Influence of heat shock on seed germination of plants from regularly burnt savanna woodlands and grasslands in Ethiopia

The effect of heat shock on the germination of seeds of 21 plant speciesfrom fire-prone wooded savanna ecosystems in western Ethiopia was analysed inorder to examine the possible implications of fire upon plant regenerationafterthis disturbance. Seeds were subjected to 6 different heat intensities (20, 60,90, 120, 150 and 200°C) for 1 or 5 minutes, in ordertosimulate the situation in the upper soil layers or on the soil surface duringfires. Germination tests were carried out in pots in a greenhouse over 20weeks.After 9 weeks no more seedlings emerged. There was wide interspecific variationin the responses of seeds to the different treatments. In all species,germination was significantly affected by the temperature treatment level.Shortexposure of seeds to high temperatures generally stimulated germination whereasprolonged exposure reduced seed germination. However, some species eventolerated 5 min treatment at 200°C. Seedheat resistance was positively correlated with seed length and mass among thespecies. Hence, production of large seeds with protective tissues promotessurvival in fire-prone savanna areas. Also, the seeds of some species showedboth a low and a high temperature optimum which ensures that at least someseedsgerminate in the absence of fire, but also that viable seeds still remain ifsubsequent late fires kill emerging seedlings. Frequent and light burning inwooded savanna grasslands seems to stimulate and enhance germination of most ofthe studied plant species.     

Nutrient resorption in tropical savanna forests and woodlands of central Brazil

Nutrient limitation in Brazilian savanna (known as cerrado) presumably causes trees to maximize nutrient resorption from senesced leaves to reduce their dependence on nutrient availability. To assess patterns between nutrient resorption and soil fertility, we measured community-level nitrogen (N), phosphorus (P), and potassium (K) concentrations in mature and senesced leaves and soil fertility in the upper 50 cm soil layer in structurally diverse cerrado ecosystems in the Cuiaba Basin (CB) and Pantanal (PAN) of Mato Grosso, Brazil. Foliar nutrient concentration data were used to estimate resorption efficiency and proficiency, and correlation was used to determine whether resorption efficiency and proficiency varied across soil fertility gradients. We found that N and P resorption proficiency (NRP and PRP, respectively) and P resorption efficiency (PRE) increased significantly as total soil N (NRP) and extractable P (PRP and PRE) declined. In contrast, K resorption efficiency (KRE) declined as soil sand content and bulk density increased, which was likely due to a reduction in soil water-holding capacity. Leaf N/P ratios indicate potential N limitation and/or N + P co-limitation for ecosystems in the PAN and P limitation and/or N + P co-limitation for ecosystems in the CB, while trends in leaf N/K ratios indicate possible K or K + P co-limitation for the CB only. Our results illustrate that cerrado forests and woodlands have highly variable nutrient resorption capacities that vary predictably across soil fertility or textural gradients and indicate that cerrado communities have flexible nutrient resorption that can reduce their dependence on soil nutrient availability.     

Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia

We investigated the fire resistance conferred by bark of seven common tree species in north Australian tropical savannas. We estimated bark thermal conductance and examined the relative importance of bark thickness, density and moisture content for protecting the cambium from lethal fire temperatures. Eucalypt and non-eucalypt species were contrasted, including the fire-sensitive conifer Callitris intratropica. Cambial temperature responses to bark surface heating were measured using a modified wick-fire technique, which simulated a heat pulse comparable to surface fires of moderate intensity. Bark thickness was a better predictor of resistance to cambial injury from fires than either bark moisture or density, accounting for 68% of the deviance in maximum temperature of the cambium. The duration of heating required to kill the cambium of a tree (τ_c) was directly proportional to bark thickness squared. Although species did not differ significantly in their bark thermal conductance (k), the thinner barked eucalypts nevertheless achieved similar or only slightly lower levels of fire resistance than much thicker barked non-eucalypts. Bark thickness alone cannot account for the latter and we suggest that lower bark moisture content among the eucalypts also contributes to their apparent fire resistance. Unique eucalypt meristem anatomy and epicormic structures, combined with their bark traits, probably facilitate resprouting after fire and ensure the dominance of eucalypts in fire-prone savannas. This study emphasises the need to take into account both the thermal properties of bark and the mechanism of bud protection in characterising the resprouting ability of savanna trees.     

Soil nitrogen and carbon heterogeneity in woodlands and grasslands: contrasts between temperate and tropical regions

Aim  Soil resource heterogeneity is linked to several ecological processes including invasion of woody species into grasslands. Studies from the temperate zone have demonstrated greater soil heterogeneity beneath woody vegetation than beneath grasslands. Woody species have a more widespread and coarser root system than herbaceous species, and may have a competitive advantage in relatively heterogeneous soils. We tested the global generality of greater soil heterogeneity beneath woody vegetation.
Location  Global.
Methods  We used data from published literature for soil nitrogen and carbon heterogeneity from paired woodland and grassland sites around the world.
Results  Woodland and grassland soil heterogeneities from paired observations were strongly correlated. There was, however, significant geographical variability in the relationship. Soils were more heterogeneous in woodlands than grasslands in temperate areas, but the opposite was true for tropical habitats. Grassland soils were more heterogeneous at lower than higher latitudes. Woodland soil heterogeneity did not vary with latitude.
Main conclusions  The previously described high soil heterogeneity in woody vegetation compared to grasslands holds only for temperate regions. Consequently, the relationship between soil resource heterogeneity and vegetation type is dependent on the study region. Macroecological studies should test the generality of relationships between soil and vegetation at the global scale.     

Distinct roles of savanna and forest tree species in regeneration under fire suppression in a Brazilian savanna

Questions: Has fire suppression relaxed barriers to the exchange of species between savanna and forest? Do all species or a subset of species participate in this exchange? Would current vegetation structure persist if fire suppression were to cease? Location: A gallery forest edge in the Cerrado region of central Brazil that burned only once in the past 35 years. Methods: Density of tree seedlings, saplings and adults, leaf area index (LAI), tree basal area and diameter were surveyed in 12, 10 m × 70 m transects centred on and perpendicular to the forest–savanna boundary. Community composition was assessed using non‐metric multi‐dimensional scaling (NMDS). Results: Basal area and LAI declined substantially from forest to savanna, with an associated shift in species composition. Savanna tree species were nearly absent in the forest, but accounted for the majority of stems in the savanna. In contrast, forest species comprised 14% of adults and more than one‐third of juveniles in the savanna. Despite the high diversity of trees (85 species) in the forest, five species play a particularly large role in this initial phase of forest expansion. Reintroduction of fire, however, would result in widespread topkill of juveniles and the majority of adult forest trees, thereby interrupting the succession towards forest. Conclusions: After 35 years during which the site burned only once, the savanna still remains dominated by savanna species. Nevertheless, the dominance of forest juveniles in border and savanna tree communities suggests that with a continued policy of fire suppression, the forest will continue to expand.     

Seasonal gas exchange and water relations in juveniles of two evergreen Neotropical savanna tree species with contrasting regeneration strategies

Colonization dynamics of woody species into grasslands in Neotropical savannas are determined by two main factors: plant-available moisture and fire. Considering seasonality of precipitation and high fire frequency in these ecosystems, vegetative reproduction has been suggested as the main regeneration strategy in woody species. This study examined seasonal variations in water relations and photosynthesis in juveniles of two tree species with contrasting regeneration strategies: Palicourea rigida (sexual reproduction) and Casearia sylvestris (asexual reproduction). The studied species showed similar transpiration rates to deep-rooted adult evergreen tree species during the rainy period, suggesting little water availability limitations on surface soil layers. P. rigida juveniles significantly decreased their leaf water potentials from wet to dry seasons. In C. sylvestris resprouts, there were no seasonal differences in their predawn water potentials and gas exchange parameters, indicating a water deficit avoidance characteristic derived from their connections to deep-rooted adult counterparts allowing access to moist soil at depth even during the drought period. P. rigida rely on strict control of water losses and turgor maintenance through elastic cell walls during the dry season. The iso-hydric behavior of gas exchange and most water relations parameters in C. sylvestris enable turgor maintenance during the dry season which also gives the possibility to achieve foliar expansion under water-stressed conditions for shallow-rooted plants. Nevertheless, in absence of water deficits, P. rigida had the advantage to be physiologically independent individuals, showing an equal or even superior photosynthetic performance that eventually could be translated into a more favorable whole-plant carbon balance and higher growth rates in wet habitats.     

Double jeopardy: bark harvest for malaria treatment and poor regeneration threaten tree population in a tropical forest of Uganda

Several forest plants known to supply medicine are under pressure worldwide. We carried out a study of four tree species (Warburgia ugandensis Sprague, Fleroya rubrostipulata (K.Schum.) Y.F.Deng, Syzygium guineense DC. and Zanthoxylum chalybeum Engl.) that are highly demanded for malaria treatment. The study was undertaken between 2006 and 2009 in the Sango Bay Forest Reserve, southern Uganda. The aim was to determine the conservation status of trees targeted for malaria treatment. We assessed the level of damage inflicted on trees during harvesting of medicinal parts and determined the population density of target species in the forest. We used 95 plots established along nineteen transects. Survival of the most preferred species, F. rubrostipulata, is of particular concern as its population suffered from a combination of extensive damage due to poor methods of harvesting and poor regeneration. The density of trees with diameter ≥5 cm differed between species and sites, demonstrating different recruitment and survival strategies. Management of Protected Areas should augment strategies to monitor the legal and illegal harvest of medicinal plants, by adopting low impact harvesting methods, and designating the temporal and spatial patterns of harvesting. This might reduce tree damage and mortality.     

Natural regeneration and population dynamics of the tree Afzelia quanzensis in woodlands in Southern Africa

The logging of tree species of high commercial value is increasing throughout the African continent, yet the ecology of these species is generally poorly known. We studied the regeneration pattern and size class distribution of Afzelia quanzensis populations in northern South Africa over a 5‐year period. Recruitment was low as the annual seedling mortality was >65%. Seedlings were located under the canopy and were affected by drought and browsing. The adults were scattered or were in a clump‐dispersed pattern, which would result in higher recruitment of offspring near parents. Individuals of 0–10 cm diameter at breast height (DBH) were few, while there were 32 trees ha^?1 at >10 cm DBH with an annual mortality of 0.8%. Annual diameter increments varied between 0.06 and 0.28 cm. It appears that the transition from the sapling into the juvenile stage could be a bottleneck in the regeneration of the species. A longer study, including more rainfall cycles, may reveal other patterns as dry and wet years have different impacts on dynamics.     

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.

     

Diversity of subaerial algae and cyanobacteria on tree bark in tropical mountain habitats

We report the species composition of subaerial epixylic algae and cyanobacteria from a South-East Asian mountain rainforest locality in Cibodas, West Java. Green algae (Trebouxiophyceae, Chlorophyceae, Trentepohliales) were dominant and Cyanobacteria were the second most frequent group. We specifically concentrated on the comparison of species composition of closed primary forest and open antropogenic spaces. Trentepohliales and Cyanobacteria dominated in open spaces with higher light intensities, whereas closed forest localities were dominated by trebouxiophycean coccal green algae. There was a significantly higher algal diversity in open spaces than in closed forest samples indicating the limiting effect of light on subaerial algal communities of closed tropical forests. A number of isolated strains and morphotypes probably represent undescribed taxa. Presented at the International Symposium Biology and Taxonomy of Green Algae V, Smolenice, June 26–29, 2007, Slovakia.     

Differences in volatile terpene composition between the bark and leaves of tropical tree species

Volatile terpenes are among the most diverse class of defensive compounds in plants, and they are implicated in both direct and indirect defense against herbivores. In terpenes, both the quantity and the diversity of compounds appear to increase the efficiency of defense as a diverse blend of compounds provides a more efficient protection against a broader range of herbivores and limits the chances that an enemy evolves resistance. Theory predicts that plant defensive compounds should be allocated differentially among tissues according to the value of the tissue, its cost of construction and the herbivore pressure on it. We collected volatile terpenes from bark and leaves of 178 individual tree belonging to 55 angiosperm species in French Guiana and compare the kind, amount, and diversity of compounds in these tissues. We hypothesized that in woody plants, the outermost part of the trunk should hold a more diverse blend of volatile terpenes. Additionally, as herbivore communities associated with the leaves is different to the one associated with the bark, we also hypothesized that terpene blends should be distinct in the bark vs. the leaves of a given species. We found that the mixture of volatile terpenes released by bark is different and more diverse than that released by leaves, both in monoterpenes and sesquiterpenes. This supports our hypothesis and further suggests that the emission of terpenes by the bark should be more important for trunk defense than previously thought.     

Seasonal,interannual and decadal drivers of tree and grass productivity in an Australian tropical savanna

Tree–grass savannas are a widespread biome and are highly valued for their ecosystem services. There is a need to understand the long‐term dynamics and meteorological drivers of both tree and grass productivity separately in order to successfully manage savannas in the future. This study investigated the interannual variability (IAV) of tree and grass gross primary productivity (GPP) by combining a long‐term (15 year) eddy covariance flux record and model estimates of tree and grass GPP inferred from satellite remote sensing. On a seasonal basis, the primary drivers of tree and grass GPP were solar radiation in the wet season and soil moisture in the dry season. On an interannual basis, soil water availability had a positive effect on tree GPP and a negative effect on grass GPP. No linear trend in the tree–grass GPP ratio was observed over the 15‐year study period. However, the tree–grass GPP ratio was correlated with the modes of climate variability, namely the Southern Oscillation Index. This study has provided insight into the long‐term contributions of trees and grasses to savanna productivity, along with their respective meteorological determinants of IAV.     

Cambial growth dynamics and climatic control of different tree life forms in tropical mountain forest in Ethiopia

Munessa Forest is a mountain forest in south-eastern Ethiopia experiencing seasonal rainfall variation. We investigated seasonal cambial activity and dormancy from increment rates of four different tree species belonging to varying life forms, namely, evergreen native conifer (Podocarpus falcatus), evergreen introduced conifer (Pinus patula), evergreen broadleaved tree (Prunus africana) and deciduous broadleaved tree (Celtis africana). Measurements of stem radius fluctuations were registered with the help of high-resolution electronic dendrometers. Daily amplitudes of stem diameter variations and daily and monthly net growth rates were determined and related to climatic variables measured at local climate stations. Thin sections of wood collected with a microcorer every 3–6 weeks allowed a visual control of newly formed wood cells during consecutive time intervals. Lack of water availability during the long dry season induced cambial dormancy of 5–7 months depending on life forms. After the onset of the short rainy season, stem swelling started quite synchronously with a variation of only single days in all studied species. Evergreen tree species were able to initiate wood formation during the short rainy season, whereas growth in the deciduous broadleaved species started in the long rainy season. The acquired data provide a basis for delineating the species-specific growth boundaries and the duration of the cambial growing season.     

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.     

Shade tolerance and regeneration of some tree species of a tropical rain forest in Western Kenya

Shadetolerance and regeneration of some heavily exploited timber species(Uvariopsis congensis, Antiaris toxicaria, Funtumia africana,Prunus africana and Aningeria altissima)were studied in two adjacent sites in Kakamega tropical rain forest. The twosites have been subjected to different logging intensities. Within one of thesites, relatively intact areas were compared with adjoining more disturbedones.Seedling presence or absence in mature forest phase, sapling proportions in gapand the mature phase and diameter distributions were used to classify thespecies into shade tolerance groups. Results indicate that except forUvariopsis, the species are non-pioneer lightdemanders and could fit into three shade tolerance groups: A groupresembling pioneers but with seedling in shade, a mid-tolerant group andone showing a high shade tolerance. Amount of regeneration and disturbance wererelated: regeneration being higher in the more disturbed site except forUvariopsis. The intersite differences were mirrored whendisturbed and adjoining more intact areas within one of the sites werecompared.Species more abundant in the disturbed site also generally scored low in shadetolerance. However, the mid-tolerant Funtumia andnot Prunus or Aningeria that hadlowershade tolerance scores, dominated regeneration in the disturbed forest parts.The shade tolerance data suggest that Uvariopsis, Funtumiaand Antiaris could be amenable to natural forestmanagement. For Prunus and Aningeria,artificial regeneration in buffer plantations around the natural forest may beabetter option.     

Interactive response of herbivores,soils and vegetation to annual burning in a South African savanna

Abstract Despite decades of research, the primary factors determining savanna structure remain elusive – a conundrum termed ‘the savanna problem’. After 47 years of annual burning in Terminalia woodland and Acacia/Combretum savanna on sandy, granite‐derived soils in the southern Kruger National Park, South Africa, a dense cover of trees and shrubs persists on some burnt plots and is largely absent from others. We postulated that intense browsing pressure by antelope and other herbivores prevents recruitment of trees in burnt plots and that herbivores concentrate on plots that are richest in nutrients. Herbivore abundance did not show a relationship with soil macronutrients and we consequently investigated micronutrient status. The reduction in tree cover as a result of annual burning was positively correlated with mass of herbivores (15–1500 kg) (r ² = 0.61, n = 8). This index of herbivore abundance was in turn positively correlated with total Zn (r ² = 0.64, n = 8). Other indices of herbivore abundance showed significant relationships with total clay content and total Mn. We suggest that herbivores concentrate on sites with greater clay content (possibly due to a greater availability of micronutrients), and that tree cover can remain relatively dense under a regime of annual burning if browsing pressure is not intense. The long‐term burn experiments in the Kruger National Park savanna provide a platform for unravelling the savanna problem. Determining possible interactions between soil properties, herbivory and fire is a step in this direction.