Review of forest landscape models: Types, methods, development and applications

Forest landscape models simulate forest change through time using spatially referenced data across a broad spatial scale (i.e. landscape scale) generally larger than a single forest stand. Spatial interactions between forest stands are a key component of such models. These models can incorporate other spatio-temporal processes such as natural disturbances (e.g. wildfires, hurricanes, outbreaks of native and exotic invasive pests and diseases) and human influences (e.g. harvesting and commercial thinning, planting, fire suppression). The models are increasingly used as tools for studying forest management, ecological assessment, restoration planning, and climate change. In this paper, we define forest landscape models and discuss development, components, and types of the models. We also review commonly used methods and approaches of modeling forest landscapes, their application, and their strengths and weaknesses. New developments in computer sciences, geographic information systems (GIS), remote sensing technologies, decision-support systems, and geo-spatial statistics have provided opportunities for developing a new generation of forest landscape models that are increasingly valuable for ecological research, restoration planning and resource management.     

Review of forest landscape models: Types, methods, development and applications

Forest landscape models simulate forest change through time using spatially referenced data across a broad spatial scale (i.e. landscape scale) generally larger than a single forest stand. Spatial interactions between forest stands are a key component of such models. These models can incorporate other spatio-temporal processes such as natural disturbances (e.g. wildfires, hurricanes, outbreaks of native and exotic invasive pests and diseases) and human influences (e.g. harvesting and commercial thinning, planting, fire suppression). The models are increasingly used as tools for studying forest management, ecological assessment, restoration planning, and climate change. In this paper, we define forest landscape models and discuss development, components, and types of the models. We also review commonly used methods and approaches of modeling forest landscapes, their application, and their strengths and weaknesses. New developments in computer sciences, geographic information systems (GIS), remote sensing technologies, decision-support systems, and geo-spatial statistics have provided opportunities for developing a new generation of forest landscape models that are increasingly valuable for ecological research, restoration planning and resource management.     

Temperate forest fragments maintain aboveground carbon stocks out to the forest edge despite changes in community composition

Edge effects are among the primary mechanisms by which forest fragmentation can influence the link between biodiversity and ecosystem processes, but relatively few studies have quantified these mechanisms in temperate regions. Carbon storage is an important ecosystem function altered by edge effects, with implications for climate change mitigation. Two opposing hypotheses suggest that aboveground carbon (AGC) stocks at the forest edge will (a) decrease due to increased tree mortality and compositional shifts towards smaller, lower wood density species (e.g., as seen in tropical systems) or, less often, (b) increase due to light/temperature-induced increases in diversity and productivity. We used field-based measurements, allometry, and mixed models to investigate the effects of proximity to the forest edge on AGC stocks, species richness, and community composition in 24 forest fragments in southern Quebec. We also asked whether fragment size or connectivity with surrounding forests altered these edge effects. AGC stocks remained constant across a 100 m edge-to-interior gradient in all fragment types, despite changes in tree community composition and stem density consistent with expectations of forest edge effects. We attribute this constancy primarily to compensatory effects of small trees at the forest edge; however, it is due in some cases to the retention of large trees at forest edges, likely a result of forest management. Our results suggest important differences between temperate and tropical fragments with respect to mechanisms linking biodiversity and AGC dynamics. Small temperate forest fragments may be valuable in conservation efforts based on maintaining biodiversity and multiple ecosystem services.     

Biodiversity and climate determine the functioning of Neotropical forests

Aim

Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community‐mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance).

Location

Neotropics.

Methods

We used data from 26 sites, 201 1‐ha plots and >92,000 trees distributed across the Neotropics. We quantified for each site water availability and soil total exchangeable bases and for each plot three key community‐weighted mean functional traits that are important for biomass stocks and productivity. We used structural equation models to test the hypothesis that all drivers have independent, positive effects on biomass stocks and dynamics.

Results

Of the relationships analysed, vegetation attributes were more frequently associated significantly with biomass stocks and dynamics than environmental conditions (in 67 vs. 33% of the relationships). High climatic water availability increased biomass growth and stocks, light disturbance increased biomass growth, and soil bases had no effect. Rarefied tree species richness had consistent positive relationships with biomass stocks and dynamics, probably because of niche complementarity, but was not related to net biomass productivity. Community‐mean traits were good predictors of biomass stocks and dynamics.

Main conclusions

Water availability has a strong positive effect on biomass stocks and growth, and a future predicted increase in (atmospheric) drought might, therefore, potentially reduce carbon storage. Forest attributes, including species diversity and community‐weighted mean traits, have independent and important relationships with AGB stocks, dynamics and ecosystem functioning, not only in relatively simple temperate systems, but also in structurally complex hyper‐diverse tropical forests.     

The variation of tree beta diversity across a global network of forest plots

Aims With the aim of understanding why some of the world's forests exhibit higher tree beta diversity values than others, we asked: (1) what is the contribution of environmentally related variation versus pure spatial and local stochastic variation to tree beta diversity assessed at the forest plot scale; (2) at what resolution are these beta‐diversity components more apparent; and (3) what determines the variation in tree beta diversity observed across regions/continents? Location World‐wide. Methods We compiled an unprecedented data set of 10 large‐scale stem‐mapping forest plots differing in latitude, tree species richness and topographic variability. We assessed the tree beta diversity found within each forest plot separately. The non‐directional variation in tree species composition among cells of the plot was our measure of beta diversity. We compared the beta diversity of each plot with the value expected under a null model. We also apportioned the beta diversity into four components: pure topographic, spatially structured topographic, pure spatial and unexplained. We used linear mixed models to interpret the variation of beta diversity values across the plots. Results Total tree beta diversity within a forest plot decreased with increasing cell size, and increased with tree species richness and the amount of topographic variability of the plot. The topography‐related component of beta diversity was correlated with the amount of topographic variability but was unrelated to its species richness. The unexplained variation was correlated with the beta diversity expected under the null model and with species richness. Main conclusions Because different components of beta diversity have different determinants, comparisons of tree beta diversity across regions should quantify not only overall variation in species composition but also its components. Global‐scale patterns in tree beta diversity are largely coupled with changes in gamma richness due to the relationship between the latter and the variation generated by local stochastic assembly processes.     

Tackling unresolved questions in forest ecology: The past and future role of simulation models

1. Understanding the processes that shape forest functioning, structure, and diversity remains challenging, although data on forest systems are being collected at a rapid pace and across scales. Forest models have a long history in bridging data with ecological knowledge and can simulate forest dynamics over spatio‐temporal scales unreachable by most empirical investigations.
2. We describe the development that different forest modelling communities have followed to underpin the leverage that simulation models offer for advancing our understanding of forest ecosystems.
3. Using three widely applied but contrasting approaches – species distribution models, individual‐based forest models, and dynamic global vegetation models – as examples, we show how scientific and technical advances have led models to transgress their initial objectives and limitations. We provide an overview of recent model applications on current important ecological topics and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.
4. Synthesis. This overview shows that forest models, due to their complementarity and mutual enrichment, represent an invaluable toolkit to address a wide range of fundamental and applied ecological questions, hence fostering a deeper understanding of forest dynamics in the context of global change.

Forest models can help understanding the processes that shape forest functioning, structure and diversity, since they can can simulate forest dynamics over spatio‐temporal scales unreachable by most empirical investigations. Here we describe the development of three widely applied but contrasting forest mo−delling approaches — species distribution models, individual‐based models and dynamic global vegetation models. We provide an overview of recent model applications and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.     

Species dynamics and colonization patterns in an abandoned forest in an urban landscape

Species dynamics in an abandoned urban forest of Central Japan is described in this paper. The dominant species in the urban plantation were Cryptomeria japonica and Chamaecyparis obtusa. A variety of eight patches of the canopy was produced by previous forest management practices. Progressive and retrogressive species dynamics within these eight patches are investigated in this paper. The study elucidates the deterministic role of patchiness in the nature of species colonization and the maintenance of species diversity in an urban forest. Altogether 139 native and/or naturalized species, including 23 shrub and 35 tree species, were recorded in the study area of 3.2 ha. The performance of species varied according to their successional attributes indicating a selective canopy influence. Twenty percent of the tree species were shade-intolerant pioneers (e.g. Cornus spp., Rhus javanica var. roxburghii) re-established under selective tree-felling. Thirty percent were shade-tolerant climax species (e.g. Neolitsea sericea, Persea thunbergii) dominant in remnant closed patches. The remaining 50% belonged to various seral types with aggressive deciduous species (e.g. Aphananthe aspera, Celtis sinensis) in most of the patches. Some ruderal herbaceous species dominated heavily disturbed clear-felled patches. This study suggests that canopy modification influences the subsequent colonization pattern. Furthermore, heterogeneous patches contribute to greater species diversity and dynamics in isolated woodlands.     

Meta-analysis of tree diversity effects on the abundance,diversity and activity of herbivores' enemies

The natural enemies hypothesis predicts that the abundance and diversity of antagonists such as predators and parasitoids of herbivores increases with the diversity of plants, which can lead to more effective top-down control of insect herbivores. However, although the hypothesis has received large support in agricultural systems, fewer studies have been conducted in forest ecosystems and a comprehensive synthesis of previous research is still lacking.We conducted a meta-analysis of 65 publications comparing the diversity, abundance or activity of various groups of natural enemies (including birds, bats, spiders and insect parasitoids) in pure vs. mixed forest stands. We tested the effects of forest biome, natural enemy taxon and type of study (managed vs experimental forest).We found a significant positive effect of forest tree diversity on natural enemy abundance and diversity but not on their activity. The effect of tree diversity on natural enemies was stronger towards lower latitudes but was not contingent on the natural enemy taxon level.Overall, our study contributes toward a better understanding of the “natural enemies hypothesis” in forest systems and provides new insights about the mechanisms involved. Furthermore, we outline potential avenues for strengthening forest resistance to the growing threat of herbivorous insects.     

Predicting prey population dynamics from kill rate, predation rate and predator-prey ratios in three wolf-ungulate systems

1.?Predation rate (PR) and kill rate are both fundamental statistics for understanding predation. However, relatively little is known about how these statistics relate to one another and how they relate to prey population dynamics. We assess these relationships across three systems where wolf-prey dynamics have been observed for 41 years (Isle Royale), 19 years (Banff) and 12 years (Yellowstone). 2.?To provide context for this empirical assessment, we developed theoretical predictions of the relationship between kill rate and PR under a broad range of predator-prey models including predator-dependent, ratio-dependent and Lotka-Volterra dynamics. 3.?The theoretical predictions indicate that kill rate can be related to PR in a variety of diverse ways (e.g. positive, negative, unrelated) that depend on the nature of predator-prey dynamics (e.g. structure of the functional response). These simulations also suggested that the ratio of predator-to-prey is a good predictor of prey growth rate. That result motivated us to assess the empirical relationship between the ratio and prey growth rate for each of the three study sites. 4.?The empirical relationships indicate that PR is not well predicted by kill rate, but is better predicted by the ratio of predator-to-prey. Kill rate is also a poor predictor of prey growth rate. However, PR and ratio of predator-to-prey each explained significant portions of variation in prey growth rate for two of the three study sites. 5.?Our analyses offer two general insights. First, Isle Royale, Banff and Yellowstone are similar insomuch as they all include wolves preying on large ungulates. However, they also differ in species diversity of predator and prey communities, exploitation by humans and the role of dispersal. Even with the benefit of our analysis, it remains difficult to judge whether to be more impressed by the similarities or differences. This difficulty nicely illustrates a fundamental property of ecological communities. Second, kill rate is the primary statistic for many traditional models of predation. However, our work suggests that kill rate and PR are similarly important for understanding why predation is such a complex process.     

Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.     

Plant growth strategy determines the magnitude and direction of drought-induced changes in root exudates in subtropical forests

Root exudates are an important pathway for plant–microbial interactions and are highly sensitive to climate change. However, how extreme drought affects root exudates and the main components, as well as species-specific differences in response magnitude and direction, are poorly understood. In this study, root exudation rates of total carbon (C) and its components (e.g., sugar, organic acid, and amino acid) were measured under the control and extreme drought treatments (i.e., 70% throughfall reduction) by in situ collection of four tree species with different growth rates in a subtropical forest. We also quantified soil properties, root morphological traits, and mycorrhizal infection rates to examine the driving factors underlying variations in root exudation. Our results showed that extreme drought significantly decreased root exudation rates of total C, sugar, and amino acid by 17.8%, 30.8%, and 35.0%, respectively, but increased root exudation rate of organic acid by 38.6%, which were largely associated with drought-induced changes in tree growth rates, root morphological traits, and mycorrhizal infection rates. Specifically, trees with relatively high growth rates were more responsive to drought for root exudation rates compared with those with relatively low growth rates, which were closely related to root morphological traits and mycorrhizal infection rates. These findings highlight the importance of plant growth strategy in mediating drought-induced changes in root exudation rates. The coordinations among root exudation rates, root morphological traits, and mycorrhizal symbioses in response to drought could be incorporated into land surface models to improve the prediction of climate change impacts on rhizosphere C dynamics in forest ecosystems.     

Spatial dynamics in model plant communities: what do we really know?

A variety of models have shown that spatial dynamics and small-scale endogenous heterogeneity (e.g., forest gaps or local resource depletion zones) can change the rate and outcome of competition in communities of plants or other sessile organisms. However, the theory appears complicated and hard to connect to real systems. We synthesize results from three different kinds of models: interacting particle systems, moment equations for spatial point processes, and metapopulation or patch models. Studies using all three frameworks agree that spatial dynamics need not enhance coexistence nor slow down dynamics; their effects depend on the underlying competitive interactions in the community. When similar species would coexist in a nonspatial habitat, endogenous spatial structure inhibits coexistence and slows dynamics. When a dominant species disperses poorly and the weaker species has higher fecundity or better dispersal, competition-colonization trade-offs enhance coexistence. Even when species have equal dispersal and per-generation fecundity, spatial successional niches where the weaker and faster-growing species can rapidly exploit ephemeral local resources can enhance coexistence. When interspecific competition is strong, spatial dynamics reduce founder control at large scales and short dispersal becomes advantageous. We describe a series of empirical tests to detect and distinguish among the suggested scenarios.     

Species pool size and rainfall account for the relationship between biodiversity and biomass production in natural forests of China

The strength of biodiversity–biomass production relationships increases with increasing environmental stress and time. However, we know little about the effects of abiotic (e.g., climate) and biotic (e.g., species pool and community composition) factors on this trend. Whether variation in biomass production is best explained by phylogenetic diversity metrics or traditional measures of species richness also remains elusive. We compiled estimates of community composition and biomass production for tree species in 111 permanent quadrats spanning three natural forests (tropical, subtropical, and temperate) in China. Based on ~10 years of data, we compared temperature, rainfall, species pool size, and community composition in each forest each year. We estimated species richness and phylogenetic diversity in each quadrat each year; the latter metric was based on the sum of branch lengths of a phylogeny that connects species in each quadrat each year. Using generalized linear mixed‐effect models, we found that top‐ranked models included the interaction between forest and biodiversity and the interaction between forest and year for both biodiversity metrics. Variation in biomass production was best explained by phylogenetic diversity; biomass production generally increased with phylogenetic diversity, and the relationship was stronger in subtropical and temperate forests. Increasing species pool size, temperature, and rainfall and decreasing inter‐quadrat dissimilarity range shifted the relationship between biomass production and phylogenetic diversity from positive to neutral. When considered alone, species pool size had the strongest influence on biomass production, while species pool size, rainfall, and their interaction with phylogenetic diversity constituted the top‐ranked model. Our study highlights the importance of species pool size and rainfall on the relationship between phylogenetic diversity and biomass production in natural forest ecosystems.     

Resistance to wildfire and early regeneration in natural broadleaved forest and pine plantation

The response of an ecosystem to disturbance reflects its stability, which is determined by two components: resistance and resilience. We addressed both components in a study of early post-fire response of natural broadleaved forest (Quercus robur, Ilex aquifolium) and pine plantation (Pinus pinaster, Pinus sylvestris) to a wildfire that burned over 6000 ha in NW Portugal. Fire resistance was assessed from fire severity, tree mortality and sapling persistence. Understory fire resistance was similar between forests: fire severity at the surface level was moderate to low, and sapling persistence was low. At the canopy level, fire severity was generally low in broadleaved forest but heterogeneous in pine forest, and mean tree mortality was significantly higher in pine forest. Forest resilience was assessed by the comparison of the understory composition, species diversity and seedling abundance in unburned and burned plots in each forest type. Unburned broadleaved communities were dominated by perennial herbs (e.g., Arrhenatherum elatius) and woody species (e.g., Hedera hibernica, Erica arborea), all able to regenerate vegetatively. Unburned pine communities presented a higher abundance of shrubs, and most dominant species relied on post-fire seeding, with some species also being able to regenerate vegetatively (e.g., Ulex minor, Daboecia cantabrica). There were no differences in diversity measures in broadleaved forest, but burned communities in pine forest shared less species and were less rich and diverse than unburned communities. Seedling abundance was similar in burned and unburned plots in both forests. The slower reestablishment of understory pine communities is probably explained by the slower recovery rate of dominant species. These findings are ecologically relevant: the higher resistance and resilience of native broadleaved forest implies a higher stability in the maintenance of forest processes and the delivery of ecosystem services.     

Predicting alpha diversity of African rain forests: models based on climate and satellite‐derived data do not perform better than a purely spatial model

Aim Our aim was to evaluate the extent to which we can predict and map tree alpha diversity across broad spatial scales either by using climate and remote sensing data or by exploiting spatial autocorrelation patterns. Location Tropical rain forest, West Africa and Atlantic Central Africa. Methods Alpha diversity estimates were compiled for trees with diameter at breast height ≥ 10 cm in 573 inventory plots. Linear regression (ordinary least squares, OLS) and random forest (RF) statistical techniques were used to project alpha diversity estimates at unsampled locations using climate data and remote sensing data [Moderate Resolution Imaging Spectroradiometer (MODIS), normalized difference vegetation index (NDVI), Quick Scatterometer (QSCAT), tree cover, elevation]. The prediction reliabilities of OLS and RF models were evaluated using a novel approach and compared to that of a kriging model based on geographic location alone. Results The predictive power of the kriging model was comparable to that of OLS and RF models based on climatic and remote sensing data. The three models provided congruent predictions of alpha diversity in well‐sampled areas but not in poorly inventoried locations. The reliability of the predictions of all three models declined markedly with distance from points with inventory data, becoming very low at distances > 50 km. According to inventory data, Atlantic Central African forests display a higher mean alpha diversity than do West African forests. Main conclusions The lower tree alpha diversity in West Africa than in Atlantic Central Africa may reflect a richer regional species pool in the latter. Our results emphasize and illustrate the need to test model predictions in a spatially explicit manner. Good OLS or RF model predictions from inventory data at short distance largely result from the strong spatial autocorrelation displayed by both the alpha diversity and the predictive variables rather than necessarily from causal relationships. Our results suggest that alpha diversity is driven by history rather than by the contemporary environment. Given the low predictive power of models, we call for a major effort to broaden the geographical extent and intensity of forest assessments to expand our knowledge of African rain forest diversity.     

Paradoxical persistence through mixed-system dynamics: towards a unified perspective of reversal behaviours in evolutionary ecology

Counterintuitive dynamics of various biological phenomena occur when composite system dynamics differ qualitatively from that of their component systems. Such composite systems typically arise when modelling situations with time-varying biotic or abiotic conditions, and examples range from metapopulation dynamics to population genetic models. These biological, and related physical, phenomena can often be modelled as simple financial games, wherein capital is gained and lost through gambling. Such games have been developed and used as heuristic devices to elucidate the processes at work in generating seemingly paradoxical outcomes across a spectrum of disciplines, albeit in a field-specific, ad hoc fashion. Here, we propose that studying these simple games can provide a much deeper understanding of the fundamental principles governing paradoxical behaviours in models from a diversity of topics in evolution and ecology in which fluctuating environmental effects, whether deterministic or stochastic, are an essential aspect of the phenomenon of interest. Of particular note, we find that, for a broad class of models, the ecological concept of equilibrium reactivity provides an intuitive necessary condition that must be satisfied in order for environmental variability to promote population persistence. We contend that further investigations along these lines promise to unify aspects of the study of a range of topics, bringing questions from genetics, species persistence and coexistence and the evolution of bet-hedging strategies, under a common theoretical purview.     

Functional diversity of carbon-gain, water-use, and leaf-allocation traits in trees of a threatened lowland dry forest in Hawaii

We examined carbon-gain, water-use, and leaf-allocation traits for six tree species of a Hawaiian dry forest to better understand the functional diversity within this threatened ecosystem. Tropical dry forests are among the most endangered ecosystems on Earth, and in Hawaii, as elsewhere, declining biodiversity threatens ecosystem processes that may depend on forest functional diversity. We found broad variation among species including a two-fold difference for mean photosynthetic rate, a greater than three-fold difference for predawn water potential, and a nearly three-fold difference for leaf life span. Principal component analysis showed a clear separation of species based on carbon-gain vs. water-use related axes, and δ(13)C analysis revealed differing limitations (supply vs. demand) on carbon assimilation. The broad functional variation not only spanned traditional classifications (avoiders vs. tolerators), but also included unusual strategies (e.g., fast growth with drought tolerance). Correlations among traits, including leaf life span, leaf mass per area, and %N, followed typical global patterns, but some exceptions appeared as a result of unique life-history characteristics, such as latex-rich sap and root parasitism. Elucidating functional variation provides important information that can be used to link plant biodiversity with ecosystem processes and also facilitate the management and preservation of tropical dry forests and other threatened communities.     

Species and structural diversity affect growth of oak,but not pine,in uneven-aged mature forests

The effects of mixing tree species on tree growth and stand production have been abundantly studied, mostly looking at tree species diversity effects while controlling for stand density and structure. Regarding the shift towards managing forests as complex adaptive systems, we also need insight into the effects of structural diversity. Strict forest reserves, left for spontaneous development, offer unique opportunities for studying the effects of diversity in tree species and stand structure. We used data from repeated inventories in ten forest reserves in the Netherlands and northern Belgium to study the growth of pine and oak. We investigated whether the diversity of a tree's local neighbourhood (i.e., species and structural diversity) is important in explaining its basal area growth. For the subcanopy oak trees, we found a negative effect of the tree species richness of the local neighbours, which – in the studied forests – was closely related to the share of shade-casting tree species in the neighbourhood. The growth of the taller oak trees was positively affected by the height diversity of the neighbour trees. Pine tree growth showed no relation with neighbourhood diversity. Tree growth decreased with neighbourhood density for both species (although no significant relationship was found for the small pines). We found no overall diversity-growth relationship in the studied uneven-aged mature forests; the relationship depended on tree species identity and the aspect of diversity considered (species vs. structural diversity).     

秦岭太白山弃耕地植物群落演替的生态学研究 Ⅱ演替系列的群落α多样性特征

应用物种丰富度、Ｓｉｍｐｓｏｎ指数、Ｓｈａｎｎｏｎ－Ｗｉｅｎｅｒ指数、Ｐｉｅｌｏｕ均匀度指数、Ａｌａｔａｌｏ均匀度指数研究了太白山弃耕地植物群落次生演替过程中的群落α多样性动态特征,结果表明,群落在由一年生草本植物群落阶段向多年生草本植物群落、灌丛、混交林阶段的演替过程中,群落多样性指数逐渐上升,至混交林阶段达到最高值。若以分布在这一垂直地带的代表性森林群落锐齿栎林为演替进一步发展的方向,则由多样性指数在锐齿栎混交林、锐齿栎林阶段呈下降趋势。不同演替阶段不同生长型的多样性指数变化规律为：在演替的初期,草本层多样性指数＞灌木层＞乔木层;在混交林阶段,这３个不同生长型的多样性指数相差无几,乔木生长型稍占优势;在锐齿栎混交林和锐齿栎林阶段,灌木生长型的物种多样性最大,草本次之,乔木生长型的物种多样性则迅速降低。首次利用群落各生长型的叶层相对厚度和相对盖度作为加权参数,对群落总体的物种多样性指数进行了测度,结果表明,这种加权处理是比较合理的。