Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.     

Environmental control of flowering periodicity in Costa Rican and Mexican tropical dry forests

Aim We analyse the proximate causes of the large variation in flowering periodicity among four tropical dry forests (TDF) and ask whether climatic periodicity or biotic interactions are the ultimate causes of flowering periodicity. Location The four TDFs in Guanacaste (Costa Rica), Yucatan, Jalisco and Sonora (Mexico) are characterized by a 5–7 month long dry season and are located along a gradient of increasing latitude (10–30°N). Methods To dissect the differences in flowering periodicity observed at the community level, individual tree species were assigned to ‘flowering types’, i.e. groups of species with characteristic flowering periods determined by similar combinations of environmental flowering cues and vegetative phenology. Results Large variation in the fraction of species and flowering types blooming during the dry and wet season, respectively, indicates large differences in the severity of seasonal drought among the four forests. In the dry upland forests of Jalisco, flowering of leafless trees remains suppressed during severe seasonal drought and is triggered by the first rains of the wet season. In the other forests, leaf shedding, exceptional rainfall or increasing daylength cause flowering of many deciduous species at various times during the dry season, well before the summer rains. The fraction of deciduous species leafing out during the summer rains and flowering when leafless during the dry season is largest in the Sonoran TDF. Main conclusions In many wide‐ranging species the phenotypic plasticity of flowering periodicity is large. The distinct temporal separation of spring flowering on leafless shoots and subsequent summer flushing represents a unique adaptation of tree development to climates with a relatively short rainy season and a long dry season. Seasonal variation in rainfall and soil water availability apparently constitutes not only the proximate, but also the ultimate cause of flowering periodicity, which is unlikely to have evolved in response to biotic adaptive pressures.     

Modification of Vegetative Phenology in a Tropical Semi‐deciduous Forest by Abnormal Drought and Rain1

The control of vegetative phenology in tropical trees is not well understood. In dry forest trees, leaf abscission may be enhanced by advanced leaf age, increasing water stress, or declining photoperiod. Normally, it is impossible to dissect the effects of each of these variables because most leaves are shed during the early dry season when day length is near its minimum and leaves are relatively old. The 1997 El‐Niño Southern Oscillation caused a ten‐week long, severe abnormal drought from June to August in the semi‐deciduous forests of Guanacaste, Costa Rica. We monitored the effect of this drought on phenology and water status of trees with young leaves and compared modifications of phenology in trees of different functional types with the pattern observed during the regular dry season. Although deciduous trees at dry sites were severely water stressed (Ψstem < ‐7MPa) and their mesic leaves remained wilted for more than two months, these and all other trees retained all leaves during the abnormal drought. Many trees exchanged leaves three to four months earlier than normal during the wet period after the abnormal drought and shed leaves again during the regular dry season. Irrigation and an exceptional 70 mm rainfall during the mid‐dry season 1998/1999 caused bud break and flushing in all leafless trees except dormant stem succulents. The complex interactions between leaf age and water stress, the principal determinants of leaf abscission, were found to vary widely among trees of different functional types.     

Productive phenology of tropical montane forests: Fertilization experiments along a moisture gradient

Phenological responses of leaves and roots were studied in the tropical montane forests of Mount Kinabalu, Borneo. Soil nutrient supply, in addition to the supply of light and water, is a potential abiotic factor influencing plant phenology in the tropics. The main objective of this study was to evaluate the contribution of soil nutrient supply to plant productive phenology. Fertilization experiments, including controls, nitrogen fertilized and nitrogen and phosphorus fertilized treatments, were conducted on three vegetation types in different moisture environments. Responses of leaves and roots were compared among treatments and among vegetation types. Leaf flushing was induced by nitrogen fertilization in the upper montane forest, where extremely wet moisture conditions are associated with cloud cover. This induction of leaf flushing by fertilization was not observed in the other forests. Root growth was suppressed by fertilization when leaf flushing was not induced by fertilization. These results indicate that soil pulsed nutrient release could be a cue for leaf flushing in a tropical wet environment, and that leaf phenology could be regulated by external abiotic factors and root phenology could be regulated by internal plant demands.     

Current issues in tropical phenology: a synthesis

We retrace the development of tropical phenology research, compare temperate phenology study to that in the tropics and highlight the advances currently being made in this flourishing discipline. The synthesis draws attention to how fundamentally different tropical phenology data can be to temperate data. Tropical plants lack a phase of winter dormancy and may grow and reproduce continually. Seasonal patterns in environmental parameters, such as rainfall, irradiance or temperature, do not necessarily coincide temporally, as they do in temperate climes. We review recent research on the drivers of phenophase cycles in individual trees, species and communities and highlight how significant innovations in biometric tools and approaches are being driven by the need to deal with circular data, the complexity of defining tropical seasons and the myriad growth and reproductive strategies used by tropical plants. We discuss how important the use of leaf phenology (or remotely‐sensed proxies of leaf phenophases) has become in tracking biome responses to climate change at the continental level and how important the phenophase of forests can be in determining local weather conditions. We also highlight how powerful analyses of plant responses are hampered at many tropical sites by a lack of contextual data on local environmental conditions. We conclude by arguing that there is a clear global benefit in increasing long term tropical phenology data collection and improving empirical collection of local climate measures, contemporary to the phenology data. Directing more resources to research in this sector will be widely beneficial.     

Fruiting phenology is linked to rainfall variability in a tropical rain forest

As the influence of climate change on tropical forests becomes apparent, more studies are needed to understand how changes in climatic variables such as rainfall are likely to affect tree phenology. Using a twelve‐year dataset (2005–2016), we studied the impact of seasonal rainfall patterns on the fruiting phenology of 69 tree species in the rain forest of southeastern Madagascar. We found that average annual rainfall in this region has increased by >800 mm (23%) during this period relative to that recorded for the previous 40 years and was highly variable both within and between years. Higher monthly measures of fruiting richness and the intensity of fruiting in our sample community were associated with significantly higher levels of rainfall. We also found that less rainfall during the dry season, but not the wet season, was associated with a significant shift toward later timing of peak richness and peak intensity of fruiting in the subsequent 12 months; however, this pattern was driven primarily by an extreme drought event that occurred during the study period. Longer time scales of phenology data are needed to see whether this pattern is consistent. Madagascar is expected to experience more extremes in rainfall and drought with increasing climate change. Thus, the linkages between variable precipitation and the fruiting phenology of forest trees will have important consequences for understanding plant reproduction and the ability of Madagascar's wildlife to cope with a changing climate.     

Litterfall seasonality and adaptive strategies of tropical and subtropical evergreen forests in China

中国热带和亚热带常绿林凋落物季节特征及适应策略 本研究收集了来自中国热带/亚热带常绿林共85个站点的凋落物量季节性变化数据,并采用线性回归、结构方程模型构建以及相位差分析等方法,综合探究中国热带/亚热带地区常绿阔叶林和针叶林叶片脱落对土壤水分、饱和水气压差和辐射强度等气候因子的响应机制。研究结果显示,在雨热同期和雨热异期两种热带/亚热带气候类型中,呈现出两种典型凋落物的物候类型(单峰季节型/双峰季节型)。在雨热同期气候条件下,光照强度和降水呈现季节性正相关,单峰的凋落物峰值和双峰的第一个峰值约出现在3–4月,不断增加的光照能促进新叶的萌发,老叶被代谢更强的新叶所替代,该类型属于一种最大程度利用光照来实现树木生长的自适应策略。双峰的第二个峰值出现在雨季末期,约在8–10月,是由不断增强的水分亏缺所导致的(常绿阔叶林：大气水分亏缺;常绿针叶林：土壤水分亏缺),这种类型是一种凋落老叶减少水分丢失来应对水分胁迫的自适应策略。在雨热异期气候条件下,光照强度和降水呈现季节性负相关,饱和水气压差与光照强度表现出一致的季节性动态变化,诱导了常绿阔叶林单峰和双峰物候的第一个凋落峰(约在3–4月),是一种权衡大气干旱胁迫和最大程度利用光照进行生长的综合自适应策略。在雨季初期,显著的土壤水分亏缺加速叶片凋落,诱导了常绿阔叶林双峰物候的第二个凋落峰(约在11月),属于凋落老叶应对土壤水分胁迫的自适应策略。这些研究结果可以为地球系统模式中热带物候的精确模拟提供重要参考。     

Flower power: tree flowering phenology as a settlement cue for migrating birds

1. Neotropical migrant birds show a clear preference for stopover habitats with ample food supplies; yet, the proximate cues underlying these decisions remain unclear. 2. For insectivorous migrants, cues associated with vegetative phenology (e.g. flowering, leaf flush, and leaf loss) may reliably predict the availability of herbivorous arthropods. Here we examined whether migrants use the phenology of five tree species to choose stopover locations, and whether phenology accurately predicts food availability. 3. Using a combination of experimental and observational evidence, we show migrant populations closely track tree phenology, particularly the flowering phenology of honey mesquite (Prosopis glandulosa), and preferentially forage in trees with more flowers. Furthermore, the flowering phenology of honey mesquite reliably predicts overall arthropod abundance as well as the arthropods preferred by migrants for food. 4. Together, these results suggest that honey mesquite flowering phenology is an important cue used by migrants to assess food availability quickly and reliably, while in transit during spring migration.     

Rainfall seasonality and pest pressure as determinants of tropical tree species' distributions

Drought and pests are primary abiotic and biotic factors proposed as selective filters acting on species distributions along rainfall gradients in tropical forests and may contribute importantly to species distributional limits, performance, and diversity gradients. Recent research demonstrates linkages between species distributions along rainfall gradients and physiological drought tolerance; corresponding experimental examinations of the contribution of pest pressure to distributional limits and potential interactions between drought and herbivory are limited. This study aims to quantitate differential performance and herbivory as a function of species range limits across a climatic and floristic transition in Southeast Asia. Khao Chong Botanical Garden, Thailand and Pasoh Forest Reserve, Malaysia straddle the Kangar‐Pattani Line. A reciprocal transplantation across a seasonality gradient was established using two groups of species (“widespread” taxa whose distributions include seasonally dry forests and “aseasonal” taxa whose distributions are limited to aseasonal forests). Growth, biomass allocation, survival, and herbivory were monitored for 19 months. Systematic differences in performance were a function of species distribution in relation to rainfall seasonality. In aseasonal Pasoh, aseasonal species had both greater growth and survivorship than widespread species. These differences were not a function of differential herbivory as widespread and aseasonal species experienced similar damage in the aseasonal forest. In seasonally dry Khao Chong, widespread species showed higher survivorship than aseasonal species, but these differences were only apparent during drought. We link this differential performance to physiological mechanisms as well as differential tolerance of biotic pressure during drought stress. Systematic decreases in seedling survival in aseasonal taxa during drought corresponded with previously documented physiological differences and may be exacerbated by herbivore damage. These results have important implications for tropical diversity and community composition in light of predicted increases in the frequency and severity of drought in hyperdiverse tropical forests.     

Climatic sensitivity of species’ vegetative and reproductive phenology in a Hawaiian montane wet forest

Understanding how tropical tree phenology (i.e., the timing and amount of seed and leaf production) responds to climate is vital for predicting how climate change may alter ecological functioning of tropical forests. We examined the effects of temperature, rainfall, and photosynthetically active radiation (PAR) on seed phenology of four dominant species and community-level leaf phenology in a montane wet forest on the island of Hawaiʻi using monthly data collected over ~ 6 years. We expected that species phenologies would be better explained by variation in temperature and PAR than rainfall because rainfall at this site is not limiting. The best-fit model for all four species included temperature, rainfall, and PAR. For three species, including two foundational species of Hawaiian forests (Acacia koa and Metrosideros polymorpha), seed production declined with increasing maximum temperatures and increased with rainfall. Relationships with PAR were the most variable across all four species. Community-level leaf litterfall decreased with minimum temperatures, increased with rainfall, and showed a peak at PAR of ~ 400 μmol/m²s⁻¹. There was considerable variation in monthly seed and leaf production not explained by climatic factors, and there was some evidence for a mediating effect of daylength. Thus, the impact of future climate change on this forest will depend on how climate change interacts with other factors such as daylength, biotic, and/or evolutionary constraints. Our results nonetheless provide insight into how climate change may affect different species in unique ways with potential consequences for shifts in species distributions and community composition.     

Arthropod Assemblages in Epiphyte Mats of Costa Rican Cloud Forests

Tropical cloud forests are functionally important ecosystems, but are severely threatened due to deforestation and fragmentation. Epiphyte mats, accumulations of live vegetation and dead organic matter on tree trunks and branches, are a conspicuous component of cloud forests and harbor diverse assemblages of meso- and microarthropods. We compared the morphospecies richness, composition, and abundance of arthropods in epiphyte mats between primary and secondary forests of Monteverde, Costa Rica, and at two nearby replicate sites. Epiphyte mats were thinner and less structurally diverse in secondary forest. We collected ca 36,000 micro- and mesoarthropods from epiphyte mats in the 2-yr study. Whereas arthropod morphospecies richness did not differ among forest types, arthropod abundance was significantly higher in secondary forest due to larger numbers of ants, especially Solenopsis spp. Arthropod assemblages showed a high degree of taxonomic overlap both within and between primary and secondary forests (Jaccard abundance-based similarity = 0.93–0.96). Although characteristics of the arthropod fauna proved to be similar among sites and between forest types, there was a significant temporal effect: arthropod morphospecies richness in epiphyte mats generally was lower in the dry season (February–May), when many taxa probably became dormant or sought shelter against desiccation in deeper portions of mats.     

Putative biotic drivers of plant phenology: With special reference to pathogens and deciduousness

Plant phenology is not only manifested in the seasonal timing of vegetative and reproductive processes but also has ontogenetic aspects. The adaptive basis of seasonal phenology has been considered mainly in terms of climatic drivers. However, some biotic factors as likely evolutionary influences on plants’ phenology appear to have been under‐researched. Several specific cases of putative biotic factors driving plant phenology are outlined, involving both herbivores and pathogens. These illustrate the diversity of likely interactions rather than any systematic coverage or review. Emphasis is on woody perennials, in which phenology is often most multifaceted and complicated by the ontogenetic aspect. The complete seasonal leaf fall that characterizes deciduous plants may be a very important defense against some pathogens. Whether biotic influences drive acquisition or long‐term persistence of deciduousness is considered. In one case, of leaf rusts in poplars, countervailing influences of the rusts and climate suggest persistence. Often, however, biotic and environmental influences likely reinforce each other. The timing and duration of shoot flushing may in at least some cases contribute to defenses against herbivores, largely through brief periods of “predator satiation” when plant tissues have highest food value. Wide re‐examination of plant phenology, accommodating the roles of biotic factors and their interplays with environments as additional adaptive drivers, is advocated toward developing and applying hypotheses that are observationally or experimentally testable.     

Changing water use and adaptive strategies along rainfall gradients in Mediterranean lupins

• There is growing interest in harnessing the genetic and adaptive diversity of crop wild relatives to improve drought resilience of elite cultivars. Rainfall gradients exert strong selection pressure on both natural and agricultural ecosystems. Understanding plant responses to these facilitates crop improvement.
• Wild and domesticated narrow‐leafed lupin (NLL) collected along Mediterranean terminal drought stress gradients was evaluated under contrasting reproductive phase water supply in controlled field, glasshouse and cabinet studies. Plant phenology, growth and productivity, water use and stress responses were measured over time.
• There is an integrated suite of adaptive changes along rainfall gradients in NLL. Low rainfall ecotypes flower earlier, accumulate lower seed numbers, biomass and leaf area, and have larger root:shoot ratios than high rainfall ecotypes. Water‐use is lower and stress onset slower in low compared to high rainfall ecotypes. Water‐use rates and ecotypic differences in stress response (Ψleaf decline, leaf loss) are lower in NLL than yellow lupin (YL). To mitigate the effects of profligate water use, high rainfall YL ecotypes maintain higher leaf water content over declining leaf water potential than low rainfall ecotypes. There is no evidence for such specific adaptation in NLL.
• The data suggests that appropriate phenology is the key adaptive trait to rainfall gradients in NLL because of the flow‐on effects on biomass production, fitness, transpiration and stress onset, and the lack of physiological adaptations as in YL. Accordingly, it is essential to match phenology with target environment in order to minimize risk and maximize yield potential.

     

Effect of prey availability on the abundance of White‐breasted Wood‐Wrens,insectivorous birds of tropical lowland forests

Some understory insectivorous birds manage to persist in tropical forest fragments despite significant habitat loss and forest fragmentation. Their persistence has been related to arthropod biomass. In addition, forest structure has been used as a proxy to estimate prey availability for understory birds and for calculating prey abundance. We used arthropod biomass and forest structural variables (leaf area index [LAI] and aerial leaf litter biomass) to explain the abundance of White‐breasted Wood‐Wrens (Henicorhina leucosticta), tropical understory insectivorous birds, in six forests in the Caribbean lowlands of Costa Rica. To estimate bird abundance, we performed point counts (100‐m radius) in two old‐growth forests, two second‐growth forests, and two selectively logged forests. Arthropod abundance was the best predictor of wood‐wren abundance (wi = 0.75). Wood‐wren abundance increased as the number of arthropods increased, and the estimated range of bird abundance obtained from the model varied from 0.51 (0.28 – 0.93 [95%CI]) to 3.70 (1.68 – 5.20 [95%CI]) within sites. LAI was positively correlated to prey abundance (P = 0.01), and explained part of the variation in wood‐wren abundance. In forests with high LAI, arthropods have more aerial leaf litter as potential habitat so more potential prey are available for wood‐wrens. Forests with a greater abundance of aerial leaf litter arthropods were more likely to sustain higher densities of wood‐wrens in a fragmented tropical landscape.     

Aboveground biomass allocation patterns within Mediterranean sub-shrubs: A quantitative analysis of seasonal dimorphism

The monthly patterns of aboveground biomass allocation were studied in the branches of six Mediterranean sub-shrubs with different leaf phenology. Four of them were seasonally dimorphic species, and the remaining two were a winter deciduous and a cushion plant with photosynthetic stems. By the analysis of these species we aimed to identify different aboveground biomass allocation patterns within seasonally dimorphic species and to understand the role of seasonal dimorphism as a strategy to avoid the main stresses of mediterranean climate: summer drought and winter cold. The biomass allocation to the different living and photosynthetic fractions of 3-year-old branches was studied monthly for a minimum of 13 months per species. Leaf area (LA, mm²) and leaf mass per area (LMA, mg cm⁻²) measurements were used to characterize the diverse types of leaves of each species. Standing dead and senescent tissues accounted for a great percentage of the branch biomass of seasonally dimorphic species both during summer and winter. Different patterns of photosynthetic biomass allocation were found within the seasonally dimorphic species analysed. These patterns ranged from the moderate photosynthetic biomass oscillation of Salvia lavandulifolia to the almost deciduousness of Lepidium subulatum, and they were achieved by keeping alive, drying out or shedding different types of branches and leaves throughout the year. The formation of stress tolerant leaves and the reduction in the amount of photosynthetic biomass responded both to the occurrence of summer drought and winter cold. These results demonstrate that seasonal dimorphism is a flexible ecological strategy, as it comprises very different leaf phenologies and enables plants to escape both summer drought and winter cold.     

Shrub shading moderates the effects of weather on arthropod activity in arctic tundra

1. Rapid warming has facilitated an increase in deciduous shrub cover in arctic tundra. Because shrubs create a cooler microclimate during the growing season, shrub cover could modulate the effects of global warming on the phenology and activity of ectotherms, including arthropods. This possibility was explored here using two dominant arthropod groups (flies and wolf spiders) in Alaskan tundra. 2. We monitored arthropods with pitfall traps over five summers at four sites that differed in shrub abundance, and used generalised additive mixed models (GAMMs) to separate the two underlying components of pitfall trap catch: the seasonal trend in arthropod density and the effects of short‐term weather variation (air temperature, wind speed, rainfall, solar radiation) on arthropod activity. 3. We found that shrub cover significantly altered the seasonal trend in the abundance of flies by reducing early‐season pitfall catch, in line with observed later snowmelt in shrub‐dominated plots at these sites. 4. Additionally, shrub cover modulated the effects of many weather variables on arthropod activity: shrub cover shifted wolf spiders' temperature–activity relationship, dampened the positive effect of solar radiation on the activity of arthropods in total, and ameliorated the negative effect of wind on the activity of flies. 5. Thus, these results indicate that shrub encroachment will probably be accompanied by altered arthropod responses to warming and other key weather variables. Because the rate of key ecological processes – herbivory, decomposition, predation – are controlled by activity at the organismal level, these effects on arthropods will have long‐term ecosystem‐level consequences.     

Effects of precipitation and soil water potential on drought deciduous phenology in the Kalahari

We utilized an ecosystem process model to investigate the influence of precipitation and soil water potential on vegetation phenology in the semi‐arid, drought‐deciduous ecosystems in the Kalahari region of South Africa. The timing of leaf flush was assumed to be the first day during which a rainfall event exceeded that day's estimate of potential evapotranspiration after a defined dry season. Leaf senescence was assumed to be a dynamic feedback between soil water potential and net plant carbon gain and was determined by dynamically modeling the effects of concomitant trends in soil water potential and net primary production on leaf area index (LAI). Model predictions of LAI were compared with satellite‐derived normalized difference vegetation indices (NDVI) for 3 years at two sites along the Kalahari transect. The mean absolute error for the prediction of modeled leaf flush date compared with leaf flush dates estimated from NDVI were 10.0 days for the Maun site and 39.3 days for the Tshane site. Correlations between model predicted 10‐day average LAI and 10‐day composite NDVI for both Maun and Tshane were high (ρ=0.67 and 0.74, respectively, P<0.001), suggesting that this method adequately predicts intra‐annual leaf area dynamics in these dry tropical ecosystems.     

Arthropod Abundance,Canopy Structure,and Microclimate in a Bornean Lowland Tropical Rain Forest1

This study applies a novel, vertically stratified fogging protocol to document arthropod abundance, density, and biomass across a vertical gradient in a primary, lowland dipterocarp forest canopy in Borneo. We fogged arthropods at 5 m vertical intervals and 20 m horizontal intervals along six full‐canopy transects and measured leaf surface areas along the same transects. The results show that arthropod biomass in the aboveground regions was 23.6 kg/ha, the abundance was 23.9 million individuals/ha, and the density on leaf surfaces was 280 individuals/m² leaf area. All three numbers are five to ten times higher than estimated by previous surveys of tropical lowland rain forest canopies using mass‐collection techniques. Arthropod abundance and biomass were analyzed in relation to canopy structure, composition, vapor pressure deficit (VPD), photosynthetic photon flux density (PPFD), and height. Using stepwise regression we found that 13 of 14 arthropod groups had significant positive relationships with one‐sided leaf area, 11 had significant negative relationships with VPD, 3 had significant relationships with height, and none showed positive relationships with light. Classifying the 14 taxa based on their responses to leaf area and VPD created three groups that corresponded roughly to the biology of these taxa. This study suggests that the biomass and abundance, and perhaps therefore—by extrapolation—the biodiversity, of tropical canopy arthropods may be very much higher than previously estimated.     

Hydraulic variability of tropical forests is largely independent of water availability

Tropical rainforest woody plants have been thought to have uniformly low resistance to hydraulic failure and to function near the edge of their hydraulic safety margin (HSM), making these ecosystems vulnerable to drought; however, this may not be the case. Using data collected at 30 tropical forest sites for three key traits associated with drought tolerance, we show that site-level hydraulic diversity of leaf turgor loss point, resistance to embolism (P₅₀), and HSMs is high across tropical forests and largely independent of water availability. Species with high HSMs (>1 MPa) and low P₅₀ values (< −2 MPa) are common across the wet and dry tropics. This high site-level hydraulic diversity, largely decoupled from water stress, could influence which species are favoured and become dominant under a drying climate. High hydraulic diversity could also make these ecosystems more resilient to variable rainfall regimes.     

Forest biomass is strongly shaped by forest height across boreal to tropical forests in China

Aims Forest height is a major factor shaping geographic biomass patterns, and there is a growing dependence on forest height derived from satellite light detecting and ranging (LiDAR) to monitor large-scale biomass patterns. However, how the relationship between forest biomass and height is modulated by climate and biotic factors has seldom been quantified at broad scales and across various forest biomes, which may be crucial for improving broad-scale biomass estimations based on satellite LiDAR.Methods We used 1263 plots, from boreal to tropical forest biomes across China, to examine the effects of climatic (energy and water availability) and biotic factors (forest biome, leaf form and leaf phenology) on biomass–height relationship, and to develop the models to estimate biomass from forest height in China.Important findings (i) Forest height alone explained 62% of variation in forest biomass across China and was far more powerful than climate and other biotic factors. (ii) However, the relationship between biomass and forest height were significantly affected by climate, forest biome, leaf phenology (evergreen vs. deciduous) and leaf form (needleleaf vs. broadleaf). Among which, the effect of climate was stronger than other factors. The intercept of biomass–height relationship was more affected by precipitation while the slope more affected by energy availability. (iii) When the effects of climate and biotic factors were considered in the models, geographic biomass patterns could be well predicted from forest height with an r 2 between 0.63 and 0.78 (for each forest biome and for all biomes together). For most biomes, forest biomass could be well predicted with simple models including only forest height and climate. (iv) We provided the first broad-scale models to estimate biomass from forest height across China, which can be utilized by future LiDAR studies. (v) Our results suggest that the effect of climate and biotic factors should be carefully considered in models estimating broad-scale forest biomass patterns with satellite LiDAR.