Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees

Water availability is a principal factor limiting the distribution of closed-canopy forest in the seasonal tropics, suggesting that forest tree species may not be well adapted to cope with seasonal drought. We studied 11 congeneric species pairs, each containing one forest and one savanna species, to test the hypothesis that forest trees have a lower capacity to maintain seasonal homeostasis in water relations relative to savanna species. To quantify this, we measured sap flow, leaf water potential (Ψ_L), stomatal conductance (g _s), wood density, and Huber value (sapwood area:leaf area) of the 22 study species. We found significant differences in the water relations of these two species types. Leaf area specific hydraulic conductance of the soil/root/leaf pathway (G _t) was greater for savanna species than forest species. The lower G _t of forest trees resulted in significantly lower Ψ_L and g _s in the late dry season relative to savanna trees. The differences in G _t can be explained by differences in biomass allocation of savanna and forest trees. Savanna species had higher Huber values relative to forest species, conferring greater transport capacity on a leaf area basis. Forest trees have a lower capacity to maintain homeostasis in Ψ_L due to greater allocation to leaf area relative to savanna species. Despite significant differences in water relations, relationships between traits such as wood density and minimum Ψ_L were indistinguishable for the two species groups, indicating that forest and savanna share a common axis of water-use strategies involving multiple traits.     

Productive leaf functional traits of Chinese savanna species

The river valleys in Southwest China are characterized by a dry?Chot climate and relatively rich soils, and host valley-type savannas that are dominated by deciduous species. However, little is known about the ecological adaptations of Chinese savanna plants to the local environments. We hypothesize that Chinese savanna species mainly possess a drought-avoiding strategy by having a deciduous leaf habit and have productive leaf traits. To test this hypothesis, we measured 26 anatomical, morphological, physiological, and chemical traits for 33 woody species from a valley savanna in Southwest China and compared them with the literature data of other dry and wet tropical tree species and a global dataset. We found that Chinese savanna species showed drought avoidance adaptations and exhibited productive leaf traits, such as thin and dense leaves with high ratio of palisade to spongy mesophyll, leaf nutrient concentrations and photosynthetic capacity. Correlations of photosynthetic capacity with N, P, and stomatal conductance across Chinese savanna species were consistent with global patterns reported for seed plants. However, the Chinese savanna species had consistently greater carbon gain at a given specific leaf area, N, P, and stomatal conductance, suggesting higher nutrient- and intrinsic water use efficiencies. These results suggest that paradoxically, Chinese savanna species are adapted to the stressful dry?Chot valley habitat by having productive leaves.     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

ADAPTIVE STRATEGIES OF WOODY SPECIES IN NEOTROPICAL SAVANNAS

1. In this review we discuss the adaptive strategy of woody species in tropical savannas. The low, evergreen, broadleaved, sclerophyllous tree is considered as the typical woody representative in these ecosystems. The discussion is largely based on data concerning four widespread neotropical species: Curatella americana, Byrsonima crassifolia, Bowdichia virgilioides and Casearia sylvestris, together with more fragmentary information available on other American and African savanna woody species. 2. Several types of savanna ecosystems with contrasting ecological features have to be distinguished. Our discussion refers to tree species in one of these types: seasonal savannas, that occur in a tropical wet and dry climate, with constantly high temperature, and on well-drained soils. Most of these savannas are normally burned once a year, towards the end of the dry season. 3. Woody species in seasonal savannas exhibit a quite distinctive morphology. They have low, tortuous trunks, deep and extensive root systems, relatively high R/S and L/S ratios, and large, highly scleromorphic leaves. Their annual phenodynamics appears somewhat puzzling since leaf renewal and expansion, as well as blooming, take place during the dry, apparently less favourable, part of the year. 4. Savanna trees maintain high leaf conductance throughout the year. Some species show a moderate midday decrease in leaf conductance suggesting partial stomatal closure, particularly under very high atmospheric water demands, or in young, developing leaves. However, given the steep vapour density gradient, transpiration flux density tends to be high, especially on clear dry-season days. 5. There is no drastic drop in leaf water potential, as might be expected with a high transpiration rate. The most negative values attained in either season only rarely exceed the leaf turgor loss point. This moderate fall in ψ permits leaf expansion in the dry season. Variable hydraulic resistance contributes to maintain high water flow when steep ψ gradients between soil and leaves are produced. 6. When all factors are taken into account, it seems that savanna trees maintain a favourable water budget all the year, thanks to their extensive root systems that may extract soil water from deep layers, thus allowing the maintenance of a high water flux through the soil-plant-atmosphere system even during the dry season. In this way, these trees have the least seasonal behaviour of all plant components in the seasonal savanna ecosystem. 7. Seasonal savannas occur on extremely poor, nutrient-deficient soils. As an apparent consequence of this nutrient stress, the concentration of nitrogen, phosphorus, potassium, calcium and magnesium in leaves tends to be significantly lower than in forest trees or in drought-deciduous species. 8. Two mechanisms contribute to improve the nutrient economy. One is the reallocation of absorbed nutrients between old and young tissues; the other, the minimization of nutrient losses due to low leaf wettability, low leaf cuticular conductance, and leaf renewal in the rainless season. 9. Savanna trees have low photosynthetic capacity. This is probably due to high internal resistance of leaves induced by their low nitrogen concentration. However, under field conditions rates of CO₂ uptake may be maintained near their optimum because leaf conductance is high all day, and leaf temperature closely matches air temperature, remaining therefore within the optimal range for photosynthesis. 10. All in all, it appears that the physiological behaviour of savanna trees favours a continuously high water flux through the plant that, even if it lowers water-use efficiency, maintains leaf temperatures near optimum for CO₂ uptake, prevents sharp drops in leaf water potential, and induces a high passive uptake of soil nutrients. In this way, the close interaction between water, carbon and nutrient economies leads to the increased fitness of these populations in the seasonal savanna environment.     

Diverging growth and resilience of Pinus tabulaeformis and Pinus massoniana to droughts in north-south transition zone,central China

Climate transition zone is a sensitive area of climate change and ecological transition where forests are vulnerable to climate extremes. Extreme droughts are increasing in frequency and magnitude under climate change, resulting in structure and function changes of forest ecosystems. Here, to analyze climate-growth relationships and quantify tree resilience to extreme droughts, we developed six tree-ring-width chronologies from P. tabulaeformis and P. massoniana sampling sites in Mt. Jigong region, Central China. The results indicated that all chronologies from the two species had good consistency, precipitation in current April and mean temperature in current August or mean minimum temperature from current August to October were the main limiting factors of the two tree species growth, but the responses of P. massoniana ring-width to climatic factors was more complex than that of P. tabulaeformis. The results also showed that tree growth of 1999–2005 was the lowest growing period during 1979–2018, and P. massoniana grew better than P. tabulaeformis before 2005 and vice versa after 2005. Comparing low growth years of trees, we identified to study tree growth resilience. The calculations from 1988, 1999–2005 and 2011 drought years indicated that P. tabulaeformis had more increased resilience to extreme droughts than that of P. massoniana, and the two species had stronger ecological recovery and resilience under global warming and non-extreme drought conditions in the recent 40 years. These results have implications for predicting tree resilience and identifying tree species in heterogeneous forest landscapes vulnerable to future climate change in climatic transition zone.     

Using phenological cameras to track the green up in a cerrado savanna and its on-the-ground validation

Plant phenology has gained new importance in the context of global change research, stimulating the development of novel technologies for phenological observations. Regular digital cameras have been effectively used as three-channel imaging sensors, providing measures of leaf color change or phenological shifts in plants. We monitored a species rich Brazilian cerrado savanna to assess the reliability of digital images to detect leaf-changing patterns. Analysis was conducted by extracting color information from selected parts of the image named regions of interest (ROIs). We aimed to answer the following questions: (i) Do digital cameras capture leaf changes in cerrado savanna vegetation? (ii) Can we detect differences in phenological changes among species crowns and the cerrado community? (iii) Is the greening pattern detected for each species by digital camera validated by our on-the-ground leafing phenology (direct observation of tree leaf changes)? We analyzed daily sequences of five images per hour, taken from 6:00 to 18:00 h, recorded during the cerrado main leaf flushing season. We defined 24 ROIs in the original digital image, including total or partial regions and crowns of six plant species. Our results indicated that: (i) for the studied period, single plant species ROIs were more sensitive to changes in relative green values than the community ROIs, (ii) three leaf strategies could be depicted from the species' ROI patterns of green color change, and (iii) the greening patterns and leaf functional groups were validated by our on-the-ground phenology. We concluded that digital cameras are reliable tools to monitor high diverse tropical seasonal vegetation and it is sensitive to inter-species differences of leafing patterns.     

Assessing the Effects of Multiple Stressors on the Recruitment of Fruit Harvested Trees in a Tropical Dry Forest,Western Ghats,India

The harvest of non-timber forest products (NTFPs), together with other sources of anthropogenic disturbance, impact plant populations greatly. Despite this, conservation research on NTFPs typically focuses on harvest alone, ignoring possible confounding effects of other anthropogenic and ecological factors. Disentangling anthropogenic disturbances is critical in regions such as India’s Western Ghats, a biodiversity hotspot with high human density. Identifying strategies that permit both use and conservation of resources is essential to preserving biodiversity while meeting local needs. We assessed the effects of NTFP harvesting (fruit harvest from canopy and lopping of branches for fruit) in combination with other common anthropogenic disturbances (cattle grazing, fire frequency and distance from village), in order to identify which stressors have greater effects on recruitment of three tropical dry forest fruit tree species. Specifically, we assessed the structure of 54 populations of Phyllanthus emblica, P. indofischeri and Terminalia chebula spread across the Nilgiri Biosphere Reserve, Western Ghats to ask: (1) How are populations recruiting? and (2) What anthropogenic disturbance and environmental factors, specifically forest type and elevation, are the most important predictors of recruitment status? We combined participatory research with an information-theoretic model-averaging approach to determine which factors most affect population structure and recruitment status. Our models illustrate that for T. chebula, high fire frequency and high fruit harvest intensity decreased the proportion of saplings, while lopping branches or stems to obtain fruit increased it. For Phyllanthus spp, recruitment was significantly lower in plots with more frequent fire. Indices of recruitment of both species were significantly higher for plots in more open-canopy environments of savanna woodlands than in dry forests. Our research illustrates an approach for identifying which factors are most important in limiting recruitment of NTFP populations and other plant species that may be in decline, in order to design effective management strategies.     

Gene flow between vicariant tree species: insights into savanna-forest evolutionary relationships

Studying the genetic structure of vicariant species (i.e., closely related species that occupy ecologically distinct yet adjacent habitats) can shed light on the evolution and divergence of species with different ecological requirements. A previous phylogeographic study identified chloroplast DNA haplotype sharing between two vicariant tree species, one from forest (Hymenaea courbaril) and one from savanna (H. stigonocarpa) habitats. These species co-occur in the Brazilian Cerrado, a biome that encompasses forest patches and riverine forests within a savanna matrix. In order to investigate the evolutionary processes involved in the genetic divergence of these trees, we used nuclear microsatellite markers, statistical methods including approximate Bayesian computation (ABC), and leaf morphology to analyze neighboring and distant populations. Bayesian analysis revealed admixture between the species. ABC analysis supported the scenarios with the occurrence of gene flow between species during the Last Glacial Maximum or from the Holocene to the present, when compared to alternative scenarios of no gene flow or constant gene flow since divergence. However, putative hybrids did not exhibit intermediate leaflet morphology, which could be related to distinct selective pressures maintaining species integrity even in the face of gene flow. Our results suggest that despite morphological differences between savanna and forest species, interspecific barriers to gene flow might not be fully developed between vicariant tree species and that interspecific hybridization in trees from Cerrado biome may be an underdiagnosed process.     

Can germination explain the distribution of tree species in a savanna wetland?

The relationship between seed germination and ecological niche is determined by matching germination characteristics with environmental features. In this study, we selected tree species occurring in the largest savanna wetland in South America – the Pantanal. Very few species are endemic or exclusively found in savanna wetlands, and the majority of tree species occurring in the Pantanal are also found in the neighbouring Brazilian Cerrado, a drier vegetation type that does not flood. We investigated the relationship between germination characteristics and occurrence of savanna trees in wetlands testing the hypothesis that such seeds are tolerant to flooding. We also addressed the question of whether seed tolerance to flood, assessed by survival analysis, explains tree distribution along a gradient of flooding intensity. In this flooding gradient, widely distributed species are those that occur in areas subjected to low as well as to high flooding intensity whereas restricted distributed species are those that occur only in areas subjected to a low level of flood. Seeds from tree species occurring in areas subjected to different flooding intensities were collected. Seed tolerance and germination during and after both one and two months of simulated flood were evaluated. Our results show that seeds of most of the studied savanna species tolerated submergence, which helps to explain their occurrence and wide distribution in wetlands. Nevertheless, germination behaviour checked by survival functions (i.e. how germination is distributed over time) partially explained tree species distribution along a flooding gradient. We conclude that seed tolerance to flooding is one of the components of the regeneration niche that determines tree occurrence and distribution at the regional scale, from savanna to wetland, but not at a local scale along a flooding gradient.     

Is the lack of leguminous savanna trees in grasslands of South Africa related to nutritional constraints?

As with many grasslands globally, the Highveld grasslands of South Africa are tree-less, despite having a climate that can support tree growth. Models predict that fire maintains these grasslands. The question arises as to why fire-tolerant savanna trees do not survive in these ecosystems? Savanna tree survival in mesic areas is restricted by demographic bottlenecks, specifically limitations to sapling-escape from fire. It was hypothesised that ancient highly leached soils from grassland areas would prevent saplings from growing fast enough to escape the fire-trap. Growth rates of savanna tree seedlings (Acacia karroo Hayne and Acacia sieberiana Burtt Davy) were measured in a common garden experiment using soils from ten sites collected along a savanna-grassland continuum. Soils from grassland sites were relatively nutrient-poor compared to those from savannas with lower pH, and associated cations. A. sieberiana growth rates responded to pH and these nutrients, whereas A. karroo growth was less strongly linked to specific nutrients. Even so, both species accumulated more biomass when grown in soils from savanna sites compared to grassland sites. An exception was a low elevation low nutrient savanna site that resulted in poor growth, yet sustains high tree biomass in situ. Differences between growth in grassland and savanna soils were small. They may contribute to, but are unlikely to explain, the treeless nature of these grasslands.     

Wildfires affect tree growth and resilience in Northeastern China natural Dahurian larch forests: A dendrochronological perspective

Wildfires are natural and ubiquitous disturbances in boreal forests. Assessing their impacts on tree growth and resilience are particularly important to recognize the adaptation strategies of fire-tolerant species and forest succession in fire conditions. To date, the growth resilience of fire-tolerant species in boreal forests remains largely unquantified, and the drivers of resilience are poorly understood. Here, we measured the tree-ring widths of 99 fire-scarred trees from three sites in natural Dahurian larch (Larix gmelinii) forests. Three moderate-severity fire events in years 1987, 1990, and 2000 occurring at three sites were detected from the records of local forestry bureau. Based on tree-ring width data, we calculated resilience components (i.e., resistance, recovery, resilience and relative resilience) to quantify the responses of growth resilience in the larch trees to fires and analyzed their drivers at three sites. Results indicated that fires significantly reduced the tree growth. With the increasing tree age, these reductions were more pronounced. As for resilience components, our study showed a limited resistance but high recovery of tree growth against fires, and resistance tended to increase northwards but recovery showed the opposite, suggesting a growth-survival tradeoff was exhibited in Dahurian larch trees. With an increasing tree age, regional resistance and resilience showed a decreasing trend, whereas recovery and relative resilience showed an increasing trend. Resilience components were mainly affected by the climatic factors in spring. An increase in moisture availability enhanced resistance, a reduction in diurnal temperature range enhanced recovery, and an increase in mean temperature enhanced resilience and relative resilience. This study reveals that Dahurian larch could be even less favorable when faced with moderate or severe fire events, but a high capacity of recovery enables this species to adapt to the fire-prone condition. Moreover, this work highlights that the resilience of tree growth should be considered to understand tree behaviors and survival strategies of boreal forests following fires across fire-prone regions under future climate warming.     

Discrimination of Deciduous Tree Species from Time Series of Unmanned Aerial System Imagery

Technology advances can revolutionize Precision Forestry by providing accurate and fine forest information at tree level. This paper addresses the question of how and particularly when Unmanned Aerial System (UAS) should be used in order to efficiently discriminate deciduous tree species. The goal of this research is to determine when is the best time window to achieve an optimal species discrimination. A time series of high resolution UAS imagery was collected to cover the growing season from leaf flush to leaf fall. Full benefit was taken of the temporal resolution of UAS acquisition, one of the most promising features of small drones. The disparity in forest tree phenology is at the maximum during early spring and late autumn. But the phenology state that optimized the classification result is the one that minimizes the spectral variation within tree species groups and, at the same time, maximizes the phenologic differences between species. Sunlit tree crowns (5 deciduous species groups) were classified using a Random Forest approach for monotemporal, two-date and three-date combinations. The end of leaf flushing was the most efficient single-date time window. Multitemporal datasets definitely improve the overall classification accuracy. But single-date high resolution orthophotomosaics, acquired on optimal time-windows, result in a very good classification accuracy (overall out of bag error of 16%).     

The role of bud protection and bark density in frost resistance of savanna trees

• Frost events occur with a significant frequency in savannas of the Southern Hemisphere, especially in the Cerrados of Brazil. One of the main strategies to deal with such events is to invest in thick and dense bark, which can insulate internal branch tissues and protect buds, essential to ensure resprouting if frost damage causes plant canopy die‐back. Such strategies may be fundamental to determine the persistence of savanna species in regions where low temperatures and frost events are recurrent.
• Here we describe bud protection and bark strategies of 53 woody species growing in typical savanna vegetation of central Brazil. In addition, we used an experimental approach exposing branches to 0 °C to measure temperature variation in internal branch tissue and test its relationship to bud protection and bark properties.
• We found that the majority of species (69%) showed medium to high bud protection against extreme temperatures; however, the degree of bud protection was not clearly related to bark properties, such as bark thickness and density. Bark density is a fundamental trait in determining protection against low temperatures (0 °C), since species with low bark density showed lower temperature variation in their internal branch tissues, independently of the bud protection degree.
• Bark properties and bud protection are two different (albeit related) strategies for the protection and persistence of savanna trees under extreme environmental temperatures and can explain ecological observations related to savanna tree responses after frost events.

     

A comparison of foliar nutrient concentration in trees from monsoon rainforest and savanna in northern Australia

Abstract Analysis of foliar nutrient concentrations revealed that two mesophyllic monsoon rainforest trees had higher concentrations of most nutrients in leaves than eight savanna species. One of the tested monsoon rainforest species with sclerophyllous leaves had similar nutrient concentrations to the savanna tree species. There were positive or no significant correlations between live foliar nutrient concentration and the percentage of nutrients withdrawn prior to litterfall. The nutrient concentration of litter was similar for most nutrients among tree species of monsoon rainforest or savanna. The results of this study suggest that the relative fertility of surface soils of monsoon rainforest compared with savanna is not determined by contrasting nutrient cycling strategies whereby monsoon rainforest tree species enrich their soils with relatively nutrient rich leaf litter relative to savanna tree species.     

Fire resilience of ant assemblages in long‐unburnt savanna of northern Australia

Abstract Tropical savannas and rainforests contrast in their flammability and the fire resilience of their associated species. While savanna species generally exhibit high resilience to burning, there is much debate about the fire resilience of forest‐associated species, and the persistence of forest patches in a flammable savanna matrix. Where fire has been excluded, savanna tends on a trajectory towards forest, with an increase in forest‐associated plants and animal species. This study tested the idea that given the high proportion of forest‐associated taxa in long‐unburnt savanna, the fauna of these areas would be expected to exhibit less resilience to fire than the fauna in frequently burnt savannas. The study investigated the immediate and short‐term effects on ant assemblages of re‐introducing fire into long‐unburnt savanna in northern Australia. The ant fauna exhibited high resistance to fires, with no significant short‐term change in mean abundance or species richness; instead, seasonality had a far stronger influence on overall ant activity. Fire caused dramatic declines in dominance of the patchily distributed forest‐associated species Oecophylla smaragdina and Papyrius sp., but had no effect on overall dominance by open savanna species of Iridomyrmex. Dominance by Iridomyrmex pallidus declined, but this was compensated for by increases in I. reburrus, while two other species of Iridomyrmex showed no change. This indicates a high level of functional redundancy among dominant species of Iridomyrmex, which universally dominate open savanna communities, but not of dominant forest‐associated species. Overall, our findings demonstrate a high degree of fire‐resilience of the long‐unburnt savanna ant fauna. Despite the occurrence of forest‐associated species, the high proportion of savanna species persisting in this habitat means that long‐unburnt savanna retains the general response characteristics of frequently burnt savanna.     

Response of leaf morphological traits of relict-endemic Symplocos species (S. coccinea and S. speciosa) to elevation and abiotic fluctuations

Abiotic fluctuations in montane ecosystems trigger changes in the hydric functional traits of tree species. These variations are better recognized in tree species inhabiting montane humid ravine slopes with different elevation, as is the case of many areas across the Mexican Neotropical montane forests. Little is known about the response of tree towards elevation gradients and abiotic changes. In this study, we analyzed the leaf morphological variation of two rare and Mexican endemic Symplocos species (S. coccinea and S. speciosa) occurring eastern Mexico on sites with different microclimate and elevation but similar floristic composition. We quantified how the abiotic factors (i.e. canopy openness, soil temperature, soil moisture, and litter depth) and site elevation influence the leaf traits of these tree species. Symplocos coccinea (with toothed leaf margins) is adapted to high humid conditions and high canopy coverage, while S. speciosa (with almost entire leaf margins) is resilient to environments with moisture deficit and high temperatures. Process-based research with fine-spatial scales at montane ecosystems are needed to understand the resilience and morphological variations of montane tree species under climate change worldwide. In this study, we confirmed that the Symplocos leaf morphological traits (i.e. leaf length, leaf width, leaf shape index, leaf base angle and vein density) are strongly influenced by abiotic conditions (i.e. canopy openness, litterfall depth, soil moisture and soil temperature).

     

Structural relationships between form factor,wood density,and biomass in African savanna woodlands

Key message

Variation in tree biomass among African savanna species of equal size is driven by a wide inter-specific variation in wood specific gravity.

Abstract

Tree form and taper is a fundamental component of tree structure and has been used for over a century in forestry to estimate timber yields and in ecological theories of scaling laws. Here, we investigate variation in form factor in the context of biomass in African savannas. Biomass is a fundamental metric of vegetation state, yet in African savannas it remains unclear whether variation in form factor F (taper) or wood specific gravity (G) is a more dominant driver of biomass differences between tree species of equal stem diameter and height. Improving our knowledge of vertical mass distribution in savanna trees provides insight into differences in life strategies, such as tradeoffs between production, disturbance avoidance, and water storage. Here, we destructively harvested 782 stems in a savanna woodland near Kruger National Park, South Africa, and measured whole tree wet mass, wood specific gravity, water content, and form factor. We found that three of four dominant species can vary in mass by over twofold, yet inter-specific variation in taper was low and taper did not vary significantly between common species (P > 0.05) (species-mean form factors ranged from F = 0.57 to 0.77, where cone F =  $0.\bar{3}$ , quadratic paraboloid F = 0.5, cylinder F = 1.0). Comparison of a general biomass allometry model to species-specific models supported the conclusion that the large difference in biomass between species of the same size was explained almost entirely (R ² = 0.97) by including species-mean G with D and H in a general allometric equation, where F was constant. Our results suggest that inter-specific variation in wood density, not form factor, is the primary driver of biomass differences between species of the same size. We also determined that a simple analytical volume-filling model accurately relates wood specific gravity of these species to their water and gas content (R ² = 0.68). These results indicate which species use a wide spectrum of water storage strategies in savanna woodlands, adhering to a trade-off between the benefits of denser wood or increased water storage.     

Termite mounds as browsing hotspots: an exception to the rule

In African savannas, termite mounds usually serve as browsing hotspots for mammals because of their soil fertility. Van der Plas et al., in this issue, describe that browsers avoid the unpalatable, evergreen tree species on mounds of Macrotermes natalensis in a mesic savanna, preferring mainly leguminous species with high leaf N and P concentrations in the matrix. This exception is probably a consequence of the fertile soils of the study area, and highlights the importance of environmental context for assessing ecological interactions.     

Termites facilitate and ungulates limit savanna tree regeneration

Both large herbivores and termites are key functional groups in savanna ecosystems, and in many savanna areas, large termite mounds (termitaria) are associated with distinct woody clusters. Studies on the effect of large mammals on tree regeneration are few, and the results are conflicting. Large herbivores have been found to be important seedling predators in some areas, but facilitate tree regeneration by outcompeting small mammals and reducing grass cover in other areas. Through the use of the experimental fencing of termite mounds and adjacent savanna areas in this study, we investigated how termites and large herbivores influence tree regeneration. Termite mounds had a higher number of seedlings, more species richness, more alpha diversity (OD) and lower evenness (E) than savanna plots. Large herbivores did not significantly affect overall seedling density, species richness, OD or E. Beta diversity was higher in savanna areas than on termitaria, and beta diversity decreased in savanna areas when herbivores were excluded. Herbivore exclusion increased the density of the 12 (40 %) most common seedling species, representing 79 % of all seedlings, and fenced plots had relatively taller seedlings than open plots. Thus, termites were the main determinants of tree regeneration in our study area, but large mammals regulated the most common species. Although our study confirms previous work suggesting that large herbivores affect tree regeneration, we found that termites were an even more important determinant. Termite impacts on tree regeneration deserve increased attention by savanna ecologists.     

When the same is not the same: phenotypic variation reveals different plant ecological strategies within species occurring in distinct Neotropical savanna habitats

Variations in abiotic characteristics such as soil water availability and fertility impose different selective pressures on plant populations. This may produce intraspecific variability in functional traits, even at a fine spatial scale. We investigated whether functional traits related to water-use efficiency, resource-retention strategy, soil nutrient acquisition, and fire tolerance differ in species that occur in two different habitats of Brazilian Cerrado: rocky savannas and savanna woodlands. Rocky savannas occur over sandstone, quartzite outcrops and have shallow nutrient-poor and low-moisture rocky soils, while savanna woodlands occur over well-drained and deep soils with frequent fire regimes. We measured nine functional traits of 40 tree species that occur in both habitats. Rocky savanna individuals exhibited a greater water-use efficiency strategy. The resource-retention strategy in rocky savanna individuals was corroborated by lower adult maximum height. However, despite the lower nutrient availability in rocky savanna soils, we only detected lower leaf phosphorus content in individuals from this habitat. Furthermore, individuals from both habitats had equally thick bark, suggesting that the fire-defense strategy is related to a stable, rather than plastic trait. Overall, our results highlight the central role of contrasting soil water availability patterns in driving phenotypic plasticity within species. We conclude that savanna species are responding to water and nutrient availabilities, via plasticity in traits related to the resource-retention strategy, and preparing for future fires, via uniformly thick bark. Wide plant distribution in contrasting habitats is possible for species that can shift ecological strategies to survive in nutrient- and water-limited habitats such as rocky savannas.