlation between species diversity and soil nutrients were explored in Hainan Island, located in southern China. Plots assigned as grassland stage (GS), shrub stage (SS), secondary forest stage (SFS), and primary forest stage (PFS) were established using a chronosequence approach. Results showed that species richness and evenness increased from GS to PFS. Species dominance/diversity curves were fitted using the lognormal distribution model (r ² ?=?0.891?C0.972). Species richness for the herb layer was maximal at SFS, whereas species richness for both the shrub layer and tree layer reached their maximum at PFS. Species turnover and soil total phosphorus decreased, whereas organic matter and total nitrogen increased from GS to PFS. Organic matter and total nitrogen were both positively correlated with species richness and total coverage, and total phosphorus was positively correlated with species turnover. The results clearly demonstrate that diversity asymptotically increases and positively correlates with increasing soil fertility, and the total phosphorus value is predicted to be an important soil factor that affects successional rate during tropical vegetation recovery processes.     

Spatial distributions of tree species in a subtropical forest of China

The spatial dispersion of individuals in a species is an important pattern that is controlled by many mechanisms. In this study we analyzed spatial distributions of tree species in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. O-ring statistic was used to measure spatial patterns of species with abundance >10. Ω_0–10, the mean conspecific density within 10 m of a tree, was used as a measure of the intensity of aggregation of a species. Our results showed: (1) aggregated distribution was the dominant pattern in the plot. The percentage of aggregated species decreased with increased spatial scale. (2) The percentages of significantly aggregated species decreased from abundant to intermediate and to rare species. Rare species was more strongly aggregated than common species. Aggregation was weaker in larger diameter classes. (3) Seed traits determined the spatial patterns of trees. Seed dispersal mode can influence spatial patterns of species, with species dispersed by both modes being less clumped than species dispersed by animal or wind, respectively. Considering these results, we concluded that seed dispersal limitation, self-thinning and habitat heterogeneity primarily contributed to spatial patterns and species coexistence in the forest.     

Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China

Spatial patterns and interspecific associations of plant species in forests are important for revealing how species interact with each other and with the environment, and hence have important implications for optimal forest management and restoration in degraded forest ecosystems. In this paper, the O-ring statistics were used to characterize the spatial patterns and interspecific associations of eight dominant tree species in two 1-ha old-growth karst forest plots in Maolan National Natural Reserve, southwestern China. We found that most of the eight dominant tree species in two forests were continuously regenerating populations. Six species (Platycarya longipes, Acer wangchii, Clausena dunniana, Castanopsis carlesii var. spinulosa, Distylium myricoides, and Rhododendron latoucheae) exhibited significant aggregations at the majority of scales while others (Celtis biondii and Cyclobalanopsis myrsinaefolia) showed a random distribution pattern at most scales. Negative association was a dominant pattern for most species pairs in the two plots, while positive associations were found at most scales for only two species pairs (Platycarya–Clausena and Castanopsis–Rhododendron). Results also indicated that the two main factors of habitat complexity and heterogeneity—the elevation and rock-bareness rate—play important roles in determining spatial distribution patterns and interspecific associations of tree species in karst forests of Maolan. Thus, the observed spatial patterns among the eight tree species are influenced by habitat heterogeneity in the context of karst topographical variations. The partitioning of habitat niches contributes to the promoting species coexistence in species-rich karst forests. The differences of species features in spatial patterns and associations should be paid more attention when planning forest management and developing restoration strategies.     

Spatial associations of tree species in a subtropical evergreen broad-leaved forest

Aims The spatial segregation hypothesis and the low-frequency hypothesis are two important proposed mechanisms that delay or prevent competitive exclusion in ecosystems. Because tree species interact with their neighbors, the importance of these potential processes can be investigated by analyzing the spatial structures of tree species.Methods The distribution of the adults of 27 common tree species in a fully mapped 5-ha subtropical forest plot in Baishanzu, eastern China, was analyzed to investigate the community-level intra- and interspecific spatial association patterns. We first tested for the overall spatial pattern in the 5- to 40-m neighborhoods and classified first-order bivariate associations with a diametric scheme based on Ripley's K and nearest-neighbor statistic (G -function). Then heterogeneous Poisson null models were used to distinguish second-order interactions from overall spatial associations (including first-order effects). Finally, we analyzed correlations between the existence of species interactions and some attributes of the species involved.Important findings Partial overlap and segregation increased with scale, whereas mixing decreased. Nearly 70% of the species pairs occurred less than expected at random, and only 3.4% of the species pairs were well mixed; 11.0% of all species pairs showed significant small-scale interactions, which was a greater frequency than expected by chance if species are abundant or prefer the same habitat, but less frequent than expected if species are highly aggregated. This suggests that both spatial segregation and low frequency of species facilitate species coexistence by reducing the opportunity that trees of two species encounter each other. The study also revealed that positive interactions were more prevalent than negative interactions in the forest, which indicates that positive interactions may have important effects on forest species assemblies.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Spatial and temporal patterns of species richness in a riparian landscape

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control). Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden. Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km² large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora. Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape. Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.     

Mycorrhizal networks and distance from mature trees alter patterns of competition and facilitation in dry Douglas-fir forests

The distribution of dry Douglas-fir forests in western North America is expected to shift northward with climate change and disappear from the grassland interface in the southern interior of British Columbia. This shift may be accentuated by clearcutting, a common harvesting practice that aims to reduce the competitive effects of residual mature trees on new regeneration, but in so doing, ignores their facilitative effects. In this study, we investigated the net effects of competition from and facilitation by mature trees retained on harvested sites on seedling establishment in the dry interface Douglas-fir forests. We demonstrate that access to a mycorrhizal network (MN) and proximity to trees have important influences on seedling performance. On six sites, we established trenched plots around 24 mature Pseudotsuga menziesii var. glauca (Douglas-fir) trees, then planted Douglas-fir seedlings into four mesh treatments that served to restrict MN access (i.e., planted into mesh bags with 0.5-, 35-, or 250-μm pores, or without mesh) or into impermeable bags (grown in isolation) at four distances (0.5, 1.0, 2.5, or 5.0 m). Seedling survival tended to be greater and water stress lower where seedlings had full access to the MN. Seedling height, shoot biomass, needle biomass, and nutrient uptake peaked at 2.5–5.0 m from mature trees. Seedlings 0.5 m from mature trees had lower CO₂ assimilation rates and wood δ¹³C compared to seedlings 5.0 m away. Competition for soil resources was highest near mature trees but facilitation was relatively greater at further distances, resulting in a zone of net benefit for seedlings. These results show that intraspecific tree-seedling interactions are both competitive and facilitative in dry Douglas-fir forests, and that they are spatially dependent. After disturbance, maintaining residual mature trees may be important for their beneficial regeneration zones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.