Disperser- vs. Establishment-Limited Distribution of a Riparian Fig Tree (Ficus insipida) in a Costa Rican Tropical Rain Forest1

Studies were conducted at the La Selva Biological Station in Costa Rica and in a greenhouse in California to determine the factors accounting for the nonrandom distribution of the riparian fig tree Ficus insipida Willd. along streams in the La Selva Biological Reserve and adjacent deforested lands. We also evaluated the potential seed dispersers of this tree relative to the role of the fruit‐eating fish Brycon guatemalensis that previously was proposed to be an important disperser of F. insipida seeds in this system. At La Selva, we recorded the fig‐foraging activities of vertebrates at fruiting F. insipida trees, surveyed for the presence or absence of F. insipida along streams of different sizes, and determined the fate of fig seedlings transplanted in different riparian habitats. In the greenhouse, we measured seed germination and seedling survival and growth under different light and soil pH conditions mimicking natural conditions. The findings provided evidence that (1) the tree occurs along the larger streams running through forest habitat and only along smaller streams with relatively high light availability; (2) bats (Artibeus spp.) and fish are the major dispersers of F. insipida seeds; (3) the seedlings are subject to mortality not only from low light conditions but also from treefalls, frequent flooding, and bank erosion; and (4) high light levels and near neutral soil pH result in relatively better seed germination, faster growth, and higher survival rates of seedlings. Overall, our results suggest that this fig tree is dispersed mainly by bats and fish and is more establishment‐limited than disperser‐limited in its local distribution in the La Selva rain forest habitat.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Colombian dry moist forest transitions in the Llanos Orientales—A comparison of model and pollen-based biome reconstructions

Colombian vegetation, at the ecological level of the biome, is reconstructed at six sites using pollen data assigned a priori to plant functional types and biomes. The chosen sites incorporate four savanna sites (Laguna Sardinas, Laguna Angel, El Piñal and Laguna Carimagua), a site on the transition between savanna and Amazon rainforest (Loma Linda) and a site within the Amazon rainforest (Pantano de Monica). The areal extent of tropical moist forest, tropical dry forest and steppe have been subject to significant change: differential responses of the vegetation to climatic shifts are related to changes in plant available moisture, duration of dry season and edaphic controls on the vegetation. The record from El Piñal shows that the present-day savanna vegetation, dominated by steppe (Poaceae) with little occurrence of woody savanna taxa (e.g. Curatella, Byrsonima), was present since the last glacial period of the northern hemisphere. Unfortunately, El Piñal is located on an edaphic savanna and is not particularly responsive to registering change. Most records cover the early Holocene; one site records the El Abra stadial (Younger Dryas equivalent), when forest expansion reflects more humid climatic conditions and higher plant available moisture. During the early and middle Holocene, the maximum expansion of steppe and tropical dry forest occurred, indicating that dry climatic conditions continued to around 4000 ¹⁴C BP. The following period, from shortly before 4000 ¹⁴C BP, is characterised by an increase in forest and gallery forests, reflecting a wetter period probably with a shorter annual dry season. Anthropogenic influence on the vegetation is recorded by all the records over the last millennial, particularly characterised by a reduction in forest cover and high amplitude changes in vegetation.Biome transitions from one type to another, and the environmental controls on this shift, are investigated by applying a vegetation model (BIOME-3). The model uses climatic data from six meteorological stations that, encompass a range of environments within lowland Colombia, which are similar to the pollen data. The signals of vegetation change can be translated to the main environmental controls of temperature and moisture to indicate the degree of change needed in these parameters to record the vegetation change depicted by the pollen data. Moisture balance is the dominant control on driving vegetation change whether under seasonal or annual control. The combined reconstruction from pollen data and model output of biome-scale vegetation dynamics for lowland Colombia allows an understanding of the environmental controls to be developed.     

Volcanic explosive eruptions of the Vesuvio decrease tree-ring growth but not photosynthetic rates in the surrounding forests

Volcanic eruptions impact the global and the hemispheric climate, but it is still unknown how and to what degree they force the climate system and in particular the global carbon cycle. In this paper, the relationships between individual eruptions (reconstructed for the past using written records), tree primary productivity (estimated using ring widths), photosynthetic rate and stomatal conductance (assessed by carbon and oxygen isotope data) are investigated, to understand the impact of volcanic eruptions on net primary production. Data from a mixed stand of Fagus sylvatica L. and Acer pseudoplatanus L. located in the area of the Vesuvio volcanic complex (Southern Italy) showed a significant decrease in ring width following each eruption. Isotope analyses indicate a change in climatic conditions after such events. Specifically, the lower oxygen isotope ratio in the tree‐ring cellulose strongly suggests an increase in relative humidity and a decrease in temperature, with the latter resulting in a strong limitation to tree‐ring growth. The carbon isotope ratio was only moderately but not significantly reduced in the years of volcanic eruption, suggesting no major changes in C fixation rates. This work is a case study on the effects of volcanic eruptions resulting in strong climatic changes on the local scale. This is an opportunity to explore the process and causal relationships between climatic changes and the response of the vegetation. Thus, we propose here a realistic model scenario, from which we can extrapolate to global scales and improve our interpretations of results of global studies.     

Observed forest sensitivity to climate implies large changes in 21st century North American forest growth

Predicting long‐term trends in forest growth requires accurate characterisation of how the relationship between forest productivity and climatic stress varies across climatic regimes. Using a network of over two million tree‐ring observations spanning North America and a space‐for‐time substitution methodology, we forecast climate impacts on future forest growth. We explored differing scenarios of increased water‐use efficiency (WUE) due to CO₂‐fertilisation, which we simulated as increased effective precipitation. In our forecasts: (1) climate change negatively impacted forest growth rates in the interior west and positively impacted forest growth along the western, southeastern and northeastern coasts; (2) shifting climate sensitivities offset positive effects of warming on high‐latitude forests, leaving no evidence for continued ‘boreal greening’; and (3) it took a 72% WUE enhancement to compensate for continentally averaged growth declines under RCP 8.5. Our results highlight the importance of locally adapted forest management strategies to handle regional differences in growth responses to climate change.     

Tree Seed Fate in a Logged and Fragmented Forest Landscape, Northeastern Costa Rica1

We compared the seed fate of two animal‐dispersed, large‐seeded timber species (Dipteryx panamensis [Fabaceae] and Carapa guianensis [Meliaceae]) in logged and fragmented forests with that for continuous forest in northeastern Costa Rica. For both species, we quantified rates of seed removal (an index of vertebrate predation) and the fate of dispersed seeds (those carried away from their original location that either germinated or were not subsequently removed within three months). We predicted that (1) fewer seeds would be dispersed by vertebrates in fragmented forest than in continuous forest due to low population abundances after hunting and/or loss of suitable habitat, and (2) seed predation rates would be higher in forest fragments than in continuous forest due to high abundance of small‐bodied seed consumers. We compared three forest fragments currently managed for timber (140–350 ha) and a large reserve of continuous forest (La Selva, 1500 ha and connected to a national park). An exclusion experiment was performed (seeds placed in the open vs. seeds within semipermeable wire cages; 5 cm mesh size) to evaluate the relative roles of large and small animals on seed removal. Seed germination capacity did not differ among all four sites for both species. Removal of Dipteryx seeds was higher in forest fragments (50% removal within 10 days and related to the activity of small rodents) compared to La Selva (50% removal after 50 days). Also, more Dipteryx seeds were dispersed at La Selva than in fragmented forests. Contrary to our predictions, removal of Carapa seeds was equally high among all four sites, and there was a trend for more seeds of Carapa to be dispersed in fragments than in La Selva. Our results suggest that fragmentation effects on tree seed fate may be specific to species in question and contingent on the animal biota involved, and that management strategies for timber production based on regeneration from seed may differ between forest patches and extensive forests.     

Quantifying mortality of tropical rain forest trees using high-spatial-resolution satellite data

Assessment of forest responses to climate change is severely hampered by the limited information on tree death on short temporal and broad spatial scales, particularly in tropical forests. We used 1‐m resolution panchromatic IKONOS and 0.7‐m resolution QuickBird satellite data, acquired in 2000 and 2002, respectively, to evaluate tree death rates at the La Selva Biological Station in old‐growth Tropical Wet Forest in Costa Rica, Central America. Using a calibration factor derived from ground inspection of tree deaths predicted from the images, we calculated a landscape‐scale annual exponential death rate of 2.8%. This corresponds closely to data for all canopy‐level trees in 18 forest inventory plots, each of 0.5 ha, for a mostly‐overlapping 2‐year period (2.8% per year). This study shows that high‐spatial‐resolution satellite data can now be used to measure old‐growth tropical rain forest tree death rates, suggesting many new avenues for tropical forest ecology and global change research.     

Climate Influences on the Radial Growth of Centrolobium microchaete,a Valuable Timber Species from the Tropical Dry Forests in Bolivia

In this study, we use tree‐ring records to determine the climate factors controlling the growth of Centrolobium microchaete, a high‐value timber species from the tropical dry Chiquitano forest in Bolivia. We present the first tree‐ring chronologies from C. microchaete for Concepción and Santa Mónica, Bolivia. Statistical analyses show that the chronologies are of good quality and have a significant common signal between trees. The growth of C. microchaete is strongly influenced by climatic conditions during late spring–early summer. Abundant precipitations concurrent with below‐average temperatures during this period of the year favor tree growth. Climate variations in late spring–early summer explain >40 percent of the total variance in C. microchaete tree growth during the interval 1943–2005. Minor differences in tree responses to climate recorded between the two stands may reflect differences in the extent of the dry season and in soil water capacity between sites. Although the chronologies cover the past 180 yr, adding samples from older individuals would permit the extension of these records further back in time. The strong climate dependency of tree growth suggests that predicted future climate changes in the region could have a significant influence on C. microchaete tree growth during the 21st century.     

Unusual forest growth decline in boreal North America covaries with the retreat of Arctic sea ice

The 20th century was a pivotal period at high northern latitudes as it marked the onset of rapid climatic warming brought on by major anthropogenic changes in global atmospheric composition. In parallel, Arctic sea ice extent has been decreasing over the period of available satellite data records. Here, we document how these changes influenced vegetation productivity in adjacent eastern boreal North America. To do this, we used normalized difference vegetation index (NDVI) data, model simulations of net primary productivity (NPP) and tree‐ring width measurements covering the last 300 years. Climatic and proxy‐climatic data sets were used to explore the relationships between vegetation productivity and Arctic sea ice concentration and extent, and temperatures. Results indicate that an unusually large number of black spruce (Picea mariana) trees entered into a period of growth decline during the late‐20th century (62% of sampled trees; n = 724 cross sections of age >70 years). This finding is coherent with evidence encoded in NDVI and simulated NPP data. Analyses of climatic and vegetation productivity relationships indicate that the influence of recent climatic changes in the studied forests has been via the enhanced moisture stress (i.e. greater water demands) and autotrophic respiration amplified by the declining sea ice concentration in Hudson Bay and Hudson Strait. The recent decline strongly contrasts with other growth reduction events that occurred during the 19th century, which were associated with cooling and high sea ice severity. The recent decline of vegetation productivity is the first one to occur under circumstances related to excess heat in a 300‐year period, and further culminates with an intensifying wildfire regime in the region. Our results concur with observations from other forest ecosystems about intensifying temperature‐driven drought stress and tree mortality with ongoing climatic changes.     

Spatial and Temporal Variation in the Ant Occupants of a Facultative Ant-Plant1

Striking variation in ant occupation of a facultative ant-plant, Conostegia setosa (Melastomataceae), was found at three scales: local spatial, geographic, and temporal. C. setosa provides housing for ants and grows in groups of stems (clones). The ant occupants of 14 C. setosa clones were censused four times over a 14-mo period at the La Selva Biological Station, Costa Rica, and twice over a 9-mo period at the Nusagandi Station, Panama. Twelve facultative ant species occupied C. setosa stems at La Selva, compared to six facultative and one obligate species at Nusagandi. Occupancy (as % of stems ever occupied/clone) was higher at Nusagandi (median = 89%) compared to La Selva (65%). Occupancy varied among clones at La Selva but not at Nusagandi. C. setosa clones differed between sites, with larger clones and more small stems/clone at La Selva. Occupancy was influenced by clone structure; larger clones contained more ant species at both sites and had lower occupancy at La Selva. Occupancy was highest in larger stems and lowest in small stems at both sites. Temporally, percent occupation/clone did not differ among censuses at either site, but overall occupancy was lower in the dry season at La Selva. Turnover in ant occupants was higher at La Selva than at Nusagandi. The variation observed in this study is likely due to a number of factors, including differences between sites in plant population structure and history, differences between and within sites in ant faunas and their nesting requirements, and changes over space and time in microclimatic variables. Such high variation at multiple scales draws attention to the importance of long-term comparative studies of facultative animal-plant interactions.     

Legacies of La Niña: North American monsoon can rescue trees from winter drought

While we often assume tree growth–climate relationships are time‐invariant, impacts of climate phenomena such as the El Niño Southern Oscillation (ENSO) and the North American Monsoon (NAM) may challenge this assumption. To test this assumption, we grouped ring widths (1900‐present) in three southwestern US conifers into La Niña periods (LNP) and other years (OY). The 4 years following each La Niña year are included in LNP, and despite 1–2 year growth declines, compensatory adjustments in tree growth responses result in essentially equal mean growth in LNP and OY, as average growth exceeds OY means 2–4 years after La Niña events. We found this arises because growth responses in the two periods are not interchangeable: Due to differences in growth–climate sensitivities and climatic memory, parameters representing LNP growth fail to predict OY growth and vice versa (decreases in R² up to 0.63; lowest R² = 0.06). Temporal relationships between growth and antecedent climate (memory) show warmer springs and longer growing seasons negatively impact growth following dry La Niña winters, but that NAM moisture can rescue trees after these events. Increased importance of monsoonal precipitation during LNP is key, as the largest La Niña‐related precipitation deficits and monsoonal precipitation contributions both occur in the southern part of the region. Decreases in first order autocorrelation during LNP were largest in the heart of the monsoon region, reflecting both the greatest initial growth declines and the largest recovery. Understanding the unique climatic controls on growth in Southwest conifers requires consideration of both the influences and interactions of drought, ENSO, and NAM, each of which is likely to change with continued warming. While plasticity of growth sensitivity and memory has allowed relatively quick recovery in the tree‐ring record, recent widespread mortality events suggest conditions may soon exceed the capacity for adjustment in current populations.     

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.     

Structure,dynamics and dendroecology of an old‐growth Fagus forest in the Apennines

Abstract. Question: Which are the structural attributes and the history of old‐growth Fagus forest in Mediterranean montane environments? What are the processes underlying their structural organization? Are these forests stable in time and how does spatial scale affect our assessment of stability? How do these forests compare to other temperate deciduous old‐growth forests? Location: 1600–1850 m a.s.l., Fagus forest near the tree line, central Apennines, Italy. Methods: An old‐growth Fagus forest was studied following historical, structural and dendroecological approaches. History of forest cover changes was analysed using aerial photographs taken in 1945, 1954, 1985 and 1994. The structural analysis was carried out in the primary old‐growth portion of the forest using 18 circular and two rectangular plots. Dendroecological analyses were conducted on 32 dominant or co‐dominant trees. Results: These primary old‐growth Fagus remnants consist of four patches that escaped logging after World War II. Both living and dead tree components are within the range of structural attributes recognized for old‐growth in temperate biomes. Dendroecological analyses revealed the roles of disturbance, competition and climate in structuring the forest. We also identified a persistent Fagus community in which gap‐phase regeneration has led to a mono‐specific multi‐aged stand at spatial scales of a few hectares, characterized by a rotated‐sigmoid diameter distribution. Conclusion: Even at the relatively small spatial scale of this study, high‐elevation Apennine Fagus forests can maintain structural characteristics consistent with those of old‐growth temperate forests. These results are important for managing old‐growth forests in the Mediterranean montane biome.     

Legacies of more frequent drought in ponderosa pine across the western United States

Despite widespread interest in drought legacies—multiyear impacts of drought on tree growth—the key implication of reported drought legacies remains unaddressed: as impaired growth and slow recovery associated with drought legacies are pervasive across forest ecosystems, what is the impact of more frequent drought conditions? We investigated the assumption that either multiple drought years occurring during a short period (multiyear droughts), or droughts occurring during the recovery period from previous drought (compounded droughts), are detrimental to subsequent growth. There is evidence that drought responses may vary among populations of widespread species, leading us to examine regional differences in responses of the conifer Pinus ponderosa to historic drought frequency in the western United States. More frequent drought conditions incurred additional growth declines and shifts in growth–climate sensitivities in the years following drought relative to single‐drought events, with ‘triple‐droughts' being worse than ‘double‐droughts'. Notably, prediction skill was not strongly reduced when ignoring compounded droughts, a consequence of the temporally comprehensive formulation of our stochastic antecedent model that accounts for the climatic memory of tree growth. We argue that incorporating drought‐induced temporal variability in tree growth sensitivities can aid inference gained from statistical models, where more simplistic models could overestimate the severity of drought legacies. We also found regional differences in response to repeated drought, and suggest plastic post‐drought sensitivities and climatic memory may represent beneficial physiological adjustments in interior regions. Within‐species variability may thus mediate forest responses to increasing drought frequency under future climate change, but experimental approaches using more species are necessary to improve our understanding of the mechanisms that underlie drought legacy effects on tree growth.     

Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests

Predicted decreases in water availability across the temperate forest biome have the potential to offset gains in carbon (C) uptake from phenology trends, rising atmospheric CO₂, and nitrogen deposition. While it is well established that severe droughts reduce the C sink of forests by inducing tree mortality, the impacts of mild but chronic water stress on forest phenology and physiology are largely unknown. We quantified the C consequences of chronic water stress using a 13‐year record of tree growth (n = 200 trees), soil moisture, and ecosystem C balance at the Morgan–Monroe State Forest (MMSF) in Indiana, and a regional 11‐year record of tree growth (n > 300 000 trees) and water availability for the 20 most dominant deciduous broadleaf tree species across the eastern and midwestern USA. We show that despite ~26 more days of C assimilation by trees at the MMSF, increasing water stress decreased the number of days of wood production by ~42 days over the same period, reducing the annual accrual of C in woody biomass by 41%. Across the deciduous forest region, water stress induced similar declines in tree growth, particularly for water‐demanding ‘mesophytic’ tree species. Given the current replacement of water‐stress adapted ‘xerophytic’ tree species by mesophytic tree species, we estimate that chronic water stress has the potential to decrease the C sink of deciduous forests by up to 17% (0.04 Pg C yr^?1) in the coming decades. This reduction in the C sink due to mesophication and chronic water stress is equivalent to an additional 1–3 days of global C emissions from fossil fuel burning each year. Collectively, our results indicate that regional declines in water availability may offset the growth‐enhancing effects of other global changes and reduce the extent to which forests ameliorate climate warming.     

Increasing Liana Abundance and Basal Area in a Tropical Forest: The Contribution of Long‐distance Clonal Colonization

Recent evidence suggests that liana abundance and biomass are increasing in Neotropical forests, representing a major structural change to tropical ecosystems. Explanations for these increases, however, remain largely untested. Over an 8‐yr period (1999–2007), we censused lianas in nine, 24 × 36 m permanent plots in old‐growth and selectively logged forest at La Selva Biological Station, Costa Rica to test whether: (1) liana abundance and basal area are increasing in this forest; (2) the increase is being driven by increased recruitment, decreased mortality, or both; and (3) long‐distance clonal colonization explains the increase in liana abundance and basal area. We defined long‐distance clonal colonization as lianas that entered and rooted in the plots as vegetative propagules of stems that originated from outside or above the plot, and were present in 2007, but not in 1999 or 2002. Our hypotheses were supported in the old‐growth forest: mean liana abundance and BA (≥1 cm diameter) increased 15 and 20 percent, respectively, and clonal colonization from outside of the plots contributed 19 and 60 percent (respectively) to these increases. Lianas colonized clonally by falling vertically from the forest canopy above or growing horizontally along the forest floor and re‐rooting—common forms of colonization for many liana species. In the selectively logged forest, liana abundance and BA did not change, and thus the pattern of increasing lianas may be restricted to old‐growth forests. In summary, our data support the hypothesis that lianas are increasing in old‐growth forests, and that long‐distance clonal colonization is a major contributor.     

Area, shape and isolation of tropical forest fragments: effects on tree species diversity and implications for conservation

Aim To quantify the influences of forest area, shape and isolation on tree species diversity in Ghana and to compare their significance with the influences of climate (average annual rainfall) and disturbance (fire burn, logging, agriculture). Location The forest zone of southern Ghana, West Africa (between 5 and 8° N). Methods For twenty‐two forest fragments (1) bivariate regression analyses of tree species diversity (number and composition) were employed with forest spatial geometry, climate and disturbance variables. (2) Multivariate regression analyses of tree species number and all seven environmental variables were used to determine the variability in tree species number that could be accounted for by these environmental variables. Results Forest area, shape and isolation accounted for sharply decreasing proportions of variability in tree species diversity. Large forest fragments contained the greatest numbers of tree species and the highest proportions of rare tree species; irregular fragments had high proportions of regenerating, light‐demanding pioneers and mature, animal‐dispersed species and isolated fragments were floristically similar to less isolated fragments. Fire burn and average annual rainfall accounted for small, but nevertheless significant, proportions of variability in tree species diversity. Logging and agriculture were non‐significant variables. Main conclusions (1) Forest area is the most important consideration when planning tropical forest reserves. (2) Management of disturbance should take priority over management of forest shape if higher levels of tree diversity and species quality are to be maintained. (3) If new reserves are to be designated, they should be located within different climatic zones in order to capture a large fraction of the regional biota. (4) Biogeographers have an important role to play in formulating and testing hypotheses at a broad spatial scale and ultimately, informing conservation management within the tropical biome.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

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.     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.