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.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.     

Environment and landscape rather than planting design are the drivers of success in long‐term restoration of riparian Atlantic forest

Question

Identifying the factors that lead to the success of restoration projects has been a major challenge in ecological restoration. Here we ask which factors, aside from time since restoration began, drive the recovery of tree biomass, density and richness of the understorey in riparian forests undergoing restoration.

Location

Semideciduous Atlantic Forest with tropical climate and deep, fertile soils, southeast Brazil.

Methods

We sampled tree basal area (DBH ≥ 5 cm), density and richness of the understorey (DBH < 5 cm) in 26 riparian forests undergoing restoration (a chronosequence spanning 4–53 years). We assessed the following variables as possible factors, besides time, influencing community attributes: (1) planting design: density and richness of seedlings planted; (2) landscape features: proximity index measuring forest cover within a 1.5‐km radius, distance and size of the nearest forest remnant; and (3) environmental factors: invasive grasses, soil fertility, drought, average annual precipitation and proportion of fine particles in the soil. We performed correlation analyses including predictor and response variables, followed by stepwise backward regression (AIC), multiple and simple linear regressions, to investigate the relationships between those factors and the community attributes.

Results

Tree basal area was primarily influenced by the proportion of small particles in the soil (+) and secondarily by rainfall (?). Understorey richness was influenced by the combination of size (+) and distance (?) of the nearest patch, rainfall (?) and soil fertility (+). Understorey density was primarily influenced by the size of the nearest forest remnant (+) and secondarily by invasive grasses (?). No influence of density or richness of the seedlings planted was observed.

Conclusion

Environmental factors and landscape configuration drive the recovery of tree biomass, density and richness in communities undergoing restoration. The most relevant ecological filters influencing restoration success are availability of soil water and nutrients and the distance and size of the nearest remnant of native vegetation. The expected influence of richness and density of seedlings planted, considered for many years as important drivers of forest restoration success, was not confirmed in this study.     

Grass Physiognomic Trait Variation in African Herbaceous Biomes

African herbaceous biomes will likely face drastic changes in the near future, due to climate change and pressures from increasing human activities. However, these biomes have been simulated only by dynamic global vegetation models and failing to include the diversity of C₄ grasses has limited the accuracy of these models. Characterizing the floristic and physiognomic diversity of these herbaceous biomes would enhance the parameterization of C₄ grass plant functional types, thereby improving simulations. To this end, we used lowermost and uppermost values of three grass physiognomic traits (culm height, leaf length, and leaf width) available in most floras to identify several grass physiognomic groups that form the grass cover in Senegal. We then checked the capacity of these groups to discriminate herbaceous biomes and mean annual precipitation domains. Specifically, we assessed whether these groups were sufficiently generic and robust to be applied to neighboring (Chad) and distant (South Africa) phytogeographic areas. The proportions of two physiognomic groups, defined by their lowermost limits, delineate steppe from savanna and forest biomes in Senegal, and nama‐karoo, savanna, and grassland biomes in South Africa. Proportions of these two physiognomic groups additionally delineate the mean annual precipitation domains <600 mm and >600 mm in Senegal, Chad, and South Africa, as well as the <250 mm and >1000 mm domains in South Africa. These findings should help to identify and parameterize new C₄ grass plant functional types in vegetation models applied to West and South Africa.     

Restoration of C4 grasses with seasonal fires in a C3/C4 grassland invaded by Prosopis glandulosa,a fire‐resistant shrub

Questions: Can prescribed fire restore C₄ perennial grasses in grassland ecosystems that have become dominated by fire‐resistant C₃ shrubs (Prosopis glandulosa) and C₃ grasses? Do fires in different seasons alter the direction of change in grass composition? Location: Texas, USA. Methods: We quantified short‐ and long‐term (12 yr post‐fire) herbaceous functional group cover and diversity responses to replicated seasonal fire treatments: (1) repeated‐winter fires (three in 5 yr), (2) repeated‐summer fires (two in 3 yr), and (3) alternate‐season fires (two winter and one summer in 4 yr), compared with a no‐fire control. Results: Summer fires were more intense than winter fires, but all fire treatments temporarily decreased Prosopis and C₃ annual grass cover. The alternate‐season fire treatment caused a long‐term increase in C₄ mid‐grass cover and functional group diversity. The repeated‐summer fire treatment increased C₄ short‐grass cover but also caused a long‐term increase in bare ground. The repeated winter fire treatment had no long‐term effects on perennial grass cover. Mesquite post‐fire regrowth had increasingly negative impacts on herbaceous cover in all fire treatments. Conclusions: Summer fire was necessary to shift herbaceous composition toward C₄ mid‐grasses. However, the repeated‐summer fire treatment may have been too extreme and caused post‐fire herbaceous composition to “over‐shift” toward less productive C₄ short‐grasses rather than C₄ mid‐grasses. This study provides some of the first long‐term data showing a possible benefit of mixing seasonal fires (i.e., the alternate‐season fire treatment) in a prescribed burning management plan to restore C₄ mid‐grass cover and enhance overall herbaceous diversity.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Root dynamics influence tree–grass coexistence in an Australian savanna

Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C₃ (trees) and C₄ (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.     

A grass–fire cycle eliminates an obligate‐seeding tree in a tropical savanna

A grass–fire cycle in Australian tropical savannas has been postulated as driving the regional decline of the obligate-seeding conifer Callitris intratropica and other fire-sensitive components of the regional flora and fauna, due to proliferation of flammable native grasses. We tested the hypothesis that a high-biomass invasive savanna grass drives a positive feedback process where intense fires destroy fire-sensitive trees, and the reduction in canopy cover facilitates further invasion by grass. We undertook an observational and experimental study using, as a model system, a plantation of C. intratropica that has been invaded by an African grass, gamba (Andropogon gayanus) in the Northern Territory, Australia. We found that high grass biomass was associated with reduced canopy cover and restriction of foliage to the upper canopy of surviving stems, and mortality of adult trees was very high (>50%) even in areas with low fuel loads (1 t·ha⁻¹). Experimental fires, with fuel loads >10 t·ha⁻¹, typical of the grass-invasion front, caused significant mortality due to complete crown scorch. Lower fuel loads cause reduced canopy cover through defoliation of the lower canopy. These results help explain how increases in grass biomass are coupled with the decline of C. intratropica throughout northern Australia by causing a switch from litter and sparse perennial grass fuels, and hence low-intensity surface fires, to heavy annual grass fuel loads that sustain fires that burn into the midstorey. This study demonstrates that changes in fuel type can alter fire regimes with substantial knock-on effects on the biota.     

Tropical grassy biomes: linking ecology,human use and conservation

Tropical grassy biomes (TGBs) are changing rapidly the world over through a coalescence of high rates of land-use change, global change and altered disturbance regimes that maintain the ecosystem structure and function of these biomes. Our theme issue brings together the latest research examining the characterization, complex ecology, drivers of change, and human use and ecosystem services of TGBs. Recent advances in ecology and evolution have facilitated a new perspective on these biomes. However, there continues to be controversies over their classification and state dynamics that demonstrate critical data and knowledge gaps in our quantitative understanding of these geographically dispersed regions. We highlight an urgent need to improve ecological understanding in order to effectively predict the sensitivity and resilience of TGBs under future scenarios of global change. With human reliance on TGBs increasing and their propensity for change, ecological and evolutionary understanding of these biomes is central to the dual goals of sustaining their ecological integrity and the diverse services these landscapes provide to millions of people.This article is part of the themed issue ‘Tropical grassy biomes: linking ecology, human use and conservation’.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

Convergence in growth responses of tropical trees to climate driven by water stress

Widely documented for temperate and cold forests in both hemispheres, variations in tree growth responses to climate along environmental gradients have rarely been investigated in the tropics. Seven tree‐ring chronologies of Centrolobium microchaete (Fabaceae) in the Cerrado tropical forests of Bolivia are used to determine the growth responses to climate along a precipitation gradient. Chronologies are distributed from the humid Guarayos forests (annual precipitation > 1600 mm) in the transition to the Amazonia to the dry‐mesic Chiquitos forests (annual precipitation < 1200 mm) in the proximity to the dry Chaco. On a large spatial scale, radial growth is positively influenced by rainfall and negatively by temperature at the end of the dry season. However, this regional pattern in climate‐tree growth relationship shows differences along the precipitation gradient. Relationships with climate are highly significant and extend over longer periods of the year in sites with low rainfall and extremely severe dry seasons. At wet sites, larger water soil capacity and endogenous forest dynamics partially mask the direct influence of climate on tree growth. Stronger similarities in tree‐growth responses to climate occur between sites in the dry Central Chiquitos and in the transition to the Guarayos forests. In contrast, the relationships show fewer similarities between sites in the humid Guarayos. We conclude that growth responses to climate in the tropics are more similar between sites with limited rainfall and severe and prolonged dry seasons. Our study points to a convergence in the patterns of growth responses of tropical trees to climate, modulated by scarce rainfall and marked seasonality. The negative impact of water deficits on tree physiological processes induces not only the documented reduction in forest species richness, but also a convergence in tree‐growth responses to climate in dry tropical forests.     

Soil organic carbon content at a range of north Australian tropical savannas with contrasting site histories

Soils play an important role in the global carbon cycle, and can be major source or sink of CO₂ depending upon land use, vegetation type and soil management practices. Natural and human impact on soil carbon concentration and storage is poorly understood in native north Australian savanna, yet this represents the largest carbon store in the ecosystem. To gain understanding of possible management impacts on this carbon pool, soil organic carbon (SOC) of the top 1m of red earth sands and sandy loams common in the region was sampled at 5 sites with different vegetation cover and site history (fire regime and tree removal). SOC was high when compared to other published values for savannas and was more comparable with dry-deciduous tropical forests. Sites sampled in this study represent high rainfall savannas of northern Australia (> 1700 mm annual rainfall) that feature frequent burning (2 in 3 years or more frequent) and a cycle of annual re-growth of tall C₄ grasses that dominate the savanna understorey. These factors may be responsible for the higher than expected SOC levels of the surface soils, despite high respiration rates. Medium term fire exclusion (15–20 years) at one of the sampled sites (Wildlife Park) dramatically reduced the grassy biomass of the understorey. This site had lower SOC levels when compared to the grass dominated and frequently burnt sites, which may be due to a reduction in detrital input to surface (0–30 cm) soil carbon pools. Exclusion of trees also had a significant impact on both the total amount and distribution of soil organic carbon, with tree removal reducing observed SOC at depth (100 cm). Soil carbon content was higher in the wet season than that in the dry season, but this difference was not statistically significant. Our results indicated that annual cycle of grass growth and wildfire resulted in small carbon accumulation in the upper region of the soil, and removal of woody plants resulted in significant carbon losses to recalcitrant, deep soil horizons greater than 80 cm depth.     

Testing the limits of resistance: a 19‐year study of Mediterranean grassland response to grazing regimes

A synthesis of a long‐term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north‐eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land‐use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long‐term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Rates of woody encroachment in African savannas reflect water constraints and fire disturbance

Aim

The aims of this study were to (1) estimate current rates of woody encroachment across African savannas; (2) identify relationships between change in woody cover and potential drivers, including water constraints, fire frequency and livestock density. The found relationships led us to pursue a third goal: (3) use temporal dynamics in woody cover to estimate potential woody cover.

Location

Sub‐Saharan African savannas.

Methods

The study used very high spatial resolution satellite imagery at sites with overlapping older (2002–2006) and newer (2011–2016) imagery to estimate change in woody cover. We sampled 596 sites in 38 separate areas across African savannas. Areas with high anthropogenic impact were avoided in order to more clearly identify the influence of environmental factors. Relationships between woody cover change and potential drivers were identified using linear regression and simultaneous autoregression, where the latter accounts for spatial autocorrelation.

Results

The mean annual change in woody cover across our study areas was 0.25% per year. Although we cannot explain the general trend of encroachment based on our data, we found that change rates were positively correlated with the difference between potential woody cover and actual woody cover (a proxy for water availability; p < .001), and negatively correlated with fire frequency (p < .01). Using the relationship between rates of encroachment and initial cover, we estimated potential woody cover at different rainfall levels.

Main conclusions

The results indicate that woody encroachment is ongoing and widespread across African savannas. The fact that the difference between potential and actual cover was the most significant predictor highlights the central role of water availability and tree–tree competition in controlling change in woody populations, both in water‐limited and mesic savannas. Our approach to derive potential woody cover from the woody cover change trajectories demonstrates that temporal dynamics in woody populations can be used to infer resource limitations.     

Liana abundance and diversity increase with rainfall seasonality along a precipitation gradient in Panama

In tropical regions, rainfall gradients often explain the abundance and distribution of plant species. For example, many tree and liana species adapted to seasonal drought are more abundant and diverse in seasonally-dry forests, characterized by long periods of seasonal water deficit. Mean annual precipitation (MAP) is commonly used to explain plant distributions across climate gradients. However, the relationship between MAP and plant distribution is often weak, raising the question of whether other seasonal precipitation patterns better explain plant distributions in seasonally-dry forests. In this study, we examine the relationship between liana abundance and multiple metrics of seasonal and annual rainfall distribution to test the hypothesis that liana density and diversity increase with increasing seasonal drought along a rainfall gradient across the isthmus of Panama. We found that a normalized seasonality index, which combines MAP and the variability of monthly rainfall throughout the year, was a significant predictor of both liana density and species richness, whereas MAP, rainfall seasonality and the mean dry season precipitation (MDP) were far weaker predictors. The strong response of lianas to the normalized seasonality index indicates that, in addition to the total annual amount of rainfall, how rainfall is distributed throughout the year is an important determinant of the hydrological conditions that favor liana proliferation. Our findings imply that changes in annual rainfall and rainfall seasonality will determine the future distribution and abundance of lianas. Models that aim to predict future plant diversity, distribution, and abundance may need to move beyond MAP to a more detailed understanding of rainfall variability at sub-annual timescales.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas

The distribution and abundance of trees can be strongly affected by disturbance such as fire. In mixed tree/grass ecosystems, recurrent grass‐fuelled fires can strongly suppress tree saplings and therefore control tree dominance. We propose that changes in atmospheric [CO₂] could influence tree cover in such metastable ecosystems by altering their postburn recovery rates relative to flammable herbaceous growth forms such as grasses. Slow sapling recovery rates at low [CO₂] would favour the spread of grasses and a reduction of tree cover. To test the possible importance of [CO₂]/fire interactions, we first used a Dynamic Global Vegetation Model (DGVM) to simulate biomass in grassy ecosystems in South Africa with and without fire. The results indicate that fire has a major effect under higher rainfall conditions suggesting an important role for fire/[CO₂] interactions. We then used a demographic model of the effects of fire on mesic savanna trees to test the importance of grass/tree differences in postburn recovery rates. We adjusted grass and tree growth in the model according to the DGVM output of net primary production at different [CO₂] relative to current conditions. The simulations predicted elimination of trees at [CO₂] typical of the last glacial period (180 ppm) because tree growth rate is too slow (15 years) to grow to a fire‐proof size of ca. 3 m. Simulated grass growth would produce an adequate fuel load for a burn in only 2 years. Simulations of preindustrial [CO₂] (270 ppm) predict occurrence of trees but at low densities. The greatest increase in trees occurs from preindustrial to current [CO₂] (360 ppm). The simulations are consistent with palaeo‐records which indicate that trees disappeared from sites that are currently savannas in South Africa in the last glacial. Savanna trees reappeared in the Holocene. There has also been a large increase in trees over the last 50–100 years. We suggest that slow tree recovery after fire, rather than differential photosynthetic efficiencies in C₃ and C₄ plants, might have been the significant factor in the Late Tertiary spread of flammable grasslands under low [CO₂] because open, high light environments would have been a prerequisite for the spread of C₄ grasses. Our simulations suggest further that low [CO₂] could have been a significant factor in the reduction of trees during glacial times, because of their slower regrowth after disturbance, with fire favouring the spread of grasses.     

Defoliation effects on basal cover and productivity in perennial grasslands of Ethiopia

This study aimed to investigate how perennial grass species in Omo National Park (ONP), Ethiopia tolerated defoliation under varying amounts of rainfall. Perennial grasses that have evolved with grazing appear to be generally tolerant to defoliation, although how rainfall influences this tolerance is unclear. Research was conducted in three perennial grasslands where there is a rainfall gradient from north to south (800 – 500 mm yr⁻¹). Grasslands were characterized as either wet, intermediate or dry sites according to their relative position along the rainfall gradient. The wet, intermediate, and dry sites were dominated by two, five, and two grass species, respectively, which comprised 98% of total plant basal cover at each site. Six exclosures containing a total of 12 defoliated and 12 non-defoliated plots (2 × 2 m) were constructed at each site. Hand-clipped defoliation treatments were imposed bimonthly for 18 months (i.e., four rainy seasons, three dry seasons). Repeated measurements of basal cover and biomass production were analyzed for overall response and by species. Basal cover increased (P < 0.05) or remained unchanged for all but one perennial grass species. Biomass production indicated trend for some species but was sensitive to annual rainfall. Overall results indicated that dominant perennial grasses of ONP were tolerant to defoliation, and this tolerance was expressed under all three rainfall levels. In addition, a decrease (P < 0.05) in basal cover was found for grasses in non-defoliated plots for five of nine cases, indicating a negative response to protection from grazing and fire. This revised version was published online in June 2006 with corrections to the Cover Date.