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.     

Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community

Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.     

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.     

Effects of past disturbance and edges on tree community structure and dynamics within a fragment of tropical semideciduous forest in south-eastern Brazil over a five-year period (1987–1992)

Tree community structure and dynamics of a 5.8 ha fragment of montane semideciduous forest in south-eastern Brazil are described based on two successive surveys of trees with dbh 5 cm carried out in 1987 and 1992 in 126 20 × 20 m quadrats. The main purpose was to assess whether the spatial variation of dynamic and structural variables were related to edge-effects and past disturbance regimes. The totals for the two surveys were, respectively, 6528 and 6770 trees, and 94.89 and 108.53 m² of basal area. The forest fragment was at an aggrading sylvigenetic phase, indicated by an overall net increase in density and basal area, and by a declining proportion of trees of smaller size. The overall annual mortality and recruitment rates were 2.6% and 3.0%, respectively. The dynamic process, however, was strongly affected by spatial heterogeneity. The more severely disturbed sectors had lower average canopy heights and basal areas per hectare, and higher tree densities, proportions of smaller trees, and relative area of canopy gaps. They also had higher mortality and recruitment rates, resulting in higher turnover rates compared to the sectors which suffered only moderate disturbance regimes. Edge sectors were also affected by different past disturbance regimes. However, they stood out by having a particularly high frequency of liana tangles and the highest rates of net increase both in density and basal area. A canonical correspondence analysis indicated that the effects of past disturbance regimes and edges were highly related to the species distribution in the area. Pioneer species were concentrated on the edges and more severely disturbed sectors, shade-tolerant species were particularly more abundant on the moderately disturbed sites, while light-demanding ones were more widely distributed. Tree community structure was also undergoing important changes, with less common species enjoying higher recruitment rates and density increase compared to the more common ones. Shade-tolerant species were increasing in relative abundance and had comparatively lower mortality rates and higher recruitment rates. On the other hand, pioneer species were under retreat, with a decreasing relative abundance, higher mortality rates and lower recruitment rates.Nomenclature: Oliveira-Filho et al. 1994a,d.     

Seedling growth strategies in Bauhinia species: comparing lianas and trees

BACKGROUND AND AIMS: Lianas are expected to differ from trees in their growth strategies. As a result these two groups of woody species will have different spatial distributions: lianas are more common in high light environments. This study determines the differences in growth patterns, biomass allocation and leaf traits in five closely related liana and tree species of the genus Bauhinia. METHODS: Seedlings of two light-demanding lianas (Bauhinia tenuiflora and B. claviflora), one shade-tolerant liana (B. aurea), and two light-demanding trees (B. purpurea and B. monandra) were grown in a shadehouse at 25% of full sunlight. A range of physiological, morphological and biomass parameters at the leaf and whole plant level were compared among these five species. KEY RESULTS: The two light-demanding liana species had higher relative growth rate (RGR), allocated more biomass to leaf production [higher leaf mass fraction (LMF) and higher leaf area ratio (LAR)] and stem mass fraction (SMF), and less biomass to the roots [root mass fraction (RMF)] than the two tree species. The shade-tolerant liana had the lowest RGR of all five species, and had a higher RMF, lower SMF and similar LMF than the two light-demanding liana species. The two light-demanding lianas had lower photosynthetic rates per unit area (A(area)) and similar photosynthetic rates per unit mass (A(mass)) than the trees. Across species, RGR was positively related to SLA, but not to LAR and A(area). CONCLUSIONS: It is concluded that the faster growth of light-demanding lianas compared with light-demanding trees is based on morphological parameters (SLA, LMF and LAR), and cannot be attributed to higher photosynthetic rates at the leaf level. The shade-tolerant liana exhibited a slow-growth strategy, compared with the light-demanding species.     

ECOLOGICAL CONSTRAINTS ON WOOD DENSITY IN A TROPICAL MONTANE RAIN FOREST

In a Costa Rican tropical lower montane rain forest the wood densities of canopy tree species are related to the windiness of their preferred habitats, and to their abilities to tolerate shade. Shade-intolerant species tend to have less dense wood than shade-tolerant species from the same habitat. Species characteristic of windy sites tend to have denser wood than species characteristic of sheltered habitats. Stand mean wood density, the average of species’ wood densities weighted by their proportional contributions to stand basal area, increases with exposure to the wind. These trends in wood density should at least partially counteract the damaging effects of wind on exposed sites. Since investment in wood must come at the expense of growth elsewhere, such trends in wood density may help explain the small stature of elfin forest and montane thicket formations in tropical mountains.     

Comparative height crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo

In rainforests, trunk size, strength, crown position, and geometry of a tree affect light interception and the likelihood of mechanical failure. Allometric relationships of tree diameter, wood density, and crown architecture vs. height are described for a diverse range of rainforest trees in Brunei, northern Borneo. The understory species follow a geometric model in their diameter-height relationship (slope, β = 1.08), while the stress-elasticity models prevail (β = 1.27-1.61) for the midcanopy and canopy/emergent species. These relationships changed with ontogeny, especially for the understory species. Within species, the tree stability safety factor (SSF) and relative crown width decreased exponentially with increasing tree height. These trends failed to emerge in across-species comparisons and were reversed at a common (low) height. Across species, the relative crown depth decreased with maximum potential height and was indistinguishable at a common (low) height. Crown architectural traits influence SSF more than structural property of wood density. These findings emphasize the importance of applying a common reference size in comparative studies and suggest that forest trees (especially the understory group) may adapt to low light by having deeper rather than wider crowns due to an efficient distribution and geometry of their foliage.     

The contribution of understory light availability and biotic neighborhood to seedling survival in secondary versus old-growth temperate forest

Seedling survival plays an important role in the maintenance of species diversity and forest dynamics. Although substantial gains have been made in understanding the factors driving patterns of seedling survival in forests, few studies have considered the simultaneous contribution of understory light availability and the local biotic neighborhood to seedling survival in temperate forests at different successional stages. Here, we used generalized linear mixed models to assess the relative importance of understory light availability and biotic neighborhood variables on seedling survival in secondary and old-growth temperate forest in north eastern China at two levels (community and guild). At the community level, biotic neighborhood effects on seedling survival were more important than understory light availability in both forests. In both the old-growth and secondary forests, conspecific basal area had a negative effect on seedling survival, consistent with negative conspecific density dependence. At guild levels, the relative importance of light and biotic neighborhood on seedling survival showed considerable variation among guilds in both forests. Available understory light tended to have positive effects on seedling survival for shrub and light-demanding species in the old-growth forest, but negative effects on survival of shade-tolerant seedlings in the secondary forest. For tree species and shade-tolerant species, the best fit models included neighborhood variables, but that was not the case for shrubs, light-demanding, or mid shade-tolerant species. Overall, our results demonstrate that the relative importance of understory light availability and biotic factors on seedling survival vary with species life-history strategy and forest successional stage.     

辽东山区次生林乔木优势树种幼苗更新动态

以辽东山区次生林4 hm²动态监测样地为平台,2014—2016年连续3年对样地进行乔木优势树种幼苗调查和半球面影像拍摄,分析森林更新过程中林下乔木幼苗和冠层结构的动态,研究幼苗的天然更新规律及其对冠层结构变化(以叶面积指数LAI的变化来表示)的响应.结果表明: 调查期间冠层LAI先减少后增加,且存在显著的年际差异.12种乔木优势树种幼苗对冠层结构变化存在明显的响应,且不同树种幼苗的响应明显不同;中性树种幼苗的相对多度或频度一般会随着冠层LAI的减小而增大;阳性树种幼苗的相对多度或频度一般会随着冠层LAI的增大而减小.2014—2016年乔木优势树种幼苗的多响应置换过程分析表明,优势树种幼苗的数量组成和分布存在显著差异,且年际间的差异逐渐减少.调查期间优势树种幼苗指示种发生明显变化,也反映出其对冠层结构变化的响应.     

Wood density and its radial variation in six canopy tree species differing in shade-tolerance in western Thailand

Background and Aims

Wood density is a key variable for understanding life history strategies in tropical trees. Differences in wood density and its radial variation were related to the shade-tolerance of six canopy tree species in seasonally dry tropical forest in Thailand. In addition, using tree ring measurements, the influence of tree size, age and annual increment on radial density gradients was analysed.

Methods

Wood density was determined from tree cores using X-ray densitometry. X-ray films were digitized and images were measured, resulting in a continuous density profile for each sample. Mixed models were then developed to analyse differences in average wood density and in radial gradients in density among the six tree species, as well as the effects of tree age, size and annual increment on radial increases in Melia azedarach.

Key Results

Average wood density generally reflected differences in shade-tolerance, varying by nearly a factor of two. Radial gradients occurred in all species, ranging from an increase of (approx. 70%) in the shade-intolerant Melia azedarach to a decrease of approx. 13% in the shade-tolerant Neolitsea obtusifolia, but the slopes of radial gradients were generally unrelated to shade-tolerance. For Melia azedarach, radial increases were most-parsimoniously explained by log-transformed tree age and annual increment rather than by tree size.

Conclusions

The results indicate that average wood density generally reflects differences in shade-tolerance in seasonally dry tropical forests; however, inferences based on wood density alone are potentially misleading for species with complex life histories. In addition, the findings suggest that a ‘whole-tree’ view of life history and biomechanics is important for understanding patterns of radial variation in wood density. Finally, accounting for wood density gradients is likely to improve the accuracy of estimates of stem biomass and carbon in tropical trees.Key words: Radial gradients, shade-tolerance, tree biomass estimates, tree rings, tropical trees, wood density     

Changes in spatial distribution during the life history of a tropical tree,Scaphium macropodum (Sterculiaceae) in borneo

We studied the spatial distribution of fruits and plants, mortality and growth rates ofScaphium macropodum (Sterculiaceae) in four 1-ha plots in a tropical rain forest in West Kalimantan, Indonesia. The species is a large deciduous tree and produces wind-dispersed fruits on defoliated twigs. The density of dispersed fruits on the ground decreased with increasing distance from a parent tree. The area under the parent's crown had the highest density of the fruits and the highest mortality of the seedlings immediately after germination. Consequently, the density of the established seedlings peaked 14 m from the tree which is outside its crown. Thick litter mainly from the parent tree seemed to physically prevent the seedlings' root from reaching the soil surface and caused the high mortality. Juvenile and mature trees distributed exclusively, suggesting that regeneration is the most successful outside of the crown of mature trees. Saplings under canopy shade did not grow well.Scaphium macropodum is hypothesized to require a gap for seedling growth and successful regeneration, whereas it can germinate and last under closed canopies as suppressed seedlings or saplings.     

Higher treefall rates on slopes and waterlogged soils result in lower stand biomass and productivity in a tropical rain forest

1.  Relationships between tropical rain forest biomass and environmental factors have been determined at regional scales, e.g. the Amazon Basin, but the reasons for the high variability in forest biomass at local scales are poorly understood. Interactions between topography, soil properties, tree growth and mortality rates, and treefalls are a likely reason for this variability.
2.  We used repeated measurements of permanent plots in lowland rain forest in French Guiana to evaluate these relationships. The plots sampled topographic gradients from hilltops to slopes to bottomlands, with accompanying variation in soil waterlogging along these gradients. Biomass was calculated for >175 tree species in the plots, along with biomass productivity and recruitment rates. Mortality was determined as standing dead and treefalls.
3.  Treefall rates were twice as high in bottomlands as on hilltops, and tree recruitment rates, radial growth rates and the abundance of light-demanding tree species were also higher.
4.  In the bottomlands, the mean wood density was 10% lower than on hilltops, the basal area 29% lower and the height:diameter ratio of trees was lower, collectively resulting in a total woody biomass that was 43% lower in bottomlands than on hilltops.
5.  Biomass productivity was 9% lower in bottomlands than on hilltops, even though soil Olsen P concentrations were higher in bottomlands.
6.   Synthesis . Along a topographic gradient from hilltops to bottomlands there were higher rates of treefall, which decreased the stand basal area and favoured lower allocation to height growth and recruitment of light-demanding species with low wood density. The resultant large variation in tree biomass along the gradient shows the importance of determining site characteristics and including these characteristics when scaling up biomass estimates from stand to local or regional scales.     

Fire affects wood formation dynamics and ecophysiology of Pinus pinaster Aiton growing in a dry Mediterranean area

The increase in frequency and intensity of wildfires is seriously affecting forest ecosystems, especially in drought-prone areas. Trees’ recovery after fire is related to direct tree damage and is influenced by climate conditions, such as warm temperature and water shortage. In this study, we evaluate the post-fire effects on a Pinus pinaster Aiton forest growing in a hot and dry area of the Mediterranean region by comparing burned trees with severe crown reduction against unburned and not-defoliated trees. Inter-annual analyses of dendrochronology and stable isotopes in tree rings were combined with xylogenesis monitoring to investigate the effects of fire on tree growth, ecophysiological processes and wood formation. Tree-ring and isotope data showed a growth reduction and a decrease in photosynthetic activity in the burned trees, compared to control individuals, in the three years after fire. Further, the monitoring of cambial activity demonstrated a negative influence of warm and dry periods on wood formation, low xylem production, a delay in phenology and a reduction in xylem plasticity in burned trees. Our findings suggest that substantial photosynthetic limitations caused by crown defoliation and recurrent drought events could lead to severe growth decrease and reduction of trees ability to regain the pre-disturbance productivity rates.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Neighbour species richness and local structural variability modulate aboveground allocation patterns and crown morphology of individual trees

Local neighbourhood interactions are considered a main driver for biodiversity–productivity relationships in forests. Yet, the structural responses of individual trees in species mixtures and their relation to crown complementarity remain poorly understood. Using a large‐scale forest experiment, we studied the impact of local tree species richness and structural variability on above‐ground wood volume allocation patterns and crown morphology. We applied terrestrial laser scanning to capture the three‐dimensional structure of trees and their temporal dynamics. We found that crown complementarity and crown plasticity increased with species richness. Trees growing in species‐rich neighbourhoods showed enhanced aboveground wood volume both in trunks and branches. Over time, neighbourhood diversity induced shifts in wood volume allocation in favour of branches, in particular for morphologically flexible species. Our results demonstrate that diversity‐mediated shifts in allocation pattern and crown morphology are a fundamental mechanism for crown complementarity and may be an important driver of overyielding.     

Mapping Tropical Forest Trees Using High‐Resolution Aerial Digital Photographs

The spatial arrangement of tree species is a key aspect of community ecology. Because tree species in tropical forests occur at low densities, it is logistically challenging to measure distributions across large areas. In this study, we evaluated the potential use of canopy tree crown maps, derived from high‐resolution aerial digital photographs, as a relatively simple method for measuring large‐scale tree distributions. At Barro Colorado Island, Panama, we used high‐resolution aerial digital photographs (~0.129 m/pixel) to identify tree species and map crown distributions of four target tree species. We determined crown mapping accuracy by comparing aerial and ground‐mapped distributions and tested whether the spatial characteristics of the crown maps reflect those of the ground‐mapped trees. Nearly a quarter (22%) of the common canopy species had sufficiently distinctive crowns to be good candidates for reliable mapping. The errors of commission (crowns misidentified as a target species) were relatively low, but the errors of omission (missed canopy trees of the target species) were high. Only 40 percent of canopy individuals were mapped on the air photographs. Despite failing to accurately predict exact abundances of canopy trees, crown distributions accurately reproduced the clumping patterns and spatial autocorrelation features of three of four tree species and predicted areas of high and low abundance. We discuss a range of ecological and forest management applications for which this method can be useful.     

Influence of canopy tree size on stand basal area may reflect uncoupling of crown expansion and trunk diameter growth

Abstract A recent article by Midgley and colleagues suggests that large trees give rise to inordinately high stand basal areas because they pack canopy space more efficiently than smaller trees. We argue that this phenomenon bears more relation to the fact that diameter increment is not necessarily accompanied by significant crown expansion during all stages of a tree's life. Using data from a canopy tree population in an old‐growth temperate forest, we found that crown area scaled as roughly the 3/5 power of trunk basal area. Rather than reflecting fixed scaling laws, we suggest that this pattern arises because of limited opportunities for crown expansion in dense stands. Old canopy trees in dense stands can thus accumulate large basal areas without occupying a commensurately large canopy area.     

Sun-shade adaptability of the red mangrove,Rhizophora mangle (Rhizophoraceae): Changes through ontogeny at several levels of biological organization

Rhizophora mangle L., the predominant neotropical mangrove species, occupies a gradient from low intertidal swamp margins with high insolation, to shaded sites at highest high water. Across a light gradient, R. mangle shows properties of both “light-demanding” and “shade-tolerant” species, and defies designation according to existing successional paradigms for rain forest trees. The mode and magnitude of its adaptability to light also change through ontogeny as it grows into the canopy. We characterized and compared phenotypic flexibility of R. mangle seedlings, saplings, and tree modules across changing light environments, from the level of leaf anatomy and photosynthesis, through stem and whole-plant architecture. We also examined growth and mortality differences among sun and shade populations of seedlings over 3 yr. Sun and shade seedling populations diverged in terms of four of six leaf anatomy traits (relative thickness of tissue layers and stomatal density), as well as leaf size and shape, specific leaf area (SLA), leaf internode distances, disparity in blade–petiole angles, canopy spread: height ratios, standing leaf numbers, summer (July) photosynthetic light curve shapes, and growth rates. Saplings showed significant sun/shade differences in fewer characters: leaf thickness, SLA, leaf overlap, disparity in bladepetiole angles, standing leaf numbers, stem volume and branching angle (first-order branches only), and summer photosynthesis. In trees, leaf anatomy was insensitive to light environment, but leaf length, width, and SLA, disparities in bladepetiole angles, and summer maximal photosynthetic rates varied among sun and shade leaf populations. Seedling and sapling photosynthetic rates were significantly depressed in winter (December), while photosynthetic rates in tree leaves did not differ in winter and summer. Seasonal and ontogenetic changes in response to light environment are apparent at several levels of biological organization in R. mangle, within constraints of its architectural baiiplan. Such variation has implications for models of stand carbon gain, and suggest that response flexibility may change with plant age.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.