Impact of a drier Early-Mid-Holocene climate upon Amazonian forests

This paper uses a palaeoecological approach to examine the impact of drier climatic conditions of the Early-Mid-Holocene (ca 8000-4000 years ago) upon Amazonia's forests and their fire regimes. Palaeovegetation (pollen data) and palaeofire (charcoal) records are synthesized from 20 sites within the present tropical forest biome, and the underlying causes of any emergent patterns or changes are explored by reference to independent palaeoclimate data and present-day patterns of precipitation, forest cover and fire activity across Amazonia. During the Early-Mid-Holocene, Andean cloud forest taxa were replaced by lowland tree taxa as the cloud base rose while lowland ecotonal areas, which are presently covered by evergreen rainforest, were instead dominated by savannahs and/or semi-deciduous dry forests. Elsewhere in the Amazon Basin there is considerable spatial and temporal variation in patterns of vegetation disturbance and fire, which probably reflects the complex heterogeneous patterns in precipitation and seasonality across the basin, and the interactions between climate change, drought- and fire susceptibility of the forests, and Palaeo-Indian land use. Our analysis shows that the forest biome in most parts of Amazonia appears to have been remarkably resilient to climatic conditions significantly drier than those of today, despite widespread evidence of forest burning. Only in ecotonal areas is there evidence of biome replacement in the Holocene. From this palaeoecological perspective, we argue against the Amazon forest 'dieback' scenario simulated for the future.     

Warming,drought, and disturbances lead to shifts in functional composition: A millennial-scale analysis for Amazonian and Andean sites

Tropical forests are changing in composition and productivity, probably in response to changes in climate and disturbances. The responses to these multiple environmental drivers, and the mechanisms underlying the changes, remain largely unknown. Here, we use a functional trait approach on timescales of 10,000 years to assess how climate and disturbances influence the community-mean adult height, leaf area, seed mass, and wood density for eight lowland and highland forest landscapes. To do so, we combine data of eight fossil pollen records with functional traits and proxies for climate (temperature, precipitation, and El Niño frequency) and disturbances (fire and general disturbances). We found that temperature and disturbances were the most important drivers of changes in functional composition. Increased water availability (high precipitation and low El Niño frequency) generally led to more acquisitive trait composition (large leaves and soft wood). In lowland forests, warmer climates decreased community-mean height probably because of increased water stress, whereas in highland forests warmer climates increased height probably because of upslope migration of taller species. Disturbance increased the abundance of acquisitive, disturbance-adapted taxa with small seeds for quick colonization of disturbed sites, large leaves for light capture, and soft wood to attain fast height growth. Fire had weak effects on lowland forests but led to more stress-adapted taxa that are tall with fast life cycles and small seeds that can quickly colonize burned sites. Site-specific analyses were largely in line with cross-site analyses, except for varying site-level effects of El Niño frequency and fire activity, possibly because regional patterns in El Niño are not a good predictor of local changes, and charcoal abundances do not reflect fire intensity or severity. With future global changes, tropical Amazonian and Andean forests may transition toward shorter, drought- and disturbance-adapted forests in the lowlands but taller forests in the highlands.     

A regional study of Holocene climate change and human occupation in Peruvian Amazonia

Aim  To investigate the influence of Holocene climatic and human-induced changes on a region of high biodiversity in southern Peruvian Amazonia.
Location  Four palaeoecological records from separate lakes within a lake district close to the modern city of Puerto Maldonado, Peru.
Results  The lakes provide a palaeoecological record spanning the last 8200 years. A mid-Holocene dry event is documented in all of the records that extend back > 6000 years. The dry event appears to have lasted from c . 7200 yr bp until c . 3300 yr bp . The onset of wetter conditions coincides with the formation of the youngest of the four lakes. The earliest occupation of these sites is inferred from the presence of charcoal at 7200 yr bp , and the first crop pollen is found at 3630 yr bp . Lakes that were regularly occupied were colonized soon after they formed. A reduction in charcoal concentration and the absence of crop pollen after c . 500  bp in all lakes is consistent with site abandonment following conquest.
Main conclusions  The mid-Holocene dry event is suggested to be part of a time-transgressive drying that tracked from north to south in both the Andes and the Amazon lowlands. The last millennium may represent the period of highest sustained lake levels within the Holocene. The proximity of the four lakes allows a landscape-scale analysis of the spatial extent of human disturbance centred on a known site of human occupation and reveals the highly localized nature of pre-Columbian anthropogenic disturbance in Amazonian landscapes. Inferences regarding widespread pre-Columbian landscape modification by indigenous peoples must take into account key site attributes, such as seasonality and proximity to rivers.     

Winter severity determines functional trait composition of phytoplankton in seasonally ice‐covered lakes

How climate change will affect the community dynamics and functionality of lake ecosystems during winter is still little understood. This is also true for phytoplankton in seasonally ice‐covered temperate lakes which are particularly vulnerable to the presence or absence of ice. We examined changes in pelagic phytoplankton winter community structure in a north temperate lake (Müggelsee, Germany), covering 18 winters between 1995 and 2013. We tested how phytoplankton taxa composition varied along a winter‐severity gradient and to what extent winter severity shaped the functional trait composition of overwintering phytoplankton communities using multivariate statistical analyses and a functional trait‐based approach. We hypothesized that overwintering phytoplankton communities are dominated by taxa with trait combinations corresponding to the prevailing winter water column conditions, using ice thickness measurements as a winter‐severity indicator. Winter severity had little effect on univariate diversity indicators (taxon richness and evenness), but a strong relationship was found between the phytoplankton community structure and winter severity when taxon trait identity was taken into account. Species responses to winter severity were mediated by the key functional traits: motility, nutritional mode, and the ability to form resting stages. Accordingly, one or the other of two functional groups dominated the phytoplankton biomass during mild winters (i.e., thin or no ice cover; phototrophic taxa) or severe winters (i.e., thick ice cover; exclusively motile taxa). Based on predicted milder winters for temperate regions and a reduction in ice‐cover durations, phytoplankton communities during winter can be expected to comprise taxa that have a relative advantage when the water column is well mixed (i.e., need not be motile) and light is less limiting (i.e., need not be mixotrophic). A potential implication of this result is that winter severity promotes different communities at the vernal equinox, which may have different nutritional quality for the next trophic level and ecosystem‐scale effects.     

Leaf‐trait plasticity and species vulnerability to climate change in a Mongolian steppe

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate‐induced biomass responses in 12 co‐occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf‐trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.     

Response of green alder (Alnus viridis subsp. fruticosa) patch dynamics and plant community composition to fire and regional temperature in north‐western Canada

Aim Feedbacks between climate warming and fire have the potential to alter Arctic and sub‐Arctic vegetation. In this paper we assess the effects and interactions of temperature and wildfire on plant communities across the transition between the Arctic and sub‐Arctic. Location Mackenzie Delta region, Northwest Territories, Canada. Methods We sampled air temperatures, green alder (Alnus viridis ssp. fruticosa) cover, growth, reproduction and age distributions, and overall plant community composition on burned and unburned sites across a latitudinal gradient. Results Mean summer temperature across the study area decreased by 3 °C per degree of increasing latitude (6 °C across the study area). In the northern part of the study area, where seed viability was low, alder was less dominant than at southern sites where seed viability was high. The age structure of alder populations across the temperature gradient was highly variable, except in the northern part of the forest–tundra transition, where populations were dominated by young individuals. Alder growth and reproduction were significantly greater on burned sites (38–51 years following fire) than on unburned sites. North to south across the temperature gradient, vegetation changed from a community dominated by dwarf shrubs and fruticose lichens to one characterized by black spruce (Picea mariana), alder and willows (Salix spp.). Regardless of the position along the temperature gradient, burned sites were dominated by tall shrubs. Main conclusions Temperature limitation of alder abundance and repro‐duction, combined with evidence of recent recruitment on unburned sites, indicates that alder is likely to respond to increased temperature. Elevated alder growth and reproduction on burned sites shows that wildfire also has an important influence on alder population dynamics. The magnitude of alder’s response to fire, combined with observations that burns at the southern margin of the low Arctic are shrub dominated, suggest that increases in the frequency of wildfire have the potential to alter northern vegetation on decadal scales. By creating new seedbeds, fire provides opportunities for colonization that may facilitate the northward movement of tall shrubs. Feedbacks between the global climate system and low Arctic vegetation make understanding the long‐term impact of increasing fire frequency critical to predicting the response of northern ecosystems to global change.     

Close anthropogenic control of Fagus sylvatica establishment and expansion in a Swedish protected landscape – implications for forest history and conservation

Aim The dominant forces behind the expansion of Fagus sylvatica (beech) in northern Europe during the late Holocene have been much debated. Palaeoecological analyses were performed for a biodiversity hotspot reserve in order to study the processes behind the local establishment of Fagus, as well as the historical vegetational development in relation to present‐day biodiversity and conservation. Location Biskopstorp Forest Reserve in south‐west Sweden. Methods Pollen and charcoal were analysed from three small‐hollow sites in the reserve. Two of the investigated wetlands were located adjacent to old‐growth stands of Fagus, and the third was located near a stand of old Quercus. Results The 2500‐year record shows that Fagus was first established around the two Fagus old‐growth sites, c. ad 900 and 1200, respectively, and that this was followed by an expansion around ad 1600. During the establishment phase, and more so in the expansion phase, there were simultaneous increases in types of pollen indicative of human activity. These indicators are also frequent throughout the record from the Quercus site, but here Fagus never became common. Main conclusions The dynamics behind the establishment and expansion of Fagus in the reserve seems to have been influenced to a large degree by human activity, for example selective cutting, human‐induced fires, and agriculture. Fagus became established in the reserve more than 1000 years after it became established regionally, making climate less probable as the dominant force behind the species’ stand‐scale establishment. The spreading of Fagus across southern Scandinavia has previously been shown to be a patchy process in time and space. Our study suggests that this patchiness is evident also at a small spatial scale (a few kilometres). At the Quercus site, relatively high amounts of pollen from the field layer throughout the record indicate open conditions that probably favoured Quercus. The degree of human impact at this site was probably too high to allow the expansion of Fagus. With the long‐term perspective provided by our study it was possible to identify the last 200–300 years as an unrepresentative period with respect to tree species composition and forest dynamics. The large increase of Picea locally and regionally over the last several hundred years, combined with dramatic levels of human impact, have altered the tree composition and forest dynamics to such an extent that active management is necessary in order to maintain biodiversity in the reserve.     

Post‐fire vegetation and climate dynamics in low‐elevation forests over the last three millennia in Yellowstone National Park

Conifer forests of the western US are historically well adapted to wildfires, but current warming is creating novel disturbance regimes that may fundamentally change future forest dynamics. Stand‐replacing fires can catalyze forest reorganization by providing periodic opportunities for establishment of new tree cohorts that set the stage for stand development for centuries to come. Extensive research on modern and past fires in the Northern Rockies reveals how variations in climate and fire have led to large changes in forest distribution and composition. Unclear, however, is the importance of individual fire episodes in catalyzing change. We used high‐resolution paleoecologic and paleoclimatic data from Crevice Lake (Yellowstone National Park, Wyoming, USA), to explore the role of fire in driving low‐elevation forest dynamics over the last 2820 yr. We addressed two questions: 1) did low‐elevation forests at Crevice Lake experience abrupt community‐level vegetation changes in response to past fire events? 2) Did the interaction of short‐term disturbance events (fire) and long‐term climate change catalyze past shifts in forest composition? Over the last 2820 yr, we found no evidence for abrupt community‐level vegetation transitions at Crevice Lake, and no evidence that an interaction of climate and fire produced changes in the relative abundance of dominant plant taxa. In part, this result reflects limitations of the datasets to detect past event‐specific responses and their causes. Nonetheless, the relative stability of the vegetation to fires over the last 2820 yr provides a local baseline for assessing current and future ecological change. Observations of climate–fire–vegetation dynamics in recent decades suggest that this multi‐millennial‐scale baseline may soon be exceeded.     

Species traits and climate velocity explain geographic range shifts in an ocean‐warming hotspot

Species' ranges are shifting globally in response to climate warming, with substantial variability among taxa, even within regions. Relationships between range dynamics and intrinsic species traits may be particularly apparent in the ocean, where temperature more directly shapes species' distributions. Here, we test for a role of species traits and climate velocity in driving range extensions in the ocean‐warming hotspot of southeast Australia. Climate velocity explained some variation in range shifts, however, including species traits more than doubled the variation explained. Swimming ability, omnivory and latitudinal range size all had positive relationships with range extension rate, supporting hypotheses that increased dispersal capacity and ecological generalism promote extensions. We find independent support for the hypothesis that species with narrow latitudinal ranges are limited by factors other than climate. Our findings suggest that small‐ranging species are in double jeopardy, with limited ability to escape warming and greater intrinsic vulnerability to stochastic disturbances.     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

The role of climate and plant functional trade‐offs in shaping global biome and biodiversity patterns

Aim Two of the oldest observations in plant geography are the increase in plant diversity from the poles towards the tropics and the global geographic distribution of vegetation physiognomy (biomes). The objective of this paper is to use a process‐based vegetation model to evaluate the relationship between modelled and observed global patterns of plant diversity and the geographic distribution of biomes. Location The global terrestrial biosphere. Methods We implemented and tested a novel vegetation model aimed at identifying strategies that enable plants to grow and reproduce within particular climatic conditions across the globe. Our model simulates plant survival according to the fundamental ecophysiological processes of water uptake, photosynthesis, reproduction and phenology. We evaluated the survival of an ensemble of 10,000 plant growth strategies across the range of global climatic conditions. For the simulated regional plant assemblages we quantified functional richness, functional diversity and functional identity. Results A strong relationship was found (correlation coefficient of 0.75) between the modelled and the observed plant diversity. Our approach demonstrates that plant functional dissimilarity increases and then saturates with increasing plant diversity. Six of the major Earth biomes were reproduced by clustering grid cells according to their functional identity (mean functional traits of a regional plant assemblage). These biome clusters were in fair agreement with two other global vegetation schemes: a satellite image classification and a biogeography model (kappa statistics around 0.4). Main conclusions Our model reproduces the observed global patterns of plant diversity and vegetation physiognomy from the number and identity of simulated plant growth strategies. These plant growth strategies emerge from the first principles of climatic constraints and plant functional trade‐offs. Our study makes important contributions to furthering the understanding of how climate affects patterns of plant diversity and vegetation physiognomy from a process‐based rather than a phenomenological perspective.     

The fate of Amazonian ecosystems over the coming century arising from changes in climate,atmospheric CO2, and land use

There is considerable interest in understanding the fate of the Amazon over the coming century in the face of climate change, rising atmospheric CO₂ levels, ongoing land transformation, and changing fire regimes within the region. In this analysis, we explore the fate of Amazonian ecosystems under the combined impact of these four environmental forcings using three terrestrial biosphere models (ED2, IBIS, and JULES) forced by three bias‐corrected IPCC AR4 climate projections (PCM1, CCSM3, and HadCM3) under two land‐use change scenarios. We assess the relative roles of climate change, CO₂ fertilization, land‐use change, and fire in driving the projected changes in Amazonian biomass and forest extent. Our results indicate that the impacts of climate change are primarily determined by the direction and severity of projected changes in regional precipitation: under the driest climate projection, climate change alone is predicted to reduce Amazonian forest cover by an average of 14%. However, the models predict that CO₂ fertilization will enhance vegetation productivity and alleviate climate‐induced increases in plant water stress, and, as a result, sustain high biomass forests, even under the driest climate scenario. Land‐use change and climate‐driven changes in fire frequency are predicted to cause additional aboveground biomass loss and reductions in forest extent. The relative impact of land use and fire dynamics compared to climate and CO₂ impacts varies considerably, depending on both the climate and land‐use scenario, and on the terrestrial biosphere model used, highlighting the importance of improved quantitative understanding of all four factors – climate change, CO₂ fertilization effects, fire, and land use – to the fate of the Amazon over the coming century.     

Both life‐history plasticity and local adaptation will shape range‐wide responses to climate warming in the tundra plant Silene acaulis

Many predictions of how climate change will impact biodiversity have focused on range shifts using species‐wide climate tolerances, an approach that ignores the demographic mechanisms that enable species to attain broad geographic distributions. But these mechanisms matter, as responses to climate change could fundamentally differ depending on the contributions of life‐history plasticity vs. local adaptation to species‐wide climate tolerances. In particular, if local adaptation to climate is strong, populations across a species’ range—not only those at the trailing range edge—could decline sharply with global climate change. Indeed, faster rates of climate change in many high latitude regions could combine with local adaptation to generate sharper declines well away from trailing edges. Combining 15 years of demographic data from field populations across North America with growth chamber warming experiments, we show that growth and survival in a widespread tundra plant show compensatory responses to warming throughout the species’ latitudinal range, buffering overall performance across a range of temperatures. However, populations also differ in their temperature responses, consistent with adaptation to local climate, especially growing season temperature. In particular, warming begins to negatively impact plant growth at cooler temperatures for plants from colder, northern populations than for those from warmer, southern populations, both in the field and in growth chambers. Furthermore, the individuals and maternal families with the fastest growth also have the lowest water use efficiency at all temperatures, suggesting that a trade‐off between growth and water use efficiency could further constrain responses to forecasted warming and drying. Taken together, these results suggest that populations throughout species’ ranges could be at risk of decline with continued climate change, and that the focus on trailing edge populations risks overlooking the largest potential impacts of climate change on species’ abundance and distribution.     

气候变化和林火干扰对大兴安岭林区地上生物量影响的动态模拟

预测森林地上生物量对气候变化和林火干扰的响应是陆地生态系统碳循环研究的重要内容,气温、降水等因素的改变和气候变暖导致林火干扰强度的变化将会影响森林生态系统的碳库动态.东北森林作为我国森林的重要组成部分,对气候变化和林火干扰的响应逐渐显现.本文运用LANDIS PRO模型,模拟气候变化对大兴安岭森林地上生物量的影响,并比较分析了气候变暖对森林地上生物量的直接影响与通过林火干扰强度改变所产生的影响.结果表明: 未来气候变暖和火干扰增强情景下,森林地上生物量增加;当前气候条件和火干扰下,研究区森林地上生物量为(97.14±5.78) t·hm^-2;在B1F2预案下,森林地上生物量均值为(97.93±5.83) t·hm^-2;在A2F3预案下,景观水平第100～150和150～200年模拟时期内的森林地上生物量均值较高,分别为(100.02±3.76)和(110.56±4.08) t·hm^-2.与当前火干扰相比,CF2预案(当前火干扰增加30%)在一定时期使景观水平地上生物量增加(0.56±1.45) t·hm^-2,CF3预案(当前火干扰增加230%)在整个模拟阶段使地上生物量减少(7.39±1.79) t·hm^-2.针叶、阔叶树种对气候变暖的响应存在差异,兴安落叶松和白桦生物量随气候变暖表现为降低趋势,而樟子松、云杉和山杨的地上生物量则随气候变暖表现出不同程度的增加;气候变暖对针阔树种的直接影响具有时滞性,针叶树种响应时间比阔叶树种迟25～50年.研究区森林对高CO₂排放情景下气候变暖和高强度火干扰的共同作用较为敏感,未来将明显改变研究区森林生态系统的树种组成和结构.