SPATIAL DYNAMICS OF A HIMANTHALIA ELONGATA (FUCALES, PHAEOPHYTA) POPULATION

In dense monospecific stands of plants intraspecific competition usually results in self-thinning, the concurrent increase in biomass and decrease in density over time. Self-thinning may also result in a change in the spatial pattern of individuals, but so far the spatial dynamics of marine plants has not been investigated. The brown alga Himanthalia elongata ( L.) S. F. Gray forms dense monospecific stands on many northern temperate rocky shores, and various attributes (including its simple form) facilitated the study of the spatial dynamics of this species .
The spatial pattern of settling zygotes was examined in the laboratory. In the absence of water movement, substratum heterogeneity, and a point source, zygotes usually settled in clumps rather than randomly. Within the clumps zygotes appeared to be regularly distributed at a scale similar to the size of the zygotes themselves. Furthermore, the clumps themselves seemed to be regularly distributed. On the shore, well-established stands of "button-stage" Himanthalia populations were examined during a period of extensive growth and self-thinning. Individual plants were initially highly regular in spatial pattern but became less so over time. The pattern of plants dying during self-thinning was also highly regular and probably reflected existing spatial regularity. However, using a hypothesis of mortality as a random event, I found that smaller plants had a less than average survival potential, while larger plants had a greater than average chance. A consideration of the spatial pattern of plants alive at the end of the study revealed regularity at a scale of 2–7 mm but a random spatial pattern at larger scales, which might indicate a small sphere of influence of competing individuals. The best predictor of mean nearest neighbor distances in the populations was mean plant diameter .     

Spatial patterns and interspecific associations among trees at different stand development stages in the natural secondary forests on the Loess Plateau,China

Quercus wutaishansea populations on the Loess Plateau are currently becoming more dominant in natural secondary forests, whereas Pinus tabulaeformis is declining. In the present paper, the diameter class (instead of age) was used to classify the different growth stages as juvenile, subadult, or adult, and the univariate function g(r) was used to analyze the dynamic changes in spatial patterns and interspecific associations in three 1‐ha tree permanent plots on the Loess Plateau, NW China. Our results suggested that the niche breadth changed with the development stage. The diameter distribution curve was consistent with the inverted “J” type, indicating that natural regeneration was common in all three plots. There was a close relationship between the spatial pattern and scale, which showed significant aggregation at small distances, and became more random as distance increased, but in the Pinus + Quercus mixed forests, the whole species were aggregated at distances up to 50 m. The degree of spatial clumping decreased from juvenile to subadult and from subadult to adult. The spatial pattern also differed at different growth stages, likely due to strong intraspecific competition. Associations among different growth stages were positively correlated at small scales. Our study is important to the understanding of the development of the Q. wutaishansea forests; thus, the spatial dynamic change features should be received greater attention when planning forest management and developing restoration strategies on the Loess Plateau.     

SPATIAL PATTERNING AND COMPETITION IN AN ASPEN-WHITE PINE SUCCESSIONAL SYSTEM

The spatial patterning of and competition between established large-toothed aspen (Populus grandidentata) and invading white pine (Pinus strobus) were examined in a 65-yr-old forest system at the AuSable Trails Institute of Environmental Studies, Kalkaska County, Michigan. Density, diameter, and distance measures were recorded for tree species on two 40 × 40 m grids containing 64 quadrats each 5 × 5 m. Several indices of dispersion were used to assess pattern and regression analysis of nearest neighbor distance to tree diameter provided an index of competition. The results indicate strong aggregation for both aspen and pine on a large scale, but regularity of pattern for pines in relation to their nearest neighbor. This regularity appears to be due to competition with established aspens and suggests that the spatial patterning of the aspens acts to control the spacing and subsequent development of the invading white pines.     

Progress in theoretical vegetation science

Book Review

Progress in theoretical vegetation scienceG. Grabhett, L. Mucina, M.B. Dale and C.J.F. ter Braak (Eds.), Vol 83. Dordrecht: Kluwer Academic Publishers, 1990, 276 pages, £95.00. ISBN 0-7923-0507-1     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

荒漠草原不同土壤条件下猪毛蒿幼苗种群的点格局分析

采用摄影定位法测定了宁夏荒漠草原3种不同土壤条件下猪毛蒿（Artemisia scoparia）幼苗种群的空间格局,并应用完全空间随机模型、泊松聚块模型和嵌套双聚块模型对其分布格局进行了分析。结果表明：（1）在灰钙土生境下,猪毛蒿幼苗种群在小尺度上（0-2.85m）表现为聚集分布,随着尺度的增大先呈现为随机分布（2.85-3.75 m）,然后又呈现为均匀分布（3.75-5m）;在风沙土生境下,猪毛蒿幼苗种群在0-1.85 m之间表现为聚集分布,在1.85-2.35 m之间表现为随机分布,当尺度大于2.35 m时表现为均匀分布;而基岩风化残积土上的猪毛蒿幼苗种群在整个尺度上均呈现随机分布。（2）猪毛蒿种群幼苗在基岩风化残积土上符合泊松聚块模型,即猪毛蒿种群空间格局的聚块中不存在较高密度的小聚块;而在风沙土和灰钙土上则符合嵌套双聚块模型,即在大聚块中分布较高密度的小聚块。猪毛蒿幼苗种群空间格局的形成与土壤异质性存在着密切的联系,种群在空间中分布格局的形成机制可以通过种群空间格局的分析加以解释。     

Structural heterogeneity and spatial autocorrelation in a natural mature Pinus sylvestris dominated forest

The tree species composition, vertical stratification and patterns of spatial autocorrelation at the tree and quadrate (25 × 25 m) scales were studied in a natural mature PinuS sylvestris dominated forest in eastern Finland. For the analyses we mapped the locations and dimensions of trees taller than 10 m in a 9 ha (300 × 300 m) area, and within this area we mapped all trees taller than 0.3 m on a core plot of 4 ha (200 × 200 m). The overall tree size distribution was bimodal. the dominant layer and the understory forming the peak frequencies. Pinus sylvestris dominated the main canopy, together with scattered Betula pendula and Picea abies. Alnus incana, Populus tremula, Salix caprea, Sorbus aucuparia and Juniperus communis occurred only in the under- and middlestories. Autocorrelation analysis (semivarianee) of tree size variation revealed spatial patterns, which were strongly dependent on the size of trees included in the analysis. When all living trees, including the understory regeneration, were taken into account, the autocorrelation pattern ranged up to 35 m inter-tree distances, reflecting the spatial scale of understory regeneration patches. Competitive interaction among middle- and upperstory trees (height>10 m) had contrasting effects on autocorrelation pattern depending on spatial scale. At the fine scale, dominant trees suppressed their smaller close neighbors (asymmetric competition), which was shown as increased tree size variation at small inter-tree distances (<2 m). At slightly larger inter-tree distances, specifically among large trees of similar size, competition was more symmetrical, which resulted in decreased tree size variation at these inter-tree distances (3–4 m). This effect was seen most clearly in the dominant trees, there being a clear autocorrelation pattern in tree size up to inter-tree distances of ～4 m. At the quadrate scale (25 × 25 m) the analysis revealed high local variation in structural characteristics such as tree height diversity (THD), tree species diversity (H) and autocorrelation of tree height. The analysis suggests that naturally developed P. sylvestris forests exhibit complex small-scale patterns of structural heterogeneity and spatial autocorrelation in tree size. These patterns may be important for stand-scale habitat diversity and can have aggregated effects on ecosystem dynamics at larger spatial scales though their influence on the spread of disturbance and regeneration after disturbance.     

Regeneration pattern of primary forest species across forest-field gradients in the subtropical Mountains of Southwestern China

Evergreen broad-leaved forest is now gradually degraded and fragmented, and there is an increase in the amount of habitat edges as a result of long-term human activity. However, the role of edges in the regeneration of primary forest species is poorly understood. After 20 years of the edge creation, we analyzed primary forest species distribution and abundance, and changes in floristic composition, vegetation structure across forest-field gradients in Ailao Mountain, SW China. Our results revealed that there was a higher abundance and richness of primary species, late secondary species and thorny lianas at the distances 0–50 m than at the distances more than 50 m from the edge into the forest exterior. At the distances >50 m, no individuals of dominant canopy trees Lithocarpus xylocarpus, Castanopsis wattii, and L. jingdongensis were found, whereas the abundance of early pioneer shrub species and herbaceous cover was significantly greater. The richness of primary species showed a decrease with increasing distances from the forest edge to the exterior, particularly of medium-seeded primary species showing a drastic decrease. Moreover, no large-seeded primary species occurred at the distances >60 m. This study indicates that the forest edge as a buffer zone may be in favor of primary species regeneration. A dense shrub and herb layer, and seed dispersal may be the major factors limiting the forest regeneration farther from the forest edge. Therefore, to facilitate forest recovery processes, management should give priority to the protection of buffer zones of this forest edge.     

BOOK REVIEWS

ter Braak, C. J. F. 1990. CANOCO Version 3.10. Agricultural Mathematics Group, Wageningen. Smilauer, P. 1991. CANODRAW: A companion program to CANOCO for publication-quality graphical output.     

桥山林区麻栎种群不同发育阶段空间格局及关联性

为探究麻栎种群不同发育阶段的空间分布特征及关联性,以黄土高原南部桥山林区麻栎天然次生林为研究对象,用L(r)函数对麻栎种群不同发育阶段空间分布特征及关联性进行分析。结果表明:(1)样地内麻栎种群各发育阶段数量结构均为不规则倒"J"型,种群结构稳定,能实现持续更新。(2)麻栎种群总体的空间分布特征受生境异质性影响较大,纯林样地中,种群在0—10 m尺度上表现为高强度聚集分布,5—35 m尺度表现为低强度聚集;混交林样地中,0—5 m尺度上表现为高强度聚集分布,5—35 m尺度上则表现为低强度聚集分布。(3)随着种群的发育及尺度的增大,麻栎种群空间聚集强度减弱,纯林样地内,麻栎幼苗在0—6 m的小尺度表现为聚集分布,6—12 m尺度上表现为随机分布;幼树在各尺度上均表现为聚集分布;中龄树与成龄树在各尺度上均表现为随机分布。混交林样地中,仅幼苗在0—6 m尺度上表现为聚集分布,其余各阶段在各尺度下均趋于随机分布。(4)种群不同发育阶段不同尺度空间关联性不同,纯林样地中,幼苗与幼树在1—2 m小尺度上呈负相关,幼树与中龄树在8 m尺度呈正相关,其余各生长阶段之间在任意尺度上均无关联性;混交林样地中,幼树与中龄树在2—12 m尺度上表现为正相关,其余各生长阶段之间在各尺度上均无关联。在未来的森林经营中,应对不同发育阶段、不同生境的种群进行不同抚育措施,促进种群良性发育。     

辽东山区次生林乔木幼苗分布格局与种间空间关联性

在辽东山区次生林建立4 hm^2样地(200 m×200 m),研究0~50 m尺度范围内乔木幼苗分布格局及种间空间关联性.结果表明:在完全随机零模型下,0~20 m尺度上,95%的树种呈现聚集分布格局;0~16 m尺度上,19个树种呈现聚集分布;随着尺度的增加,聚集分布树种的比例逐渐减少,50 m尺度上,随机分布成为树种分布的主要形式;在异质性泊松过程零模型下,0~24 m尺度上,5%的树种呈现聚集分布,26~50 m尺度上,42%和58%的树种呈现随机和均匀分布.在完全随机零模型下,正相关树种对比例最高,且在50 m尺度下呈现正相关、负相关、无相关3种相关性的树种对比例相同;在异质性泊松过程零模型下,树种对主要呈现负相关,且随尺度增大,负相关的树种对比例逐渐升高.种子扩散限制和生境异质性在某种程度上解释了乔木幼苗的聚集分布格局,乔木幼苗强烈的聚集分布又促使种间空间关联性密切,更新群落稳定性较差.     

Local and regional spatial distribution of an eruptive and a latent herbivore insect species

Abstract In this work, we investigated the spatial distribution of two sessile insect herbivores over the entire range of their host plant, Coccoloba cereifera, a sclerophyllous shrub endemic to Serra do Cipó, Brazil. The two insects have very distinct life histories and dispersal behaviours and we hypothesized that their classification into behavioural syndromes could be used to predict their spatial distribution patterns. Abgrallaspis cyanophylli (Homoptera) is an armoured scale insect that fits well into the eruptive syndrome. Stenapion aff. contrarium (Coleoptera) is a petiole borer with wide search capabilities, which fits into the latent syndrome. We expected that Abgrallaspis would follow the host plant aggregation pattern whereas Stenapion would be distributed more uniformly through the region and be less affected by host aggregation. We counted the number of attacked and non‐attacked ramets within two perpendicular belt transects as well as within a 20 m × 20 m quadrat placed over a dense shrub aggregation. Local quadrat covariance methods were used to estimate the spatial pattern of each insect. At fine scales, we found Stenapion evenly distributed over the host plant and Abgrallaspis with a significantly aggregated pattern. This finding is in accordance with our hypothesis. At larger scales, however, this pattern was lost and the results were largely variable. We conclude that the classification of insects into behavioural syndromes may be useful to predict distribution patterns at fine scales. At larger scales, however, history and chance events may be more important.     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

The regeneration ecology of Kunzea ericoides (A. Rich.) J. Thompson at Coranderrk Reserve,Healesville

The spatial structure, age structure and population dynamics of the shrub Kunzea ericoides (A. Rich.) J. Thompson were investigated at Coranderrk Reserve, near Healesville, Victoria, Australia. The shrub is known to be invasive in many areas and, although indigenous to the reserve, has greatly increased its population size and distribution within the reserve in the past 30 years. Dendrochronology showed a constant relationship between age and stem diameter for K. ericoides in the reserve. The spatial structure of the variable stem diameter was investigated by the use of spatial correlograms and results suggested that K. ericoides is spreading via the formation of discrete clumps and gap-phase regeneration. The two main populations in the reserve recruited continuously in time but patchily in space. A simple statistical model for size–frequency data showed that K. ericoides is expanding faster in some areas of the reserve than others. The trend of expansion of K. ericoides may be irreversible. Intensive management of Coranderrk Reserve will be required if K. ericoides is to be controlled and the conservation value of the reserve maintained.     

Testing the Stress‐Gradient Hypothesis During the Restoration of Tropical Degraded Land Using the Shrub Rhodomyrtus tomentosa as a Nurse Plant

The relative importance of facilitation and competition between pairwise plants across abiotic stress gradients as predicted by the stress‐gradient hypothesis has been confirmed in arid and temperate ecosystems, but the hypothesis has rarely been tested in tropical systems, particularly across nutrient gradients. The current research examines the interactions between a pioneer shrub Rhodomyrtus tomentosa (the nurse plant) and seedlings of a transplanted native woody Schima superba (the target species) in a tropical system in which position on a slope corresponds with a nutrient gradient; high soil nutrients at the slope bottom and relatively low soil nutrients at the slope top. In contrast, soil physical traits were more favorable for seedling growth under the shrub than in open spaces. The effect of R. tomentosa on S. superba survival was positive (facilitation) at the top of the slope, as indicated by the relative interaction index (RII), but negative in the bottom (competition). RII indicated a positive effect on seedling height at the top of the slope but was not at the bottom. Seedling survival was positively related to soil nutrient level and negatively related to soil acidity, but seedling growth of S. superba seemed to be enhanced by the shrub canopy. Thus, the results seem to support stress‐gradient hypothesis in terms of target species survival but not growth. We suggest using the shrub as a nurse plant in forest restoration in tropical degraded land with caution because not all of its effects on target species are positive .     

Competition by herbs as a limiting factor in shrub invasion in grassland: a test with different growth forms

Abstract. We tested the hypothesis that seedling establishment, the critical stage in the invasion of grassland by shrubs, is limited by competition with perennial grasses in seasonally wet/dry savannas. We placed seeds of two invasive exotic shrubs – Cryptostegia grandiflora, a woody vine, and Acacia nilotica, an arborescent legume – into pots with a wide range of existing above- and below-ground herbaceous biomass provided by either a tussock or a stoloniferous perennial grass. We also imposed different levels of watering frequency (5, 10 and 21 d), nutrient addition (+ and -) and grass clipping intensity (no clipping, clipped to 5 cm and clipped to 25 cm). There was no effect of any treatment on shrub seedling emergence or survival and all of the seedlings that emerged survived the 90-d growing period. Herbaceous competition also failed to have an effect on biomass accumulation in shrub seedlings. More frequent watering significantly increased above- and below-ground biomass accumulation for both shrub species and nutrient addition significantly increased Cryptostegia biomass accumulation. Based on these results, we question the proposition that reduction in competition by herbs via livestock grazing has been a significant factor in determining the rate or pattern of exotic shrub increase in the seasonally wet/dry tropics. We also question the suitability of the two-layer soil moisture hypothesis as a basis for management practices to control the ingress of woody species into grasslands and open savannas.     

Multi-proxy evidence for competition between savanna woody species

Coexistence of trees and grasses in savannas should be possible if competition between the woody and the grassy components is less intense than the competition within each component. Although several studies have investigated competition between trees and grasses, little is known about tree–tree interactions. We used a multi-proxy approach to examine the spatial pattern of Acacia mellifera and other savanna woody species in a semi-arid savanna in South Africa. Spatial analysis of the point patterns of young and reproductively mature shrubs detected decreasing aggregation with size/age over all spatial scales. This indicated the prevalence of competition although the overall spatial shrub pattern was aggregated. In contrast to point pattern statistics that detect changes only when competition has led to the death of the inferior competitor, we also applied methods identifying the competitive effect on sizes of individual trees. Competition should lead to a negative spatial autocorrelation in size, which we observed in half of the studied cases. Quantile regressions show that nearest-neighbour distance increased steeply with combined size of the target shrub and its neighbours indicating strong competitive effects. The medians of the distributions of maximum root lengths of A. mellifera, of the scale of regular patterns, and of negative autocorrelations were not significantly different, suggesting that overlapping root systems mediate competitive interactions. A competitor removal experiment did not lead to increased shrub sizes, which may be due to the limited duration of the experiment. From the nearest neighbour and autocorrelation analyses, we conclude that competition had a strong impact on growth rates of savanna woody species. Competition-induced mortality only becomes obvious when analysing the shift towards less aggregated spatial patterns when shrubs become reproductively mature. As the overall clustered spatial pattern masks the perceptible effect of competition, a time component should always be included in spatial pattern-based inference of competition.     

Spatial analysis of a subalpine heath woodland

The horizontal spatial patterns of heath species have implications in dynamic community modelling, fire behaviour modelling and ecological interpretation. The distribution pattern of the dominant woody species in a subalpine heath woodland near Kiandra, New South Wales was analysed as spatial point processes. The species analysed were Bossiaea foliosa, Grevillea australis, Hakea lissosperma and Oxylobium ellipticum, and both large (>2m) and small (<2 m) snowgums (Eucalyptus pauciflora) They all exhibited significant aggregation at scales ranging from 1 to 15 m. Bivariate spatial analyses of these species revealed significant negative association between G. australis and H. lissosperma, between G. australis and O. ellipticum, and between O. ellipticum and H. lissosperma, B. foliosa was independently distributed with respect to other shrub species. There was some evidence for negative association between small snowgums and shrub species, though small snowgums were positively associated with large snowgums. The joint spatial distribution of the individuals of all shrub species was also aggregated. A Poisson cluster process was developed and tested to model the joint spatial pattern of the shrub stratum.     

塔里木荒漠绿洲过渡带主要种群生态位与空间格局分析

于新疆塔里木盆地荒漠绿洲过渡带设置3条样带(长900 m×宽50 m)进行植物群落调查,运用生态位测度指标和点格局法对种群空间格局、空间关联性及生态位特征进行研究。结果显示,胡杨(Populus euphratica Oliv.)、多枝柽柳(Tamarix ramosissima Ledeb.)和甘草(Glycyrrhiza inflate Bat.)具有较大的重要值和生态位宽度,对其它生态位较窄的物种具有资源竞争和扩张优势,生态适应能力强,对群落结构和环境变化起决定性作用;生态过渡带植物的种间生态位重叠普遍较高,尤其是草本植物间更明显,此分配格局反映出荒漠植物长期适应旱化生境而发生趋同适应,生态位分化程度低,种间资源竞争激烈;优势种群空间格局呈明显的空间变异性,0~25 m尺度内多枝柽柳和甘草分别呈随机分布和聚集分布,胡杨种群在≤5 m尺度内呈聚集分布,并随尺度增大转为随机分布;胡杨与多枝柽柳、胡杨与甘草、多枝柽柳与甘草分别在4 m、≤16 m和≤25 m尺度内呈显著空间负关联,种间相互排斥。表明在匮乏环境资源条件下植物趋同适应与资源竞争是驱动荒漠植物群落演替和限制物种共存的主要原因。