海南岛霸王岭热带低地雨林树木的空间格局

树木空间格局及其形成过程是物种共存及生物多样性维持机制研究的一个重要方面。该文以海南岛两个1 hm ²的典型热带低地雨林老龄林森林动态样地为基础, 通过4个点格局模型(均质Poisson过程、异质Poisson过程、均质Thomas过程和异质Thomas过程)模拟扩散限制和生境异质性作用对树木空间分布格局的影响, 并分析不同空间尺度下(< 2 m, 2-5 m, 5-10 m, 10-15 m, 15-20 m和20-25 m)不同作用的相对重要性。结果表明: 热带低地雨林的所有树木总体上呈现聚集分布的空间格局, 随着尺度的增大, 聚集强度逐渐减小。树种在模拟空间分布格局最优模型中的比例由高到低分别是: 均质Thomas过程, 均质Poisson过程、异质Thomas过程和异质Poisson过程。扩散限制作用是形成热带低地雨林树木空间分布格局最重要的生态过程, 其次是完全随机作用以及生境异质性和扩散限制的联合作用, 而生境异质性的作用最小。不同空间尺度上模拟各树种空间分布格局的最优模型比例差异显著, 扩散限制作用能够在多数空间尺度上模拟多个树种的空间分布格局, 其次为随机作用; 生境异质性和扩散限制的联合作用主要在小尺度(0-5 m)影响树种分布, 而生境异质性在较大尺度(15-25 m)上影响树种的空间分布格局。     

Habitat niche partitioning by 16 species of Myristicaceae in Amazonian Ecuador

The distribution and spatial pattern of plants in tropical forests have important implications for how species interact with each other and their environments. In this article we use a large-scale permanent census plot to address if the coexistence of 16 co-occurring species of Myristicaceae is aided by topographic and light gradient niche partitioning. We used a second order spatial pattern analysis based on Ripley’s K function to describe species’ distributions and associations among species, and a torus translation procedure to test for associations with three topographically defined habitats. A majority of species displayed spatial aggregation and over half had one or more significant habitat associations. Four species were associated with the ridge habitat, four species with slope habitat, and two with valley habitat. Seven other species showed no habitat association. Within each habitat group, species exhibited a variety of distributions in relation to light availability. However, habitat associations were largely unexplained by differential rates of mortality, growth or recruitment over a 5-year interval. We conclude that although in principle partitioning of the topographic and light environments may double or treble the number of species able to coexist, there is no evidence that partitioning of physical habitats can explain the coexistence of all 16 of these closely related species.     

海南尖峰岭热带山地雨林林冠层树种功能多样性特征

以海南尖峰岭热带山地雨林3块1 hm²样地为研究对象,利用11个林冠功能性状结合样地地形及林冠乔木树种样地清查数据,分别基于单维性状和多维性状比较物种多度加权对群落功能离散度指数——平均成对距离(MPD)和平均最近类群距离(MNTD)的影响;同时分析林冠层功能丰富度(FRic)与物种丰富度之间的关系,最后利用零模型探讨不同生境类型下标准化效应MPD和MNTD(经过物种多度加权且剔除群落物种丰富度差异影响)的变化,进而评价林冠层群落水平功能多样性格局及其对局域生境异质性的响应.结果表明: 功能性状维度和物种多度对MPD的影响强烈,不同维度功能性状多度加权前后MPD相关性较弱(R=0.359～0.628);但对MNTD的影响相对较弱,不同维度功能性状多度加权前后MNTD相关性较强(R=0.746～0.820);未经物种多度加权的MPD和MNTD均普遍高估了林冠层的功能离散度.林冠层功能丰富度与物种丰富度基本呈指数相关关系(F=128.20;R²=0.632;AIC=97.72;P＜0.001),且功能丰富度很有可能存在一定的物种丰富度阈值.基于不同维度功能性状的林冠层功能多样性格局及其生境响应存在一定的差异性.在生物竞争激烈的低沟生境中,林冠层功能多样性倾向于比预期零模型随机产生的功能多样性高,林冠树种功能性状表现出离散分布;而在其他生境类型中,林冠层功能多样性倾向于接近或低于随机产生的功能多样性,林冠树种功能性状随机或聚集分布.     

Survival and growth of dominant tree seedlings in seasonally tropical dry forests of Yucatan: site and fertilization effects

Aims Seasonally tropical dry forests of the Yucatan Peninsula are typically found in sites with nutrient-poor soils because of the recent geological origin of the region. The landscape is dominated by extensive karstic plates that shape environments where vegetation regeneration through seed germination may be limited by the availability of suitable microsites. In this study, we documented the survival and growth of seedlings from three dominant tree species (Bursera simaruba, Piscidia piscipula and Lysiloma latisiliquum) in seasonally tropical dry forests in Yucatan. Specifically, we evaluated the effect of nutrient addition (N and P, separately and in combination) on seedling survival and growth across three sites with differing levels of precipitation.Methods We conducted a nutrient addition experiment, whereby we established 12 plots of dimensions 10×10 m (100 m 2) at each site, from which three plots were randomly selected to receive one of four treatments: N addition, P addition, N and P addition and no nutrient addition (controls). Prior to treatment application, in each plot, we planted 10 seedlings of each species in October 2010 and subsequently conducted surveys of plant growth and survival every 20 days from November 2010 to April 2011.Important findings Overall, nutrient addition increased seedling survival and the magnitude of this effect was similar among sites. We did not observe an additive effect of the N + P treatment on survival. Similarly, we observed a positive effect of nutrient addition on seedling growth, but this effect was contingent upon site; regarding survival, the effects of N and P on seedling growth were not additive. These results suggest that seedling recruitment and growth in the three dominant species of trees in Yucatan are limited by nutrient availability but that the magnitude of this effect, particularly on seedling growth, is specific for species and site.     

海南五指山热带山地雨林植物物种多样性研究

王指山热带山地雨林的物种种类十分丰富,在１ｈｍ＾２样地中共出现５４个科,１００个属的乔木树种１７７种,１３３７个个体。区系地理分析表明属的分布区类型以热带分布型成分占优,为８９．０％,充分体现了五指山山地雨林的热带性质。多样性指数分析结果为：样地１Ｍａｒｇａｌｅｆ指数１７．８２２,Ｓｈａｎｎｏｎ－Ｗｉｅｎｅｒ指数５．６２１,均匀度０．８２３,Ｓｉｍｐｓｏｎ指数０．０５０,均匀度６．７７５,表明五指     

西双版纳热带季节雨林的树种组成和群落结构动态

研究了西双版纳热带季节雨林1 hm2(hectare)动态监测样地1993年与2007年之间树种组成和群落结构的变化。对样地中胸径≥5 cm的乔木进行了每木调查。目前其树种组成的热带分布科、属所占比例分别为91%和94%,具有较高比例的热带植物区系性质。在1993年与2007年两次调查之间,树种数量由145种增至179种,仅有1到2个个体的稀有树种所占比例从54%降为51.1%。从森林的垂直结构来看,A、B、C三层的个体死亡率分别为12.8%、12.9%和19.0%,各层树木的增长率分别为-8.5%、-1.4%和44.8%。与此相对应,C层小径级的树木所占比例有较大提高。虽然小径级的树木在种类和数量上比例增大,但个体数量和种类组成相对稳定的A、B层优势树种变化不大,维持了群落结构的稳定性。14 a间,群落中新增加的具有先锋性质的树种不超过5个。1993年时,A、B两层尚有先锋树种存在,2007年已经从A、B两层中退出。因此,从14 a间树种组成和群落结构的变化来看,虽然具有树木的死亡和增补,但其物种成分和群落结构的总体格局没有明显的变化,处于动态平衡过程中。     

热带森林中的斑块动态与物种多样性维持

斑块作为景观要素之一,直接到景观结构的空间格局及其内部各要素之间的相互关系,其动态也将导致景观格局的变化。异质性的斑块在自然森林中是普遍存在的,这就是说,顶极森林中仍然存在着由不同种类或不同生长时期的植物种群组成的森林斑块。由自然或人为干扰所驱动的森林生长循环导致敢这些斑块在空间上的镶嵌,对于持定的地域片段,也导致了不同生长时期的森林斑块的周期性循环。在热带森林中,森林的生长循环由林窗期（ｇａｐ     

滇南热带雨林物种多样性的取样面积探讨

通过不同面积样方种数的比较、物种多样性指数等的计算和逐步扩大样地面积的调查, 本文研究了滇南热带雨林种数/面积关系、个体/种数关系、物种多样性及树种的频度分布规律, 认为滇南热带雨林群落学调查的最适取样(样方)面积是0.25 hm², 这个面积接近该类型热带雨林理论上的群落最小表现面积。 为能体现一个具体森林(群落)类型的基本的植物区系组成, 需设置4～5个这样的样方(总面积累计1 hm²以上)。从与世界不同地区热带雨林的比较亦可见,云南热带雨林有类似的树种频度分布规律,单位面积上的物种多样性比典型的东南亚低地热带雨林稍低,但比非洲的热带雨林要高。     

Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain

Aims Our study aimed to understand the effects and the relative importance of biotic neighborhood and habitat heterogeneity for tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain, north-eastern China.
Methods The generalized mixed linear model was used to examine the relative effects of biotic neighborhood and habitat heterogeneity on seedling survival over two years.
Important findings Our results showed that both biotic neighborhood and habitat heterogeneity had significant effects on the seedling survival at community level. The local environment suitable for the adult growth was also suitable for seedling survival. The soil moisture and soil available nitrogen exhibited significant positive effects on seedling survival. On the other hand, seedling density had significant negative effects on seedling survival due to the individual competition. Particularly, we found significant negative density-dependent effects on seedling survival which was caused by conspecific adult and seedling neighbors. As expected, with the increasing of seedling survival age, the habitat heterogeneity became more important on seedling survival. These results suggest that both local biotic neighborhood and habitat heterogeneity drive seedling survival in this temperate forest, and their relative importance varies with different seedling age classes and species traits.     

长白山次生针阔混交林乔木幼苗存活的影响因素分析

基于长白山次生针阔混交林样地, 以520个1 m × 1 m幼苗样方中胸径小于1 cm的乔木幼苗为研究对象, 选取2013年和2014年的幼苗调查数据, 运用广义线性混合模型(GLMM)分析了生物邻体和生境异质性对幼苗存活的影响, 探讨了次生针阔混交林幼苗存活影响因素及物种共存机制。结果表明: (1)适宜大树生长的局域生境同样也适宜幼苗的存活, 幼苗存活率与土壤含水量和有效氮等土壤养分显著正相关。(2)幼苗个体之间存在明显的竞争, 较多的幼苗邻体显著降低幼苗的存活率。同种大树邻体和同种幼苗邻体与幼苗存活显著负相关, 表明存在负密度制约效应。(3)随着幼苗年龄的增加, 生境异质性对幼苗存活的影响逐渐增大。该研究证实了密度制约效应和生境异质性对幼苗存活有着重要影响, 其相对重要性随着幼苗年龄级、功能群以及物种种类而变化。     

Evolutionary history and distance dependence control survival of dipterocarp seedlings

One important hypothesis to explain tree-species coexistence in tropical forests suggests that increased attack by natural enemies near conspecific trees gives locally rare species a competitive advantage. Host ranges of natural enemies generally encompass several closely related plant taxa suggesting that seedlings should also do poorly around adults of closely related species. We investigated the effects of adult Parashorea malaanonan on seedling survival in a Bornean rain forest. Survival of P. malaanonan seedlings was highest at intermediate distances from parent trees while heterospecific seedlings were unaffected by distance. Leaf herbivores did not drive this relationship. Survival of seedlings was lowest for P. malaanonan , and increased with phylogenetic dissimilarity from this species, suggesting that survival of close relatives of common species is reduced. This study suggests that distance dependence contributes to species coexistence and highlights the need for further investigation into the role of shared plant enemies in community dynamics.
Ecology Letters (2010) 13: 51–59     

Commonness,rarity, and intraspecific variation in traits and performance in tropical tree seedlings

One of the few rules in ecology is that communities are composed of many rare and few common species. Trait‐based investigations of abundance distributions have generally focused on species‐mean trait values with mixed success. Here, using large tropical tree seedling datasets in China and Puerto Rico, we take an alternative approach that considers the magnitude of intraspecific variation in traits and growth as it relates to species abundance. We find that common species are less variable in their traits and growth. Common species also occupy core positions within community trait space indicating that they are finely tuned for the available conditions. Rare species are functionally peripheral and are likely transients struggling for success in the given environment. The work highlights the importance of considering intraspecific variation in trait‐based ecology and demonstrates asymmetry in the magnitude of intraspecific variation among species is critical for understanding of how traits are related to abundance.     

Phylogenetic turnover in tropical tree communities: impact of environmental filtering,biogeography and mesoclimatic niche conservatism

Aim We addressed the roles of environmental filtering, historical biogeography and evolutionary niche conservatism on the phylogenetic structure of tropical tree communities with the following questions. (1) What is the impact of mesoclimatic gradients and dispersal limitation on phylogenetic turnover and species turnover? (2) How does phylogenetic turnover between continents compare in intensity with the turnover driven by climatic gradients at a regional scale? (3) Are independent phylogenetic reconstructions of the mesoclimatic niche of clades congruent between continents? Location Panama Canal Watershed and Western Ghats (India), two anciently divergent biogeographic contexts but with comparable rainfall gradients. Methods Using floristic data for 50 1‐ha plots in each region, independent measures of phylogenetic turnover (Π_ST) and species turnover (Jaccard) between plots were regressed on geographic and ecological distances. Mesoclimatic niches were reconstructed for each node of the phylogeny and compared between the two continents. Results (1) The phylogenetic turnover within each region is best explained by mesoclimatic differences (environmental filtering), while species turnover depends both on mesoclimatic differences and geographic distances (dispersal limitation). (2) The phylogenetic turnover between continents (Π_ST = 0.009) is comparable to that caused by mesoclimatic gradients within regions (Π_ST = 0.010) and both effects seem cumulative. (3) Independent phylogenetic reconstructions of the mesoclimatic niches were strongly correlated between the two continents (r = 0.61), despite the absence of shared species. Main conclusions Our results demonstrate a world‐wide deep phylogenetic signal for mesoclimatic niche within a biome, indicating that positive phylogenetic turnover at a regional scale reflects environmental filtering in plant communities.     

Effects of local biotic neighbors and habitat heterogeneity on seedling survival in a spruce‐fir valley forest,northeastern China

Seedlings are vulnerable to many biotic and abiotic agents, and studying seedling dynamics helps understand mechanisms of species coexistence. In this study, the relative importance of biotic neighbors and habitat heterogeneity to seedling survival was examined by generalized linear mixed models for 33 species in a spruce‐fir valley forest in northeastern China. The results showed that the relative importance of these factors varied with species and functional groups. Conspecific negative density dependence (CNDD) was important to the survival of Abies nephrolepis and Picea koraiensis seedling, whereas phylogenetic negative density dependence (PNDD) was critical to Pinus koraiensis and Betula platyphylla, as well as functional groups of tree, deciduous, and shade‐intolerant seedlings. For shrubs and Acer ukurunduense, habitat heterogeneity was significant. Despite of the significance of CNDD, PNDD, and habitat heterogeneity on seedling survival, large proportions of the total variance were not accounted for by the studied variables, suggesting the needs to examine the influences of other factors such as pests, diseases, herbivores, forest structure, species functional traits, and microclimatic conditions on seedling survival in the future.     

Distance‐dependent effects of soil‐derived biota on seedling survival of the tropical tree legume Ormosia semicastrata

Question: How do seed germination and subsequent seedling survival of O. semicastrata (Hance forma litchiifolia How) vary with respect to distance from parent trees and conspecific density in different types of tropical forest? Are there effects of soil biota on O. semicastrata that systematically depend on distance from parent trees and conspecific density? Do soil pathogens differently affect survival of O. semicastrata in different types of tropical forest? Location: Tropical lowland rain forest and tropical montane rain forest in Jianfengling National Nature Reserve, Hainan Island, China. Methods: Individual adult O. semicastrata trees were selected in lowland rain forest and montane rain forest. Soil was collected at a distance of 0‐5 m or 15‐20 m from the parent tree. Soil samples from each distance were combined into a bulk sample. Half of the soil sample was sterilized by autoclaving. Surface‐sterilized seeds were then added to the soil material in shade‐houses at both forests. Results: Germination of O. semicastrata seeds at low‐ or high‐seed density was barely affected by the sterilization procedure. In both forests, seedlings grown in non‐sterilized soil collected close to parent trees had significantly higher mortality compared to those in sterilized soil. In contrast, seedling survival with soil collected far from parent trees was not affected by the soil sterilization procedure. Conclusions: Host‐specific pathogens concentrated in the soil around parent trees may regulate community structure of tropical trees at the stage of seedling development.     

Spatial and temporal distribution of chloroplast DNA polymorphism in a tropical tree species

The level and the spatial organization of chloroplast DNA polymorphism were investigated in Dicorynia guianensis Hamshoff (Caesalpiniaceae) at different spatial and temporal scales. D. guianensis is a canopy tree of the rain forest that is distributed throughout the Guiana plateau in small aggregates. Twelve different haplotypes were identified using restriction analysis of polymerase chain reaction (PCR) amplified fragments of the chloroplast genome. When populations from different areas of French Guiana were compared, a clear geographical pattern of haplotype frequencies was identified along the Atlantic coast. This pattern is most likely the result of the restriction-expansion dynamics of the tropical forest during the Quaternary. At the local level, D. guianensis was characterized by a high level of within population diversity. Maintenance of within population diversity results from the dynamics of the aggregates; stochastic demography associated with the turnover of aggregates generates genetic differentiation among them. At the stand level, a strong spatial aggregation of haplotypes persisted from the adult to the seedling cohort indicating limited seed flow. There was also a strong difference in levels of diversity between the cohorts which suggested that recruitment over several years is needed in order to maintain genetic diversity during regeneration.     

Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity

Neutral and niche theories give contrasting explanations for the maintenance of tropical tree species diversity. Both have some empirical support, but methods to disentangle their effects have not yet been developed. We applied a statistical measure of spatial structure to data from 14 large tropical forest plots to test a prediction of niche theory that is incompatible with neutral theory: that species in heterogeneous environments should separate out in space according to their niche preferences. We chose plots across a range of topographic heterogeneity, and tested whether pairwise spatial associations among species were more variable in more heterogeneous sites. We found strong support for this prediction, based on a strong positive relationship between variance in the spatial structure of species pairs and topographic heterogeneity across sites. We interpret this pattern as evidence of pervasive niche differentiation, which increases in importance with increasing environmental heterogeneity.