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1.
Fungal pathogens are implicated in driving tropical plant diversity by facilitating strong, negative density‐dependent mortality of conspecific seedlings (C‐NDD). Assessment of the role of fungal pathogens in mediating coexistence derives from relatively few tree species and predominantly the Neotropics, limiting our understanding of their role in maintaining hyper‐diversity in many tropical forests. A key question is whether fungal pathogen‐mediated C‐NDD seedling mortality is ubiquitous across diverse plant communities. Using a manipulative shadehouse experiment, we tested the role of fungal pathogens in mediating C‐NDD seedling mortality of eight mast fruiting Bornean trees, typical of the species‐rich forests of South East Asia. We demonstrate species‐specific responses of seedlings to fungicide and density treatments, generating weak negative density‐dependent mortality. Overall seedling mortality was low and likely insufficient to promote overall community diversity. Although conducted in the same way as previous studies, we find little evidence that fungal pathogens play a substantial role in determining patterns of seedling mortality in a SE Asian mast fruiting forest, questioning our understanding of how Janzen‐Connell mechanisms structure the plant communities of this globally important forest type.  相似文献   

2.
Forest community structure may be influenced by seedling density dependence, however, the effect is loosely coupled with population dynamics and diversity in the short term. In the long term the strength of conspecific density dependence may fluctuate over time because of seedling abundance, yet few long‐term studies exist. Based on 11 years of seedling census data and tree census data from a 25‐ha temperate forest plot in Northeast China, we used generalized linear mixed models to test the relative effects of local neighborhood density and abiotic factors on seedling density and seedling survival. Spatial point pattern analysis was used to determine if spatial patterns of saplings and juveniles, in relation to conspecific adults, were in accordance with patterns uncovered by conspecific negative density dependence at the seedling stage. Our long‐term results showed that seedling density was mainly positively affected by conspecific density, suggesting dispersal limitation of seedling development. The probability of seedling survival significantly decreased over 1 year with increasing conspecific density, indicating conspecific negative density dependence in seedling establishment. Although there was variation in conspecific negative density dependence at the seedling stage among species and across years, a dispersed pattern of conspecific saplings relative to conspecific adults at the local scale (<10 m) was observed in four of the 11 species examined. Overall, sapling spatial patterns were consistent with the impacts of conspecific density on seedling dynamics, which suggests that conspecific negative density dependence is persistent over the long term. From the long‐term perspective, conspecific density dependence is an important driver of species coexistence in temperate forests.  相似文献   

3.
Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4‐year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat‐specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.  相似文献   

4.
The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO2‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO2 would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.  相似文献   

5.
The species‐area relationship (SAR) has proven to be one of the few strong generalities in ecology. The temporal analog of the SAR, the species‐time relationship (STR), has received considerably less attention. Recent work primarily from the temperate zone has aimed to merge the SAR and the STR into a synthetic and unified species‐time‐area relationship (STAR) as originally envisioned by Preston (1960). Here we test this framework using two tropical tree communities and extend it by deriving a phylogenetic‐time‐area relationship (PTAR). The work finds some support for Preston's prediction that diversity‐time relationships, both species and phylogenetic, are sensitive to the spatial scale of the sampling. Contrary to the Preston's predictions we find a decoupling of diversity‐area and diversity‐time relationships in both forests as the time period used to quantify the diversity‐area relationship changes. In particular, diversity‐area and diversity‐time relationships are positively correlated using the initial census to quantify the diversity‐area relationship, but weakly or even negatively correlated when using the most recent census. Thus, diversity‐area relationships could forecast the temporal accumulation of biodiversity of the forests, but they failed to “back‐cast” the temporal accumulation of biodiversity suggesting a decoupling of space and time.  相似文献   

6.
A life‐history trade‐off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged‐over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species‐specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed‐effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed‐effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade‐off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade‐off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.  相似文献   

7.
Precipitation patterns are changing across the globe causing more severe and frequent drought for many forest ecosystems. Although research has focused on the resistance of tree populations and communities to these novel precipitation regimes, resilience of forests is also contingent on recovery following drought, which remains poorly understood, especially in aseasonal tropical forests. We used rainfall exclusion shelters to manipulate the interannual frequency of drought for diverse seedling communities in a tropical forest and assessed resistance, recovery and resilience of seedling growth and mortality relative to everwet conditions. We found seedlings exposed to recurrent periods of drought altered their growth rates throughout the year relative to seedlings in everwet conditions. During drought periods, seedlings grew slower than seedlings in everwet conditions (i.e., resistance phase) while compensating with faster growth after drought (i.e., recovery phase). However, the response to frequent drought was species dependent as some species grew significantly slower with frequent drought relative to everwet conditions while others grew faster with frequent drought due to overcompensating growth during the recovery phase. In contrast, mortality was unrelated to rainfall conditions and instead correlated with differences in light. Intra‐annual plasticity of growth and increased annual growth of some species led to an overall maintenance of growth rates of tropical seedling communities in response to more frequent drought. These results suggest these communities can potentially adapt to predicted climate change scenarios and that plasticity in the growth of species, and not solely changes in mortality rates among species, may contribute to shifts in community composition under drought.  相似文献   

8.
Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.  相似文献   

9.
Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N‐limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old‐growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N‐mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha?1 yr?1 (Low‐N); 100 kg N ha?1 yr?1 (Medium‐N), and 150 kg N ha?1 yr?1 (High‐N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low‐to‐medium levels of N addition (≤100 kg N ha?1 yr?1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine‐root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition‐based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high‐N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.  相似文献   

10.
Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.  相似文献   

11.
Climate and other global environmental changes are major threats to ecosystem functioning and biodiversity. However, the importance of plant diversity in mitigating the responses of functioning of natural ecosystems to long‐term environmental change remains unclear. Using inventory data of boreal forests of western Canada from 1958 to 2011, we found that aboveground biomass growth increased over time in species‐rich forests but decreased in species‐poor forests, and importantly, aboveground biomass loss from tree mortality was smaller in species‐rich than species‐poor forests. A further analysis indicated that growth of species‐rich (but not species‐poor) forests was statistically positively associated with rising CO2, and that mortality in species‐poor forests increased more as climate moisture availability decreased than it did in species‐rich forests. In contrast, growth decreased and mortality increased as the climate warmed regardless of species diversity. Our results suggest that promoting high tree diversity may help reduce the climate and environmental change vulnerability of boreal forests.  相似文献   

12.
The effects of global glaciations on the distribution of organisms is an essential element of many diversification models. However, the empirical evidence supporting this idea is mixed, in particular with respect to explaining tropical forest evolution. In the present study, we evaluated the impacts of range shifts associated with Pleistocene global glacial cycles on the evolution of tropical forests. In particular, we tested the predictions: (1) that population genetic structure increases with fragmentation variation between the present and the Last Glacial Maximum (LGM) and also (2) with geographical range instability; and (3) that genetic diversity increases with range stability and (4) decreases with fragmentation variation between periods. To address our predictions, we studied population genetic structures and modelled present and past distributions of 15 Atlantic Forest (AF) endemic birds. Afterwards, we evaluated the relationship of population genetic parameters with metrics of species range shifts between the present and the LGM. We found that geographical ranges of AF birds changed in concert with Pleistocene glacial cycles but, unexpectedly, our findings suggest the novel idea that ranges during glacial maxima were slightly larger on average, as well as equally fragmented and displaced from the interglacial ranges. Our findings suggest that range shifts over the late Pleistocene impacted on the diversification of forest organisms, although they did not show that those range shifts had a strong effect. We found that a combination of fragmentation variation across time, small current range size, and range stability increased population genetic structure. However, neither fragmentation, nor range stability affected genetic diversity. Our study showed that evolutionary responses to range shifts across AF birds have a high variance, which could explain the mixed support given by single‐species studies to the action of Pleistocene range shifts on population evolution.  相似文献   

13.
This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.  相似文献   

14.
Climatic changes have profound effects on the distribution of biodiversity, but untangling the links between climatic change and ecosystem functioning is challenging, particularly in high diversity systems such as tropical forests. Tropical forests may also show different responses to a changing climate, with baseline climatic conditions potentially inducing differences in the strength and timing of responses to droughts. Trait‐based approaches provide an opportunity to link functional composition, ecosystem function and environmental changes. We demonstrate the power of such approaches by presenting a novel analysis of long‐term responses of different tropical forest to climatic changes along a rainfall gradient. We explore how key ecosystem's biogeochemical properties have shifted over time as a consequence of multi‐decadal drying. Notably, we find that drier tropical forests have increased their deciduous species abundance and generally changed more functionally than forests growing in wetter conditions, suggesting an enhanced ability to adapt ecologically to a drying environment.  相似文献   

15.
Density‐dependent mortality has been recognized as an important mechanism that underpins tree species diversity, especially in tropical forests. However, few studies have attempted to explore how density dependence varies with spatial scale and even fewer have attempted to identify why there is scale‐dependent differentiation. In this study, we explore the elevational variation in density dependence. Three 1‐ha permanent plots were established at low and high elevations in the Heishiding subtropical forest, southern China. Using data from 1200 1 m2 seedling quadrats, comprising of 200 1 m2 quadrats located in each 1‐ha plot, we examined the variation in density dependence between elevations using a generalized linear mixed model with crossed random effects. A greenhouse experiment also investigated the potential effects of the soil biota on density‐dependent differentiation. Our results demonstrated that density‐dependent seedling mortality can vary between elevations in subtropical forests. Species found at a lower elevation suffered stronger negative density dependence than those found at a higher elevation. The greenhouse experiment indicated that two species that commonly occur at both elevations suffered more from soilborne pathogens during seed germination and seedling growth when they grew at the lower elevation, which implied that soil pathogens may play a crucial role in density‐dependent spatial variation.  相似文献   

16.
For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade‐offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade‐offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade‐offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51–53%) followed by substrate type (35–37%) and light (11–12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade‐offs among species during regeneration are common in low‐diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.  相似文献   

17.
Climate change is predicted to result in warmer and drier Neotropical forests relative to current conditions. Negative density‐dependent feedbacks, mediated by natural enemies, are key to maintaining the high diversity of tree species found in the tropics, yet we have little understanding of how projected changes in climate are likely to affect these critical controls. Over 3 years, we evaluated the effects of a natural drought and in situ experimental warming on density‐dependent feedbacks on seedling demography in a wet tropical forest in Puerto Rico. In the +4°C warming treatment, we found that seedling survival increased with increasing density of the same species (conspecific). These positive density‐dependent feedbacks were not associated with a decrease in aboveground natural enemy pressure. If positive density‐dependent feedbacks are not transient, the diversity of tropical wet forests, which may rely on negative density dependence to drive diversity, could decline in a future warmer, drier world.  相似文献   

18.
Negative density‐dependent seedling mortality has been widely detected in tropical, subtropical and temperate forests, with soil pathogens as a major driver. Here we investigated how host density affects the composition of soil pathogen communities and consequently influences the strength of plant‐soil feedbacks. In field censuses of six 1‐ha permanent plots, we found that survival was much lower for newly germinated seedlings that were surrounded by more conspecific adults. The relative abundance of pathogenic fungi in soil increased with increasing conspecific tree density for five of nine tree species; more soil pathogens accumulated around roots where adult tree density was higher, and this greater pathogen frequency was associated with lower seedling survival. Our findings show how tree density influences populations of soil pathogens, which creates plant‐soil feedbacks that contribute to community‐level and population‐level compensatory trends in seedling survival.  相似文献   

19.
Forest degradation accounts for ~70% of total carbon losses from tropical forests. Substantial emissions are from selective logging, a land‐use activity that decreases forest carbon density. To maintain carbon values in selectively logged forests, climate change mitigation policies and government agencies promote the adoption of reduced‐impact logging (RIL) practices. However, whether RIL will maintain both carbon and timber values in managed tropical forests over time remains uncertain. In this study, we quantify the recovery of timber stocks and aboveground carbon at an experimental site where forests were subjected to different intensities of RIL (4, 8, and 16 trees/ha). Our census data span 20 years postlogging and 17 years after the liberation of future crop trees from competition in a tropical forest on the Guiana Shield, a globally important forest carbon reservoir. We model recovery of timber and carbon with a breakpoint regression that allowed us to capture elevated tree mortality immediately after logging. Recovery rates of timber and carbon were governed by the presence of residual trees (i.e., trees that persisted through the first harvest). The liberation treatment stimulated faster recovery of timber albeit at a carbon cost. Model results suggest a threshold logging intensity beyond which forests managed for timber and carbon derive few benefits from RIL, with recruitment and residual growth not sufficient to offset losses. Inclusion of the breakpoint at which carbon and timber gains outpaced postlogging mortality led to high predictive accuracy, including out‐of‐sample R2 values >90%, and enabled inference on demographic changes postlogging. Our modeling framework is broadly applicable to studies that aim to quantify impacts of logging on forest recovery. Overall, we demonstrate that initial mortality drives variation in recovery rates, that the second harvest depends on old growth wood, and that timber intensification lowers carbon stocks.  相似文献   

20.
Tropical forest responses to climate and atmospheric change are critical to the future of the global carbon budget. Recent studies have reported increases in estimated above‐ground biomass (EAGB) stocks, productivity, and mortality in old‐growth tropical forests. These increases could reflect a shift in forest functioning due to global change and/or long‐lasting recovery from past disturbance. We introduce a novel approach to disentangle the relative contributions of these mechanisms by decomposing changes in whole‐plot biomass fluxes into contributions from changes in the distribution of gap‐successional stages and changes in fluxes for a given stage. Using 30 years of forest dynamic data at Barro Colorado Island, Panama, we investigated temporal variation in EAGB fluxes as a function of initial EAGB (EAGBi) in 10 × 10 m quadrats. Productivity and mortality fluxes both increased strongly with initial quadrat EAGB. The distribution of EAGB (and thus EAGBi) across quadrats hardly varied over 30 years (and seven censuses). EAGB fluxes as a function of EAGBi varied largely and significantly among census intervals, with notably higher productivity in 1985–1990 associated with recovery from the 1982–1983 El Niño event. Variation in whole‐plot fluxes among census intervals was explained overwhelmingly by variation in fluxes as a function of EAGBi, with essentially no contribution from changes in EAGBi distributions. The high observed temporal variation in productivity and mortality suggests that this forest is very sensitive to climate variability. There was no consistent long‐term trend in productivity, mortality, or biomass in this forest over 30 years, although the temporal variability in productivity and mortality was so strong that it could well mask a substantial trend. Accurate prediction of future tropical forest carbon budgets will require accounting for disturbance‐recovery dynamics and understanding temporal variability in productivity and mortality.  相似文献   

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