Foliage color contrasts and adaptive fruit color variation in a bird-dispersed plant community

The color of vertebrate-dispersed fruits has been a source of inquiry for over 150 years, yet the ecological and evolutionary processes responsible for fruit color diversity remain elusive. We tested the hypothesis that fruit color varies temporally, to maximize conspicuousness against seasonal changes in foliage coloration, in a bird-dispersed plant community in western North America. Field observations showed that while red fruits predominate during summer periods of green foliage coloration, black fruits are produced during flushes of red-orange foliage coloration in autumn. Although two species did not conform to this pattern, one produced its own contrasting background color, via colored bracts. We conducted experimental manipulations of both fruit color and the color of artificial backgrounds to test whether both factors had a synergistic effect on fruit removal rates. Interactions between fruit and background color explained most of the variation in experimental fruit removal rates. Although red fruits were removed rapidly on green backgrounds, preference for black fruits on red-orange backgrounds was less pronounced. Consequently, the temporal pattern in fruit color appears to result from elevated fruit conspicuousness against seasonal changes in foliage coloration. Support for this hypothesis suggests a temporal connection between fruit color diversity, foliage color contrasts and avian color preferences.     

Reproductive phenology, life-forms, and habitats of the Venezuelan Central Plain

Reproductive phenology of 171 plant species belonging to 57 families of angiosperms was studied according to life-forms in four habitat types in a savanna-forest mosaic on the Venezuelan Central Plain. Flowering, unripe fruit, and mature fruit patterns were affected significantly according to life-forms and habitats respectively. Production of flowers, unripe fruits, and mature fruits showed marked seasonality for all habitats except for the forest. Flowering peaked during the rainy season, and fruiting peaked toward the end of the rainy season. The savanna and the disturbed area had similar proportions of species that flowered over the year. The percentage of species with unripe fruits produced throughout the year was more seasonal for the disturbed area than for the other habitats. Mature fruit patterns showed an increase during the late rainy season for the ecotone and savanna. A large number of herbaceous (annual and perennial) and liana species flowered during the wet season, and a smaller fraction flowered during the dry season; and trees, shrubs, and epiphytes increased flowering activity during the dry season. Unripe fruit patterns were similar to those of flowering for all life-forms, however, tree species were less seasonal. Mature fruit production by shrubs peaked in the period of maximum rainfall, while the peak for perennial herbs was in the late rainy season and the peak for annual herbs was during the transition between the rainy season and the dry season. The largest proportion of tree and liana species with ripe fruits occurred during the dry season. Differences among phenological patterns in habitats were caused mainly by life-forms and promote a wider distribution of reproductive events in habitats and overall community in the Venezuelan Central Plain.     

Why are fruits colorful? The relative importance of achromatic and chromatic contrasts for detection by birds

The colors of fruits and flowers are traditionally viewed as an adaptation to increase the detectability of plant organs to animal vectors. The detectability of visual signals increases with increasing contrasts between target and background. Contrasts consist of a chromatic aspect (color) and an achromatic aspect (light intensity), which are perceived separately by animals. To evaluate the relative importance of fruits’ chromatic and achromatic contrasts for the detection by avian fruit consumers we conducted an experiment with artificial fruits of four different colors in a tropical forest. We displayed the fruits against two different backgrounds, an artificial background and a natural one, because they differed in achromatic properties. We found no effect of the type of background on fruit detection rates. Detection rates differed for the four fruit colors. The probability of detection was explained by the chromatic contrast between fruits and their background, not by the achromatic contrasts. We suggest that birds attend primarily to chromatic contrast probably because these are more reliably detected under variable light conditions. Consistent with this hypothesis, we found habitat-specific differences in the conspicuousness of natural fruit colors in the study area. Fruits of understory species that are subjected to the variable light conditions within a forest displayed higher chromatic contrasts than species growing in the open restinga forest with constant bright illumination. There was no such difference for achromatic contrasts. In sum, we suggest that fruit colors differ between habitats because fruit colors that have strong chromatic contrasts against background can increase plants’ reproductive success, particularly under variable light conditions.     

The role of chromatic and achromatic signals for fruit detection by birds

Fruit color changes during ripening are typically viewed asan adaptation to increase signal efficacy to seed dispersers.Plants can increase signal efficacy by enhancing chromatic (wavelengthrelated) and/or achromatic (intensity related) contrasts betweenfruit and background. To assess how these contrasts determinethe detectability of fruit signals, we conducted 2 experimentswith free-flying crows (Corvus ossifragus) under seminaturalconditions in a 2025 m² aviary. Crows searched first for artificialred and black fruits and detected red fruits from a larger distance.Because artificial red fruits had higher chromatic and lowerachromatic contrasts against foliage than artificial black fruits,crows apparently prioritized chromatic contrasts. Thus, thecommon change in fruit color from red to black during ripeningdoes not increase signal efficacy to crows. In a second trial,crows searched for UV-reflecting and black blueberries (Vaccinummyrtillus) against backgrounds of foliage and sand. Againstfoliage, UV-reflecting berries had higher chromatic and achromaticcontrasts than black berries, and crows detected them from alarger distance. Against sand, UV-reflecting berries had lowachromatic contrasts and black berries low chromatic contrasts.Crows detected both fruit types equally, suggesting that theyused chromatic contrasts to detect UV-reflecting berries andachromatic contrasts to detect black berries. Birds prioritizedchromatic contrasts when searching for artificial red fruitsin foliage but not when searching for blueberries on sand. Wesuggest that the relative importance of chromatic and achromaticcontrasts is contingent on the chromatic and achromatic varianceof the background. Models of signal perception can be improvedby incorporating background-specific effects.     

Diet of the maned wolf, Chrysocyon brachyurus, in central Brazil

Analysis of 105 scats of the maned wolf ( Chrysocyon brachyurus ) collected in central Brazil yielded 304 occurrences of food items of which fruits of Solanum lycocarpum , rodents, and birds accounted for 61.5%. By analysis of minimum number of individual animals, rodents and birds were 75% of a total of 156 prey. On the other hand, fruits of S. lycocarpum and armadillos ( Dasypus spp.) were the bulk of the total estimated biomass consumption, yielding 63.7% in a total of 73.5 kg. Rodents were mostly captured during the dry season, while miscellaneous fruits were consumed mostly in the wet season. The consumption of S. lycocarpum fruits and armadillos was aseasonal. Small rodents were taken in about the ranks of abundance in the study area, but S. lycocarpum fruits were actively searched by maned wolves, for its occurrence is limited to secondary savanna. Prey and fruits typical of savanna ('cerrado') and grassland ('campo') were the bulk of species, occurrences, prey numbers, and biomass consumed. These findings reinforce the importance of conservation of cerrado and campo in central Brazil. Despite maned wolf being a vulnerable species, its feeding habits are opportunistic, and the main reasons for its decline are likely to be habitat destruction and human disturbance.     

Impact on fruit removal and seed predation of a secondary metabolite, emodin, in Rhamnus alaternus fruit pulp

There are two contradictory approaches to explaining the presence of secondary metabolites in ripe fruits. One holds that they evolved toward enhancing dispersal success (adaptive approach); the other claims that they evolved primarily to deter herbivores from eating leaves and seeds and that their presence in ripe fruits is a byproduct of that function (non‐adaptive approach). We tested the validity of three hypotheses of the adaptive approach that explain the presence of secondary metabolites in ripe fruits. We explored the current function of a secondary metabolite, emodin, in Mediterranean buckthorn (Rhamnus alaternus, Rhamnaceae) fruits by relating intraspecific variation and seasonal patterns of concentration to fruit removal and seed damage and by conducting feeding trials with captive birds presented artificial fruits that varied in emodin concentration. The concentration of emodin in the pulp of 10–13 Rhamnus plants from the same population was determined by HPLC every month during two fruiting seasons. Fruit removal by birds and seed predation by invertebrates and microbes were determined for the same plants. Emodin concentration rose during the first stages of ripening, reaching a peak before the fruits were ripe, and then decreased to a minimum when the fruits were ripe. No significant correlation between emodin concentration and ripe fruit removal rate among trees was observed in the first year, whereas in the second year the correlation was positive and significant. Thus, the impact of emodin on fruit selection varied between years, suggesting that emodin concentration does not solely govern fruit selection. A significant negative correlation was found in the first year between emodin concentration and seed predation during the first fruiting month. The yellow‐vented bulbul (Pycnonotus xanthopygos), a seed dispersing bird, distinguished between artificial foods that differed in emodin concentration (control, 0.001% and 0.002%), always preferring the lower concentration. In contrast, house sparrows, (Passer domesticus), a seed predator, did not detect such differences in emodin concentration but did distinguish between control foods and food with 0.005% and 0.001% emodin. We suggest that emodin has an ecological role, preventing seed predation by invertebrates and microbes without decreasing fruit removal by avian dispersers.     

Soil-vegetation relationships in hyperseasonal cerrado, seasonal cerrado, and wet grassland in Emas National Park (central Brazil)

In South America, the largest savanna region is the Brazilian cerrado, in which there are few areas that become waterlogged in the rainy season. However, we found a small cerrado area in which the soil is poorly drained and becomes waterlogged at the end of the rainy season, allowing the appearance of a hyperseasonal cerrado. We investigated the soil–vegetation relationships in three vegetation forms: hyperseasonal cerrado, seasonal cerrado, and wet grassland. We collected vegetation and soil samples in these three vegetation forms and submitted obtained data to a canonical correspondence analysis. Our results showed a distinction among hyperseasonal cerrado, seasonal cerrado and wet grassland, which presented different floristic compositions and species abundances. The edaphic variables best related to the hyperseasonal and seasonal cerrados were sand, base saturation, pH, and magnesium. The wet grassland was related to higher concentrations of clay, organic matter, aluminium saturation, aluminium, phosphorus, and potassium. Although it is not possible to infer causal relationships based on our results, we hypothesize that the duration of waterlogging in the hyperseasonal cerrado may not be long enough to alter most of its soil characteristics, such as organic matter, phosphorus, and potassium, but may be long enough to alter some, such as pH and base saturation, as the soils under both cerrados were more similar to one another than to the soil under the wet grassland. Since waterlogging may alter soil characteristics and since these characteristics were enough to explain the plant community variation, we may conclude that water excess—permanent or seasonal—is one of the main factors to distinguish the three vegetation forms, which presented different floristic compositions and species abundances.     

Structure and biomass along an Acacia zanzibarica woodland–savanna gradient in a former ranching area in coastal Tanzania

Questions: What factors influence the density, size and growth form of trees in secondary Acacia zanzibarica woodlands on a former humid savanna rangeland? How does tree density relate to variation in tree foliage and spines, and woody and grass biomass? Location: Tropical coastal Tanzania (former Mkwaja Ranch, now in Saadani National Park). Methods: We surveyed 97 circular plots (4‐m radius) representing a gradient from open savanna to dense woodland. Within each plot, we measured all trees and estimated the biomass of spines. Foliage biomass of tree and grass layers was estimated on three occasions, twice during the wet season and once in the dry season. Soil samples were taken from each plot and analysed for texture and nutrient content. Interrelationships among various variables were investigated using linear multiple regression and mixed effects models. Results: Tree densities were highest on more nutrient‐rich, heavy soils. Spinescence was highest on trees in open savanna. Biomass of tree foliage in the wet season was best explained by numbers of ant nests and tree live‐wood ratio. Foliage biomass in the dry season was less than half that in the wet season and best predicted by grass biomass. Variables related to biomass of the grass layer were strongly influenced by fire; living grass biomass also decreased with increasing tree density. Conclusions: A. zanzibarica is a tree with a high water demand, and the association with heavy soils is probably due to greater availability of water on these sites. Establishment of A. zanzibarica woodlands significantly reduced grazing resources at Mkwaja Ranch. Under post‐ranching conditions, however, fires and soil conditions predominate. The woodlands may, therefore, represent a transient state of woody density in a still resilient humid savanna.     

Soil chemical factors and grassland species density in Emas National Park (central Brazil)

Studies of grasslands on specific soil types suggest that different nutrients can limit biomass production and, hence, species composition and number. The Brazilian cerrado is the major savanna region in America and once covered about 2 million km(2), mainly in the Brazilian Central Plateau, under seasonal climate, with wet summer and dry winter. In view of the importance of soil chemical factors in the distribution of the vegetation forms within the Cerrado domain and which may influence the number of species, we analyzed some soil characteristics in three herbaceous vegetation forms -- hyperseasonal cerrado, seasonal cerrado, and wet grassland -- in Emas National Park, a core cerrado site, to investigate the relationship between number of species and soil characteristics. We collected vegetation and soil samples in these three vegetation forms and submitted the obtained data to multiple linear regression. We found out that aluminum and pH were the best predictors of species density, the former positively related to species density and the latter negatively related. Since the predictable variation in species density is important in determining areas of conservation, we can postulate that these two soil factors are indicators of high species density areas in tropical grasslands, which could be used in selecting priority sites for conservation.     

Latitudinal and elevational variation in fruiting phenology among western European bird-dispersed plants

I try to test the prediction that bird-dispersed plants should produce fruits when fruit-eating birds are most abundant by reviewing some phenological data of fleshy fruit production in western Europe The prediction that fruit ripening dates in populations of the same species should occur later at lower latitudes and elevations, to coincide with the maximum abundance of fruit-eating birds, is not supported by the data The patterns of seasonal variation in the total number and biomass of fruits, but not in the proportion of species in fruit, in communities at different latitudes and elevations do coincide with patterns of seasonal abundances of avian frugivores 1 suggest that this coincidence is due to the greater relative abundance (and contribution to total fruit production) in each locality of species that fruit at times of the year when birds are most abundant These species may have achieved a demographic advantage by getting more seeds dispersed than species that ripen fruits in other seasons     

Intra‐ and inter‐individual variation show distinct trends as drivers of seasonal changes in the resource use of a neotropical marsupial

Individual specialization can influence important ecological and evolutionary traits and both inter‐ and intra‐individual variation in resource use can drive niche shifts in natural populations. We evaluated the predominance of these two factors for determining seasonal differences in the trophic niche of the didelphid marsupial Gracilinanus agilis (Burmeister, 1854) in the highly seasonal Brazilian savanna. In the three sampled sites, the population of G. agilis increased its dietary niche width in the warm–wet season, when food resources are more abundant, and there were no differences between sexes and no interaction between season and sex. However, the evaluation of intra‐individual variation indicated that females reduce the number of items consumed during the warm–wet season, whereas males show no seasonal differences. Inter‐individual variation nonetheless followed the overall population pattern because both sexes increased their spread with respect to food‐item consumption in the warm–wet season. Additionally, we found positive relationships between body length and diet only in the warm–wet season, when larger animals fed more on invertebrates and less on fruits than the small ones. Our results show a previously unknown pattern for mammals, in which the trophic niche is wider during the high‐resource season as a result of inter‐individual variation along the body‐size axis. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 737–747.     

Predispersal reproductive biology of female Osyris quadripartita (Santalaceae), a hemiparasitic dioecious shrub of Mediterranean scrublands

Female Osyris quadripartita plants exhibit uninterrupted reproductive activity throughout the year, due to the long duration of successive stages in the cycle and marked within-crop developmental asynchrony. Cycles corresponding to the flowering seasons of consecutive years overlap in each individual. Flowering takes place in spring, and fruits develop in the dry summer season and ripen at any time of the year. Variation in flowering time explains a negligible proportion of variation in ripening time. The greatest reproductive losses are incurred in the phase extending from closed flowers through unripe fruits, mostly due to ovary abortion. Only 30% of closed flowers eventually reach this latter stage. In contrast, 75% of unripe fruits complete their development, with subsequent dispersal of seeds. The probability of the setting of ripe fruit steadily decreases from early to late season flowers, due to increased ovary abortion rates. Resource limitation in the dry summer season seems responsible for this pattern of selective fruit maturation.     

Transfer of Mass and Energy in a Cassava (Manihot esculenta Crantz cv. Cubana) Community 1. Microclimate and Water Vapour Exchange in a Savanna Climate

The energy balance approach was used to evaluate the transferprocesses which occur between a cassava community, growing duringa savanna wet season, and its environment. Crop performanceunder these conditions was also analysed. During this season, the soil profile remained near saturation,hence net radiation absorbed by the cassava crop during typicallow radiation days, was mainly dissipated as latent heat (78–86per cent); a process which depends on the growth stage and onthe foliage cover of the canopy. However, the seasonal rootenvironment, with a sufficient water supply, contrasted withthe extremely variable shoot environment, because the dailyflux density of short wave radiation input reached values similarto those reported for typical dry season days. On these days,cassava responded to environmental stress mainly by a reductionin transpiration. At the end of the wet season soil water content markedly decreased,an additional stress factor on the roots, which produced a notablereduction of cassava leaf area index. Therefore, the moderatechanges in energy partitioning determined by morphological andphysiological adaptive responses of cassava, produce a hindrancein the equilibrium between water vapour transfer and the surroundingsavanna environment. Manihot esculenta Crantz, cassava, microclimate, energy balance, heat fluxes     

Reproduction and growth in three species of West African fruit bats

This study outlines the reproduction periods of the African fruit bats Epomops buettikoferi, Epomophorus garnbianus , and Micropteropus pusillus at two West African savanna sites: a wet southern Guinea savanna and a drier southern Sudanese savanna. At both sites the two annual birth periods were timed such that both lactation by only one weaning period coincided with the rainy season peaks in fruit availability. On this basis we propose that lactation rather than weaning was the important determinant of the timing of reproduction. There was no evidence of cycling of testes size in males corresponding with the seasonal mating periods. E. buettikoferi and M. pusillus females mated at six months and gave birth at the age of 12 months. Males of the two species reached puberty by 11 months and seven months, respectively. Growth rates did not differ between cohorts growing through the wet and dry season and E. buettikoferi and M. pusillus grew at rates of 276 mg/day and 116 mg/day, respectively.     

Taxonomic distinctness and diversity of a hyperseasonal savanna in central Brazil

Savannas are characterized by a sharp seasonality, in which the water shortage defines the community functioning. Hyperseasonal savannas, however, experience additionally waterlogging in the rainy season. Since waterlogging may cause local extinctions of intolerant species, we asked whether waterlogging constricts the phylogenetic structure of a hyperseasonal savanna. We studied a hyperseasonal cerrado, comparing it with a nearby seasonal cerrado, never waterlogged, in Emas National Park, central Brazil. In each vegetation form, we sampled all vascular plants by placing fifty 1-m² quadrats in five surveys. We compared the phylogenetic structure of both vegetation forms, calculating their taxonomic distinctness, taxonomic diversity, expected taxonomic distinctness, and species, genus, and family similarities. The taxonomic distinctness of both cerrados was similar and the values of similarities were high, but taxonomic diversity and expected taxonomic distinctness were lower in the hyperseasonal cerrado than in the seasonal one. Assuming that phenotypic attraction is the major process organizing local communities, the waterlogging in hyperseasonal cerrado assembles phylogenetically unrelated species that have converged on similar habitat use. As a consequence, the habitat use of hyperseasonal cerrado species is a trait widespread in the phylogeny of seasonal cerrado. Waterlogging constrains the phylogenetic structure of the hyperseasonal cerrado, especially by reducing species diversity. In more ecological terms, we can only fully assess the phylogenetic structure of a community if we consider the species abundance.     

Nutritional ecology of a browsing ruminant, the kudu (Tragelaphus strepsiceros), through the seasonal cycle

The activities and food selection of four hand-reared kudus were recorded in a large fenced enclosure containing natural savanna vegetation in the Nylsvley Nature Reserve, South Africa. Leaves of selected species were analysed chemically for crude protein, fibre constituents, phosphorus, condensed tannin and total polyphenols. Available protein and metabolizable energy were estimated allowing for potential antinutritional effects of tannins.
Leaves of palatable deciduous woody plants and herbaceous forbs formed the main dietary constituents during the late wet season. Foliage from palatable evergreens and robust forbs were added to the diet during the dry season. Towards the end of the dry season unpalatable species of evergreens were eaten. At the start of the growing season new leaves of otherwise unpalatable woody species formed the staple food source, together with fruits of Strychnos spp. Correspondingly, protein and digestible dry matter concentrations in the diet declined to reach a low at the end of the dry season.
Total daily food intake increased to compensate for reduced dietary quality during the dry season, until little edible foliage remained. While the estimated daily intake of protein remained well above maintenance requirements, the estimated metabolizable energy intake fell below requirements during the late dry season. Phosphorus intake may have been submaintenance in the dry season. Nutritional balance was dependent on the availability of particular vegetation components to serve as nutritional stepping stones during crucial times of the year. These included forbs during the late wet season, palatable evergreens in the dry season, and Strychnos fruits plus early-flushing woody plants during the dry season-wet season transition.     

Relationships between leaf deciduousness and flowering traits of woody species in the Brazilian neotropical savanna

The relationships between foliage permanence and flowering throughout the year were analyzed in 92 woody species of Cerrado vegetation categorized as either deciduous (DE), semideciduous (SD) or evergreen (EV). Flowering of DE, SD and EV species was investigated via three variables, measured over the course of the year: flowering duration (FLD), calculated as the number of months in flower in each species; flowering distribution (FDI), calculated as the number of species in flower per month; and flowering peak (FPE), defined as the four consecutive months yielding the highest number of species in flower. The months with the highest numbers of species in flower were October (52 species), September (50) and August (49). These months correspond to the period of transition from the dry season to the wet season. In the majority of species studied, seasonal climatic factors were strong enough to induce fruit formation in the dry season and seed dispersal in the following wet season, when sufficient water was available to support germination and plantlet growth. However, significant differences in FLD, FDI and FPE were found among the leaf phenological groups. High FLD in EV species is likely favored by the continuous input of resources from the year-round foliage. In contrast, DE species employ reserves of carbon, water and nutrients to form new leaves and flowers on a crown free of foliage at the end of the dry season. In DE species, their low FLD may reduce the impact of flowering on reserve consumption. SD species showed an intermediate level of foliage persistence, resulting in intermediate FLD values. In addition, SD species exhibited a different pattern of flowering distribution from those of DE and EV species. Many SD species have two flowering periods per year. The first period occurs when the crowns are full of leaves, in the middle of the dry season in June, similar to EV species. The second occurs when only half of the original foliage area is present, near the peak of the dry season in September, similar to DE species. Therefore, despite a strong influence of seasonal climatic conditions on the flowering behavior of DE, SD and EV woody species of Cerrado vegetation, these leaf phenological groups differ significantly in FLD, FDI and FPE.     

Effects of seasonal precipitation change on soil respiration processes in a seasonally dry tropical forest

Precipitation is projected to change intensity and seasonal regime under current global projections. However, little is known about how seasonal precipitation changes will affect soil respiration, especially in seasonally dry tropical forests. In a seasonally dry tropical forest in South China, we conducted a precipitation manipulation experiment to simulate a delayed wet season (DW) and a wetter wet season (WW) over a three‐year period. In DW, we reduced 60% throughfall in April and May to delay the onset of the wet season and irrigated the same amount water into the plots in October and November to extend the end of the wet season. In WW, we irrigated 25% annual precipitation into plots in July and August. A control treatment (CT) receiving ambient precipitation was also established. Compared with CT, DW significantly increased soil moisture by 54% during October to November, and by 30% during December to April. The treatment of WW did not significantly affect monthly measured soil moisture. In 2015, DW significantly increased leaf area index and soil microbial biomass but decreased fine root biomass. In contrast, WW significantly decreased fine root biomass and forest floor litter stocks. Soil respiration was not affected by DW, which could be attributed to the increased microbial biomass offsetting the decrease in fine root biomass. In contrast, WW significantly increased soil respiration from 3.40 to 3.90 μmol m^?2 s^?1 in the third year, mainly due to the increased litter decomposition and soil pH (from 4.48 to 4.68). The present study suggests that both a delayed wet season and a wetter wet season will have significant impacts on soil respiration‐associated ecosystem components. However, the ecosystem components can respond in different directions to the same change in precipitation, which ultimately affected soil respiration.     

Influence of rainfall and grazing on herbage dynamics in a seasonally dry tropical savanna

Species composition and herbage dynamics in relation to rainfall variability and cattle grazing were studied in permanently protected, grazed, and temporarily fenced treatments on three sites in a seasonally dry tropical savanna. Permanently protected sites, established between 1979 and 1984, were 55–79% similar with each other in species composition, and 14–25% similar with grazed sites during the period 1986–1988. Similarity among grazed sites was only 36–43%. Number of species was greater in the grazed treatment than in the permanently protected treatment. The percentages of annual grasses and non-leguminous forbs were greater in grazed savanna than in permanently protected savanna. Species diversity was higher in grazed savanna than in the corresponding permanently protected savanna. Species the two annual cycles studied, peak live shoot biomass was 614 g m^-2 in permanently protected savanna, 109 g m^-2 in grazed savanna, and 724 g m^-2 in temporarily fenced savanna. Live shoot biomass in temporarily fenced savanna was 18 to 44% greater than in permanently protected savanna. Peak canopy biomass ranged from 342 to 700 g m^-2 in permanently protected savanna. It was related with total rainy season rainfall, and was particularly sensitive to late rainy season rainfall. On the other hand, peak canopy biomass in grazed savanna ranged from 59 to 169 g m^-2 and was related to grazing intensity rather than either total rainy season rainfall or late rainy season rainfall. Coefficient of variation of green biomass in permanently protected savanna was related with rainfall variability indicating it to be a pulsed system which responds quickly to rainfall events. Biomass of woody species ranged from 2466 to 5298 g m^-2 in permanently protected savanna and from 744 to 1433 g m^–2 in the grazed savanna. Green foliage biomass was 3.7 to 6.4% of the woody biomass in permanently protected and 5.6 to 5.9% in grazed savanna, and supplements substantially the fodder resource during the dry periods of the year.     

Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna

In frequently burnt mesic savannas, trees can get trapped into a cycle of surviving fire-induced stem death (i.e. topkill) by resprouting, only to be topkilled again a year or two later. The ability of savanna saplings to resprout repeatedly after fire is a key component of recent models of tree–grass coexistence in savannas. This study investigated the carbon allocation and biomass partitioning patterns that enable a dominant savanna tree, Acacia karroo, to survive frequent and repeated topkill. Root starch depletion and replenishment, foliage recovery and photosynthesis of burnt and unburnt plants were compared over the first year after a burn. The concentration of starch in the roots of the burnt plants (0.08 ± 0.01 g g⁻¹) was half that of the unburnt plant (0.16 ± 0.01 g g⁻¹) at the end of the first growing season after topkill. However, root starch reserves of the burnt plants were replenished over the dry season and matched that of unburnt plants within 1 year after topkill. The leaf area of resprouting plants recovered to match that of unburnt plants within 4–5 months after topkill. Shoot growth of resprouting plants was restricted to the first few months of the wet season, whereas photosynthetic rates remained high into the dry season, allowing replenishment of root starch reserves. ¹⁴C labeling showed that reserves were initially utilized for shoot growth after topkill. The rapid foliage recovery and the replenishment of reserves within a single year after topkill implies that A. karroo is well adapted to survive recurrent topkill and is poised to take advantage of unusually long fire-free intervals to grow into adults. This paper provides some of the first empirical evidence to explain how savanna trees in frequently burnt savannas are able to withstand frequent burning as juveniles and survive to become adults. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.