Tropical Dry Forest Tree Species Research Articles

Above and belowground functional trait response to biochar addition in seedlings of six tropical dry forest tree species

Above and belowground functional trait response to biochar addition in seedlings of six tropical dry forest tree species

Variations in the plasticity of functional traits indicate the differential impacts of abiotic and biotic factors on the structure and growth of trees in tropical dry forest fragments

Abiotic and biotic factors have considerable impact on the plasticity of plant functional traits, which influences forest structure and productivity; however, their inter-relationships have not been quantified for fragmented tropical dry forest (TDF) ecosystems. We asked the following questions: (1) what are the variations in the plasticity of functional traits due to soil moisture availability in TDF fragments? (2) what are the roles of soil nutrients and forest disturbances in influencing variations in the plasticity of functional traits in the TDF fragments? and (3) how do the variations in the plasticity of functional traits influence the structure and productivity of TDF fragments? Based on linear mixed-effects results, we observed significant variations among tree species for soil moisture content (SMC) under the canopy and selected functional traits across forest fragments. We categorized tree species across fragments by principal component analysis (PCA) and hierarchical clustering on principal components (HCPC) analyses into three functional types, viz., low wood density high deciduous (LWHD), high wood density medium deciduous (HWMD), and high wood density low deciduous (HWLD). Assemblage of functional traits suggested that the LWHD functional type exhibits a drought-avoiding strategy, whereas HWMD and HWLD adopt a drought-tolerant strategy. Our study showed that the variations in functional trait plasticity and the structural attributes of trees in the three functional types exhibit contrasting affinity with SMC, soil nutrients, and disturbances, although the LWHD functional type was comparatively more influenced by soil resources and disturbances compared to HWMD and HWLD along the declining SMC and edge distance gradients. Plasticity in functional traits for the LWHD functional type exhibited greater variations in traits associated with the conservation of water and resources, whereas for HWMD and HWLD, the traits exhibiting greater plasticity were linked with higher productivity and water transport. The cumulative influence of SMC, disturbances, and functional trait variations was also visible in the relative abundance of functional types in large and small sized fragments. Our analysis further revealed the critical differences in the responses of functional trait plasticity of the coexisting tree species in TDF, which suggests that important deciduous endemic species with drought-avoiding strategies might be prone to strategic exclusion under expected rises in anthropogenic disturbances, habitat fragmentation, and resource limitations.

Throughfall exclusion and fertilization effects on tropical dry forest tree plantations, a large-scale experiment

Abstract. Across tropical ecosystems, global environmental change is causing drier climatic conditions and increased nutrient deposition. Such changes represent large uncertainties due to unknown interactions between drought and nutrient availability in controlling ecosystem net primary productivity (NPP). Using a large-scale manipulative experiment, we studied for 4 years whether nutrient availability affects the individual and integrated responses of aboveground and belowground ecosystem processes to throughfall exclusion in 30-year-old mixed plantations of tropical dry forest tree species in Guanacaste, Costa Rica. We used a factorial design with four treatments: control, fertilization (F), drought (D), and drought + fertilization (D + F). While we found that a 13 %–15 % reduction in soil moisture only led to weak effects in the studied ecosystem processes, NPP increased as a function of F and D + F. The relative contribution of each biomass flux to NPP varied depending on the treatment, with woody biomass being more important for F and root biomass for D + F and D. Moreover, the F treatment showed modest increases in maximum canopy cover. Plant functional type (i.e., N fixation or deciduousness) and not the experimental manipulations was the main source of variation in tree growth. Belowground processes also responded to experimental treatments, as we found a decrease in nodulation for F plots and an increase in microbial carbon use efficiency for F and D plots. Our results emphasize that nutrient availability, more so than modest reductions in soil moisture, limits ecosystem processes in tropical dry forests and that soil fertility interactions with other aspects of drought intensity (e.g., vapor pressure deficit) are yet to be explored.

Phosphorus limitation of early growth differs between nitrogen-fixing and nonfixing dry tropical forest tree species.

Tropical forests are often characterized by low soil phosphorus (P) availability, suggesting that P limits plant performance. However, how seedlings from different functional types respond to soil P availability is poorly known but important for understanding and modeling forest dynamics under changing environmental conditions. We grew four nitrogen (N)-fixing Fabaceae and seven diverse non-N-fixing tropical dry forest tree species in a shade house under three P fertilization treatments and evaluated carbon (C) allocation responses, P demand, P-use, investment in P acquisition traits, and correlations among P acquisition traits. Nitrogen fixers grew larger with increasing P addition in contrast to non-N fixers, which showed fewer responses in C allocation and P use. Foliar P increased with P addition for both functional types, while P acquisition strategies did not vary among treatments but differed between functional types, with N fixers showing higher root phosphatase activity (RPA) than nonfixers. Growth responses suggest that N fixers are limited by P, but nonfixers may be limited by other resources. However, regardless of limitation, P acquisition traits such as mycorrhizal colonization and RPA were nonplastic across a steep P gradient. Differential limitation among plant functional types has implications for forest succession and earth system models.

Effects of Light Availability on Survival and Growth of Seedling and Sapling of Tropical Dry Deciduous Forest Tree Species in Ballavpur Wild ife anctuary, West Bengal, India

Effects of Light Availability on Survival and Growth of Seedling and Sapling of Tropical Dry Deciduous Forest Tree Species in Ballavpur Wild ife anctuary, West Bengal, India

Tropical wet and dry forest tree species exhibit contrasting hydraulic architecture

Forest tree species in wet and dry habitats are generally considered functionally divergent in leaf and stem functional traits such as leaf area, leaf mass per area, wood density and tree height. Yet, these traits have limited utility for characterizing plant water transport adaptations and strategies. We tested the hypothesis that wet and dry forest trees are functionally divergent in their water conducting apparatus. To assess trait differences and adaptations, we sampled branch wood from nine same-genus species-pairs, each species-pair occurring respectively in the wet (>1500 mm annual rainfall) and dry forest (<800 mm annual rainfall) in tropical Queensland, Australia. From branch wood sections, we measured anatomical traits involved in water conduction (stem vessel dimensions, fractions and their spatial distributions, theoretical water conductivities), storage (parenchyma), and providing hydraulic safety functions (fibres fractions, vulnerability index). Relative to wet forest species, we found on overall that dry forest trees had trait combinations showing adaptations to aridity such as more storage tissue and greater vessel connectivity which may provide alternative pathways for water transport should vessel embolism occur. Habitat is an environmental filter that influences trait behaviour across related species. However, depending on the genera, species in both dry and wet forest habitats also exhibit various tradeoffs in trait values, highlighting the existence of diverse hydraulic strategies within wet forest and dry forest trees.

Beyond leaf habit: generalities in plant function across 97 tropical dry forest tree species.

Leaf habit has been hypothesized to define a linkage between the slow-fast plant economic spectrum and the drought resistance-avoidance trade-off in tropical forests ('slow-safe vs fast-risky'). However, variation in hydraulic traits as a function of leaf habit has rarely been explored for a large number of species. We sampled leaf and branch functional traits of 97 tropical dry forest tree species from four sites to investigate whether patterns of trait variation varied consistently in relation to leaf habit along the 'slow-safe vs fast-risky' trade-off. Leaf habit explained from 0% to 43.69% of individual trait variation. We found that evergreen and semi-deciduous species differed in their location along the multivariate trait ordination when compared to deciduous species. While deciduous species showed consistent trait values, evergreen species trait values varied as a function of the site. Last, trait values varied in relation to the proportion of deciduous species in the plant community. We found that leaf habit describes the strategies that define drought avoidance and plant economics in tropical trees. However, leaf habit alone does not explain patterns of trait variation, which suggests quantifying site-specific or species-specific uncertainty in trait variation as the way forward.

Identification of spectral features in the longwave infrared (LWIR) spectra of leaves for the discrimination of tropical dry forest tree species

With the emergence of longwave hyperspectral imaging systems, studies are revealing the potential of these data for discriminating tree species. However, few studies have applied statistical methods of band selection to select and characterize features at the species level that can then be used for improved classification. A dataset of leaf spectra was recently collected in-situ from twenty-six tree species in a Costa Rican tropical dry forest. The spectra of the species present overall low contrast and a range in spectral shapes, with some species displaying spectral similarity. This motivates our study to explore the performance of band selection tools to help identify key spectral features for the classification of these species.The bands selected using an ensemble of multiple methods improved the Logistic Regression classification performance by 3% in comparison to a result without band selection. The multiple methods encompassed the random forest, minimum redundancy maximum relevance and n-dimensional spectral solid angle methods. Bands selected by the ensemble methods agree well with the features previously identified based on expert knowledge and can be understood in the context of leaf constitutional compounds and related spectral features. The longwave hyperspectral bands or features identified in this study can potentially assist the future image mapping of tree species at large scales. The ensemble strategy is recommended for the band analysis of vegetation for its highest accuracy and stability.

Effects of grass competition on tree seedlings growth under different light and nutrient availability conditions in tropical dry forests in India

AbstractThe interactive effects of light, nutrient availability and grass competition on seedlings of four tropical dry forest tree species, namely, Indian jujube (Ziziphus mauritiana), arjun (Terminalia arjuna), gum arabic (Acacia nilotica) and catechu (Acacia catechu), were evaluated in an experimental study at Banaras Hindu University, Varanasi, India. Full‐ and reduced‐ (20% of the full) sunlight, no‐nutrient and nutrient supplementation, and presence of grass and no‐grass conditions were taken in a split‐plot design to observe the effects on the plant growth and leaf nutrient parameters. We observed a significant interaction between light and nutrient availability on total dry weight and relative growth rate for the tree seedlings of all the species. Seedling growth was lower in presence of grass under full sunlight condition with/without nutrient addition, and also under reduced sunlight condition with nutrient addition. However, leaf‐N and ‐P concentration showed slight increase following nutrient addition across the species and light conditions. All the species responded positively to nutrient addition irrespective of light and grass conditions. Our study further indicates that nutrient addition under reduced light condition in tropical dry forests may be a possible strategy to ameliorate the negative impact of grasses on tree seedling growth. Overall, a better understanding of tree seedling establishment in the presence of grasses is of utmost importance for the development of management plans that may improve the diversity of tree species in tropical dry forests.

Using large-scale tropical dry forest restoration to test successional theory.

Microclimatic conditions change dramatically as forests age and impose strong filters on community assembly during succession. Light availability is the most limiting environmental factor in tropical wet forest succession; by contrast, water availability is predicted to strongly influence tropical dry forest (TDF) successional dynamics. While mechanisms underlying TDF successional trajectories are not well understood, observational studies have demonstrated that TDF communities transition from being dominated by species with conservative traits to species with acquisitive traits, the opposite of tropical wet forest. Determining how functional traits predict TDF tree species' responses to changing environmental conditions could elucidate mechanisms underlying tree performance during TDF succession. We implemented a 6-ha restoration experiment on a degraded Vertisol in Costa Rica to determine (1) how TDF tree species with different resource-use strategies performed along a successional gradient and (2) how ecophysiological functional traits correlated with tree performance in simulated successional stages. We used two management treatments to simulate distinct successional stages including: clearing all remnant vegetation (early-succession), or interplanting seedlings with no clearing (mid-succession). We crossed these two management treatments (cleared/interplanted) with two species mixes with different resource-use strategies (acquisitive/conservative) to examine their interaction. Overall seedling survival after 2 yr was low, 15.1-26.4% in the four resource-use-strategy×management-treatment combinations, and did not differ between the management treatments or resource-use-strategy groups. However, seedling growth rates were dramatically higher for all species in the cleared treatment (year 1, 69.1% higher; year 2, 143.3% higher) and defined resource-use strategies had some capacity to explain seedling performance. Overall, ecophysiological traits were better predictors of species' growth and survival than resource-use strategies defined by leaf and stem traits such as specific leaf area. Moreover, ecophysiological traits related to water use had a stronger influence on seedling performance in the cleared, early-successional treatment, indicating that the influence of microclimatic conditions on tree survival and growth shifts predictably during TDF succession. Our findings suggest that ecophysiological traits should be explicitly considered to understand shifts in TDF functional composition during succession and that using these traits to design species mixes could greatly improve TDF restoration outcomes.

Tree height predicts the shape of radial sap flow profiles of Costa-Rican tropical dry forest tree species

Sap flow measurements are a crucial tool for studies of plant water relations, but upscaling to tree or stand transpiration requires accounting for the radial distribution of sap flow across the stem. Because radial sap flow profiles cannot be measured with single-point measuring methods, predicting their shape based on easier to measure structural or functional traits is a potential route to cost-efficiently improve water-use estimates.We used heat field deformation (HFD) sensors to measure sap flux density at different sapwood depths on 38 trees of eight tropical tree species in secondary dry forest patches in north-west Costa Rica. Based on a Bayesian hierarchical modeling framework, we developed a flexible model for radial profiles that expresses the average profile depth and the spread of the profile around this depth as functions of tree height, wood density and stem growth rate, while allowing for random tree and species effects. From the model output, we scale up to approximate whole-tree daily water use (DWU).Our model revealed pronounced tree height effects, with taller trees having narrower sap flow profiles peaking closer to the cambium. While it explained 96% of the variance in the dataset, large fractions were attributed to random species and tree individual effects. Including functional traits as predictors improved the accuracy of predictions considerably both for new trees and new species. DWU responded positively to tree height, but not to annual stem increment and wood density. Extrapolating the average sap flux in the outermost 2 cm over the entire sapwood area overestimated DWU on average by 26% compared to estimates based on radial profiles.Our model provides a starting point for future studies aiming to improve landscape-scale water-use estimates integrating single-point sap flow measurements and radial profiles from a subset of trees.

Wood density, deposits and mineral inclusions of successional tropical dry forest species

Wood water content, wood dry mass fraction, wood deposits and mineral inclusions separately affect wood mechanical properties, as well as plant survival and growth rates. Tree species occurring in recovering vegetation from deforestation face water stress, which is a particularly strong environmental filter in tropical dry forest regions. In this study, we analyze wood density and its associated variables of tree species present in early and late successional communities. We assessed and compared their wood density, wood water content, wood dry mass fraction, as well as wood cellular deposits, secretory elements and mineral inclusions which were described through anatomical sections. Wood density ranged from 0.10 g/cm3 (Jacaratia mexicana in late successional community) to 0.92 g/cm3 (Lysiloma divaricatum in early successional community). Successional species showed low intraspecific variability, despite large differences in the environmental conditions in the communities where they grow. Species with high wood water content were restricted to late successional communities, and pioneers showed higher wood density than non-pioneer species, which supports the linkage between high wood density and harsh environmental filtering through hydraulic safety. Wood density did not differ between columnar cacti (Pachycereus pecten-aboriginum and Pilosocereus collinsii) and tree species such as Bursera simaruba. Starch, tannins and crystals were abundant in most successional species and some species had gums; these deposits could potentially influence wood density and hydraulic, defense and storage plant functions. Our results suggest that modifications in the life cycle of successional tropical dry forest tree species due to anthropogenic activities may not be an important source of wood density variation.

Edaphic factors, successional status and functional traits drive habitat associations of trees in naturally regenerating tropical dry forests

Abstract Many studies have examined individual environmental drivers of tropical tree species distributions, but edaphic and successional gradients have not been considered simultaneously. Furthermore, determining how functional traits influence species distributions along these gradients may help to elucidate mechanisms behind community assembly. To assess the influence of environmental filtering on tropical dry forest (TDF) tree species distributions, we used forest inventory data from sites with large edaphic and successional gradients in NW Costa Rica. Our goals were to determine (1) whether edaphic or successional factors are more important determinants of the abundance of individual tree species in regenerating TDF, (2) how species‐level functional traits are related to edaphic and/or successional niche associations of tree species and (3) correlations between species‐level edaphic and successional niche associations. The distributions of 82 focal tree species were strongly driven by both edaphic and successional gradients. Overall, 94% of species responded to soil chemistry, 89% to soil texture and 94% to stand age gradients. Some functional traits were correlated with the edaphic and successional niche associations of TDF tree species. Species that specialized on soils with high total nutrient concentrations had higher foliar nutrient concentrations (nitrogen and phosphorus) and lower leaf dry matter content (LDMC). Species with wider stand age niches had lower LDMC and wood density. There were no correlations between edaphic and successional niche optima of TDF tree species. Our results indicate that successional and edaphic gradients concurrently drive community assembly in regenerating TDF. Moreover, our work underscores the importance of considering how the functional characteristics of TDF trees dictate species distributions across environmental gradients. A plain language summary is available for this article.

Using soil amendments and plant functional traits to select native tropical dry forest species for the restoration of degraded Vertisols

Abstract Tropical dry forests (TDFs) are critically endangered, and their restoration is understudied. Large‐scale passive restoration efforts in north‐west (NW) Costa Rica have catalysed TDF regeneration but are not effective on degraded Vertisols, where active restoration is necessary due to high content of shrink–swell clays that impede regeneration following degradation. We established a large‐scale restoration experiment in degraded former pastures in NW Costa Rica to determine (1) the restoration potential of native TDF tree species on Vertisols, (2) if plant functional traits elucidate mechanisms behind interspecific variability in species performance and (3) if affordable and readily available soil amendments increase seedling survivorship and growth. We planted 1,710 seedlings of 32 native species coupled with five amendments aimed at ameliorating root‐zone microclimatic conditions: sand, rice hulls, rice hull ash, hydrogel and unamended controls. For each species, we quantified a suite of resource‐acquisition and ecophysiological functional traits, and monitored survival and growth seasonally over 2 years. Interspecific survivorship after 2 years ranged widely (0%–92.5%). Functional traits including wood density, photosynthetic parameters and upregulation of integrated water‐use efficiency, explained interspecific variation in survivorship and growth at distinct ontogenetic stages. Easily measured leaf traits, however, were not good predictors of restoration potential. Hydrogel and sand amendments increased initial seedling survival, but after 2 years no differences among treatments were found. Synthesis and applications. We have shown it is possible, albeit challenging, to restore tropical dry forest (TDF) on degraded Vertisols. Our results support the use of functional trait‐based screenings to select tree species for restoration projects as tree species with high survivorship and growth in this stressful environment have overlapping ecophysiological functional traits. Furthermore, practitioners should consider water‐use and photosynthetic traits when designing initial species mixes for TDF restorations.

Effects of nutrient addition and soil drainage on germination of N-fixing and non-N-fixing tropical dry forest tree species

To develop generalised predictions regarding the effects of atmospheric nitrogen (N) and phosphorus (P) deposition on vegetation communities, it is necessary to account for the impacts of increased nutrient availability on the early life history stages of plants. Additionally, it is important to determine if these responses (a) differ between plant functional groups and (b) are modulated by soil drainage, which may affect the persistence of added nutrients. We experimentally assessed seed germination responses (germination proportion and germination energy, i.e. time to germination) of commonly occurring N-fixing and non-N-fixing tropical dry forest tree species found in India to simulated N and P deposition in well-drained soils, as well as soils with impeded drainage. When soils were not allowed to drain, germination proportion declined with nutrient addition, while germination energy remained unchanged. Stronger declines in germination proportion were observed for N-fixing species. In free-draining soils, nutrient addition did not affect germination proportion in either functional group. However, we detected a trend of delayed germination with nutrient addition, especially in N-fixers. Our results suggest that nutrient deposition can lead to potential shifts in functional dominance and tree community composition of tropical dry forests in the long term through its effects on early life stages of trees, although the mechanisms underlying the observed germination responses remain unclear. Further, such effects are likely to be spatially variable across the geographic range in which tropical dry forests occur depending on soil drainage properties.

Is the inverse leafing phenology of the dry forest understory shrub Jacquinia nervosa (Theophrastaceae) a strategy to escape herbivory?

In the dry forest of Santa Rosa National Park, Costa Rica, the understory shrub Jacquinia nervosa presents an inverse pattern of phenology that concentrates vegetative growth and reproduction during the dry season. In this study, we tested the "escape from herbivory" hypothesis as a potential explanation for the inverse phenological pattern of J. nervosa. We monitored leaf, flower and fruit production in 36 adult plants from October 2000 to August 2001. Leaves of six randomly selected branches per plant were marked and monitored every two weeks to measure the cumulative loss in leaf area. To analyze pre-dispersal seed predation we collected 15 fruits per plant and counted the total number of healthy and damaged seeds, as well as the number and type of seed predators found within the fruits. Leaf, flower, and fruit production occurred during the first part of the dry season (end of November to February). The cumulative herbivory levels were similar to those observed in other tropical dry forest tree species that concentrate leaf production during the wet season, and were concentrated on young leaves, which lost an average of 36.77 % of their area (SD = 34.35 %, N = 195). Chewing beetles of the genus Epicauta (Meloidae) were the most important herbivores. In mature leaves, most of the damage was caused by the beetle Coptocycla rufonotata (Chrysomelidae). Fruits took 4 months to develop during the dry season (January-March 2001) but continue increasing in size well into the first 3 months of the wet season (May-July). Average seed number per ripe fruit was 9 (SD = 5, N = 500). Seed predation in mature fruits was 42 % (SD = 47 %, N = 122). Most seeds were damaged by moth larvae of the family Tortricidae. Only 3 % of the flowers became fruits. This was influenced by the low level of flower synchrony (0.38+/-0.26, N = 36 plants), but neither leaf synchrony (0.88+/-0.06, N = 36 plants) nor plant size influenced fruit numbers. The significant damaged produced by insect herbivores in young leaves, fruits, and seeds, as well as the low reproductive index observed in J. nervosa, shows that the inverse leafing phenology of this species is not consistent with the "escape hypothesis" since J. nervosa was considerably attacked during the dry season. Considering the strong seasonality of the tropical dry forest and the heliophyte character of J. nervosa, it is more likely that this phenological strategy evolved in response to seasonal fluctuations in light availability, light quality, and daylength.

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients? Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (MD), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-toshoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.

Source water, phenology and growth of two tropical dry forest tree species growing on shallow karst soils

Seasonally dry tropical forests are dominated by deciduous and evergreen tree species with a wide range of leaf phenology. We hypothesized that Piscidia piscipula is able to extend leaf senescence until later in the dry season due to deeper and more reliable water sources than Gymnopodium floribundum, which loses leaves earlier in the dry season. Physiological performance was assessed as timing of leaf production and loss, growth, leaf water potential, depth of water uptake determined by stable isotopes, and leaf stable isotopic composition of carbon (δ13C) and oxygen (δ18O). P. piscipula took water primarily from shallow sources, whereas G. floribundum took water from shallow and deep sources. The greatest variation in water sources occurred during the onset of the dry season, when G. floribundum was shedding old leaves and growing new leaves, but P. piscipula maintained its leaves from the previous wet season. P. piscipula showed greater relative growth rate, greater leaf expansion rates, and more negative predawn and midday water potentials than G. floribundum. P. piscipula also exhibited greater leaf organic δ13C and lower δ18O values, indicating that the decrease in photosynthetic carbon isotope discrimination was associated with greater stomatal conductance and greater photosynthesis. Our results indicate that the contrasting early and late dry season leaf loss phenology of these two species is not simply determined by rooting depth, but rather a more complicated suite of characteristics based on opportunistic use of dynamic water sources, maximizing carbon gain, and maintenance of water potential during the dry season.

From seedlings to trees: Using ontogenetic models of growth and survivorship to assess long-term (>100 years) dynamics of a neotropical dry forest

Long-term tree and seedling growth and survivorship data from permanent sample plots established in a neotropical dry forest in Jamaica from 1998 to 2008 were used to (1) model growth (periodic annual increment) and survivorship dynamics, (2) cluster structural and functional types, and (3) estimate the age of selected tropical dry forest tree species. A statistical comparison of parameter estimates derived from a generalized linear model (GLM) of each species to a reference species was used to group individuals based on size (DBH and height), and demographic dynamics (periodic annual increment and survivorship). We identified two groups of species based on structural types (canopy and sub-canopy species), three groups of species based on periodic annual increment (fast, intermediate, and slow growing) and four groups of species based on the probability of survivorship of seedlings and trees (very low probability of seedling survivorship but high tree survivorship (two groups); high survivorship throughout the DBH classes; very low survivorship, regardless of stem size). The composition of the groups was mixed, and included individuals of both structural types, and with different periodic annual increment and survival probabilities. The dichotomy of guilds found in tropical rainforests (pioneer and climax species) was not found in this forest. Individual and group GLMs incorporating empirical relationships between periodic annual increment and survivorship, across a spectrum of ontogenies and DBH’s, were also generated. The periodic annual increment models were then used to estimate the time taken by a newly germinated seedling to reach the largest recorded DBH. The fastest growing species was the hemi-cryptophyte Clusia flava which was estimated to take 74 years to reach its maximum recorded size (12.1 cm DBH), whereas the slowest growing species, Ziziphus sarcomphalus, was estimated to take 399 years to reach its maximum size (24.4 cm DBH). These dry forest trees were estimated to reach their maximum size (which was one-half or one-third of the largest DBH recorded for tropical rainforests) in a time similar to tropical rainforest trees. Some of the tree species are ubiquitous to other neotropical dry forests; therefore, our equations for periodic annual increment and survivorship can be applied elsewhere in the region.

Hydraulics and life history of tropical dry forest tree species: coordination of species’ drought and shade tolerance

Plant hydraulic architecture has been studied extensively, yet we know little about how hydraulic properties relate to species' life history strategies, such as drought and shade tolerance. The prevailing theories seem contradictory. We measured the sapwood (K(s) ) and leaf (K(l) ) hydraulic conductivities of 40 coexisting tree species in a Bolivian dry forest, and examined associations with functional stem and leaf traits and indices of species' drought (dry-season leaf water potential) and shade (juvenile crown exposure) tolerance. Hydraulic properties varied across species and between life-history groups (pioneers vs shade-tolerant, and deciduous vs evergreen species). In addition to the expected negative correlation of K(l) with drought tolerance, we found a strong, negative correlation between K(l) and species' shade tolerance. Across species, K(s) and K(l) were negatively correlated with wood density and positively with maximum vessel length. Consequently, drought and shade tolerance scaled similarly with hydraulic properties, wood density and leaf dry matter content. We found that deciduous species also had traits conferring efficient water transport relative to evergreen species. Hydraulic properties varied across species, corresponding to the classical trade-off between hydraulic efficiency and safety, which for these dry forest trees resulted in coordinated drought and shade tolerance across species rather than the frequently hypothesized trade-off.