How widespread is woody plant encroachment in temperate Australia? Changes in woody vegetation cover in lowland woodland and coastal ecosystems in Victoria from 1989 to 2005

Aim Encroachment or densification by woody plants affects natural ecosystems around the world. Many studies have reported encroachment in temperate Australia, particularly in coastal ecosystems and grassy woodlands. However, the degree to which published studies reflect broad-scale changes is unknown because most studies intentionally sampled areas with conspicuous densification. We aimed to estimate changes in woody vegetation cover within lowland grassy woodland and coastal ecosystems in Victoria from 1989 to 2005 to determine whether published reports of recent encroachment are representative of broad-scale ecosystem changes. Location All lowland grassy woodland and coastal ecosystems (c. 6.11 × 10⁵ ha) in Victoria, Australia. Four major ecosystems were analysed: Plains woodlands, Herb-rich woodlands, Riverine woodlands and Coastal vegetation. Methods Changes in woody vegetation cover from 1989 to 2005 were assessed based on state-wide vegetation maps and Landsat analyses of woody vegetation cover conducted by the Australian Greenhouse Office’s National Carbon Accounting System. The results show changes in woody cover within mapped patches of native vegetation, rather than changes in the extent of woody vegetation resulting from clearing and revegetation. Results When pooled across all ecosystems, woody vegetation increased by 18,730 ha from 1989 to 2005. Woody cover within Riverine woodlands and within Plains woodlands each increased by >7000 ha. At the patch scale, the mean percentage cover of woody vegetation in each polygon increased by >5% in all four ecosystems: Riverine woodlands (+9.2% on average), Herb-rich woodlands (+7.6%), Plains woodlands (+6.7%) and Coastal vegetation (+5.9%). Regression models relating degree of encroachment to geographic and climatic variables were extremely weak (r² ≤ 0.026), indicating that most variation occurred at local scales rather than across broad geographic gradients. Main conclusions At the scale of observation, woody vegetation cover increased in all lowland woodland and coastal ecosystems over the 16-year period. Thus, published examples of encroachment in selected coastal and woodland patches do appear to reflect widespread increases in woody vegetation cover in these ecosystems. This densification appears to be associated with changes in land management rather than with post-fire vegetation recovery and is likely to be ongoing and long-lasting, with substantial implications for biodiversity conservation and ecosystem services.     

Soil nutrients distribution and moisture dynamics on upper catena in a semi-arid nature reserve

The distribution of soil nutrients and soil moisture dynamics on slopes of catenary landscape, were recorded in a semi-arid nature reserve located in the Eastern Transvaal, South Africa. Soil nutrients were accumulating in the slopes of the catena whereas the crests were highly leached. Open grassland bands segregating between the two soil types and their associated plant species, did not show clear stratification of nutrient deposition amongst ground levels down to 1.5 m below the surface. There was however, a tendency of soils with high nutrient contents to correlate with areas of high woody vegetation density. Patterns of soil moisture levels during two annual rainfall periods showed some segregation between soil depths, although the trends indicated that the duration of saturation and dry out of soils levels differed more between rainy seasons than between soil levels during a particular season.     

Drought-induced tree death in savanna

Increasing densities of woody plants in savannas has been attributed to both elevated atmospheric CO₂ and reduced burning with grazing management, such that the biome could represent a substantial carbon sink. However, we show that extreme droughts (less than two-thirds expected rainfall over 3 years) occur in the drier half of the savanna biome and can cause substantial tree death. An Australian case study reveals that a net increase in tree cover over five decades of above-average rainfall was offset by sudden tree death during drought. The relationship between woody cover change and rainfall is moderated by competition with growth being facilitated by low woody cover and drought-induced death more likely as the woody component of savanna increases. The results are not supportive of a sustained increase in the woody component of xeric savannas resulting from CO₂ fertilization or land management. Extensive tree death in savanna regions will become a stark consequence of climate change if predictions of increasing severity and frequency of drought are realized.     

Carbon dioxide and the uneasy interactions of trees and savannah grasses

Savannahs are a mixture of trees and grasses often occurring as alternate states to closed forests. Savannah fires are frequent where grass productivity is high in the wet season. Fires help maintain grassy vegetation where the climate is suitable for woodlands or forests. Saplings in savannahs are particularly vulnerable to topkill of above-ground biomass. Larger trees are more fire-resistant and suffer little damage when burnt. Recruitment to large mature tree size classes depends on sapling growth rates to fire-resistant sizes and the time between fires. Carbon dioxide (CO(2)) can influence the growth rate of juvenile plants, thereby affecting tree recruitment and the conversion of open savannahs to woodlands. Trees have increased in many savannahs throughout the world, whereas some humid savannahs are being invaded by forests. CO(2) has been implicated in this woody increase but attribution to global drivers has been controversial where changes in grazing and fire have also occurred. We report on diverse tests of the magnitude of CO(2) effects on both ancient and modern ecosystems with a particular focus on African savannahs. Large increases in trees of mesic savannahs in the region cannot easily be explained by land use change but are consistent with experimental and simulation studies of CO(2) effects. Changes in arid savannahs seem less obviously linked to CO(2) effects and may be driven more by overgrazing. Large-scale shifts in the tree-grass balance in the past and the future need to be better understood. They not only have major impacts on the ecology of grassy ecosystems but also on Earth-atmosphere linkages and the global carbon cycle in ways that are still being discovered.     

Termite‐induced heterogeneity in African savanna vegetation: mechanisms and patterns

Objectives: To (1) assess the strength of evidence for the role of termites in vegetation heterogeneity in African savannas, and (2) identify the mechanisms by which termites induce such heterogeneity. Location: African savannas. Methods: We conducted a review of the literature, a meta‐analysis and qualitative systems analysis to identify mechanisms to explain the observed patterns. Results: The review provided evidence for termite‐induced heterogeneity in floristic composition and vegetation patterning in savannas across Africa. Termites induced vegetation heterogeneity directly or indirectly through their nest‐building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire. The literature reviewed indicated that termite mounds essentially act as islands of fertility, which are responsible for ecosystem‐level spatial heterogeneity in savannas. This was supported by the meta‐analysis, which demonstrated that mounds of Ancistrotermes, Macrotermes, Odontotermes (family Macrotermitinae), Cubitermes (family Termitinae) and Trinervitermes (Nasutitermitinae) are significantly enriched in clay (75%), carbon (16%), total nitrogen (42%), calcium (232%), potassium (306%) and magnesium (154%) compared to the surrounding savanna soil. Conclusions: Termite activity is one of the major factors that induce vegetation patterning in African savannas. The implications of this are discussed and research questions for future studies and modelling efforts are indicated.     

Net changes in regional woody vegetation cover and carbon storage in Texas Drylands, 1937–1999

Although local increases in woody plant cover have been documented in arid and semiarid ecosystems worldwide, there have been few long‐term, large‐scale analyses of changes in woody plant cover and aboveground carbon (C) stocks. We used historical aerial photography, contemporary Landsat satellite data, field observations, and image analysis techniques to assess spatially specific changes in woody vegetation cover and aboveground C stocks between 1937 and 1999 in a 400‐km² region of northern Texas, USA. Changes in land cover were then related to topo‐edaphic setting and historical land‐use practices. Mechanical or chemical brush management occurred over much of the region in the 1940–1950s. Rangelands not targeted for brush management experienced woody cover increases of up to 500% in 63 years. Areas managed with herbicides, mechanical treatments or fire exhibited a wide range of woody cover changes relative to 1937 (?75% to + 280%), depending on soil type and time since last management action. At the integrated regional scale, there was a net 30% increase in woody plant cover over the 63‐year period. Regional increases were greatest in riparian corridors (33%) and shallow clay uplands (26%) and least on upland clay loams (15%). Allometric relationships between canopy cover and aboveground biomass were used to estimate net aboveground C storage changes in upland (nonriparian) portions of regional landscapes. Carbon stocks increased from 380 g C m^?2 in 1937 to 500 g C m^?2 in 1999, a 32% net increase across the 400 km² region over the 63‐year period. These plant C storage change estimates are highly conservative in that they did not include the substantial increases in woody plant cover observed within riparian landscape elements. Results are discussed in terms of implications for ‘carbon accounting’ and the global C cycle.     

Savanna tree–grass competition is modified by substrate type and herbivory

Question: Woody plant and grass interactions in savannas have frequently been studied from the perspective of the response of one growth form on the other but seldom evaluated as two‐way interactions. What causes woody plant encroachment in semi‐arid savannas and what are the competitive responses of tree seedlings and grasses on rocky and sandy substrates? Methods: In this greenhouse study, we investigated the influence of substrate and grazing on responses to interspecific competition by tree seedlings and grasses. We measured competitive/facilitative responses on biomass and nutrient status of tree seedlings and grasses grown together. Results: Interspecific competition suppressed growth of trees and grasses. Tree seedlings and uncut grass accumulated double the biomass when grown without competition relative to when they competed. Competitive responses varied on different substrates. Grass biomass on rocky substrate showed no response to tree competition, but appeared to be facilitated by trees on sandy substrate. Grass clipping resulted in higher tree seedling biomass on rocky substrate, but not on sandy substrate. There was a positive response of grass nutrient status to competition from tree seedlings. Conclusion: Selective grass herbivory in the absence of browsing or suppression of shade‐intolerant grasses by trees are commonly cited reasons behind bush encroachment in savannas. We show that grazing may confer a competitive advantage to tree seedlings and promote bush encroachment more readily on rocky substrates. This may be due to the imposed sharing of the soil depth niche on rocky substrates, whereas possible niche separation on sandy substrates minimizes the advantage conferred by reduced competition.