Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

Description of Tree Distribution Patterns and Their Development Through Marked Gibbs Processes

Tree distribution patterns in forest stands can be considered as marked point processes in the plane. Gibbs processes are a natural tool for modelling such point processes because the interaction between trees can be taken into account and described by pair potential functions. Consequently, tree distribution patterns can be characterized with few parameters and be compared quantitatively. By means of forest stand examples it is shown how these parametric models can be used for the analysis of tree distribution patterns at different time points.     

How does forest landscape structure explain tree species richness in a Mediterranean context?

Although the strong relationship between vegetation and climatic factors is widely accepted, other landscape composition and configuration characteristics could be significantly related with vegetation diversity patterns at different scales. Variation partitioning was conducted in order to analyse to what degree forest landscape structure, compared to other spatial and environmental factors, explained forest tree species richness in 278 UTM 10 × 10 km cells in the Mediterranean region of Catalonia (NE Spain). Tree species richness variation was decomposed through linear regression into three groups of explanatory variables: forest landscape (composition and configuration), environmental (topography and climate) and spatial variables. Additionally, the forest landscape characteristics which significantly contributed to explain richness variation were identified through a multiple regression model. About 60% of tree species richness variation was explained by the whole set of variables, while their joint effects explained nearly 28%. Forest landscape variables were those with a greater pure explanatory power for tree species richness (about 15% of total variation), much larger than the pure effect of environmental or spatial variables (about 2% each). Forest canopy cover, forest area and land cover diversity were the most significant composition variables in the regression model. Landscape configuration metrics had a minor effect on forest tree species richness, with the exception of some shape complexity indices, as indicators of land use intensity and edge effects. Our results highlight the importance of considering the forest landscape structure in order to understand the distribution of vegetation diversity in strongly human-modified regions like the Mediterranean.     

Distribution of functional traits in subtropical trees across environmental and forest use gradients

The relationship between functional traits and environmental factors contribute to understanding community structure and predicting which species will be able to elude environmental filters in different habitats. We selected 10 functional traits related to morphology, demography and regeneration niche in 54 subtropical premontane tree species to describe their main axes of functional differentiation. We derived species traits, environmental variables and species abundance data from 20 1-ha permanent plots established in a seasonal subtropical premontane forest in northwestern Argentina. We analyzed the relationship between species functional traits and environmental factors through RLQ and fourth-corner analyzes. We found an axis of structural differentiation that segregates understory from canopy species, and an axis of functional differentiation that segregates species that maximize resource acquisition from those that promote resource conservation. Environmental and forest use gradients operate hierarchically over subtropical premontane tree species influencing the distribution of demographic and morphological traits. The interaction between climatic and topographic factors influences the distribution of species functional traits at the regional scale. In addition, the history of forest use seems to operate at the landscape scale and explains the distribution of species traits reflecting a trade-off between resource acquisition and resource conservation strategies in secondary forests across different successional stages. Our results support the idea that functional traits may be used to analyze community structure and dynamics through niche differentiation and environmental filtering processes.     

High population differentiation and unusual haplotype structure in a shade-intolerant pioneer tree species, Zanthoxylum ailanthoides (Rutaceae) revealed by analysis of DNA polymorphism at four nuclear loci

Differences in demographic history, life-history traits, and breeding systems affect nucleotide variation patterns. It is expected that shade-intolerant pioneer tree species have different patterns of genetic polymorphism and population structure than climax species. We studied patterns of nucleotide polymorphism at four putative starch pathway loci (agpSA, agpSB, agpL, and GBSSI) in Zanthoxylum ailanthoides , a shade-intolerant pioneer tree species that occupies forest gaps in warm-temperate forests of East Asia. Genetic diversity was lower within each population than among populations, and differentiation among populations was high across the loci ( F _ST = 0.32–0.64), as expected from the insect-pollinated breeding system and the metapopulation structure of this pioneer species. Numbers of haplotypes were smaller than those expected from the observed numbers of segregating sites. Single haplotypes accounted for more than 47% of all the sampled genes at the respective loci. These variation patterns were incompatible with neutral predictions for populations of a finite island model. Complex population dynamics, such as bottleneck and/or admixture, in the history of this pioneer tree species might have resulted in the observed patterns of genetic variation and population structure, which are different from those of climax wind-pollinated tree species, such as conifers. In contrast to the other loci investigated in this study, agpL showed nearly no variation in Z. ailanthoides (one singleton only), but there was some extent of variation in a closely related species, Zanthoxylum schinifolium . This suggests possibly a recent selective sweep at or near the locus in Z. ailanthoides .     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Effects of topography on rainforest tree community structure and diversity in American Samoa, and implications for frugivore and nectarivore populations

Aim We investigated the spatial variation of rainforest tree community structure and composition to determine if forest structure and diversity varied as a function of topography; and in turn if this could influence patterns of habitat use by native forest birds and pteropodid bats. Location The study was undertaken on the island of Tutuila, American Samoa, located in the South Pacific Ocean. Methods All trees ≥10 cm diameter were censused in sixty 200 m² plots in ridge, slope and valley forest across the island of Tutuila. Results Forest structure varied significantly across topographical space. Ridge forest was shortest and had the highest stem densities, and valley forest was tallest with the fewest stems per unit area. Species richness was highest on ridges, and slope and valley forest were more similar in composition with each other than they were with ridge forest. Of the fifty-two tree species encountered in the plots, nine showed a statistical affiliation to one of the three topographical positions. Main conclusions We explain patterns of forest structure and diversity in the context of chronic and catastrophic disturbances. Higher stem densities in ridge forest suggested a higher degree of disturbance on ridges, and this was supported by the fact that the height/diameter ratio of the forest was lowest on ridges, which indicated wind-cropping. We hypothesize the potential effects of topographical variation and known phenological patterns on wildlife abundances. We predict that flowering episodes of ridge-affiliated, bird-visited species (particularly Syzygium inophylloides (A. Gray) C. Muell.; Myrtaceae) will concentrate honeyeater densities on ridges, and that fruiting of the tree Canarium vitiense A. Gray (Burseraceae) could localize populations of the Pacific pigeon (Ducula pacifica). Overall (i.e. net) bat foraging patterns are unlikely to be affected by either flowering or fruiting events. Most of the tree species on Tutuila are generalist in their demographic patterns, and the island is depauperate in wildlife fauna; the evolutionary and conservation implications are discussed. We conclude with the argument that conservation of vertebrate species is essential to maintain the current generalist demographic patterns of Samoan trees.     

Tree size distributions in an old-growth temperate forest

Despite the wide variation in the structural characteristics in natural forests, tree size distribution show fundamental similarities that suggest general underlying principles. The metabolic ecology theory predicts the number of individual scales as the −2 power of tree diameter. The demographic equilibrium theory predicts tree size distribution starting from the relationship of size distributions with growth and mortality at demographic equilibrium. Several analytic predictions for tree size distributions are derived from the demographic equilibrium theory, based on different growth and mortality functions. In addition, some purely phenomenological functions, such as polynomial function, have been used to describe the tree size distributions. In this paper, we use the metabolic ecology theory, the demographic equilibrium theory and the polynomial function to predict the tree size distribution for both the whole community and each species in an old-growth temperate forest in northeastern China. The results show that metabolic ecology theory predictions for the scaling of tree abundance with diameter were unequivocally rejected in the studied forest. Although these predictions of demographic theory are the best models for most of the species in the temperate forest, the best models for some species ( Tilia amurensis , Quercus mongolica and Fraxinus mandshurica ) are compound curves (i.e. rotated sigmoid curves), best predicted by the polynomial function. Hence, the size distributions of natural forests were unlikely to be invariant and the predictive ability of general models was limited. As a result, developing a more sophisticated theory to predict tree size distributions remains a complex, yet tantalizing, challenge.     

Variation in reproduction and growth in declining Scots pine populations

Disentangling how variation in reproduction and growth is linked in plants across different ecological scales, and how allocation rules change in response to stress are fundamental aspects of life history theory. Although it is known that reproductive allocation is an allometric process and that environmental conditions can influence demographic traits, patterns of variation in vegetative and reproductive functions across and within individuals of tree species suffering drought-induced decline have rarely been documented. In this study we use Scots pine (Pinus sylvestris L.) as a model species to explore patterns of variation in cone production and growth in two declining populations at the southern edge of its distribution. A Bayesian approach was used to assess how these demographic traits vary as a function of drought effects and competition and covary across different ecological scales. The allometric trajectories relating tree size with cone production and growth differed along gradients of drought impacts and biotic interactions. Although reproduction and growth increased with tree size, cone production reached a maximum at intermediate sized trees and stabilized or decreased at larger sizes. Drought stress effects (defoliation at the tree level and overall decline at the plot level) and competition for resources reduced cone production and growth. Our results also showed differential effects of defoliation on cone production depending on tree size, with stronger effects on larger individuals. After accounting for these effects, much of the variation of demographic traits and correlations among them occurred at small ecological scales across individuals (i.e. within plots) and within individuals across years. This resulted in covariations between demographic traits among nearby individuals and within individuals through time, suggesting a consistent advantage in resource acquisition of some individuals within plots, and trade-offs between growth and cone production within trees across years. In conclusion, this study reports that drought-induced forest decline is associated with lower growth and cone production in Scots pine, which could contribute to explain the long-term impacts of drought in southern populations of this species and, in particular, its low regeneration capacity after severe drought.     

Habitat structure and disturbance affect small mammal populations in Mediterranean forests

Habitat structure, the physical arrangement of objects in space, affects animal populations, ecological interactions, species diversity, and ecosystem functioning. Conventional forest management practices that reduce habitat structural complexity are thus under reappraisal, and there are increasing efforts to understand how habitat structure affects functionally important species. We investigate here the effect of habitat structures on small mammal dens and their interactions with widespread antagonistic species (wild boars). Among habitat structures we considered tree dimension (diameter), shrub cover, rock cover, dead wood volume, and stump area. Ground cover with rooting signs measured wild boar disturbance. The number of small mammal dens was related positively to rock cover and tree diameter, and negatively to wild boar disturbance. Additionally, a positive interaction between wild boar disturbance and tree diameter emerged, showing that the effect of big trees on small mammal nesting was more evident with higher levels of disturbance. This study suggests that habitat structures that can be affected by forest management provide effective refuges for functionally important species. The effect of habitat structure on species survival and interactions thus needs to be addressed to understand ecosystem management and functioning.     

Forest decline: Endogenous dynamics,tree defenses,and the elimination of spurious correlation

Two important unresolved problems in pollution impact research are early detection of changes in tree and forest growth and determination of the causes of observed growth declines. Consideration of tree defensive systems can shed light on dieback phenomena. Trees possess both chemical and structural defenses, with both types having both active and passive dimensions. When stress or pollution causes a reduction in vigor, defenses may become ineffective, resulting in insect pest and pathogen attack. This leads to a positive feedback cycle of declining vigor. Level of defenses differs by species, site, and life-stage. Consideration of these factors leads to better statistical methods for correlating pollution dose with tree growth response. It is shown that several recent studies are potentially flawed by inadequate consideration of species type, tree age, or stand density. It is shown that forest growth models can be used to correct for endogenous changes in tree growth rates as long as the trees used to derive parameters for the model were not themselves affected by pollution.     

Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.     

Determinants of tree survival at local scale in a sub-tropical forest

Tree survival is a critical driver of stand dynamics, influencing forest structure and composition. Many local-scale drivers (tree size, abiotic and biotic factors) have been proposed as being important in explaining patterns of tree survival, but their contributions are still unknown. We examined the relative importance of these local drivers on tree survival using generalized linear mixed models in an old-growth sub-tropical forest in south China at three levels (community, guild, and species). Among the variables tested, tree size was typically the most important driver of tree survival, followed by abiotic and then biotic variables. Tree size has a strongly positive effect on tree survival for small trees (10–30 cm dbh) and shade-tolerant tree species. Of the abiotic factors tested, elevation tended to be more important in affecting tree survival than other topographic variables. Abiotic factors generally influenced survival of species with relatively high abundances, for individuals in smaller size classes and for mid-tolerant species. Among biotic factors, we found that the mortality of tree species was not driven by density- and frequency-dependent effects in this sub-tropical forest, as indicated by the results of both total basal area of neighbors and the proportion of conspecific neighbors in our study. We conclude that the relative importance of variables driving patterns of tree survival varied greatly among tree size classes, species guilds, shade tolerance, density, and abundance classes in this sub-tropical forest. These results also provide critical information for future studies of forest dynamics and offer insight into forest management in this region.     

The effect of deforestation on the genetic diversity and structure in <Emphasis Type="Italic">Acer saccharum</Emphasis> (Marsh): Evidence for the loss and restructuring of genetic variation in a natural system

The level and distribution of genetic diversity can be influenced by species life history traits and demographic factors, including perturbations that might produce population bottlenecks. Deforestation and forest fragmentation are common sources of population disturbance in contemporary populations of forest ecosystems. Although the genetic effects of forest fragmentation and deforestation have been examined by assessing levels of genetic variation in forest fragments that remain after logging, few considerations have been made of the populations that re-colonize once-cleared areas. Here we examine the effects of human-mediated population bottlenecks on the level and distribution of genetic diversity in natural populations of the long-lived forest tree species, Acer saccharum (sugar maple). We compared genetic variation and structure for populations of sugar maple found within old-growth forested area and in area that has re-colonized since logging. In this study the percent polymorphic loci and allelic richness estimates were reduced in the logged populations compared to old-growth populations. Jackknifed estimates of population genetic differentiation showed significantly higher differentiation among logged populations, with this result being consistently seen when individuals within populations were grouped according to diameter at breast height. The result of decreased genetic variation and higher levels of genetic structure among logged populations suggests that even one extensive bout of logging can alter the level and distribution of genetic variation in this forest tree species.     

Latitudinal comparison of altitudinal changes in forest structure,leaf-type,and species richness in humid monsoon Asia

A new template for mountain vegetation zonation along latitudinal gradients is proposed for examining geographical pattern of various forest attributes in humid monsoon Asia. The contrasting temperature regime in tropical and temperate mountains, i.e., the former is a non-seasonal, temperature-sum controlled environment, and the latter is a seasonal, low temperature limiting environment, leads to different altitudinal patterns of tree height distribution and species richness. In the tropical mountains, both tree height and species richness decrease steeply, and the tree height often stepwise. The decline of tree height and species diversity in the temperate mountains is far less pronounced except near the forest limit. Both trends are explained by their temperature regime.     

后河自然保护区珍稀濒危植物群落乔木层结构特征

对1hm^2固定样地内的亚热带常绿落叶阔叶混交林乔木层树种的胸径级结构和高度级结构以及按径经对乔木树种多样性和均匀度进行了研究。结果显示;群落内乔木树种数和个体数堕落有径级和高度级增加呈下降趋势。树种数和个体数之间存在显著正相关关系。胸径面积之和随胸径级增加呈波动状态,而随高度级的增加呈先升后降的变化。群落的多相关性一指数和均匀度指数都较高,群落内优势种明显,各种的多度差异较小,群落结构较为稳定,其中珍稀植物在各个胸径级和高度级内的树种数和个体数都较高,群落的更新良好。幼苗补充充足,珍稀树种幼苗丰富度和多度较高。     

Lack of demographic equilibrium indicates natural,large‐scale forest dynamics,not a problematic forest conservation policy – a reply to Brzeziecki et al.

Brzeziecki et al. 2016 (Journal of Vegetation Science 27: 460–467.) describe a decrease in population densities and proportion of younger individuals for several tree species in permanent research plots in the core zone of the Bia?owie?a National Park. They attribute insufficient tree recruitment inter alia to the strict protection. Although the authors performed a thorough analysis of tree population dynamics, the scales of the study mean that their far‐reaching conclusions on the causes and consequences of the lack of demographic equilibrium cannot be supported by the data. The inadequate spatial and temporal scales of the study did not allow for the observation of representative population dynamics. Furthermore, they did not compare the results obtained in the strictly protected area with known demographic dynamics of trees in surrounding managed forests or under other forms of nature conservation. Looking from the wider ecosystem perspective, it is clear that strict protection is not a cause for concern and, instead, that such a near‐natural forest manifests population dynamics at rather larger scales.     

Species–habitat associations and demographic rates of forest trees

Niche‐driven effects on demographic processes generated in response to habitat heterogeneity partly shape local distributions of species. Thus, tree distributions are commonly studied in relation to habitat conditions to understand how niche differentiation contributes to species coexistence in forest communities. Many such studies implicitly assume that local abundance reflects habitat suitability, and that abundance is relatively stable over time. We compared models based on abundance with those based on demographic performance for making inferences about habitat association for 287 tree species from three large dynamic plots located in tropical, subtropical and temperate forests. The correlation between the predictions of the abundance‐based models and the demography‐based models varied widely, with correlation coefficients ranging nearly from ?1 to 1.This suggests that the two types of models capture different information about species–habitat associations. Demography‐based models evaluate habitat quality by focusing on population processes and thus should be preferred for understanding responses of tree species to habitat conditions, especially when habitat conditions are changing and species–habitat interactions cannot be considered to be at equilibrium.     

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。     

台风对森林的影响

台风通过树枝折断、吹落叶果、产生倒木和折干等许多途径影响林分结构和动态。森林受害程度随树种、林龄、森林类型、树高和地形而异。高密度的森林通常具有较差的根系和较大的树高/胸径比值,在台风袭击下,往往具有较高的受损和死亡的风险。台风疏开郁闭的林冠层,促进了先锋树种的大量增加、生长和成熟,形成的林隙也为个体更新提供了机会。强风造成了土壤基质的多样化,从而促进了实生苗和幼树的更新和生物多样性的增加。台风也通过改变粗木质残体,枯枝落叶层,地洞和土墩,以及繁殖可用性来影响生物多样性。台风产生的粗死木和枯枝落叶使森林的碳储量迅速归还土壤,并影响土壤的养分分布。台风减少了动物的食物供应和恶化栖息地的环境,减少鸟的数量,促进昆虫扩散。受害森林给害虫滋生提供了场所。今后的研究热点是受台风干扰森林的长期监测,不同森林土壤的有机碳贮藏,土壤和养分流失规律,台风和其他自然灾害的交叉影响,改进数学模型以准确预测台风损害。