Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Alternative fire resistance strategies in savanna trees

Bark properties (mainly thickness) are usually presented as the main explanation for tree survival in intense fires. Savanna fires are mild, frequent, and supposed to affect tree recruitment rather than adult survival: trunk profile and growth rate of young trees between two successive fires can also affect survival. These factors and fire severity were measured on a sample of 20 trees near the recruitment stage of two savanna species chosen for their contrasted fire resistance strategies (Crossopteryx febrifuga and Piliostigma thonningii). Crossopteryx has a higher intrinsic resistance to fire (bark properties) than Piliostigma: a 20-mm-diameter stem of Crossopteryx survives exposure to 650°C, while Piliostigma needs a diameter of at least 40 mm to survive. Crossopteryx has a thicker trunk than Piliostigma: for two trees of the same height, the basal diameter of Crossopteryx will be 1.6 times greater. Piliostigma grows 2.26 times faster than Crossopteryx between two successive fires. The two species have different fire resistance strategies: one relies on resistance of aboveground structures to fire, while the other relies on its ability to quickly re-build aboveground structures. Crossopteryx is able to recruit in almost any fire conditions while Piliostigma needs locally or temporarily milder fire conditions. In savannas, fire resistance is a complex property which cannot be assessed simply by measuring only one of its components, such as bark thickness. Bark properties, trunk profile and growth rate define strategies of fire resistance. Fire resistance may interact with competition: we suggest that differences in fire resistance strategies have important effects on the structure and dynamics of savanna ecosystems. Received: 16 August 1996 / Accepted: 4 January 1997     

Fire-tolerance mechanisms of common woody plant species in a semiarid savanna in south-western Zimbabwe

The frequency of fire has increased in savannas yet few studies have assessed how plants persist when subjected to long‐term disturbance by fire. We investigated the contributions of bark thickness and resprouting to the persistence of woody plants in two fire trials that were started in 1948 and 1949. The number of resprouts per individual, bark thickness, basal diameter and height of woody plants were measured in unburnt plots and those burnt annually, triennially and quinquennially during the late dry season. Changes in tree density, number of resprouts and individuals in different height classes between 1963 and 2002 were assessed. Bark thickness varied among species and also increased with increases in basal diameter. Generally, plants with thick bark survived fire more than those with thin bark. Resprouting was the major fire survival strategy for most species. The number of resprouts produced per plant ranged from 4 ± 3 (Acacia rehmanniana) to 14 ± 9 (Pseudolachnostylis maprouneifolia). Fire reduced species richness in plots burnt annually and triennially by 47% and 6% respectively. Species richness increased in unburnt plots (5%) and those burnt quinquennially (16%). Most woody species survived fire through a combination of traits.     

The protective role of bark against fire damage: a comparative study on selected introduced and indigenous tree species in the Western Cape,South Africa

The objective of this study was to compare the protective role of bark against fire for three selected indigenous and five exotic species in the Western Cape, South Africa. Bark thickness, compass direction, stem diameter at breast height, bark moisture content and relative height of the sample in the stem were tested for their effect on heat insulation capacity of bark. Trees were felled and heating experiments were conducted at 400 ºC on fresh billets with intact bark. Time to heat the cambium to lethal 60 ºC was determined. Statistical analysis based on correlation, multi-model inference and multiple regression revealed no significant influence of compass direction and diameter at breast height. Heat resistance was mainly determined by bark thickness, to a lesser degree by moisture content. In several species relative height at the stem modulated the bark thickness effect. Higher up the stem bark of the same thickness offered less protection against heat. Significant species-specific differences in heat resistance were apparent in the results, which could not be explained by bark thickness thus indicating further need for research in scrutinising these factors, which might help to explain the relative higher fire tolerance of certain species compared to others.     

Convergence of bark investment according to fire and climate structures ecosystem vulnerability to future change

Fire regimes in savannas and forests are changing over much of the world. Anticipating the impact of these changes requires understanding how plants are adapted to fire. In this study, we test whether fire imposes a broad selective force on a key fire‐tolerance trait, bark thickness, across 572 tree species distributed worldwide. We show that investment in thick bark is a pervasive adaptation in frequently burned areas across savannas and forests in both temperate and tropical regions where surface fires occur. Geographic variability in bark thickness is largely explained by annual burned area and precipitation seasonality. Combining environmental and species distribution data allowed us to assess vulnerability to future climate and fire conditions: tropical rainforests are especially vulnerable, whereas seasonal forests and savannas are more robust. The strong link between fire and bark thickness provides an avenue for assessing the vulnerability of tree communities to fire and demands inclusion in global models.     

Does rapid utilization of elevated nutrient availability allow eucalypts to dominate in the tropical savannas of Australia?

Northern Australia's savannas are among the most fire‐prone biomes on Earth and are dominated by eucalypts (Eucalyptus and Corymbia spp.). It is not clear what processes allow this group to dominate under such extreme fire frequencies and whether a superior ability to compete for nutrients and water might play a role. There is evidence that eucalypts are adapted to frequent fires; juvenile eucalypts escape the fire trap by growing rapidly in height between fires. However, non‐eucalypts are less able to escape the fire trap and tend to have stand structures strongly skewed toward suppressed juveniles. The mechanisms that drive these contrasting fire responses are not well understood. Here, we describe the results of a controlled glasshouse seedling experiment that evaluated the relative importance of nutrient and water availability in determining height growth and biomass growth of two eucalypt and one noneucalypt tree species, common in northern Australian savannas. We demonstrate that growth of eucalypt seedlings is particularly responsive to nutrient addition. Eucalypt seedlings are able to rapidly utilize soil nutrients and accumulate biomass at a much greater rate than noneucalypt seedlings. We suggest that a seasonal spike in nutrient availability creates a nutrient‐rich microsite that allows eucalypt seedlings to rapidly gain height and biomass, increasing their likelihood of establishing successfully and reaching a fire‐resistant size. Our results extend our understanding of how eucalypts dominate northern Australian savannas under extremely high fire frequencies.     

Effects of habitat and growing season fires on resprouting of shrubs in longleaf pine savannas

The effects of habitat and timing of growing season fires on resprouting of shrubs were studied in second-growth longleaf pine savannas of the west Gulf coastal plain in the southeastern United States. Within the headwaters of three different drainages of the Calcasieu River in the Kisatchie National Forest in western Louisiana, replicated permanent transects were established that extended from xeric upland longleaf pine savannas into downslope hydric seepage savannas. All shrubs were mapped and tagged, and numbers of stems were counted prior to any fires. Replicated prescribed fires were set early (June) and late (August) during the 1990 growing season; maximum fire temperatures were measured within both upland and seepage habitats within each transect. Shrubs were relocated; stems were recensused two and twelve months after the fires. At least some shrubs of all species resprouted from underground organs; none regenerated solely from seed banks in the soil. There was no reduction in total numbers of stems one year after fires compared to before fires, either in the upland or in seepage savannas. In addition, there was no reduction in total numbers of stems one year after early or late growing season fires. Fire-related mortality was restricted to small shrubs (< 18 stems) and was not associated with high fire temperatures. The rate of resprouting varied among species and between habitats. Resprouting occurred more rapidly in seepage than upland savannas, but more resprouts were produced in upland than seepage savannas one year after fires. In contrast to other upland species, Vaccinium arboreum and V. elliottii delayed resprouting more than two months following fire. Stems of Rhus copallina and Pyrus arbutifolia, species with long rhizomes, increased more after fires in June than fires in August. We suggest that growing season fires may block further recruitment of shrubs into longleaf pine savannas, but reduction in numbers of large shrubs may require additional management.     

Water relations of host trees and resistance to the phloem-boring beetle Phoracantha semipunctata F. (Coleoptera: Cerambycidae)

Environmental stresses, particularly water deficit, predispose eucalypt trees to attack by the eucalyptus longhorned borer, Phoracantha semipunctata F. (Coleoptera: Cerambycidae). Our experiments with potted eucalypts revealed that reduced tree water potential was associated with lower resistance to colonization by neonate P. semipunctata, but the linear relationship between water potential and colonization success was reversed at higher larval densities. There was no indication that the bark exudate “kino” served to defend trees from borer attack. Larvae were not able to colonize the cambium of eucalypt logs with high bark moisture, and survival was low under high moisture conditions in artificial hosts composed of pure cellulose. In trees and cut logs with moist bark, larvae failed to reach the cambium, feeding instead in poorer-quality tissues just beneath the bark surface. Our findings suggest that variation in resistance of eucalypts to attack by the borer is associated with moisture content of the bark. Received: 2 September 1998 / Accepted: 8 January 1999     

Fire and the demography of camelthorn (Acacia erioloba Meyer) in the southern Kalahari – evidence for a bonfire effect?

Fires in arid environments are rare, so are not deemed as important as in mesic savannas. We investigated mortality and resprouting amongst camelthorn (Acacia erioloba) after two fires (at Vaalbos National Park and Susanna farm) in semi‐arid savanna near Kimberley, South Africa. Resprouting response 18 months after a fire was the greatest amongst <6.5 m high trees; extent of foliage damage by fire and bark thickness were the next best predictors of resprouting vigour amongst that size class. The largest size class (8–12 m height) of A. erioloba suffered the greatest mortality rates (40% and 83% at Vaalbos and Susanna respectively), with damage either severe or minor. We hypothesize that large tree mortality rates are partly attributable to well‐developed assemblages of flammable subcanopy plants producing a bonfire beneath trees. These mortality rates indicate that fire reduces both tree abundance and relative representation of large trees, and although able to resprout, A. erioloba is fire‐sensitive, which may explain its restriction to Kalahari sands where rainfall is less than 900 mm year^?1. Therefore, although relatively infrequent, fires shape Kalahari woodland structure, particularly as A. erioloba is long lived and slow growing. Large trees have been shown to be important to biodiversity in the southern Kalahari, so frequent fires could impact biodiversity.     

Beating the blaze: Fire survival in the fan aloe (Kumara plicatilis), a succulent monocotyledonous tree endemic to the Cape fynbos,South Africa

Fire is central to the ecology of Mediterranean‐type climate ecosystems, but little is known about the fire ecology of succulent plants therein. This study investigated the fire ecology of an arborescent succulent monocot, Kumara plicatilis (L.) G. D. Rowley (Asphodelaceae), a Cape fynbos endemic. Habitat suitability was assessed to determine whether the species tolerates or ‘avoids’ fire, and fire survival traits (bark thickness and tissue water content) were measured. The population size structure and density of three K. plicatilis populations were assessed after natural fires, and resprouting potential was investigated. Kumara plicatilis adopts a dual fire survival strategy, occupying rocky sites to ‘avoid’ fire and possessing morphological features that afford fire tolerance, e.g. well‐protected apical meristems and thick corky bark. Bark thickness of burned individuals in situ was similar to unburned plants, suggesting that K. plicatilis bark provides effective insulation against fire. Mortality rates were 64%, 40% and 11%, and decreased as rock cover at the population level increased. All three populations showed reduced plant density post‐fire, with greater density reductions associated with lower rock cover. Small plants appear most vulnerable to fire damage due to lower absolute bark thickness and plant heights within the flame zone. Kumara plicatilis is an apical sprouter, recovering after fire or mechanical stem damage by onward growth from surviving stem apices, rather than resprouting. Post‐fire population recovery therefore likely depends on inter‐fire recruitment.     

Responses of root‐crown bearing shrubs to differences in fire regimes in Pinus palustris (longleaf pine) savannas: Exploring old‐growth questions in second‐growth systems

Question: What are the effects of fire season and intensity on resprouting of different root‐crown bearing shrub species in second‐growth Pinus palustris (longleaf pine) savannas? Location: northern Florida and eastern Louisiana, USA. Methods: In Florida, quadrats were burned biennially either during the dormant season or the growing season. In Louisiana, we applied intensity treatments to quadrats by manipulating ground‐cover fuels, just prior to biennial growing season fires. Maximum fire temperatures were measured, and stem densities were censused before and after fires in both regions. Results: After dormant season fires in Florida, stem densities were seven times greater than initial levels for Hypericum spp. In contrast, growing season fires reduced densities of H. brachyphyllum by 65%, but did not change densities of H. microsepalum. Only resprouting of H. microsepalum decreased with increased fire intensity. In Louisiana, fire intensity influenced Ilex vomitoria, but not Quercus spp. Following fires, stem densities oil. vomitoria were five times greater in fuel removal than fuel addition areas. Conclusions: Past use of dormant season fires likely contributed to increased abundances of some species of root‐crown bearing shrubs observed today in old‐growth savannas. Reintroduction of growing season fires will be effective in maintaining or decreasing stem densities, depending on species and fuel type. Genet mortality and stem density reductions appear most likely in areas at localized scales where tree falls and needle coverage create hotspots in Pinus palustris savannas.     

Eucalyptus regnans (Myrtaceae): A fire-sensitive eucalypt with a resprouter epicormic structure

Determining the location of buds and bud-forming meristems and hence the level of protection from heat is essential to understanding plant response to fire. Most eucalypts resprout readily from the stem (epicormic resprouting) and the base after felling or high intensity fire. In contrast, Eucalyptus regnans is one of the few eastern Australian fire-sensitive, obligate seeder eucalypts. Some authors have suggested that the relatively weak epicormic resprouting is due to a lack of bud-forming structures. Epicormic strands from the bark and outer xylem of three very large trees and two saplings were examined anatomically. Epicormic bud-forming structures were found in all samples examined. The bud-forming capacity consisted of narrow, radially elongated strips of cells of meristematic appearance. These strips were continuous from the outermost secondary xylem through to the outer bark. Bark was relatively thick at the base of the large trees, but remarkably thin above this basal skirt. Eucalyptus regnans was found to possess the apparently fire-adapted epicormic strands previously described in other eucalypts, thus showing its fire-adapted lineage. However, this fire-sensitive species apparently directs much of its resources to rapid height-growth rates in younger trees, rather than to vegetative fire survival.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

Numerous predictions indicate rising CO₂ will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought‐fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire‐driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2‐million km² Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait‐based differences in fire tolerance is critical for determining the climate‐carbon‐fire feedback in tropical savanna and forest biomes.     

Thinner bark increases sensitivity of wetter Amazonian tropical forests to fire

Understory fires represent an accelerating threat to Amazonian tropical forests and can, during drought, affect larger areas than deforestation itself. These fires kill trees at rates varying from < 10 to c. 90% depending on fire intensity, forest disturbance history and tree functional traits. Here, we examine variation in bark thickness across the Amazon. Bark can protect trees from fires, but it is often assumed to be consistently thin across tropical forests. Here, we show that investment in bark varies, with thicker bark in dry forests and thinner in wetter forests. We also show that thinner bark translated into higher fire‐driven tree mortality in wetter forests, with between 0.67 and 5.86 gigatonnes CO₂ lost in Amazon understory fires between 2001 and 2010. Trait‐enabled global vegetation models that explicitly include variation in bark thickness are likely to improve the predictions of fire effects on carbon cycling in tropical forests.     

Ants,fire, and bark traits affect how African savanna trees recover following damage

Bark damage resulting from elephant feeding is common in African savanna trees with subsequent interactions with fire, insects, and other pathogens often resulting in tree mortality. Yet, surprisingly little is known about how savanna trees respond to bark damage. We addressed this by investigating how the inner bark of marula (Sclerocarya birrea), a widespread tree species favoured by elephants, recovers after bark damage. We used a long‐term fire experiment in the Kruger National Park to measure bark recovery with and without fire. At 24 months post‐damage, mean wound closure was 98, 92, and 72%, respectively, in annual and biennial burns and fire‐exclusion treatments. Fire exclusion resulted in higher rates of ant colonization of bark wounds, and such ant colonization resulted in significantly lower bark recovery. We also investigated how ten common savanna tree species respond to bark damage and tested for relationships between bark damage, bark recovery, and bark traits while accounting for phylogeny. We found phylogenetic signal in bark dry matter content, bark N and bark P, but not in bark thickness. Bark recovery and damage was highest in species which had thick moist inner bark and low wood densities (Anacardiaceae), intermediate in species which had moderate inner bark thickness and wood densities (Fabaceae) and lowest in species which had thin inner bark and high wood densities (Combretaceae). Elephants prefer species with thick, moist inner bark, traits that also appear to result in faster recovery rates.     

Oak Bark Allometry and Fire Survival Strategies in the Chihuahuan Desert Sky Islands,Texas, USA

Trees may survive fire through persistence of above or below ground structures. Investment in bark aids in above-ground survival while investment in carbohydrate storage aids in recovery through resprouting and is especially important following above-ground tissue loss. We investigated bark allocation and carbohydrate investment in eight common oak (Quercus) species of Sky Island mountain ranges in west Texas. We hypothesized that relative investment in bark and carbohydrates changes with tree age and with fire regime: We predicted delayed investment in bark (positive allometry) and early investment in carbohydrates (negative allometry) under lower frequency, high severity fire regimes found in wetter microclimates. Common oaks of the Texas Trans-Pecos region (Quercus emoryi, Q. gambelii, Q. gravesii, Q. grisea, Q. hypoleucoides, Q. muehlenbergii, and Q. pungens) were sampled in three mountain ranges with historically mixed fire regimes: the Chisos Mountains, the Davis Mountains and the Guadalupe Mountains. Bark thickness was measured on individuals representing the full span of sizes found. Carbohydrate concentration in taproots was measured after initial leaf flush. Bark thickness was compared to bole diameter and allometries were analyzed using major axis regression on log-transformed measurements. We found that bark allocation strategies varied among species that can co-occur but have different habitat preferences. Investment patterns in bark were related to soil moisture preference and drought tolerance and, by proxy, to expected fire regime. Dry site species had shallower allometries with allometric coefficients ranging from less than one (negative allometry) to near one (isometric investment). Wet site species, on the other hand, had larger allometric coefficients, indicating delayed investment to defense. Contrary to our expectation, root carbohydrate concentrations were similar across all species and sizes, suggesting that any differences in below ground storage are likely to be in total volume of storage tissue rather than in carbohydrate concentration.     

Post-fire regeneration strategies and tree bark resistance to heating in frequently burning tropical savanna woodlands and grasslands in Ethiopia

Regeneration mechanisms of vegetation and the role of tree bark resistance to frequent fire were studied in savanna woodlands and grasslands in Gambella, Western Ethiopia. Data were collected from four sites, each with three replicate plots. The variation between sites in species composition and biomass correlated with the differences in fire intensity. Foliar cover was recorded for individual plant species regenerating by sprouting from older parts of plants that had survived fire or by seedlings; records were made during the dry season and at the beginning of the wet season. Data on bark thickness and tree diameters of 12 dominant tree species were also recorded. Both facultative and obligate sprouters significantly contributed to post‐fire recovery, comprising 98.5 % of total vegetation cover. The contribution of seedlings to cover and abundance immediately following fire was negligible, but seedling density increased in the beginning of the rainy season, 4 to 5 months after fire. The importance of the sprouting and seeding strategies varied between the different plant growth forms. The highest contribution to cover and frequency was made by the most abundant grass species, which reproduced in both ways. Facultative sprouters made up 67.3 % of the vegetation cover, out of which 54 % consisted of grasses. Broad‐leaved herbs and trees/shrubs regenerating mainly by sprouting made up 31.3 % of the vegetation cover. Adaptations to fire in tree species seemed to include the development of a thick bark, once the tree has passed seedling stages. Tree bark thickness and tree diameter at breast height were strongly correlated with the time taken for cambium to reach an assumed lethal temperature of 60°C when exposed to fire, which indicated that mature trees with thick barks might resist stronger fire better than, e.g., small or young trees and trees with thin bark. However, for a given bark thickness the cambium resistance to heat varied three‐fold among species. Hence, site differences in fire intensity seemed to influence the distribution of trees depending on their bark characteristics and resistance to fire.     

Effects of phorophyte determinants on lichen abundance in the <Emphasis Type="Italic">cerrado</Emphasis> of central Brazil

Previous studies have indicated that epiphytic lichens can be good bioindicators of fire history in tropical savannas. A Lichen Fire History (LFH) Key was developed to assess fire history in areas of cerrado (savanna) in central Brazil. However, the effectiveness and reliability of the LFH Key is much influenced by other lichen determinants. The aim of this study, therefore, was to investigate some of these factors in more detail, thereby allowing better estimates of fire history using the LFH Key. Fieldwork was carried out at the Reserva Ecológica do IBGE, 33 km outside Brasilia D.F., in plots of cerrado denso of varying fire history. Vegetation sampling took place in 20 × 20 m-quadrats within which measurements of the lichen abundance, scorch and phorophyte characteristics, including height, girth and tortuosity, were recorded for all the phorophytes encountered. Bark samples were collected from common cerrado phorophytes and tested for pH, conductivity, moisture content and absorbing capacity, texture and nutrient content. The results show that the greater the impact of fire, the lower the influence of other factors, such as bark characteristics, on the lichens. The strongest determinants of lichens in areas subjected to rare fires or protected from fire are bark aluminium content, bark pH, and microclimatic factors. Using the information gathered from the study, phorophyte species are grouped in terms of their reliability for use in the LFH Key. This study highlights the range of factors which can affect lichen abundance in the tropics, and the relationships between them.     

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.