Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

Anthropogenic impacts upon plant species richness and net primary productivity in California

We assess the importance of anthropogenic land‐use, altered productivity, and species invasions for observed productivity–richness relationships in California. To this end, we model net primary productivity (NPP) c. 1750 AD and at present (1982–1999) and map native and exotic vascular plant richness for 230 subecoregions. NPP has increased up to 105% in semi‐arid areas and decreased up to 48% in coastal urbanized areas. Exotic invasions have increased local species diversity up to 15%. Human activities have reinforced historical gradients in species richness but reduced the spatial heterogeneity of NPP. Structural equation modelling suggests that, prior to European settlement, NPP and richness were primarily controlled by precipitation and other abiotic variables, with NPP mediating richness. Abiotic variables remain the strongest predictors of present NPP and richness, but intermodel comparisons indicate a significant anthropogenic impact upon statewide distributions of NPP and richness. Exotic and native species each positively correlate to NPP after controlling for other variables, which may help explain recent reports of positively associated native and exotic richness.     

The relationships between net primary productivity, human population density and species conservation

Aim In this study, I determine the relationships between net primary productivity (NPP), human population density, species richness and land use. I also examine the implications of human settlement patterns for species conservation. Location Australia. Methods I document the associations between NPP, human population density and the species richness of birds, butterflies and mammals using correlations and spatial regressions. I also assess changes in land‐use with NPP and population density, focussing particularly on protected areas. An initial exploration into the implications of the NPP‐population density relationship for regional conservation strategies is provided. Results Human population density increases with NPP suggesting that available energy may be a key driving force of human settlement patterns. The species richness of each taxonomic group and geographically restricted species also increases with NPP leading to substantial overlap between species diversity and populated regions. The percentage of land designated as minimal use decreases considerably with increasing human population density and NPP, while intensive agriculture is confined entirely to areas of high NPP. There are strong negative relationships between the size of Australia's National Parks and human population density and NPP. Small parks are often surrounded by relatively dense settlements, but have high average NPP, while large parks are mostly isolated and characterized by low productivity. There are no areas in the highest quartile of NPP that also occur in the most sparsely populated regions, presenting challenges for conservation strategies wanting to protect productive areas under the least threat of human development. Main conclusions Human population density and species richness respond similarly to variation in NPP, leading to spatial congruence between human settlements and productive, species rich regions. Planning strategies are required that minimize the potential threat posed by human development to diverse ecosystems and maximize the underlying productivity of protected areas. Reducing the level of threat may require stabilizing the size of the human population, while capturing larger areas of relatively high productivity in the conservation reserve system would lead to greater protection of local diversity.     

深圳福田红树林植被碳储量和净初级生产力

红树林是滨海湿地“蓝碳”的主要类型之一.准确和定位评估不同植物群落的固碳能力,对于红树林保育管理和恢复造林具有指导作用.本研究对深圳福田红树林4种代表性群落(白骨壤群落、秋茄群落、海桑群落、无瓣海桑群落)的各个植被碳库组分(乔木植物生物量碳库、林下灌丛碳库、呼吸根碳库、枯立木碳库、枯倒木碳库和枯枝落叶层碳库等)进行调查,计算各群落的植被碳储量,并通过生长增量-凋落物产量法计算得到各群落的净初级生产力.结果表明: 白骨壤群落、秋茄群落、海桑群落和无瓣海桑群落的植被碳储量分别为28.7、127.6、100.1、73.6 t C·hm^-2,各群落的净初级生产力分别为8.75、7.67、9.60、11.87 t C·hm^-2·a^-1.位于深圳市中心的福田红树林,每年固定大气CO₂高达4000 t.本研究结果将为红树林“蓝碳”碳汇功能的评估提供理论指导,并为我国红树林碳汇林建设提供依据.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Tree species richness and density affect parasitoid diversity in cacao agroforestry

In Brazil, cacao is mostly planted beneath shade trees. The diversity of shade trees varies from monospecific to highly diverse canopies, characteristic of pristine Atlantic Forest. This study evaluates the relationships between family richness of Hymenoptera-Parasitica and Chrysidoidea, and tree species richness and density, the species richness of herbaceous understorey, and the area and age of the cacao agroforestry system. We sampled 16 cacao agroforestry systems, with canopy diversity ranging from one to 22 tree species per hectare, in three seasons: summer (March), winter (August) and spring (November). Parasitoids were sampled using eight Malaise-Townes traps per site. Tree species richness and density were enumerated within 1 ha at each site, and herbaceous plant species richness was calculated in eight 1 m² plots, within the hectare. The number of parasitoid families increased with tree species richness and density in spring and summer, but decreased in winter. Neither species richness of herbaceous plants nor area and age of the system affected parasitoid family richness. We suggest that the increase of parasitoid diversity with tree species richness and density in warmer seasons reflects increasing heterogeneity and availability of resources. The decrease in parasitoid family number with tree density in winter may be due to local impoverishment of resources, leading to parasitoid emigration to neighbouring forest remnants. This result implies that a higher diversity of shade trees will help to maintain high parasitoid levels and, in consequence, higher levels of natural enemies of cacao pests, particularly in the warmer seasons. This prediction is borne out in the experience of cacao producers. The proper management of shade tree diversity will play a vital role in maintaining the sustainability of cacao agroforestry production systems in the tropics and, concurrently, will maintain high biodiversity values in these locations.     

Intraspecific trait variation improves the detection of deterministic community assembly processes in early successional forests,but not in late successional forests

Aims Intraspecific trait variation (ITV) has been increasingly recognized to play an important role in understanding the underlying processes influencing community assembly. However, gaps remain in our understanding of how incorporating ITV will influence the relative importance of deterministic (e.g. habitat filtering, limiting similarity) and stochastic processes in driving community assembly at different successional stages.     

Modeling analysis of primary controls on net ecosystem productivity of seven boreal and temperate coniferous forests across a continental transect

Process‐based models are effective tools to synthesize and/or extrapolate measured carbon (C) exchanges from individual sites to large scales. In this study, we used a C‐ and nitrogen (N)‐cycle coupled ecosystem model named CN‐CLASS (Carbon Nitrogen‐Canadian Land Surface Scheme) to study the role of primary climatic controls and site‐specific C stocks on the net ecosystem productivity (NEP) of seven intermediate‐aged to mature coniferous forest sites across an east–west continental transect in Canada. The model was parameterized using a common set of parameters, except for two used in empirical canopy conductance–assimilation, and leaf area–sapwood relationships, and then validated using observed eddy covariance flux data. Leaf Rubisco‐N dynamics that are associated with soil–plant N cycling, and depend on canopy temperature, enabled the model to simulate site‐specific gross ecosystem productivity (GEP) reasonably well for all seven sites. Overall GEP simulations had relatively smaller differences compared with observations vs. ecosystem respiration (RE), which was the sum of many plant and soil components with larger variability and/or uncertainty associated with them. Both observed and simulated data showed that, on an annual basis, boreal forest sites were either carbon‐neutral or a weak C sink, ranging from 30 to 180 g C m^?2 yr^?1; while temperate forests were either a medium or strong C sink, ranging from 150 to 500 g C m^?2 yr^?1, depending on forest age and climatic regime. Model sensitivity tests illustrated that air temperature, among climate variables, and aboveground biomass, among major C stocks, were dominant factors impacting annual NEP. Vegetation biomass effects on annual GEP, RE and NEP showed similar patterns of variability at four boreal and three temperate forests. Air temperature showed different impacts on GEP and RE, and the response varied considerably from site to site. Higher solar radiation enhanced GEP, while precipitation differences had a minor effect. Magnitude of forest litter content and soil organic matter (SOM) affected RE. SOM also affected GEP, but only at low levels of SOM, because of low N mineralization that limited soil nutrient (N) availability. The results of this study will help to evaluate the impact of future climatic changes and/or forest C stock variations on C uptake and loss in forest ecosystems growing in diverse environments.     

Species richness and species turnover in a successional heathland

Abstract. Changes in species richness and species turnover during secondary succession following experimental disturbance were studied in eight permanent plots in a species‐rich dry heathland in the southern part of the Czech Republic. The treatments applied were sod‐cutting, burning, cutting of above‐ground biomass, and control. The plots were sampled annually between 1992 and 2000; species richness was analysed at three scales, 25 cm × 25 cm, 1 m × 1 m, and 3 m × 3 m. Disturbances resulted in increased species richness. The highest vascular plant richness was attained during the secondary succession after sod‐cutting, where species established on exposed bare ground. Less severe disturbances by burning and cutting also caused a slight increase in the number of vascular plant species. For bryophytes and lichens, the highest increase in the number of species was also found in the sod‐cut plots, where all cryptogams were removed by the disturbance. At the scale of 3 m × 3 m, species richness of both vascular plants and cryptogams peaked in 1995–1996, i.e. 3–4 yrs after the disturbance, and slowly decreased or slightly fluctuated without any trend thereafter. At smaller scales it either peaked later or constantly increased over the entire observation period of 9 yrs. Species mobility, expressed as species accumulation over time, was lower than reported from grasslands. Rates of species turnover, calculated as Jaccard dissimilarity between pairs of consecutive years, corresponded across different scales. This implies that successional dry heathlands have a higher small‐scale mobility than heathlands which are apparently stable at larger scale.     

Biogeographic historical legacies in the net primary productivity of Northern Hemisphere forests

It has been suggested that biogeographic historical legacies in plant diversity may influence ecosystem functioning. This is expected because of known diversity effects on ecosystem functions, and impacts of historical events such as past climatic changes on plant diversity. However, empirical evidence for a link between biogeographic history and present‐day ecosystem functioning is still limited. Here, we explored the relationships between Late‐Quaternary climate instability, species‐pool size, local species and functional diversity, and the net primary productivity (NPP) of Northern Hemisphere forests using structural equation modelling. Our study confirms that past climate instability has negative effects on plant functional diversity and through that on NPP, after controlling for present‐day climate, soil conditions, stand biomass and age. We conclude that global models of terrestrial plant productivity need to consider the biogeographical context to improve predictions of plant productivity and feedbacks with the climate system.     

Some autecological characteristics of early to late successional tree species in Venezuela

The breadth of the continuum concept of strategy with respect to succession was tested on 21 tree and shrub species common in either unlogged or logged stands, respectively, in the Forest Reserve of Caparo, Venezuela, by examining morphological, physiological and population characteristics. Based on a preliminary abundance analysis, `early', `mid' and `late' successional species as well as `generalists' were distinguished. Early successional species, i.e. Ochroma lagopus, Heliocarpus popayanensis and Cecropia peltata were similar in many autecological aspects, e.g. monolayered leaf arrangement, orthotropic architectural models, no adaptive reiteration, clumped distribution, but differed in gap association and distribution along a drainage gradient. Mid-successional species established themselves both in large and small gaps (> 300 m[sup2 ]; 80–300 m[sup2 ]) and showed a clumped to regular distribution pattern in logged areas; they exhibited more diverse crown and leaf characteristics than early successional species. Late successional species established themselves only in small gaps and understorey, and showed a regular spatial pattern in undisturbed areas. All late successional species displayed architectural models with plagiotropic lateral axes and showed a multilayered leaf arrangement. Adaptive reiteration was a common feature of late successional species which could be further subdivided into large, medium-sized and small trees, indicating different light requirements at maturity. Generalists were common treelet and shrub species in both disturbed and undisturbed sites where they are also capable of completing their life cycle. The light compensation point (LCP) of an individual plant was strongly influenced by its crown illuminance. Large late successional species showed the widest range of LCP values, reflecting the increasing light availability with increasing height in mature forest. On the basis of many autecological characteristics, it was found (i) that there is in fact a continuum of species strategies with respect to succession even among early and mid-successional species and (ii) that the latter group of species showed the widest breadth of autecological traits, reflecting the heterogeneous environment in which they establish and mature.     

Factors influencing epiphytic bryophyte and lichen species richness at different spatial scales in managed temperate forests

The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.     

Small-scale dynamics and species richness in successional alvar plant communities

Species richness and small-scale spatial dynamics (turnover) were compared in four sites during 1990-1993 1) former alvar grassland, now young Pinus sylvestris forest, 2) clear-cut in 1990, 3) clear-cut and grazed by sheep, 4) nearby grassland It was hypothesized that in communities which show large-scale dynamics, small-scale dynamics is also more rapid to support succession Area-based species richness and shoot density increased in the cut and grazed site No significant dynamics was found in shoot-based species richness Therefore the forest was considered to be stable and the cut and grazed site large-scale dynamical (successional) Almost equally high small-scale turnover values were found in all sites, which supports the carousel model for successional communities, but means we had to reject the initial hypothesis The successional and intrinsic small-scale dynamics of the communities studied were two separate processes which acted at different scales, but the scale could not be only spatially determined, it depended also on the shoot density The overgrowing of this former alvar grassland has resulted in the decrease of shoot density and thus the decrease of species richness per unit area, but the total number of species has not changed much Consequently the possibility to restore grassland communities remains     

Bird species richness patterns of northern Taiwan: primary productivity, human population density, and habitat heterogeneity

Energy, climate, habitat heterogeneity, and human activity are important correlates of spatial variation in species richness. We examined the correlation between species richness and these variables using the birds that breed in northern Taiwan. We conducted general linear models (GLMs) and spatial correlation models to examine the relationship between bird species richness (BSR) and environmental variables. We found that normalized difference vegetation index (NDVI) was the most important predictor of BSR. We suggest productivity is the primary process of BSR. Additionally, we hypothesized that scale dependency might exist in the relationship between BSR and NDVI in Taiwan. Human population density, the second most important factor, was inversely correlated with BSR. The factor and BSR did not have similar response to NDVI, which contradicted observations in most of the previous studies on human population vs. species richness. We proposed that the human population density had an effect on NDVI, which in turn had an effect on BSR. Moreover, we hypothesized that the contradiction between our study and the previous studies might arise from a higher level of human disturbance in Taiwan than in other areas. The necessity of conserving native species in intensively developed lowlands of Taiwan cannot be overemphasized. Number of land cover type was another significant predictor of BSR. Habitat heterogeneity may have an effect on BSR in Taiwan.     

On the relationships between leaf-litter lignin and net primary productivity in tropical rain forests

We investigated if tropical rainforest trees produced more-lignified leaves in less productive environments using forests on Mount Kinabalu, Borneo. Our investigation was based on two earlier suggestions that slower litter decomposition occurs under less productive forests and that trees under resource limitation invest a large amount of carbon as lignin as a defense substance to minimize the loss from herbivores. When nine forests at different altitudes (700–3100 m) and soil conditions (derived from sedimentary or ultrabasic rocks) but with the same gentle relief position were compared, the concentrations of leaf-litter lignin were positively correlated with litterfall rates and leaf-litter nitrogen concentrations. These patterns would be reinforced in intact leaves if the effects of resorption at the time of leaf shedding were taken into account, because greater magnitude of resorption of mobile elements but not of lignin would occur in less productive environments (i.e. dilution of lignin in intact leaves). These results did not support earlier suggestions to explain the variation of leaf-litter lignin. Instead, we suggest that lower lignin contents are adaptive to recycle minerals without retarding decomposition in less productive environments.     

Soil microbiome mediates positive plant diversity‐productivity relationships in late successional grassland species

Which processes drive the productivity benefits of biodiversity remain a critical, but unanswered question in ecology. We tested whether the soil microbiome mediates the diversity‐productivity relationships among late successional plant species. We found that productivity increased with plant richness in diverse soil communities, but not with low‐diversity mixtures of arbuscular mycorrhizal fungi or in pasteurised soils. Diversity‐interaction modelling revealed that pairwise interactions among species best explained the positive diversity‐productivity relationships, and that transgressive overyielding resulting from positive complementarity was only observed with the late successional soil microbiome, which was both the most diverse and exhibited the strongest community differentiation among plant species. We found evidence that both dilution/suppression from host‐specific pathogens and microbiome‐mediated resource partitioning contributed to positive diversity‐productivity relationships and overyielding. Our results suggest that re‐establishment of a diverse, late successional soil microbiome may be critical to the restoration of the functional benefits of plant diversity following anthropogenic disturbance.