Elevated atmospheric CO2: a nurse plant substitute for oak seedlings establishing in old fields

It has been hypothesized that elevated levels of atmospheric CO₂ (eCO₂) may facilitate the encroachment of woody plants into grasslands by reducing water stress. In east‐central Minnesota, sandy soils frequently create drought conditions for plants, and water limitation inhibits the establishment of oaks into old fields situated on these soils. Some have argued that eCO₂ should slow secondary succession by favoring fast‐growing early successional species. However, if oak encroachment into old fields is being inhibited by water stress, then eCO₂ could accelerate old‐field succession in this region. The purpose of this study was to test the hypothesis that eCO₂ will increase the establishment success of oak seedlings in an old field environment. The study was conducted with CO₂ levels controlled by free air CO₂ enrichment (FACE). In May 1999, four oak (Quercus ellipsoidalis) acorns were planted in each of 24 plots in each of six experimental FACE rings (n= 576), three of which received elevated levels (550 ppm) of CO₂. Half the plots in each ring were weeded during the first three summers of the experiment. In summer 2000, water input was manipulated during a 3.5‐week period, during which half the plots received regular watering while the other half received no water. Summer 2001 was dry, receiving 35% less rainfall than the mean level. Under hot and dry conditions, eCO₂ increased soil water levels in unweeded plots and enhanced oak establishment (survival and growth) in weeded plots. In 2006, after the eighth growing season following planting, survival was five times greater under elevated than ambient CO₂. The results showed that under hot and dry conditions, eCO₂ can act like a nurse plant for tree seedlings growing in bare and unshaded areas, increasing seedling survival and growth, and thereby expanding the establishment window for trees encroaching into a grassland environment.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

Effects of moisture,nitrogen, grass competition and simulated browsing on the survival and growth of Acacia karroo seedlings

The effects of irrigation, nitrogen fertilization, grass competition and clipping were investigated for one growing season at the research farm of the University of Fort Hare in the Eastern Cape Province of South Africa. The aim of the experiment was to assess the short‐term performance of Acacia karroo seedlings under different environmental conditions and the implications of such factors on the long‐term recruitment of plant species in savanna rangelands. There were no significant treatment effects on the survival of A. karroo seedlings. Using stem length and basal diameter as growth parameters, it was observed that irrigation enhanced both variables, while nitrogen fertilization did not have any significant effects. Clipping, grass competition and their interaction greatly suppressed the growth of the seedlings. Clipping increased the mean stem length when they were irrigated and fertilized. Control and fertilized plants had the highest stem length in the absence of grass competition, while grass competition combined with clipping resulted in the lowest stem length in both irrigated and nonirrigated plants. It was concluded that in the presence of grass competition, controlled browsing could be a viable solution to the problem of bush encroachment in savanna rangelands.     

Fire responses of three co-occurring Asteraceae shrubs in a temperate savanna in South America

South American temperate savannas have undergone significant woody plant encroachment through changes in their disturbance regimes. We studied fire strategies and the spatial variation in individual fire responses of Eupatorium buniifolium, Baccharis medullosa and B. dracunculifolia, the dominant shrub species at El Palmar National Park, in areas with different fire history. In recently burnt sites, all Baccharis dracunculifolia individuals died, whereas all E. buniifolium and B. medullosa individuals sprouted. The relative growth rate was higher in E. buniifolium, which invested more biomass in shoots, and in B. medullosa, which invested more biomass in leaves. At the unburnt site, B. dracunculifolia showed the highest growth rate and leaf biomass. B. medullosa produced capitula immediately after fire, whereas E. buniifolium reproduction was delayed for 1 year. In both species, production of capitula and reproductive effort were higher in recently burnt sites. For B. dracunculifolia, production of capitula was lower and the reproductive effort was higher in burnt than in unburnt sites. Seedling establishment was extremely low for E. buniifolium, low for B. medullosa and high for B. dracunculifolia in both recently burnt sites. E. buniifolium behaved as a sprouter, B. medullosa as facultative sprouter and B. dracunculifolia as obligate seeder species. In order to control the increase and/or attain desirable population levels of these species, fire management practices in these savannas should consider individual species’ responses, particularly to the time since the last fire.     

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.     

Land‐use changes as major drivers of mountain pine (Pinus uncinata Ram.) expansion in the Pyrenees

Aim To assess the spatial patterns of forest expansion (encroachment and densification) for mountain pine (Pinus uncinata Ram.) during the last 50 years at a whole mountain range scale by the study of different topographic and socio‐economic potential drivers in the current context of global change. Location The study area includes the whole distributional area of mountain pine in the Catalan Pyrenees (north‐east Spain). This represents more than 80 municipalities, covering a total area of 6018 km². Methods Forest cover was obtained by image reclassification of more than 200 pairs of aerial photographs taken in 1956 and 2006. Encroachment and densification were determined according to changes in forest cover, and were expressed as binary variables on a 150 × 150 m cell‐size grid. We then used logistic regression to analyse the effects of several topographic and socio‐economic variables on forest expansion. Results In the period analysed, mountain pine increased its surface coverage by 8898 ha (an increase of more than 16%). Mean canopy cover rose from 31.0% in 1956 to 55.6% in 2006. Most of the expansion was found on north‐facing slopes and at low altitudes. Socio‐economic factors arose as major factors in mountain pine expansion, as encroachment rates were higher in municipalities with greater population losses or weaker primary sector development. Main conclusions The spatial patterns of mountain pine expansion showed a good match with the main patterns of land‐use change in the Pyrenees, suggesting that land‐use changes have played a more important role than climate in driving forest dynamics at a landscape scale over the period studied. Further studies on forest expansion at a regional scale should incorporate patterns of land‐use changes to correctly interpret drivers of forest encroachment and densification.     

Water uptake and hydraulic redistribution across large woody root systems to 20 m depth

Deep water uptake and hydraulic redistribution (HR) are important processes in many forests, savannas and shrublands. We investigated HR in a semi‐arid woodland above a unique cave system in central Texas to understand how deep root systems facilitate HR. Sap flow was measured in 9 trunks, 47 shallow roots and 12 deep roots of Quercus, Bumelia and Prosopis trees over 12 months. HR was extensive and continuous, involving every tree and 83% of roots, with the total daily volume of HR over a 1 month period estimated to be approximately 22% of daily transpiration. During drought, deep roots at 20 m depth redistributed water to shallow roots (hydraulic lift), while after rain, shallow roots at 0–0.5 m depth redistributed water among other shallow roots (lateral HR). The main driver of HR appeared to be patchy, dry soil near the surface, although water may also have been redistributed to mid‐level depths via deeper lateral roots. Deep roots contributed up to five times more water to transpiration and HR than shallow roots during drought but dramatically reduced their contribution after rain. Our results suggest that deep‐rooted plants are important drivers of water cycling in dry ecosystems and that HR can significantly influence landscape hydrology.     

Regeneration of indigenous trees in Mgahinga Gorilla National Park, Uganda

This study examines the regeneration of indigenous tree species in the formerly encroached area in Mgahinga Gorilla National Park (MGNP), south‐western Uganda. Before gazetting in 1992, MGNP had basically been agricultural land for well over 50 years. The distribution of exotic vegetation was established using a Geographical Positioning System receiver and indigenous vegetation was sampled by establishment of quadrats along transect lines. Observations indicated that approximately 2% of the old cropland was covered by exotic woodlots. Black wattle (Acacia mearnsii) and Eucalyptus trees were found to be the most widely distributed and Pinus patula the least distributed species in the park. Species numbers of indigenous trees (n = 26) were high in the old cropland, compared with twelve species observed in exotic woodlots. The natural forest supported the highest (75%) stem density and the lowest (4%) stem density was recorded in exotic woodlots. Seedling class (< 2 cm, d.b.h.) accounted for the majority of juveniles, with the lowest stem density (1350 seedlings ha^?1) recorded in exotic woodlots compared with 6609 seedlings ha^?1 in the old cropland and 24,625 seedlings ha^?1 in the natural forest. The levels of tree diversity and stocking characteristics recorded under the exotic species suggest that a low diverse community of native species may exploit this environment.