Decelerating growth in tropical forest trees

The impacts of global change on tropical forests remain poorly understood. We examined changes in tree growth rates over the past two decades for all species occurring in large (50-ha) forest dynamics plots in Panama and Malaysia. Stem growth rates declined significantly at both forests regardless of initial size or organizational level (species, community or stand). Decreasing growth rates were widespread, occurring in 24–71% of species at Barro Colorado Island, Panama (BCI) and in 58–95% of species at Pasoh, Malaysia (depending on the sizes of stems included). Changes in growth were not consistently associated with initial growth rate, adult stature, or wood density. Changes in growth were significantly associated with regional climate changes: at both sites growth was negatively correlated with annual mean daily minimum temperatures, and at BCI growth was positively correlated with annual precipitation and number of rainfree days (a measure of relative insolation). While the underlying cause(s) of decelerating growth is still unresolved, these patterns strongly contradict the hypothesized pantropical increase in tree growth rates caused by carbon fertilization. Decelerating tree growth will have important economic and environmental implications.     

Cool habitats support darker and bigger butterflies in Australian tropical forests

Morphology mediates the relationship between an organism's body temperature and its environment. Dark organisms, for example, tend to absorb heat more quickly than lighter individuals, which could influence their responses to temperature. Therefore, temperature‐related traits such as morphology may affect patterns of species abundance, richness, and community assembly across a broad range of spatial scales. In this study, we examined variation in color lightness and body size within butterfly communities across hot and cool habitats in the tropical woodland–rainforest ecosystems of northeast Queensland, Australia. Using thermal imaging, we documented the absorption of solar radiation relative to color lightness and wingspan and then built a phylogenetic tree based on available sequences to analyze the effects of habitat on these traits within a phylogenetic framework. In general, darker and larger individuals were more prevalent in cool, closed‐canopy rainforests than in immediately adjacent and hotter open woodlands. In addition, darker and larger butterflies preferred to be active in the shade and during crepuscular hours, while lighter and smaller butterflies were more active in the sun and midday hours—a pattern that held after correcting for phylogeny. Our ex situ experiment supported field observations that dark and large butterflies heated up faster than light and small butterflies under standardized environmental conditions. Our results show a thermal consequence of butterfly morphology across habitats and how environmental factors at a microhabitat scale may affect the distribution of species based on these traits. Furthermore, this study highlights how butterfly species might differentially respond to warming based on ecophysiological traits and how thermal refuges might emerge at microclimatic and habitat scales.     

热带森林中的斑块动态与物种多样性维持

斑块作为景观要素之一,直接到景观结构的空间格局及其内部各要素之间的相互关系,其动态也将导致景观格局的变化。异质性的斑块在自然森林中是普遍存在的,这就是说,顶极森林中仍然存在着由不同种类或不同生长时期的植物种群组成的森林斑块。由自然或人为干扰所驱动的森林生长循环导致敢这些斑块在空间上的镶嵌,对于持定的地域片段,也导致了不同生长时期的森林斑块的周期性循环。在热带森林中,森林的生长循环由林窗期（ｇａｐ     

Testing metabolic ecology theory for allometric scaling of tree size, growth and mortality in tropical forests

The theory of metabolic ecology predicts specific relationships among tree stem diameter, biomass, height, growth and mortality. As demographic rates are important to estimates of carbon fluxes in forests, this theory might offer important insights into the global carbon budget, and deserves careful assessment. We assembled data from 10 old-growth tropical forests encompassing censuses of 367 ha and > 1.7 million trees to test the theory's predictions. We also developed a set of alternative predictions that retained some assumptions of metabolic ecology while also considering how availability of a key limiting resource, light, changes with tree size. Our results show that there are no universal scaling relationships of growth or mortality with size among trees in tropical forests. Observed patterns were consistent with our alternative model in the one site where we had the data necessary to evaluate it, and were inconsistent with the predictions of metabolic ecology in all forests.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

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.     

Biodiversity in species,traits, and structure determines carbon stocks and uptake in tropical forests

Impacts of climate change require that society urgently develops ways to reduce amounts of carbon in the atmosphere. Tropical forests present an important opportunity, as they take up and store large amounts of carbon. It is often suggested that forests with high biodiversity have large stocks and high rates of carbon uptake. Evidence is, however, scattered across geographic areas and scales, and it remains unclear whether biodiversity is just a co‐benefit or also a requirement for the maintenance of carbon stocks and uptake. Here, we perform a quantitative review of empirical studies that analyzed the relationships between plant biodiversity attributes and carbon stocks and carbon uptake in tropical forests. Our results show that biodiversity attributes related to species, traits or structure significantly affect carbon stocks or uptake in 64% of the evaluated relationships. Average vegetation attributes (community‐mean traits and structural attributes) are more important for carbon stocks, whereas variability in vegetation attributes (i.e., taxonomic diversity) is important for both carbon stocks and uptake. Thus, different attributes of biodiversity have complementary effects on carbon stocks and uptake. These biodiversity effects tend to be more often significant in mature forests at broad spatial scales than in disturbed forests at local spatial scales. Biodiversity effects are also more often significant when confounding variables are not included in the analyses, highlighting the importance of performing a comprehensive analysis that adequately accounts for environmental drivers. In summary, biodiversity is not only a co‐benefit, but also a requirement for short‐ and long‐term maintenance of carbon stocks and enhancement of uptake. Climate change policies should therefore include the maintenance of multiple attributes of biodiversity as an essential requirement to achieve long‐term climate change mitigation goals.     

Tree demography dominates long‐term growth trends inferred from tree rings

Understanding responses of forests to increasing CO₂ and temperature is an important challenge, but no easy task. Tree rings are increasingly used to study such responses. In a recent study, van der Sleen et al. (2014) Nature Geoscience, 8, 4 used tree rings from 12 tropical tree species and find that despite increases in intrinsic water use efficiency, no growth stimulation is observed. This challenges the idea that increasing CO₂ would stimulate growth. Unfortunately, tree ring analysis can be plagued by biases, resulting in spurious growth trends. While their study evaluated several biases, it does not account for all. In particular, one bias may have seriously affected their results. Several of the species have recruitment patterns, which are not uniform, but clustered around one specific year. This results in spurious negative growth trends if growth rates are calculated in fixed size classes, as ‘fast‐growing’ trees reach the sampling diameter earlier compared to slow growers and thus fast growth rates tend to have earlier calendar dates. We assessed the effect of this ‘nonuniform age bias’ on observed growth trends and find that van der Sleen's conclusions of a lack of growth stimulation do not hold. Growth trends are – at least partially – driven by underlying recruitment or age distributions. Species with more clustered age distributions show more negative growth trends, and simulations to estimate the effect of species’ age distributions show growth trends close to those observed. Re‐evaluation of the growth data and correction for the bias result in significant positive growth trends of 1–2% per decade for the full period, and 3–7% since 1950. These observations, however, should be taken cautiously as multiple biases affect these trend estimates. In all, our results highlight that tree ring studies of long‐term growth trends can be strongly influenced by biases if demographic processes are not carefully accounted for.     

Patterns of tree growth in relation to environmental variability in the tropical dry deciduous forest at Mudumalai, southern India

Tree diameter growth is sensitive to environmental fluctuations and tropical dry forests experience high seasonal and inter-annual environmental variation. Tree growth rates in a large permanent plot at Mudumalai, southern India, were examined for the influences of rainfall and three intrinsic factors (size, species and growth form) during three 4-year intervals over the period 1988–2000. Most trees had lowest growth during the second interval when rainfall was lowest, and skewness and kurtosis of growth distributions were reduced during this interval. Tree diameter generally explained <10% of growth variation and had less influence on growth than species identity or time interval. Intraspecific variation was high, yet species identity accounted for up to 16% of growth variation in the community. There were no consistent differences between canopy and understory tree growth rates; however, a few subgroups of species may potentially represent canopy and understory growth guilds. Environmentally-induced temporal variations in growth generally did not reduce the odds of subsequent survival. Growth rates appear to be strongly influenced by species identity and environmental variability in the Mudumalai dry forest. Understanding and predicting vegetation dynamics in the dry tropics thus also requires information on temporal variability in local climate.     

西双版纳热带季节雨林的树种组成和群落结构动态

研究了西双版纳热带季节雨林1 hm2(hectare)动态监测样地1993年与2007年之间树种组成和群落结构的变化。对样地中胸径≥5 cm的乔木进行了每木调查。目前其树种组成的热带分布科、属所占比例分别为91%和94%,具有较高比例的热带植物区系性质。在1993年与2007年两次调查之间,树种数量由145种增至179种,仅有1到2个个体的稀有树种所占比例从54%降为51.1%。从森林的垂直结构来看,A、B、C三层的个体死亡率分别为12.8%、12.9%和19.0%,各层树木的增长率分别为-8.5%、-1.4%和44.8%。与此相对应,C层小径级的树木所占比例有较大提高。虽然小径级的树木在种类和数量上比例增大,但个体数量和种类组成相对稳定的A、B层优势树种变化不大,维持了群落结构的稳定性。14 a间,群落中新增加的具有先锋性质的树种不超过5个。1993年时,A、B两层尚有先锋树种存在,2007年已经从A、B两层中退出。因此,从14 a间树种组成和群落结构的变化来看,虽然具有树木的死亡和增补,但其物种成分和群落结构的总体格局没有明显的变化,处于动态平衡过程中。     

Onward but not always upward: individualistic elevational shifts of tree species in subtropical montane forests

Ongoing global climate change is driving widespread shifts in species distributions. Trends show frequent upwards shifts of treelines, but information on changes in montane forest below the treeline and in the tropics and subtropics is limited, despite the importance of these areas for biodiversity and ecosystem function. Patterns of species shifts in tropical and subtropical regions are likely to be more complex and individualistic than global averages suggest due to high species diversity and strong influence of competition, alongside direct climatic limitations on distributions. To address the question of how subtropical montane tree species are likely to move as climate changes, we used an extensive national forest inventory to estimate distribution shifts of 75 tree species in Taiwan by comparing the optimum elevation and range edges of adults and juveniles within species. Overall there was a significant difference in optimum elevation of adults and juveniles. Life stage mismatches suggested upward shifts in 35% of species but downward shifts of over half (56%), while 8% appeared stable. Upward elevation shifts were disproportionately common in high elevation species, whilst mid to low elevation species suggested greater variation in shift direction. Whilst previous research on mountain forest range shifts has been dominated by work addressing changes in treeline position, we show that although high elevation species shift up, below the treeline species may shift individualistically, heralding widespread changes in forest communities over coming decades. The wide variation of responses indicated is likely driven by individual species responses to interacting environmental factors such as competition, topography and anthropogenic influences across the broad range of forest types investigated. As global environmental changes continue, more detailed understanding of tree range shifts across a wide spectrum of forests will allow us to prepare for the implications of such changes for biodiversity, ecosystem function and dependent human populations.     

Apparent climatically induced increase of tree mortality rates in a temperate forest

We provide a first detailed analysis of long-term, annual-resolution demographic trends in a temperate forest. After tracking the fates of 21 338 trees in a network of old-growth forest plots in the Sierra Nevada of California, we found that mortality rate, but not the recruitment rate, increased significantly over the 22 years of measurement (1983–2004). Mortality rates increased in both of two dominant taxonomic groups ( Abies and Pinus ) and in different forest types (different elevational zones). The increase in overall mortality rate resulted from an increase in tree deaths attributed to stress and biotic causes, and coincided with a temperature-driven increase in an index of drought. Our findings suggest that these forests (and by implication, other water-limited forests) may be sensitive to temperature-driven drought stress, and may be poised for die-back if future climates continue to feature rising temperatures without compensating increases in precipitation.     

Ecosystem carbon storage does not vary with mean annual temperature in Hawaiian tropical montane wet forests

Theory and experiment agree that climate warming will increase carbon fluxes between terrestrial ecosystems and the atmosphere. The effect of this increased exchange on terrestrial carbon storage is less predictable, with important implications for potential feedbacks to the climate system. We quantified how increased mean annual temperature (MAT) affects ecosystem carbon storage in above‐ and belowground live biomass and detritus across a well‐constrained 5.2 °C MAT gradient in tropical montane wet forests on the Island of Hawaii. This gradient does not systematically vary in biotic or abiotic factors other than MAT (i.e. dominant vegetation, substrate type and age, soil water balance, and disturbance history), allowing us to isolate the impact of MAT on ecosystem carbon storage. Live biomass carbon did not vary predictably as a function of MAT, while detrital carbon declined by ~14 Mg of carbon ha^?1 for each 1 °C rise in temperature – a trend driven entirely by coarse woody debris and litter. The largest detrital pool, soil organic carbon, was the most stable with MAT and averaged 48% of total ecosystem carbon across the MAT gradient. Total ecosystem carbon did not vary significantly with MAT, and the distribution of ecosystem carbon between live biomass and detritus remained relatively constant across the MAT gradient at ~44% and ~56%, respectively. These findings suggest that in the absence of alterations to precipitation or disturbance regimes, the size and distribution of carbon pools in tropical montane wet forests will be less sensitive to rising MAT than predicted by ecosystem models. This article also provides needed detail on how individual carbon pools and ecosystem‐level carbon storage will respond to future warming.     

Understanding and predicting frost‐induced tropical tree mortality patterns

Extreme climatic and weather events are increasing in frequency and intensity across the world causing episodes of widespread tree mortality in many forested ecosystems. However, we have a limited understanding about which local factors influence tree mortality patterns, restricting our ability to predict tree mortality, especially within topographically complex tropical landscapes with a matrix of mature and secondary forests. We investigated the effects of two major local factors, topography and forest successional type, on climate‐induced tropical tree mortality patterns using an observational and modeling approach. The northernmost Neotropical dry forest endured an unprecedented episode of frost‐induced tree mortality after the historic February 2011 cold wave hit northwestern Mexico. In a moderately hilly landscape covering mature and secondary tropical dry forests, we surveyed 454 sites for the presence or absence of frost‐induced tree mortality. In addition, across forty‐eight 1 ha plots equally split into the two forest types, we examined 6,981 woody plants to estimate a frost‐disturbance severity metric using the density of frost‐killed trees. Elevation is the main factor modulating frost effects regardless of forest type. Higher occurrence probabilities of frost‐induced tree mortality at lowland forests can be explained by the strong influence of elevation on temperature distribution since heavier cold air masses move downhill during advective frosts. Holding elevation constant, the probability of frost‐induced tree mortality in mature forests was twice that of secondary forests but severity showed the opposite pattern, suggesting a cautious use of occurrence probabilities of tree mortality to infer severity of climate‐driven disturbances. Extreme frost events, in addition to altering forest successional pathways and ecosystem services, likely maintain and could ultimately shift latitudinal and altitudinal range margins of Neotropical dry forests.     

Population responses to observed climate variability across multiple organismal groups

A major challenge in ecology is to understand how populations are affected by increased climate variability. Here, we assessed the effects of observed climate variability on different organismal groups (amphibians, insects, mammals, herbaceous plants and reptiles) by estimating the extent to which interannual variation in the annual population growth rates (CV_λ) and the absolute value of the long-term population growth rate (|log λ|) were associated with short-term climate variability. We used empirical data (≥ 20 consecutive years of annual abundances) from 59 wild populations in the Northern Hemisphere, and quantified variabilities in population growth rates and climatic conditions (temperature and precipitation in active and inactive seasons) calculated over four- and eight-year sliding time windows. We observed a positive relationship between the variability of growth rate (CV_λ) and the variability of temperature in the active season at the shorter timescale only. Moreover, |log λ| was positively associated with the variability of precipitation in the inactive season at both timescales. Otherwise, the direction of the relationships between population dynamics and climate variability (if any) depended largely on the season and organismal group in question. Both CV_λ and |log λ| correlated negatively with species' lifespan, indicating general differences in population dynamics between short-lived and long-lived species that were not related to climate variability. Our results suggest that although temporal variation in population growth rates and the magnitude of long-term population growth rates are partially associated with short-term interannual climate variability, demographic responses to climate fluctuations might still be population-specific rather than specific to given organismal groups, and driven by other factors than the observed climate variability.     

Detecting long‐term growth trends using tree rings: a critical evaluation of methods

Tree‐ring analysis is often used to assess long‐term trends in tree growth. A variety of growth‐trend detection methods (GDMs) exist to disentangle age/size trends in growth from long‐term growth changes. However, these detrending methods strongly differ in approach, with possible implications for their output. Here, we critically evaluate the consistency, sensitivity, reliability and accuracy of four most widely used GDMs: conservative detrending (CD) applies mathematical functions to correct for decreasing ring widths with age; basal area correction (BAC) transforms diameter into basal area growth; regional curve standardization (RCS) detrends individual tree‐ring series using average age/size trends; and size class isolation (SCI) calculates growth trends within separate size classes. First, we evaluated whether these GDMs produce consistent results applied to an empirical tree‐ring data set of Melia azedarach, a tropical tree species from Thailand. Three GDMs yielded similar results – a growth decline over time – but the widely used CD method did not detect any change. Second, we assessed the sensitivity (probability of correct growth‐trend detection), reliability (100% minus probability of detecting false trends) and accuracy (whether the strength of imposed trends is correctly detected) of these GDMs, by applying them to simulated growth trajectories with different imposed trends: no trend, strong trends (?6% and +6% change per decade) and weak trends (?2%, +2%). All methods except CD, showed high sensitivity, reliability and accuracy to detect strong imposed trends. However, these were considerably lower in the weak or no‐trend scenarios. BAC showed good sensitivity and accuracy, but low reliability, indicating uncertainty of trend detection using this method. Our study reveals that the choice of GDM influences results of growth‐trend studies. We recommend applying multiple methods when analysing trends and encourage performing sensitivity and reliability analysis. Finally, we recommend SCI and RCS, as these methods showed highest reliability to detect long‐term growth trends.     

Time lag and negative responses of forest greenness and tree growth to warming over circumboreal forests

The terrestrial forest ecosystems in the northern high latitude region have been experiencing significant warming rates over several decades. These forests are considered crucial to the climate system and global carbon cycle and are particularly vulnerable to climate change. To obtain an improved estimate of the response of vegetation activity, e.g., forest greenness and tree growth, to climate change, we investigated spatiotemporal variations in two independent data sets containing the dendroecological information for this region over the past 30 years. These indices are the normalized difference vegetation index (NDVI3g) and the tree‐ring width index (RWI), both of which showed significant spatial variability in past trends and responses to climate changes. These trends and responses to climate change differed significantly in the ecosystems of the circumarctic (latitude higher than 67°N) and the circumboreal forests (latitude higher and lower than 50°N and 67°N, respectively), but the way in which they differed was relatively similar in the NDVI3g and the RWI. In the circumarctic ecosystem, the climate variables of the current summer were the main climatic drivers for the positive response to the increase in temperatures showed by both the NDVI3g and the RWI indices. On the other hand, in the circumboreal forest ecosystem, the climate variables of the previous year (from summer to winter) were also important climatic drivers for both the NDVI3g and the RWI. Importantly, both indices showed that the temperatures in the previous year negatively affected the ecosystem. Although such negative responses to warming did not necessarily lead to a past negative linear trend in the NDVI3g and the RWI over the past 30 years, future climate warming could potentially cause severe reduction in forest greenness and tree growth in the circumboreal forest ecosystem.     

Drought Impacts Survivorship and Reproductive Strategies of an Epiphyllous Leafy Liverwort in Central Amazonia

Understanding how changing precipitation patterns impact the population dynamics of Amazonian plants is necessary for predicting their long‐term survival. Most terrestrial plants are characterized by life cycles intractably slow for evaluating such demographic consequences. Due to the demands of surviving on an ephemeral substrate such as a leaf surface, epiphyllous (leaf‐inhabiting) bryophytes have some of the fastest generation times for terrestrial plants. Considering these advantages, we investigated the demographic consequences of seasonality in precipitation during one‐year of the epiphyllous bryophyte Radula flaccida Gott. (Radulaceae) in a central Amazonian rain forest. In a 1‐ha sized study area, 154 epiphyll colonies from 18 phorophytes of the understory shrub Naucleopsis ulei (Warb.) Ducke (Moraceae) were marked and colony growth, extinction, recolonization, as well as rates of sexual and asexual expression were measured in nine demographic censuses. The probability of survivorship decreased in the dry season due to the synergistic effects of both increased leaf fall (e.g., loss of occupied patches) and colony mortality on persisting leaves. Asexual expression also declined significantly during the driest months; however, sexual expression, as measured by sporophyte density, was not related to seasonality. Logistic regression probabilities also indicate a minimum threshold colony size for optimal sporophyte expression. Lower survivorship and colony growth in the dry season along with reduced sporophyte production in smaller colonies implies that both demographic stability as well as population structure of Amazonian epiphyll species may be compromised whether climate models projecting prolonged droughts for the region are accurate.