SELF‐THINNING AND SIZE INEQUALITY DYNAMICS IN A CLONAL SEAWEED (SARGASSUM LAPAZEANUM,PHAEOPHYCEAE)1

Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.     

纯林自然稀疏研究综述

自然稀疏是林分内的个体由于部分有限的资源而引起的一部分个体死亡的现象。评述40a来世界上有关纯林自然稀疏的研究内容,包括了自然稀疏法则,异速生长模型、平均个体重与密度的时间轨线,种的自然稀疏线和自然稀疏动力线的关系,自然稀疏线的斜率与生物量的生物量的关系,自然稀疏的机理。     

Radial and stand‐level thinning treatments: 15‐year growth response of legacy ponderosa and Jeffrey pine trees

Restoration efforts to improve vigor of large, old trees and decrease risk to high‐intensity wildland fire and drought‐mediated insect mortality often include reductions in stand density. We examined 15‐year growth response of old ponderosa pine (Pinus ponderosa) and Jeffrey pine (Pinus jeffreyi) trees in northeastern California, U.S.A. to two levels of thinning treatments compared to an untreated (control) area. Density reductions involved radial thinning (thinning 9.1 m around individual trees) and stand thinning. Annual tree growth in the stand thinning increased immediately following treatment and was sustained over the 15 years. In contrast, radial thinning did not increase growth, but slowed decline compared to control trees. Available soil moisture was higher in the stand thinning than the control for 5 years post‐treatment and likely extended seasonal tree growth. Our results show that large, old trees can respond to restoration thinning treatments, but that the level of thinning impacts this response. Stand thinning must be sufficiently intensive to improve old tree growth and health, in part due to increasing available soil moisture. Importantly, focusing stand density reductions around the immediate neighborhood of legacy trees was insufficient to elicit a growth response, calling into question treatments attempting to increase vigor of legacy trees while still maintaining closed canopies in dry, coniferous forest types. Although radial thinning did not affect tree growth rates, this treatment may still achieve other resource objectives not studied here, such as protecting wildlife habitat, reducing the risk of severe fire injury, and decreasing susceptibility to bark beetle attacks.     

Comparison of dwarf bamboos (Indocalamus sp.) leaf parameters to determine relationship between spatial density of plants and total leaf area per plant

The relationship between spatial density and size of plants is an important topic in plant ecology. The self‐thinning rule suggests a ?3/2 power between average biomass and density or a ?1/2 power between stand yield and density. However, the self‐thinning rule based on total leaf area per plant and density of plants has been neglected presumably because of the lack of a method that can accurately estimate the total leaf area per plant. We aimed to find the relationship between spatial density of plants and total leaf area per plant. We also attempted to provide a novel model for accurately describing the leaf shape of bamboos. We proposed a simplified Gielis equation with only two parameters to describe the leaf shape of bamboos one model parameter represented the overall ratio of leaf width to leaf length. Using this method, we compared some leaf parameters (leaf shape, number of leaves per plant, ratio of total leaf weight to aboveground weight per plant, and total leaf area per plant) of four bamboo species of genus Indocalamus Nakai (I. pedalis (Keng) P.C. Keng, I. pumilus Q.H. Dai and C.F. Keng, I. barbatus McClure, and I. victorialis P.C. Keng). We also explored the possible correlation between spatial density and total leaf area per plant using log‐linear regression. We found that the simplified Gielis equation fit the leaf shape of four bamboo species very well. Although all these four species belonged to the same genus, there were still significant differences in leaf shape. Significant differences also existed in leaf area per plant, ratio of leaf weight to aboveground weight per plant, and leaf length. In addition, we found that the total leaf area per plant decreased with increased spatial density. Therefore, we directly demonstrated the self‐thinning rule to improve light interception.     

Restoration of Old Forest Features in Coast Redwood Forests Using Early‐stage Variable‐density Thinning

To accelerate development of old forest features in coast redwood, two thinning treatments and an unthinned control were compared in three treatment areas in north coastal California. One thinning treatment was designed to restore old forest densities of 125 trees/ha and the other 250 trees/ha representing a one‐step and partial treatments to the desired stand density. Four years after treatment, numbers of trees had increased in the thinning treatments due to recruitment of new trees, but had decreased in the control due to self‐thinning. Residual trees increased in stem volume following thinning by 128% in low‐density thinning compared to 70% in the controls indicating thinning accelerated stand development. The thinning treatments also moved the species composition of these stands to a greater proportion of redwood. Considerable slash was produced by the thinning treatments but was decomposing rapidly. Black bears damaged approximately 15% of all trees and more than 38% of residual trees in the thinned treatments compared to less than 2% of all trees in the control. This damage included killing some trees and damaging other trees that survived. Decisions over restoration densities in these stands are complicated by prolonged stand development, and balancing risks and costs. In this case, the bears represent a stochastic factor that dramatically increases risk. Thinning appears to be an effective means of enhancing old forest development by accelerating tree growth, modifying species composition, and increasing stand‐level variability. Continued monitoring will be necessary to evaluate long‐term trends in density relative to effects of bear damage.     

Relationship between size hierarchy and density of trees in a tropical dry deciduous forest of western India

Questions: Density dependence is thought to restrict exponential growth as well as give rise to size structure in populations. Size hierarchy in trees from tropical dry deciduous forests is studied to ask (1) whether nature of competition is symmetric or asymmetric and (2) what is the self thinning trajectory under a natural gradient of tree density. Location: Western India. Methods: Density was measured as the number of trees in 10‐m radius circular plots (n= 96) and size was measured at DBH. Size variation was evaluated by the Gini coefficient (n= 1239 trees). Results: Size inequality between neighbours decreased with density but in a non‐linear manner. In the backdrop of existing theory this indirectly suggests that competitive interactions may be symmetric over a ‘depletive’ resource such as below‐ground water (rather than a ‘pre‐emptive’ resource such as light), which is very plausible in a semi‐arid environment. The self thinning coefficient derived from the relationship between stem diameter and density (γ～?1/4), is higher than expected from existing models of allometric plant growth (γ=?1/3) which are based on above‐ground interactions alone. Seen in conjunction, these results suggest that above‐ground structures, such as stem size, do not adequately represent the outcome of competitive interactions when below‐ground resources, such as water, may be more important under semi‐arid conditions. Conclusions: The non‐linear relationship between size inequality and density indicates that there exists a density threshold beyond which investment in above‐ground biomass becomes sluggish in semi‐arid, deciduous forests. Since current allometric models do not incorporate below‐ground biomass for trees, these data suggest that a more comprehensive allometric growth model may have higher predictive power and wider applicability.     

SEASONAL CHANGES IN SIZE STRUCTURE OF SARGASSUM AND TURBINARIA POPULATIONS (PHAEOPHYCEAE) ON TROPICAL REEF FLATS IN THE SOUTHERN RED SEA1

Seasonal variation in density, thallus length and biomass, population size structure, and allometric length‐biomass relationships was investigated in populations of Sargassum ilicifolium (Turner) C. Agardh, Sargassum subrepandum (Forssk.) C. Agardh, and Turbinaria triquetra (J. Agardh) Kütz. (Phaeophyceae) on shallow reef flats in the southern Red Sea. Thallus length and biomass varied strongly with season, with the highest values occurring in the cooler months. Thallus densities showed no significant temporal variation. Log‐total biomass versus log‐density relationships were positive throughout the growth season without any decrease in the slope of the relationship. In two populations, biomass‐density combinations approached the interspecific biomass‐density line, but the massive annual shedding of modules occurred before self‐thinning would set in. Allometric length‐biomass relationships varied with season in all populations and were associated with seasonal module initiation, growth, and shedding. Evidence of a strong asymmetric competition was found in two high‐density populations. These populations showed a predominance of small thalli during peak development, asymmetrical Lorenz curves, increasing Gini coefficients, and increasing thallus length relative to biomass during the main growth phase. In two other less crowded populations, small thalli were absent during peak development, Lorenz curves were symmetrical, and Gini coefficients decreased during the main growth phase. In these populations, size equalization appears to be due to responses at the modular level rather than size‐dependent mortality. We conclude that changes in size structure in this highly seasonal environment are determined by module dynamics, modified by asymmetric competition in some populations, with a minor role of recruitment and no regulatory effect of self‐thinning.     

沙地樟子松人工林自然稀疏规律

林分在自然稀疏过程中,与密度相关的死亡是由林木大小的增加引起的。因此,很自然地要选择反映林木大小的指标,来表示林木株数与林木大小的关系。选取林分总断面积作为林大小的指标,建立了沙地樟子松人工林自然稀疏过程中林分密度与断面积的相关关系,旨在为沙地樟子松人工林的合理经营提供依据。对自然稀疏－３／２法则进行的评述,认为尽管没有充分证据表明斜率－３／２存在的精确性,但它可以作为一个理想的常数,并且考虑到各     

间伐强度对秦岭南坡栓皮栎天然林种群更新的影响

2006年5月和2011年8月,以秦岭南坡商洛地区近自然经营的不同间伐强度(30%、20%、10%)下栓皮栎天然林为对象,未间伐林分为对照,分析了间伐前后林分的生长情况,以及间伐5年后种群更新、物种多样性和土壤肥力等指标的变化,并对抚育间伐后的林地更新、群落发育等效果进行综合评价.结果表明: 1～6年生栓皮栎幼苗个体数量随着间伐强度的增加而增加,6年生以后的幼苗数量在各间伐强度下无显著差异;间伐10%、20%和30%样地的种群更新潜力分别比对照提高10.8 %、28.5%和32.9%;间伐促进了乔灌层植株(尤其是喜光植物)胸径、树冠以及灌木层植株树高的增加,且随着间伐强度的增加,其促进作用越来越显著;间伐后林地的物种多样性和土壤肥力得到改善,表现为间伐30%>20%>10%>对照样地.在栓皮栎郁闭度≥0.85的天然林中,间伐强度30%(郁闭度保留0.6)更有利于其可持续发育.     

南平溪后杉木林取代杂木林后土壤肥力变化的研究

 本文对福建省南平溪后70年生杉木丰产林与杂木林的土壤肥力进行对比研究。结果表明：与前茬杂木林相比,70年生杉木林下土体较紧实,土壤有机质和营养元素含量降低,土壤酶活性和氧化代谢能力减弱。目前70年生杉木丰产林之所以仍保持较高的土壤肥力水平,主要是杉木林林冠疏开后林下植物多样性提高及生物量增加的缘故。     

Relationship between the virtual dynamic thinning line and the self-thinning boundary line in simulated plant populations

The self-thinning rule defines a straight upper boundary line on log-log scales for all possible combinations of mean individual biomass and density in plant populations. Recently, the traditional slope of the upper boundary line, -3/2, has been challenged by -4/3 which is deduced from some new mechanical theories, like the metabolic theory. More experimental or field studies should be carried out to identify the more accurate self-thinning exponent. But it＇s hard to obtain the accurate self-thinning exponent by fitting to data points directly because of the intrinsic problem of subjectivity in data selection. The virtual dynamic thinning line is derived from the competition-density （C-D） effect as the initial density tends to be positive infinity, avoiding the data selection process. The purpose of this study was to study the relationship between the virtual dynamic thinning line and the upper boundary line in simulated plant stands. Our research showed that the upper boundary line and the virtual dynamic thinning line were both straight lines on log-log scales. The slopes were almost the same value with only a very little difference of 0.059, and the intercept of the upper boundary line was a little larger than that of the virtual dynamic thinning line. As initial size and spatial distribution patterns became more uniform, the virtual dynamic thinning line was more similar to the upper boundary line. This implies that, given appropriate parameters, the virtual dynamic thinning line may be used as the upper boundary line in simulated plant stands.     

Size structure and dynamics of an invasive population of lineage 2 of Asparagopsis taxiformis (Florideophyceae) in the Alboran Sea

In this study, we present basic population data of the red macroalga Asparagopsis taxiformis, widely recognized as invasive in the Mediterranean Sea. A 13‐month field study was carried out on a population located in southern Spain, addressing its phenology, population dynamics and demography. We further tested whether biomass variations were related to environmental variables at the study site. Gametophytes were present year‐round while tetrasporophytes were only found in spring and summer. Recruitment capacity and vegetative growth of the gametophytes are discussed as important modulators for the population structure and enhancers of its persistence. Thallus size‐time histograms revealed a high prevalence of small shoots that showed high mortality that was not related to self‐thinning. Biomass of A. taxiformis was higher from March to July. Vegetative growth was the main way the gametophyte population was maintained, although the presence of tetrasporophytes and fertile gametophytes in the field confirms that sexual reproduction also occurs. Its continuous and high recruitment, in terms of the number of smallest shoots, makes this southern population of A. taxiformis a source of future invasive populations due to the intensive maritime traffic in the region.     

Does soil moisture availability explain liana seedling distribution across a tropical rainfall gradient?

Liana density tends to increase with decreasing rainfall and increasing seasonality. However, the pattern of liana distribution may be due to differences in soil water retention capacity, not rainfall and seasonality per se. We tested the effect of rainfall and soil substrate with respect to the distribution of liana seedlings in six sites across a rainfall gradient from the wet Atlantic to the dry Pacific in central Panama. Soils were either limestone, with low water‐holding capacity, or laterite, with higher water‐holding capacity. We sampled liana seedlings at each site using three 1 × 100 m transect. We found that relative liana seedling density was higher on limestone soils compared to laterite soils regardless of the amount of rainfall. Furthermore, liana community composition on limestone soils was more similar to dry forest sites than to adjacent wet and moist forest sites. Liana seedling species diversity relative to trees was significantly higher in a low‐fertility dry forest site compared to a high‐fertility forest, but did not differ from the other sites. Thus, liana seedling density and community structure may be driven more by soil type and thus by soil moisture availability than strictly by mean annual rainfall and the seasonality of rainfall.     

Mechanical Restoration of California Mixed‐Conifer Forests: Does it Matter Which Trees Are Cut?

The montane ecosystems of northern California have been subjected to repeated manipulation and active fire suppression for over a century, resulting in changes in community structure that contribute to increased wildfire hazard. Ecosystem restoration via reduction of stand density for wildfire hazard mitigation has received substantial attention in recent years; however, many ecological questions remain unanswered. This study compares belowground effects of two alternative forest thinning treatments designed to restore the large, old tree component of late‐seral structure, one of which focuses on restoring Pinus ponderosa dominance (Pine‐preference) and the other of which promotes development of large trees regardless of species (Size‐preference). We evaluated forest floor and soil chemical and microbial parameters in six experimental thinning treatment units of 40 ha each in the Klamath National Forest of northern California 5–6 years after thinning. Inorganic N availability, soil organic C content, phenol oxidase activity, and forest floor C:N ratio were greater in the Size‐preference treatment, whereas forest floor N and soil pH were greater in the Pine‐preference treatment. Our results indicate that these two thinning strategies produce differences in the soil environment that has the potential to affect growth rates of trees that remain, as well as the growth and survivorship of newly established seedlings. Thus, which species/individuals are removed during structural restoration of these mixed‐conifer forests matters both to the belowground components of the ecosystem today and the vegetation and productivity of the ecosystem in future decades.     

疏伐对黄土丘陵区刺槐林蒸腾的影响

黄土高原地区植被建设已达到土壤水分承载力的阈值,需要对现有林分进行结构优化并提升其生态功能。不合理的林分密度是导致黄土丘陵区刺槐林土壤干化、生长衰退的主要原因之一。疏伐可以优化林分结构,并能够通过控制蒸腾耗水来调控土壤水分,是促进刺槐林可持续生长的有效手段。疏伐对黄土丘陵区刺槐林蒸腾有何影响,目前并不清楚。研究基于树干液流法估算了4个不同疏伐强度（样地1：52%、样地2：48%、样地3：35%、样地4：未疏伐）下刺槐单株尺度的液流速率与林分尺度的日平均蒸腾量,并分析了不同时间尺度下液流速率与环境因子的关系,以阐明疏伐对黄土丘陵区刺槐林蒸腾的影响。结果表明：（1） 单株尺度刺槐蒸腾速率（即液流速率）随疏伐强度减小（林分密度增大）呈现下降趋势（样地1：0.53 kg cm^-2 d^-1、样地2：0.41 kg cm^-2 d^-1、样地3：0.31 kg cm^-2 d^-1、样地4：0.33 kg cm^-2 d^-1）;（2） 观测期林分尺度日平均蒸腾量随疏伐强度减小呈现上升趋势（样地1：0.90 mm/d、样地2：1.18 mm/d、样地3：1.04 mm/d、样地4：1.44 mm/d）;（3） 在半小时尺度与日尺度上,各样地液流速率与环境因子的关系没有显著差异,半小时尺度单株液流速率均与太阳辐射相关性最高（相关系数0.883-0.908）,液流速率日变化过程与环境因子日变化过程存在时滞现象;日尺度单株液流速率与饱和水汽压亏缺相关性最高（相关系数0.843-0.913）,样地间日尺度单株液流速率的差异性随着饱和水汽压亏缺增大而增大。研究结果初步反映了疏伐导致的林分密度变化对刺槐蒸腾的影响,将为黄土丘陵区刺槐林的结构改造、功能提升和土壤水分调控提供理论支持。     

抚育间伐对侧柏人工林及林下植被生长的影响

抚育间伐是森林经营管理措施之一,是对森林生态系统的人为干扰。研究不同抚育方式、强度和频率对森林生态系统的干扰对重建、恢复森林生态系统固有功能及实现森林可持续经营管理有重要意义。以31块北京地区侧柏人工林样地对不同抚育强度后侧柏林生长、林下植被生长及林下植物多样性变化进行了研究。研究结果表明:侧柏林分经不同强度抚育后单木胸径、树高、冠幅和林分蓄积生长均增加,不同抚育强度下单木胸径、树高、冠幅生长量和林分蓄积生长量与立地条件有关,好的立地条件宜采取强度抚育,较差立地条件宜采取中弱度抚育;侧柏林分经弱度、中度抚育后林下植物种类增加,经强度抚育后减少;不同强度抚育后林下植物Simpson多样性指数和Shannon-Wiener多样性指数随时间增大,后者对抚育强度更为敏感;林下灌草生物量随抚育强度增大而增加,并随时间增加;侧柏人工林生态系统种间竞争程度与立地条件有关,好的立地条件能够经受较强的人为干扰,差的立地条件受弱度干扰就会打破种间生长平衡。     

Comparing forest structure and biodiversity on private and public land: secondary tropical dry forests in Costa Rica

Secondary forests constitute a substantial proportion of tropical forestlands. These forests occur on both public and private lands and different underlying environmental variables and management regimes may affect post‐abandonment successional processes and resultant forest structure and biodiversity. We examined whether differences in ownership led to differences in forest structure, tree diversity, and tree species composition across a gradient of soil fertility and forest age. We collected soil samples and surveyed all trees in 82 public and 66 private 0.1‐ha forest plots arrayed across forest age and soil gradients in Guanacaste, Costa Rica. We found that soil fertility appeared to drive the spatial structure of public vs. private ownership; public conservation lands appeared to be non‐randomly located on areas of lower soil fertility. On private lands, areas of crops/pasture appeared to be non‐randomly located on higher soil fertility areas while forests occupied areas of lower soil fertility. We found that forest structure and tree species diversity did not differ significantly between public and private ownership. However, public and private forests differed in tree species composition: 11 percent were more prevalent in public forest and 7 percent were more prevalent in private forest. Swietenia macrophylla, Cedrela odorata, and Astronium graveolens were more prevalent in public forests likely because public forests provide stronger protection for these highly prized timber species. Guazuma ulmifolia was the most abundant tree in private forests likely because this species is widely consumed and dispersed by cattle. Furthermore, some compositional differences appear to result from soil fertility differences due to non‐random placement of public and private land holdings with respect to soil fertility. Land ownership creates a distinctive species composition signature that is likely the result of differences in soil fertility and management between the ownership types. Both biophysical and social variables should be considered to advance understanding of tropical secondary forest structure and biodiversity.     

Passive Restoration of Atlantic Forest Following Pinus taeda Harvesting in Southern Brazil

The understory of exotic tree plantations can have non‐negligible native species richness. Ecological restoration of these sites may include the harvest of trees, depending on the tradeoff between timber income and harvest impacts on biodiversity. This study aimed to investigate how a site can recover from harvest disturbance, by comparing the regeneration of woody species in the understory of two types of 37‐year‐old Pinus taeda plantation (P1 and P2, high and low relative density of pine seedlings in the understory, respectively), with stands that were similar to P2 but subjected to harvest and then abandoned for 15 years (R sites). Secondary forests (SF) were used as references. We sampled three different sites for each stand condition; soil chemical properties, estimations of litter mass, and canopy cover were measured. P1 had low species diversity, and P2 and R had 50 and 46% of SF richness, respectively. The R site contained few pine saplings and was floristically similar to P2; this indicated that 15 years was sufficient for the recovery of plant diversity to near pre‐harvesting levels. Soil fertility was highest in SF and lowest in P1. Thus old plantations of P. taeda with low relative density of pine juveniles can be cost‐effective starting points for restoration. Despite the destructive effects of pine harvest, recovery of native species can occur rapidly. In situations in which clearcutting of pine stands is not planned or possible, modest thinning of P. taeda adults and/or intensive thinning of juveniles could expedite restoration.     

落叶松人工林凋落物与土壤肥力变化的研究

以东北东部山区帽儿山实验林场落叶松人工林作为研究对象,从林地凋落量、林地凋落物归还量、凋落物层现存量以及土壤理化性质的变化等方面,研究了经抚育间伐后不同年龄阶段同一落叶松林分土壤肥力的变化以及间伐与未间伐林分土壤理化性质的差异．结果表明,落叶松人工林凋落量和以凋落物形式归还于林地的营养元素呈现出随年龄增加而增长的总趋势．经抚育间伐后,由于阔叶树种的引入使凋落物的组成结构发生变化,凋落物层的分解率逐步提高,加速了凋落物中营养物质的释放和归还,减缓了凋落物积累与分解的矛盾,提高了落叶松人工林的土壤肥力．１５年间土壤有机质、全Ｎ和全Ｐ２Ｏ５平均值分别增长了３５２．００％、２２５．５３％和３４．９６％．间伐后的落叶松人工林土壤的理化性质得到不同程度的改善．     

A Hierarchical Bayesian Model to Predict Self-Thinning Line for Chinese Fir in Southern China

Self-thinning is a dynamic equilibrium between forest growth and mortality at full site occupancy. Parameters of the self-thinning lines are often confounded by differences across various stand and site conditions. For overcoming the problem of hierarchical and repeated measures, we used hierarchical Bayesian method to estimate the self-thinning line. The results showed that the self-thinning line for Chinese fir (Cunninghamia lanceolata (Lamb.)Hook.) plantations was not sensitive to the initial planting density. The uncertainty of model predictions was mostly due to within-subject variability. The simulation precision of hierarchical Bayesian method was better than that of stochastic frontier function (SFF). Hierarchical Bayesian method provided a reasonable explanation of the impact of other variables (site quality, soil type, aspect, etc.) on self-thinning line, which gave us the posterior distribution of parameters of self-thinning line. The research of self-thinning relationship could be benefit from the use of hierarchical Bayesian method.