Functional diversity and traits assembly patterns of lichens as indicators of successional stages in a tropical rainforest

Understanding community assembly patterns with regard to functional traits, which may be common to different species, allows us to compare ecological communities in a wider range of environmental and phytogeographic conditions. Moreover, a functional approach may facilitate the comprehension of the relation between biotic changes and ecosystem functions in complex systems such as tropical forests. Considering the lack of information in relation to the influence of environmental conditions on lichen functional traits, on lichen functional patterns in tropical forests and the potential usage of this approach in bioindication studies, this paper aims at (i) determining the functional structure of the lichen communities along the forest succession gradient, (ii) assessing the relation of lichen functional traits to this gradient and (iii) verifying the potential of using lichen functional traits as indicators of successional stages in tropical rainforests. Lichens were sampled in 24 sampling units of three successional stages (6–10; 12–20; and 40–60 years of recovering). The results corroborated our main hypothesis that the functional structure of the lichen communities responds to structural changes along a forest regeneration gradient. Growth forms (foliose, fruticose and crustose) as well as some reproductive trait states (lirella, isidia and perithecia) were the most suitable lichen traits to be used as indicators of forest succession in the southern edge of the Atlantic Rainforest. Lirella, fruticose and foliose were more related to earlier successional stages; isidia was more related to intermediate stages, while perithecia and crustose were more related to older stages. These results reinforce the ability of lichens to be used as bioindicators of forest conditions.     

The weight of the crust: Biomass of crustose lichens in tropical dry forest represents more than half of foliar biomass

In recent years, our ecological knowledge of tropical dry forests has increased dramatically. However, the functional contributions of whole ecosystem components, such as lichens, remain mostly unknown. In these forests, the abundance of epiphyte crustose lichens is responsible for the characteristic white bark on most woody plants, conspicuous during the dry season, but the amount of resources that the lichen component represents remains unexplored. We estimated lichen biomass in a Mexican tropical dry forest using the bark area of trees, the dry mass of lichens per unit area and the percentage of bark covered by lichens, together with previously known tree densities. The lowest 2.5 m of the forests main trunks contained 188 kg/ha of lichen biomass, with lichens covering 85% of the available bark for trees <12 cm DBH and 38% for trees >12 cm. Total epiphytic lichen biomass was 1.34–1.99 Mg/ha. Lichen biomass represented 61% of the foliar biomass in the forest. To our knowledge, this is the first time that a lichen biomass estimate is provided for an ecosystem in which crustose lichens are the dominant lichen growth form. Crustose lichens are typically considered to contribute little to the total lichen biomass and to be difficult to include in ecological analyses. The high lichen biomass in this ecosystem implies a significant ecological role which so far is unexplored. We suggest the crustose lichen component should not be underestimated a priori in ecological studies, especially in ecosystems with abundant lichen cover.     

Field observed relationships between biodiversity and ecosystem functioning during secondary succession in a tropical lowland rainforest

The relationship between biodiversity and ecosystem functioning (BEF) is one of the most concerned topics in ecology. However, most of the studies have been conducted in controlled experiments in grasslands, few observational field studies have been carried out in forests. In this paper, we report variations of species diversity, functional diversity and aboveground biomass (AGB) for woody plants (trees and shrubs) along a chronosequence of four successional stages (18-year-old fallow, 30-year-old fallow, 60-year-old fallow, and old-growth forest) in a tropical lowland rainforest recovered after shifting cultivation on Hainan Island, China. Fifty randomly selected sample plots of 20 m × 20 m were investigated in each of the four successional stages. Four functional traits (specific leaf area, wood density, maximum species height and leaf dry matter content) were measured for each woody plants species and the relationships between species/functional diversity and AGB during secondary succession were explored. The results showed that both plant diversity and AGB recovered gradually with the secondary succession. AGB was positively correlated with both species and functional diversity in each stage of succession. Consistent with many controlled experimental results in grasslands, our observational field study confirms that ecosystem functioning is closely related to biodiversity during secondary succession in species rich tropical forests.     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

Modeling the Sensitivity of the Seasonal Cycle of GPP to Dynamic LAI and Soil Depths in Tropical Rainforests

The seasonality of pan-tropical wet forests has been highlighted by recent remote sensing and eddy flux measurements that have recorded both increased and sustained dry-season gross primary productivity (GPP). These observations suggest that wet tropical forests are primarily light limited and that the mechanisms for resilience to drought and projected climate change must be considered in ecosystem model development. Here we investigate two proposed mechanisms for drought resilience of tropical forests, deep soil water access and the seasonality of phenology, using the LPJmL Dynamic Global Vegetation Model. We parameterize a new seasonal phenology module for tropical evergreen trees using remotely sensed leaf area index (LAI) and incoming solar radiation data from the Terra Earth Observing System. Simulations are evaluated along a gradient of dry-season length (DSL) in South America against MODIS GPP estimates. We show that deep soil water access is critical for maintaining dry-season GPP, whereas implementing a seasonal LAI did not enhance simulated dry-season GPP. The Farquhar-Collatz photosynthesis scheme used in LPJmL optimizes leaf nitrogen allocation according to light conditions, causing maximum photosynthetic capacity in the dry season. High LAI, characteristic of tropical forests, also dampens the seasonal amplitude of the fraction of photosynthetically active radiation (FPAR). Given the relatively high uncertainty in tropical phenology observations and their corresponding proximate drivers, we recommend that ecosystem model development focus on belowground processes. An improved representation of soil depths and rooting distributions is necessary for modeling the dynamics of dry-season tropical forest functioning and may have important impacts for modeling tropical forest vulnerability to climate change. Author Contributions  BP conceived of the study, analyzed data, and wrote the paper. UH designed study and contributed new methods. WC designed study and contributed to paper.     

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.     

Multiple successional pathways in human‐modified tropical landscapes: new insights from forest succession,forest fragmentation and landscape ecology research

Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.     

Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ_L) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although Ψ_L did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ¹⁸O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar ¹⁸O (∆¹⁸O_l) and ¹³C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.     

Regeneration of Juvenile Woody Plants after Fire in a Seasonally Dry Tropical Forest of Southern India

Woody tree species in seasonally dry tropical forests are known to have traits that help them to recover from recurring disturbances such as fire. Two such traits are resprouting and rapid post‐fire growth. We compared survival and growth rates of regenerating small‐sized individuals (juveniles) of woody tree species after dry season fire (February–March) at eight adjacent pairs of burnt and unburnt transects in a seasonally dry tropical forest in southern India. Juveniles were monitored at 3‐mo intervals between August 2009 and August 2010. High juvenile survivorship (>95%) was observed in both burnt and unburnt areas. Growth rates of juveniles, analyzed at the community level as well as for a few species individually (especially fast‐growing ones), were distinctly higher in burnt areas compared to unburnt areas after a fire event, particularly during the pre‐monsoon season immediately after a fire. Rapid growth by juveniles soon after a fire may be due to lowered competition from other vegetative forms such as grasses, possibly aided by the availability of resources stored belowground. Such an adaptation would allow a juvenile bank to be retained in the understory of a dry forest, from where individuals can grow to a possible fire‐tolerant size during favorable conditions.     

Plant functional traits with particular reference to tropical deciduous forests: a review

Functional traits (FTs) integrate the ecological and evolutionary history of a species, and can potentially be used to predict its response as well as its influence on ecosystem functioning. Study of inter-specific variation in the FTs of plants aids in classifying species into plant functional types (PFTs) and provides insights into fundamental patterns and trade-offs in plant form and functioning and the effect of changing species composition on ecosystem functions. Specifically, this paper focuses on those FTs that make a species successful in the dry tropical environment. Following a brief overview, we discuss plant FTs that may be particularly relevant to tropical deciduous forests (TDFs). We consider the traits under the following categories: leaf traits, stem and root traits, reproductive traits, and traits particularly relevant to water availability. We compile quantitative information on functional traits of dry tropical forest species. We also discuss trait-based grouping of plants into PFTs. We recognize that there is incomplete knowledge about many FTs and their effects on TDFs and point out the need for further research on PFTs of TDF species, which can enable prediction of the dynamics of these forests in the face of disturbance and global climate change. Correlations between structural and ecophysiological traits and ecosystem functioning should also be established which could make it possible to generate predictions of changes in ecosystem services from changes in functional composition.     

Evidence for strong seasonality in the carbon storage and carbon use efficiency of an Amazonian forest

The relative contribution of gross primary production and ecosystem respiration to seasonal changes in the net carbon flux of tropical forests remains poorly quantified by both modelling and field studies. We use data assimilation to combine nine ecological time series from an eastern Amazonian forest, with mass balance constraints from an ecosystem carbon cycle model. The resulting analysis quantifies, with uncertainty estimates, the seasonal changes in the net carbon flux of a tropical rainforest which experiences a pronounced dry season. We show that the carbon accumulation in this forest was four times greater in the dry season than in the wet season and that this was accompanied by a 5% increase in the carbon use efficiency. This seasonal response was caused by a dry season increase in gross primary productivity, in response to radiation and a similar magnitude decrease in heterotrophic respiration, in response to drying soils. The analysis also predicts increased carbon allocation to leaves and wood in the wet season, and greater allocation to fine roots in the dry season. This study demonstrates implementation of seasonal variations in parameters better enables models to simulate observed patterns in data. In particular, we highlight the necessity to simulate the seasonal patterns of heterotrophic respiration to accurately simulate the net carbon flux seasonal tropical forest.     

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。     

The Effects of Established Trees on Woody Regeneration during Secondary Succession in Tropical Dry Forests

Understanding the mechanisms controlling secondary succession in tropical dry forests is important for the conservation and restoration of this highly threatened biome. Canopy‐forming trees in tropical forests strongly influence later stages of succession through their effect on woody plant regeneration. In dry forests, this may be complex given the seasonal interplay of water and light limitations. We reviewed observational and experimental studies to assess (1) the relative importance of positive and negative effects of established trees on regeneration; (2) the mechanisms underlying these effects; and (3) to test the ‘stress gradient hypothesis’ in successional tropical dry forests. The effects of established trees on seed dispersal, seed survival, and seed germination—either through direct changes to moisture and temperature regimes or mediated by seed dispersers and predators—are mainly positive. The balance between positive and negative effects on seedling establishment is more complex and depends on the season and leaf phenology of both trees and seedlings. Seedling survival is generally enhanced by established trees mitigating dry conditions. Established trees have counteracting effects on water and light availability that influence seedling growth. The probability of a positive effect of established trees on seedling survival decreases with increased rainfall, which supports the stress gradient hypothesis. Priorities for future research are experiments to test for facilitation and competition and their underlying mechanisms, long‐term studies evaluating how these effects change with ontogeny, and studies focusing on the species‐specificity of interactions.     

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.     

Fruiting and flushing phenology in Asian tropical and temperate forests: implications for primate ecology

In order to understand the ecological adaptations of primates to survive in temperate forests, we need to know the general patterns of plant phenology in temperate and tropical forests. Comparative analyses have been employed to investigate general trends in the seasonality and abundance of fruit and young leaves in tropical and temperate forests. Previous studies have shown that (1) fruit fall biomass in temperate forest is lower than in tropical forest, (2) non-fleshy species, in particular acorns, comprise the majority of the fruit biomass in temperate forest, (3) the duration of the fruiting season is shorter in temperate forest, and (4) the fruiting peak occurs in autumn in most temperate forests. Through our comparative analyses of the fruiting and flushing phenology between Asian temperate and tropical forests, we revealed that (1) fruiting is more annually periodic (the pattern in one year is similar to that seen in the next year) in temperate forest in terms of the number of fruiting species or trees, (2) there is no consistent difference in interannual variations in fruiting between temperate and tropical forests, although some oak-dominated temperate forests exhibit extremely large interannual variations in fruiting, (3) the timing of the flushing peak is predictable (in spring and early summer), and (4) the duration of the flushing season is shorter. The flushing season in temperate forests (17–28 % of that in tropical forests) was quite limited, even compared to the fruiting season (68 %). These results imply that temperate primates need to survive a long period of scarcity of young leaves and fruits, but the timing is predictable. Therefore, a dependence on low-quality foods, such as mature leaves, buds, bark, and lichens, would be indispensable for temperate primates. Due to the high predictability of the timing of fruiting and flushing in temperate forests, fat accumulation during the fruit-abundant period and fat metabolization during the subsequent fruit-scarce period can be an effective strategy to survive the lean period (winter).     

Fruiting phenology in a tropical dry evergreen forest on the Coromandel coast of India in relation to plant life-forms,physiognomic groups,dispersal modes,and climatic constraints

The fruiting phenology of 22 woody plant species belonging to 19 families was studied with respect to life-forms, physiognomic groups and dispersal modes, for 1 year at monthly intervals, in a tropical dry evergreen forest at Oorani (12°11′N, 79°57′E) on the Coromandel coast of India. At the community level, bimodal fruiting pattern prevailed, with a major peak in the dry season and a minor one in the early rainy season. An annual fruiting pattern was observed in many species and among the studied species fruiting lasted for 2–9 months. There was no significant difference in the frequency of species at three fruiting stages across the life-form categories and many species of upper and lower canopy trees and lianas were in the ripe fruiting phase during the late dry season. Plant physiognomic groups displayed distinct seasonality in fruiting pattern. The fruit maturation period was much longer for the wet season fruiting brevi-deciduous species than evergreen and deciduous species that fruited during the dry season. The variation in timing of fruiting behaviour among zoochorous species demonstrated less seasonality and zoochorous fruits were available throughout the year. Fruiting in anemochorous species peaked during the driest months and dryness favoured the dissemination of seeds. The fruiting patterns observed in the studied tropical dry evergreen forest across various plant traits were comparable with patterns recorded in other tropical seasonal forests.     

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.     

Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀‐values ranged from ?0.8 to ?6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade‐tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber‐value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade‐off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade‐tolerants and evergreens. A trade‐off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.     

Sensitivity of Bolivian seasonally-dry tropical forest to precipitation and temperature changes over glacial–interglacial timescales

We used fossil pollen to investigate the response of the eastern Chiquitano seasonally-dry tropical forest (SDTF), lowland Bolivia, to high-amplitude climate change associated with glacial–interglacial cycles. Changes in the structure, composition and diversity of the past vegetation are compared with palaeoclimate data previously reconstructed from the same record, and these results shed light on the biogeographic history of today’s highly disjunct blocks of SDTF across South America. We demonstrate that lower glacial temperatures limited tropical forest in the Chiquitanía region, and suggest that SDTF was absent or restricted at latitudes below 17°S, the proposed location of the majority of the hypothesized ‘Pleistocene dry forest arc’ (PDFA). At 19500 yrs b.p., warming supported the establishment of a floristically-distinct SDTF, which showed little change throughout the glacial–Holocene transition, despite a shift to significantly wetter conditions beginning ca. 12500–12200 yrs b.p. Anadenanthera colubrina, a key SDTF taxon, arrived at 10000 yrs b.p., which coincides with the onset of drought associated with an extended dry season. Lasting until 3000 yrs b.p., Holocene drought caused a floristic shift to more drought-tolerant taxa and a reduction in α-diversity (shown by declining palynological richness), but closed-canopy forest was maintained throughout. In contrast to the PDFA, the modern distribution of SDTF most likely represents the greatest spatial coverage of these forests in southern South America since glacial times. We find that temperature is a key climatic control upon the distribution of lowland South American SDTF over glacial-interglacial timescales, and seasonality of rainfall exerts a strong control on their floristic composition.