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.     

Water-use efficiency and whole-plant performance of nine tropical tree species at two sites with contrasting water availability in Panama

Across their natural distributions, tropical tree species are regularly exposed to seasonal droughts of varying intensities. Their ability to tolerate drought stress plays a vital role in determining growth and mortality rates, as well as shaping the functional composition of tropical forests. In order to assess the ability of species to acclimate to contrasting levels of drought stress, physiological and structural traits involved in drought adaptation—wood C isotope discrimination (δ¹³C), wood specific gravity, and wood C content—of 2-year-old saplings of nine tropical tree species were evaluated in common garden experiments at two study sites in Panama with contrasting seasonality. We assessed co-variation in wood traits with relative growth rates (RGR_BD), aboveground biomass, and basal diameter and the plasticity of wood traits across study sites. Overall, species responded to lower water availability by increasing intrinsic water-use efficiency, i.e., less negative wood δ¹³C, but did not exhibit a uniform, directional response for wood specific gravity or wood C content. Trait plasticity for all wood traits was independent of RGR_BD and tree size. We found that the adaptive value of intrinsic water-use efficiency varied with water availability. Intrinsic water-use efficiency increased with decreasing RGR_BD at the more seasonal site, facilitating higher survival of slower growing species. Conversely, intrinsic water-use efficiency increased with tree size at the less seasonal site, which conferred a competitive advantage to larger individuals at the cost of greater susceptibility to drought-induced mortality. Our results illustrate that acclimation to water availability has negligible impacts on tree growth over short periods, but eventually could favor slow-growing species with conservative water-use strategies in tropical regions experiencing increasingly frequent and severe droughts.     

Geographical distributions in tropical trees: can geographical range predict performance and habitat association in co‐occurring tree species?

Aim A major floristic and climatic transition from aseasonal to seasonal evergreen tropical forest (the Kangar–Pattani Line; KPL) exists in the Indo‐Sundaic region of Southeast Asia. Mechanisms constraining species distribution here are at present poorly understood, but it is hypothesized that species differ in their tolerances of abiotic factors, in particular water availability. Under this hypothesis, we anticipate differences in performance or habitat preferences, or both, of species differing in distribution with respect to the KPL. The aim of this study is to test whether geographical distributions can be used to explain variation in growth, mortality and habitat preferences in co‐occurring tree species differing in their distribution in relation to the KPL. Location Pasoh Forest Reserve, Negeri Sembilan, Malaysia; south of the KPL. Methods All tree species within a 50‐ha forest dynamics plot were classified as widespread or southern based upon their distributions in relation to the KPL and as habitat specialists or generalists based on spatial association with soil‐based habitat categories. Growth and mortality rates, variation in growth and mortality with respect to soil type, and levels of habitat association were quantified for species with different geographical distributions. Results Differences existed in species performance based upon geographical distributions. Specifically, widespread species had lower growth rates than did species restricted to the aseasonal forests. Mortality rates did not differ as a function of geographical distribution. The growth responses of species to soil habitats also diverged, such that differences in performance of widespread species among soil types were more conservative than those of species restricted in their distribution to the aseasonal forests. However, the proportion of species showing positive habitat associations did not differ significantly between widespread and southern species. Main conclusions Distribution‐based differences in species performance and response to soil type support the hypothesis that species tolerant of wider climatic variation perform less well in any given environment due to limitations on plasticity. These performance differences provide quantitative evidence of the role of climatic transitions in determining tree species distributions in relation to the Kangar–Pattani Line in the Indo‐Malay region. Such differences in performance have important implications for our understanding of biodiversity gradients and responses of Indo‐Sundaic forests to climate change.     

Distribution of seed dormancy classes across a fire-prone continent: effects of rainfall seasonality and temperature

Background and AimsDifferent seed dormancy classes control the timing of germination via different cues. The ecological dissimilarities between classes therefore suggest that they are likely to be subject to different selective pressures, and that species within each class will have diverse functional responses. We aimed to investigate this by assessing how variation in the distribution of dormancy classes is correlated with regional environmental factors, in particular rainfall seasonality and temperature. Additionally, we compare the relative proportions of species with physiological (PD) or physical (PY) dormancy to assess whether dormancy class influences their ability to persist under different rainfall seasonality regimes.MethodsDormancy class was assigned for 3990 species from 281 genera occurring across two climate regions, with either winter or aseasonal rainfall, across temperate fire-prone Australia. All regions have similar vegetation and fire regimes. Using a Bayesian framework, we compared the distribution of dormancy classes across temperature and rainfall climate gradients, for threatened and common species.Key ResultsA high dormant:non-dormant species ratio highlighted the critical role of dormancy across our study regions. Critically, species showing PD were more likely to be threatened in aseasonal rainfall climate regions.ConclusionsOur results support the assumption that dormancy is favoured in environments with stochastic disturbance     

Markedly Divergent Tree Assemblage Responses to Tropical Forest Loss and Fragmentation across a Strong Seasonality Gradient

We examine the effects of forest fragmentation on the structure and composition of tree assemblages within three seasonal and aseasonal forest types of southern Brazil, including evergreen, Araucaria, and deciduous forests. We sampled three southernmost Atlantic Forest landscapes, including the largest continuous forest protected areas within each forest type. Tree assemblages in each forest type were sampled within 10 plots of 0.1 ha in both continuous forests and 10 adjacent forest fragments. All trees within each plot were assigned to trait categories describing their regeneration strategy, vertical stratification, seed-dispersal mode, seed size, and wood density. We detected differences among both forest types and landscape contexts in terms of overall tree species richness, and the density and species richness of different functional groups in terms of regeneration strategy, seed dispersal mode and woody density. Overall, evergreen forest fragments exhibited the largest deviations from continuous forest plots in assemblage structure. Evergreen, Araucaria and deciduous forests diverge in the functional composition of tree floras, particularly in relation to regeneration strategy and stress tolerance. By supporting a more diversified light-demanding and stress-tolerant flora with reduced richness and abundance of shade-tolerant, old-growth species, both deciduous and Araucaria forest tree assemblages are more intrinsically resilient to contemporary human-disturbances, including fragmentation-induced edge effects, in terms of species erosion and functional shifts. We suggest that these intrinsic differences in the direction and magnitude of responses to changes in landscape structure between forest types should guide a wide range of conservation strategies in restoring fragmented tropical forest landscapes worldwide.     

How much have we learned about seasonality in tropical insect abundance since Wolda (1988)?

Seasonal patterns in climatic conditions affect the life cycles and temporal patterns in the abundance of most temperate insect species. In tropical regions where there is no winter season, the situation may be different. For a better understanding of the evolution of seasonal life cycles, and the dynamics affecting temporal patterns in abundance of tropical insect populations and assemblages, it is important to study the life cycles of tropical insects and the presence or absence of seasonality in relation to climatic conditions. By reviewing studies on temporal patterns of abundance, this article examines the patterns of seasonality in adult tropical forest insects and discusses the variation in such patterns in various forest types. Seasonal and aseasonal patterns were found to be common in tropical dry and wet regions, respectively. In wet regions, which lack a distinctive dry season, there exists a wide variety of temporal patterns in addition to aseasonal patterns: distinctively seasonal and supra‐annual fluctuations in some insect species. Some of the problems of hidden ecological mechanisms underlying seasonal patterns in abundance are discussed, and the definition of seasonality in temporal patterns of insect abundance at a particular stage in the life cycle is considered. Methodological problems are also discussed.     

Geological substrates shape tree species and trait distributions in african moist forests

Background

Understanding the factors that shape the distribution of tropical tree species at large scales is a central issue in ecology, conservation and forest management. The aims of this study were to (i) assess the importance of environmental factors relative to historical factors for tree species distributions in the semi-evergreen forests of the northern Congo basin; and to (ii) identify potential mechanisms explaining distribution patterns through a trait-based approach.

Methodology/Principal Findings

We analyzed the distribution patterns of 31 common tree species in an area of more than 700,000 km² spanning the borders of Cameroon, the Central African Republic, and the Republic of Congo using forest inventory data from 56,445 0.5-ha plots. Spatial variation of environmental (climate, topography and geology) and historical factors (human disturbance) were quantified from maps and satellite records. Four key functional traits (leaf phenology, shade tolerance, wood density, and maximum growth rate) were extracted from the literature. The geological substrate was of major importance for the distribution of the focal species, while climate and past human disturbances had a significant but lesser impact. Species distribution patterns were significantly related to functional traits. Species associated with sandy soils typical of sandstone and alluvium were characterized by slow growth rates, shade tolerance, evergreen leaves, and high wood density, traits allowing persistence on resource-poor soils. In contrast, fast-growing pioneer species rarely occurred on sandy soils, except for Lophira alata.

Conclusions/Significance

The results indicate strong environmental filtering due to differential soil resource availability across geological substrates. Additionally, long-term human disturbances in resource-rich areas may have accentuated the observed patterns of species and trait distributions. Trait differences across geological substrates imply pronounced differences in population and ecosystem processes, and call for different conservation and management strategies.     

Testing the low latitude/high defense hypothesis for broad-leaved tree species

We tested the hypothesis that leaves of broad-leaved tree species are more highly defended at low latitudes than at high latitudes. We used canonical discriminant analysis to compare tree species from Panama (9°N, 39 species), Missouri, USA (38°N, 37 species), and southern Ontario, Canada (44°N, 34 species) with respect to two structural and five nutritional traits, taking into account each species' tolerance to shade. Trees from the three locations differed significantly, with Panamanian species the most distinct. Defenses of shade-tolerant species were significantly greater than those of shade-intolerant species, but only for the Panamanian sample, which is consistent with the low latitude/high defense hypothesis. Because we sampled many of the same tree species from Missouri and southern Ontario, and many tree species in the same taxonomic families in Missouri and Panama, we were able to control for the potential confounding effects of phylogeny. Overall defense levels, calculated by summing the z-scores for individual traits in each location, were significantly higher for Panama compared to Missouri, and marginally so for Missouri compared to southern Ontario, again consistent with the low latitude/high defense hypothesis. Traits contributing to these differences were mostly structural factors (e.g., fiber) and to a lesser degree nutritional traits, while secondary compounds made no independent contribution to differences in overall defense levels (four traits compared between Panama and Missouri). Contrary to our expectation, the number and types of secondary compounds per species reported in the literature for our species did not differ between temperate and tropical locations, while the diversity of these compounds was greater for the temperate species. Overall, our results provide some support for the hypothesis that leaf defenses against herbivory are better developed in tropical than in temperate trees, but the differences were due to structural and nutritional factors rather than secondary compounds.     

Functional traits of lianas in an Australian lowland rainforest align with post‐disturbance rather than dry season advantage

Lianas are an important component of tropical forests; they alter tree mortality and recruitment and impact biogeochemical cycling. Recent evidence suggests that the abundance of lianas in tropical forests is increasing. To understand and predict the effect of lianas on ecosystem processes in tropical forests, it is important to understand the mechanisms through which they compete with trees. In this study, we investigated the functional traits of lianas and trees in a lowland tropical forest in northeast Queensland, Australia. The site is located at 16.1° south latitude and experiences significant seasonality in rainfall, with pronounced wet and dry seasons. It is also subject to relatively frequent disturbance by cyclones. We asked the question of whether the canopy liana community at this site would display functional traits consistent with a competitive advantage over trees in response to disturbance, or in response to dry season water stress. We found that traits that we considered indicative of a dry season advantage (xylem water δ¹⁸O as an indicator of rooting depth; leaf and stem tissue δ¹³C and instantaneous gas exchange as measures of water‐use efficiency) did not differ between canopy lianas and canopy trees. On the other hand, lianas differed from trees in traits that should confer an advantage in response to disturbance (low wood density; low leaf dry matter content; high leaf N concentration; high mass‐based photosynthetic rates). We conclude that the liana community at the study site expressed functional traits geared towards rapid resource acquisition and growth in response to disturbance, rather than outcompeting trees during periods of water stress. These results contribute to a body of literature which will be useful for parameterising a liana functional type in ecosystem models.     

Liana abundance and diversity increase with rainfall seasonality along a precipitation gradient in Panama

In tropical regions, rainfall gradients often explain the abundance and distribution of plant species. For example, many tree and liana species adapted to seasonal drought are more abundant and diverse in seasonally-dry forests, characterized by long periods of seasonal water deficit. Mean annual precipitation (MAP) is commonly used to explain plant distributions across climate gradients. However, the relationship between MAP and plant distribution is often weak, raising the question of whether other seasonal precipitation patterns better explain plant distributions in seasonally-dry forests. In this study, we examine the relationship between liana abundance and multiple metrics of seasonal and annual rainfall distribution to test the hypothesis that liana density and diversity increase with increasing seasonal drought along a rainfall gradient across the isthmus of Panama. We found that a normalized seasonality index, which combines MAP and the variability of monthly rainfall throughout the year, was a significant predictor of both liana density and species richness, whereas MAP, rainfall seasonality and the mean dry season precipitation (MDP) were far weaker predictors. The strong response of lianas to the normalized seasonality index indicates that, in addition to the total annual amount of rainfall, how rainfall is distributed throughout the year is an important determinant of the hydrological conditions that favor liana proliferation. Our findings imply that changes in annual rainfall and rainfall seasonality will determine the future distribution and abundance of lianas. Models that aim to predict future plant diversity, distribution, and abundance may need to move beyond MAP to a more detailed understanding of rainfall variability at sub-annual timescales.     

Monthly survival of African Sylvia warblers in a seasonally arid tropical environment

Latitudinal gradients of life-history traits in animals are thought to be shaped by environmental variables. For example, it has been suggested that the increase in avian clutch size from the tropics towards the northern temperate regions is caused by a reduced survival of adult birds in the north due to increasing environmental seasonality. However, the tropical savannahs of East Africa show pronounced seasonality in resources caused by distinct rainy and dry seasons. This raises the question of whether survival and other life-history traits of birds living in these tropical savannahs are influenced by this seasonality, making them more similar to northern temperate species. We used 2-year monthly resighting data, a multistate modelling approach and the program MARK to test whether survival, transition probabilities between breeding states and other life-history traits of two resident Kenyan Sylvia species (Aves: Passeriformes: Sylviidae) are shaped by seasonality of rainfall in their environment. Contradicting our hypotheses, the two species showed only very slight influence of seasonality of rainfall on their survival. Survival in the dry months was hardly lower than in the rainy months. The species in the more seasonal environment ( S. boehmi , annual survival 71%) survived as well as the one in the more constant environment ( S. lugens , 56%). The observed survival rates correspond well to other life-history traits of the two species and are of similar magnitude to survival rates of other tropical passerines. This implies that either seasonality is not the driving force behind the life-history traits of the two species or the birds do not experience their environment as seasonal, as might be suggested by fluctuations in rainfall.     

Testing the divergent adaptation of two congeneric tree species on a rainfall gradient using eco‐physio‐morphological traits

In tropical Africa, evidence of widely distributed genera transcending biomes or habitat boundaries has been reported. The evolutionary processes that allowed these lineages to disperse and adapt into new environments are far from being resolved. To better understand these processes, we propose an integrated approach, based on the eco‐physio‐morphological traits of two sister species with adjacent distributions along a rainfall gradient. We used wood anatomical traits, plant hydraulics (vulnerability to cavitation, wood volumetric water content, and hydraulic capacitance), and growth data from the natural habitat, in a common garden, to compare species with known phylogeny, very similar morphologically, but occupying contrasting habitats: Erythrophleum ivorense (wet forest) and Erythrophleum suaveolens (moist forest and forest gallery). We identified some slight differences in wood anatomical traits between the two species associated with strong differences in hydraulics, growth, and overall species distribution. The moist forest species, E. suaveolens, had narrower vessels and intervessel pits, and higher vessel cell‐wall reinforcement than E. ivorense. These traits allow a high resistance to cavitation and a continuous internal water supply of the xylem during water shortage, allowing a higher fitness during drought periods, but limiting growth. Our results confirm a trade‐off between drought tolerance and growth, controlled by subtle adaptations in wood traits, as a key mechanism leading to the niche partitioning between the two Erythrophleum species. The generality of this trade‐off and its importance in the diversification of the African tree flora remains to be tested. Our integrated eco‐physio‐morpho approach could be the way forward.     

Seedling Traits Determine Drought Tolerance of Tropical Tree Species

Water availability is the most important factor determining tree species distribution in the tropics, but the underlying mechanisms are still not clear. In this study, we compared functional traits of 38 tropical tree species from dry and moist forest, and quantified their ability to survive drought in a dry‐down experiment in which wilting and survival were monitored. We evaluated how seedling traits affect drought survival, and how drought survival determines species distribution along the rainfall gradient. Dry forest species tended to have compound leaves, high stem dry matter content (stem dry mass/fresh mass), and low leaf area ratio, suggesting that reduction of transpiration and avoidance of xylem cavitation are important for their success. Three functional groups were identified based on the seedling traits: (1) drought avoiders with a deciduous leaf habitat and taproots; (2) drought resisters with tough tissues (i.e., a high dry matter content); and (3) light‐demanding moist forest species with a large belowground foraging capacity. Dry forest species had a longer drought survival time (62 d) than moist forest species (25 d). Deciduousness explained 69 percent of interspecific variation in drought survival. Among evergreen species, stem density explained 20 percent of the drought survival. Drought survival was not related to species distribution along the rainfall gradient, because it was mainly determined by deciduousness, and species with deciduous seedlings are found in both dry and moist forests. Among evergreen species, drought survival explained 28 percent of the variation in species position along the rainfall gradient. This suggests that, apart from drought tolerance, other factors such as history, dispersal limitation, shade tolerance, and fire shape species distribution patterns along the rainfall gradient.     

Effects of Drought,Pest Pressure and Light Availability on Seedling Establishment and Growth: Their Role for Distribution of Tree Species across a Tropical Rainfall Gradient

Tree species distributions associated with rainfall are among the most prominent patterns in tropical forests. Understanding the mechanisms shaping these patterns is important to project impacts of global climate change on tree distributions and diversity in the tropics. Beside direct effects of water availability, additional factors co-varying with rainfall have been hypothesized to play an important role, including pest pressure and light availability. While low water availability is expected to exclude drought-intolerant wet forest species from drier forests (physiological tolerance hypothesis), high pest pressure or low light availability are hypothesized to exclude dry forest species from wetter forests (pest pressure gradient and light availability hypothesis, respectively). To test these hypotheses at the seed-to-seedling transition, the potentially most critical stage for species discrimination, we conducted a reciprocal transplant experiment combined with a pest exclosure treatment at a wet and a dry forest site in Panama with seeds of 26 species with contrasting origin. Establishment success after one year did not reflect species distribution patterns. However, in the wet forest, wet origin species had a home advantage over dry forest species through higher growth rates. At the same time, drought limited survival of wet origin species in the dry forest, supporting the physiological tolerance hypothesis. Together these processes sort species over longer time frames, and exclude species outside their respective home range. Although we found pronounced effects of pests and some effects of light availability on the seedlings, they did not corroborate the pest pressure nor light availability hypotheses at the seed-to-seedling transition. Our results underline that changes in water availability due to climate change will have direct consequences on tree regeneration and distributions along tropical rainfall gradients, while indirect effects of light and pests are less important.     

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.     

Does breeding season variation affect evolution of a sexual signaling trait in a tropical lizard clade?

Sexually selected traits can be expected to increase in importance when the period of sexual behavior is constrained, such as in seasonally restricted breeders. Anolis lizard male dewlaps are classic examples of multifaceted signaling traits, with demonstrated intraspecific reproductive function reflected in courtship behavior. Fitch and Hillis found a correlation between dewlap size and seasonality in mainland Anolis using traditional statistical methods and suggested that seasonally restricted breeding seasons enhanced the differentiation of this signaling trait. Here, we present two tests of the Fitch–Hillis Hypothesis using new phylogenetic and morphological data sets for 44 species of Mexican Anolis. A significant relationship between dewlap size and seasonality is evident in phylogenetically uncorrected analyses but erodes once phylogeny is accounted for. This loss of strong statistical support for a relationship between a key aspect of dewlap morphology and seasonality also occurs within a species complex (A. sericeus group) that inhabits seasonal and aseasonal environments. Our results fail to support seasonality as a strong driver of evolution of Anolis dewlap size. We discuss the implications of our results and the difficulty of disentangling the strength of single mechanisms on trait evolution when multiple selection pressures are likely at play.     

Stem growth rhythms in trees of a tropical rainforest in Southern Brazil

Key message

We demonstrate that tropical trees growing in wet climates can have a marked seasonality in cambium activity and stem growth associated with high temperature and day length of summer.

Abstract

Monitoring the rhythm of tree growth associated with the growth rings can contribute substantially to understanding forest dynamics and the management of tropical forests. In this study, we monitored the girth increment rhythm and described the wood characteristics (anatomy of growth rings, wood specific gravity) in 10 tropical tree species (103 individuals) naturally occurring in a wet and weakly seasonal region of Atlantic Forest in southern Brazil. We aimed to verify whether tree growth dynamics are associated with climate and woody anatomy in tropical trees with contrasting ecological characteristics. We installed permanent dendrometer bands and monthly assessed the girth increment for 22 months. We collected wood samples (non-destructive method), measured wood specific gravity and prepared permanent slides to characterize the growth ring markers. We found growth rings in all species (distinct in six species); deciduous species produced more distinguishable tree rings compared with semi-deciduous and evergreen tree species. Species varied in their accumulated girth growth (in average, from 1.83 to 62.64 mm), growth rates (1–15 %), and annual radial increment (0.16–5.44 mm). Girth increment was positively related to temperature and day length in five out of ten tree species, indicating the possible effects of these climatic variables in triggering cambial activity in these species. The growth pattern varied among species and was marginally associated to the tree deciduousness. We concluded that even in wet and less seasonal climates, there can be an association in the cambium activity and stem growth with the hotter and longer days of summer months.

     

Exploring wood anatomy,density and chemistry profiles to understand the tree-ring formation in Amazonian tree species

Long-term analysis of tree growth using annual tree rings is increasingly in demand for tropical tree species. The basis of these studies has traditionally been the anatomical identification of the annual ring boundary. However, the structure of these annual rings has been sparsely explored for complementary physical and chemical wood traits. Here, we explore the relationships among wood density features and chemical elements (S, K, Ca, Mn) involved in the annual tree ring formation of 12 tropical tree species from non-flooded forest in the southern Amazon basin. Transverse wood sections were used for each species to determine: 1) macroscopic distinction (radial growth and wood density), 2) microscopic analyse of vessels, axial and ray parenchyma (anatomy) and 3) X-ray densitometry (physical) and X-ray fluorescence (chemical). For some species, the profiles of wood density, and Ca and Mn content showed intra- and inter-annual patterns that allowed to define and characterize the growth boundary of tree rings. Ca, K and S were mainly distributed in axial parenchyma cells, and around vessels, whereas, Mn was mainly distributed in fibres. Our results showed significant species-specific correlations between tree-ring width, density and concentrations of Ca, K and Mn. The anatomical characterization and the complementary information provided by the density and chemical profiles in some Amazonian species can represent a valuable proxy to improve the definition of annual ring-boundaries and improve the understanding of long-term growth and physiological patterns.     

Tree and Liana Enumeration and Diversity on a One-Hectare Plot in Papua New Guinea1

Enumeration of a one hectare plot at 900 m a.s.l in Papua New Guinea revealed 693 individuals of 228 tree and liana species ≥ 10 cm DBH. A 0.1 hectare subplot contained 302 individuals of 106 species 2.5 ≥ DBH < 10 cm. Lauraceae, Moraceae, and Myristicaceae were the most important families in both size classes. This site is very diverse compared with other tropical forests, and like other species-rich sites worldwide, it has high aseasonal rainfall and high rates of natural disturbance.     

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

One of the key hypothesized drivers of gradients in species richness is environmental filtering, where environmental stress limits which species from a larger species pool gain membership in a local community owing to their traits. Whereas most studies focus on small‐scale variation in functional traits along environmental gradient, the effect of large‐scale environmental filtering is less well understood. Furthermore, it has been rarely tested whether the factors that constrain the niche space limit the total number of coexisting species. We assessed the role of environmental filtering in shaping tree assemblages across North America north of Mexico by testing the hypothesis that colder, drier, or seasonal environments (stressful conditions for most plants) constrain tree trait diversity and thereby limit species richness. We assessed geographic patterns in trait filtering and their relationships to species richness pattern using a comprehensive set of tree range maps. We focused on four key plant functional traits reflecting major life history axes (maximum height, specific leaf area, seed mass, and wood density) and four climatic variables (annual mean and seasonality of temperature and precipitation). We tested for significant spatial shifts in trait means and variances using a null model approach. While we found significant shifts in mean species’ trait values at most grid cells, trait variances at most grid cells did not deviate from the null expectation. Measures of environmental harshness (cold, dry, seasonal climates) and lower species richness were weakly associated with a reduction in variance of seed mass and specific leaf area. The pattern in variance of height and wood density was, however, opposite. These findings do not support the hypothesis that more stressful conditions universally limit species and trait diversity in North America. Environmental filtering does, however, structure assemblage composition, by selecting for certain optimum trait values under a given set of conditions.