Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

Woody encroachment and forest degradation in sub-Saharan Africa's woodlands and savannas 1982–2006

We review the literature and find 16 studies from across Africa''s savannas and woodlands where woody encroachment dominates. These small-scale studies are supplemented by an analysis of long-term continent-wide satellite data, specifically the Normalized Difference Vegetation Index (NDVI) time series from the Global Inventory Modeling and Mapping Studies (GIMMS) dataset. Using dry-season data to separate the tree and grass signals, we find 4.0% of non-rainforest woody vegetation in sub-Saharan Africa (excluding West Africa) significantly increased in NDVI from 1982 to 2006, whereas 3.52% decreased. The increases in NDVI were found predominantly to the north of the Congo Basin, with decreases concentrated in the Miombo woodland belt. We hypothesize that areas of increasing dry-season NDVI are undergoing woody encroachment, but the coarse resolution of the study and uncertain relationship between NDVI and woody cover mean that the results should be interpreted with caution; certainly, these results do not contradict studies finding widespread deforestation throughout the continent. However, woody encroachment could be widespread, and warrants further investigation as it has important consequences for the global carbon cycle and land–climate interactions.     

Response of green alder (Alnus viridis subsp. fruticosa) patch dynamics and plant community composition to fire and regional temperature in north‐western Canada

Aim Feedbacks between climate warming and fire have the potential to alter Arctic and sub‐Arctic vegetation. In this paper we assess the effects and interactions of temperature and wildfire on plant communities across the transition between the Arctic and sub‐Arctic. Location Mackenzie Delta region, Northwest Territories, Canada. Methods We sampled air temperatures, green alder (Alnus viridis ssp. fruticosa) cover, growth, reproduction and age distributions, and overall plant community composition on burned and unburned sites across a latitudinal gradient. Results Mean summer temperature across the study area decreased by 3 °C per degree of increasing latitude (6 °C across the study area). In the northern part of the study area, where seed viability was low, alder was less dominant than at southern sites where seed viability was high. The age structure of alder populations across the temperature gradient was highly variable, except in the northern part of the forest–tundra transition, where populations were dominated by young individuals. Alder growth and reproduction were significantly greater on burned sites (38–51 years following fire) than on unburned sites. North to south across the temperature gradient, vegetation changed from a community dominated by dwarf shrubs and fruticose lichens to one characterized by black spruce (Picea mariana), alder and willows (Salix spp.). Regardless of the position along the temperature gradient, burned sites were dominated by tall shrubs. Main conclusions Temperature limitation of alder abundance and repro‐duction, combined with evidence of recent recruitment on unburned sites, indicates that alder is likely to respond to increased temperature. Elevated alder growth and reproduction on burned sites shows that wildfire also has an important influence on alder population dynamics. The magnitude of alder’s response to fire, combined with observations that burns at the southern margin of the low Arctic are shrub dominated, suggest that increases in the frequency of wildfire have the potential to alter northern vegetation on decadal scales. By creating new seedbeds, fire provides opportunities for colonization that may facilitate the northward movement of tall shrubs. Feedbacks between the global climate system and low Arctic vegetation make understanding the long‐term impact of increasing fire frequency critical to predicting the response of northern ecosystems to global change.     

Low functional redundancy among mammalian browsers in regulating an encroaching shrub (Solanum campylacanthum) in African savannah

Large herbivorous mammals play an important role in structuring African savannahs and are undergoing widespread population declines and local extinctions, with the largest species being the most vulnerable. The impact of these declines on key ecological processes hinges on the degree of functional redundancy within large-herbivore assemblages, a subject that has received little study. We experimentally quantified the effects of three browser species (elephant, impala and dik-dik) on individual- and population-level attributes of Solanum campylacanthum (Solanum incanum sensu lato), an encroaching woody shrub, using semi-permeable exclosures that selectively removed different-sized herbivores. After nearly 5 years, shrub abundance was lowest where all browser species were present and increased with each successive species deletion. Different browsers ate the same plant species in different ways, thereby exerting distinct suites of direct and indirect effects on plant performance and density. Not all of these effects were negative: elephants and impala also dispersed viable seeds and indirectly reduced seed predation by rodents and insects. We integrated these diffuse positive effects with the direct negative effects of folivory using a simple population model, which reinforced the conclusion that different browsers have complementary net effects on plant populations, and further suggested that under some conditions, these net effects may even differ in direction.     

Bush encroachment under three contrasting land-use practices in a mesic South African savanna

This study determined the effects of land-use practice had on the rate and extent of bush encroachment in a mesic savanna in KwaZulu-Natal, South Africa. Changes in woody cover were measured for 1 km² sites in areas under communal, commercial and conservation land-use systems for the period between 1937 and 2000. Land users from each area were interviewed to gain the histories of each area and to determine how the changes in woody cover had impacted them and whether anything was being done to counteract the spread of trees and shrubs on their land. Bush encroachment occurred across all three of the land-use types in the 67-year period between 1937 and 2000. The results showed that land-use practice had enormous impacts on the process of bush encroachment. The communal site showed a decrease in grass (21%) and tree (5%) cover and an increase in shrub cover (13%). At the commercial site, there was a considerable decrease in grass cover (46%) and moderate increase in shrub cover (10%) and a massive increase in tree cover (36%). The area under conservation showed a substantial decrease in grass cover (47%), a slight decrease in shrub cover (19%) and a massive increase in tree cover (66%). The perceived causes of these changes were fairly similar amongst the different land users. The changes were mostly not perceived to be a problem for the communal land users. The main advantages mentioned were increased woody resources for building and firewood and increased browse availability. The commercial and conservation land users perceived the changes to have significant negative connotations including the loss of grazing land and biodiversity and secondary invasion of encroached areas by alien plant species. Despite these perceptions, very little has been done to combat bush encroachment in the commercial and conservation land use systems.     

Relative impacts of disturbance and temperature: persistent changes in microenvironment and vegetation in retrogressive thaw slumps

In the Low Arctic, a warming climate is increasing rates of permafrost degradation and altering vegetation. Disturbance associated with warming permafrost can change microclimate and expose areas of ion-rich mineral substrate for colonization by plants. Consequently, the response of vegetation to warming air temperatures may differ significantly from disturbed to undisturbed tundra. Across a latitudinal air temperature gradient, we tested the hypothesis that the microenvironment in thaw slumps would be warmer and more nutrient rich than undisturbed tundra, resulting in altered plant community composition and increased green alder ( Alnus viridis subsp. fruticosa ) growth and reproduction. Our results show increased nutrient availability, soil pH, snow pack, ground temperatures, and active layer thickness in disturbed terrain and suggest that these variables are important drivers of plant community structure. We also found increased productivity, catkin production, and seed viability of green alder at disturbed sites. Altered community composition and enhancement of alder growth and reproduction show that disturbances exert a strong influence on deciduous shrubs that make slumps potential seed sources for undisturbed tundra. Overall, these results indicate that accelerated disturbance regimes have the potential to magnify the effects of warming temperature on vegetation. Consequently, understanding the relative effects of temperature and disturbance on Arctic plant communities is critical to predicting feedbacks between northern ecosystems and global climate change.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Fire frequency influences composition and structure of the shrub layer in an Australian subcoastal temperate grassy woodland

Little is known about the relationship between fire regimes and plant diversity in Australia's temperate grassy woodlands. The effect of fire frequency on shrubs in grassy woodland remnants across Western Sydney's Cumberland Plain was examined. Shrub species richness and composition were compared in sites that had experienced a high, moderate or low frequency of fire over the previous 20 years. Nine sites were surveyed, three in each fire frequency category; most sites, including all low‐fire‐frequency sites, had burnt 9–36 months prior to sampling. Fire frequency had a profound effect on the composition and structure of the shrub layer. Per cent frequency and density of the prickly shrub Bursaria spinosa (Pittosporaceae) was considerably higher in low‐fire‐frequency sites than where fires had occurred at least once a decade. In sites where fire had been absent for decades prior to a recent fire, this species dominated the landscape, while elsewhere it occurred as clumps in a grassy matrix. Per cent frequency of other native shrubs, particularly obligate seeders, was greatest at moderate fire frequencies. Exotic shrubs were recorded most often where fire had been rare. While ordination clearly separated out the low‐fire‐frequency sites, complete separation between high‐ and moderate‐fire‐frequency blocks was not achieved. The increase in Bursaria in the absence of fire mirrors the encroachment of woody plants into a range of grassy ecosystems around the world. The sensitivity of obligate seeder species, many of them short‐lived legumes with fire‐cued seeds, to both very frequent and very infrequent fire shows the vulnerability of these species to extreme fire regimes, despite the safeguards conferred by hard‐seededness. Competition from Bursaria, as well as loss of viable seed in the soil, may have contributed to the low frequency of these species after a long inter‐fire interval.     

Impact of storm‐burning on Melaleuca viridiflora invasion of grasslands and grassy woodlands on Cape York Peninsula,Australia

This paper examines invasion of grasslands on Cape York Peninsula, Australia, by Melaleuca viridiflora and other woody species, and the role of storm‐burning (lighting fires after the first wet season rains) in their maintenance. Trends in disturbance features, fuel characteristics, ground layer composition, and woody plants dynamics under combinations of withholding fire and storm‐burning over a 3‐year period were measured on 19 plots in three landscape settings. Population dynamics of M. viridiflora are described in detail and 20‐year population projections based on transition matrices under different fire regimes generated. Numerous M. viridiflora suckers occurred within the grass layer, increasing each year regardless of fire regime, and were rapidly recruited to the canopy in the absence of fire. Storm‐burning had little impact on fuel, ground layer or woody plant composition, but maintained open vegetation structure by substantially reducing recruitment of M. viridiflora suckers to the sapling layer, and by reducing the above‐grass‐layer abundance of several other invasive woody species. Population projections indicated that withholding fire for 20 years could cause a sevenfold increase of M. viridiflora density on Ti‐tree flats, and that annual to triennial storm‐burning should be effective at maintaining a stable open vegetation structure. These findings argue against vegetation thickening being an inevitable consequence of climate change. We conclude that a fire regime that includes regular storm‐burning can be effective for maintaining grasslands and grassy woodlands being invaded by M. viridiflora.