Multi‐scale patterns and bush encroachment in an arid savanna with a shallow soil layer

Abstract. Question: Bush encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and threatens the livelihood of many pastoralists. Can we derive a better understanding of the factors causing bush encroachment by investigating the scale dependency of patterns and processes in savannas? Location: An arid savanna in the Khomas Hochland, Namibia. Methods: Patterns of bush, grass, and soil nutrient distribution were surveyed on several scales along a rainfall gradient, with emphasis on intraspecific interactions within the dominant woody species, Acacia reficiens. Results: Savannas can be interpreted as patch‐dynamic systems where landscapes are composed of many patches (a few ha in size) in different states of transition between grassy and woody dominance. Conclusions: In arid savannas, this patchiness is driven both by rainfall that is highly variable in space and time and by inter‐tree competition. Within the paradigm of patch‐dynamic savannas, bush encroachment is part of a cyclical succession between open savanna and woody dominance. The conversion from a patch of open savanna to a bush‐encroached area is initiated by the spatial and temporal overlap of several (localized) rainfall events sufficient for Acacia germination and establishment. With time, growth and self‐thinning will transform the bush‐encroached area into a mature Acacia stand and eventually into open savanna again. Patchiness is sustained due to the local rarity (and patchiness) of rainfall sufficient for germination of woody plants as well as by plant‐soil interactions.     

Relationship between small-scale structural variability and simulated vegetation productivity across a regional moisture gradient in southern Africa

The observed variability in vegetation structure within landscapes was used as the basis for model estimates of the range of annual productivity of landscape patches at four sites along a moisture gradient in southern Africa ranging from 879 to 365 mm mean annual rainfall. Principal components of patch‐scale variability in leaf area, woody biomass and vertical leaf profiles were derived from intensive characterization of the small‐scale spatial structure of woody vegetation at each site. For each site, the mean and extremes of the principal component distribution parameterized an ecophysiology model of vegetation productivity. Vegetation was most heterogeneous at intermediate locations along the rainfall gradient. Variability in vegetation structure led to a range of annual productivity within one site (600 mm) that accounted for 68% of the total range in mean productivity across all sites. Patch‐scale estimates of tree productivity were found to be primarily correlated to annual rainfall (r²=0.66, P=0.001) and not woody leaf area (r²=0.01, P=0.75), while grass productivity was found to be related to values of woody leaf area (r²=0.77, P<0.001) and not annual rainfall (r²=0.11, P=0.29). This result indicates that life‐form interactions have a significant role in controlling vegetation productivity across the rainfall gradient. The findings of this study emphasize the importance of considering heterogeneity rather than mean structure when modeling productivity, particularly when considering dynamic vegetation structure, where differences between landscape patches may not be well represented in the mean structure.     

Stable forest–savanna mosaic in north‐western Tanzania: local‐scale evidence from δ13C signatures and 14C ages of soil fractions

Aim The spatio‐temporal dynamics of dry evergreen forest patches in the savanna biome of the Kagera region (north‐western Tanzania) are largely unknown owing to a lack of pollen and macrofossil evidence. Our aims were to reconstruct local‐scale shifts of the forest–savanna boundary in order to determine whether the forests have been expanding or retreating on a centennial and millennial time‐scale. Location The Kagera region of north‐western Tanzania, East Africa. Methods The vegetation reconstruction was based on analysing δ¹³C signatures in soils along a transect spanning both C₄ open savanna and C₃ forest vegetation. Furthermore, we fractionated soil organic matter (SOM) according to density and chemical stability to analyse δ¹³C values of soil fractions with distinct radiocarbon ages. Results We found sharp changes in δ¹³C signatures in bulk SOM from the forest to the savanna, within a few metres along the transect. The forest soil profiles carried a persistent C₃‐dominated signature. Radiocarbon dating of the oldest, most recalcitrant forest soil fraction yielded a mean age of 5500 cal. yr bp , demonstrating that the forest has existed since at least the mid‐Holocene. The savanna sites showed a typical C₄ isotopic signature in SOM of topsoils, but subsoils and more recalcitrant SOM fractions also contained signals of C₃ plants. The dense soil fraction (ρ > 1.6 g cm^?3) carrying a pure C₄ label had a mean age of c. 1200 cal. yr bp , indicating the minimum duration of the dominance of grass vegetation on the savanna site. At the forest edge, the older C₄ grass signature of SOM has steadily been replaced by the more negative δ¹³C fingerprint of the forest trees. As this replacement has occurred mainly in the 10‐m‐wide forest–savanna ecotone over the last c. 1200 years, the forest expansion must be very slow and is very likely less than 15 m century^?1. Main conclusions Our results suggest that forest patches in the Kagera savanna landscape are very stable vegetation formations which have persisted for millennia. During the last millennium, they have been expanding very slowly into the surrounding savanna at a rate of less than 15 m century^?1.     

Vegetation dynamics in a Quercus‐Juniperus savanna: An isotopic assessment

Abstract. Woody plants are increasing in many grassland and savanna ecosystems around the world. As a case in point, the Edwards Plateau of Texas, USA, is a vast region (93 000 km²) in which rapid woody encroachment appears to be occurring. The native vegetation (prior to the Anglo‐European settlement 150–200 yr ago) and the biogeochemical consequences of woody encroachment in this region, however, are poorly understood. To assess these matters we measured plant and soil δ¹³C, soil organic C and soil N content from grasslands and two important woody patch types (mature Quercus virginiana clusters and Juniperus ashei woodlands) in this region. Soil δ¹³C values showed that relative productivity of C₃ species has increased in grassland and both woody habitats in recent times. δ¹³C of SOC in grasslands and Q. virginiana clusters increased with depth from the litter layer to 30 cm (grasslands =?21 to ?13‰Q. virginiana clusters =?27 to ?17‰) and were significantly different between habitats at all depths, indicating that Q. virginiana has been a long‐term component of the landscape. In J. ashei woodlands, soil δ¹³C values (at 20–30 cm depth) near the woodland edge (‐13‰) converged with those of an adjacent grassland (‐13‰) while those from the woodland interior (‐15‰) remained distinct, indicating that the woodland has been present for many years but has recently expanded. Concentrations and densities of SOC and total N were generally greater in woody patches than in grasslands. However, differences in the amount of SOC and N stored beneath the two woody patch types indicates that C and N sequestration potentials are species dependent.     

Climate warming and precipitation redistribution modify tree–grass interactions and tree species establishment in a warm‐temperate savanna

Savanna tree–grass interactions may be particularly sensitive to climate change. Establishment of two tree canopy dominants, post oak (Quercus stellata) and eastern redcedar (Juniperus virginiana), grown with the dominant C₄ perennial grass (Schizachyrium scoparium) in southern oak savanna of the United States were evaluated under four climatic scenarios for 6 years. Tree–grass interactions were examined with and without warming (+1.5 °C) in combination with a long‐term mean rainfall treatment and a modified rainfall regime that redistributed 40% of summer rainfall to spring and fall, intensifying summer drought. The aim was to determine: (1) the relative growth response of these species, (2) potential shifts in the balance of tree–grass interactions, and (3) the trajectory of juniper encroachment into savannas, under these anticipated climatic conditions. Precipitation redistribution reduced relative growth rate (RGR) of trees grown with grass. Warming increased growth of J. virginiana and strongly reduced Q. stellata survival. Tiller numbers of S. scoparium plants were unaffected by warming, but the number of reproductive tillers was increasingly suppressed by intensified drought each year. Growth rates of J. virginiana and Q. stellata were suppressed by grass presence early, but in subsequent years were higher when grown with grass. Quercus stellata had overall reduced RGR, but enhanced survival when grown with grass, while survival of J. virginiana remained near 100% in all treatments. Once trees surpassed a threshold height of 1.1 m, both tiller number and survival of S. scoparium plants were drastically reduced by the presence of J. virginiana, but not Q. stellata. Juniperus virginiana was the only savanna dominant in which neither survival nor final aboveground mass were adversely affected by the climate scenario of warming and intensified summer drought. These responses indicate that climate warming and altered precipitation patterns will further accelerate juniper encroachment and woody thickening in a warm‐temperate oak savanna.     

Vegetation state changes in the course of shrub encroachment in an African savanna since about 1850 CE and their potential drivers

1. Shrub encroachment has far‐reaching ecological and economic consequences in many ecosystems worldwide. Yet, compositional changes associated with shrub encroachment are often overlooked despite having important effects on ecosystem functioning.
2. We document the compositional change and potential drivers for a northern Namibian Combretum woodland transitioning into a Terminalia shrubland. We use a multiproxy record (pollen, sedimentary ancient DNA, biomarkers, compound‐specific carbon (δ¹³C) and deuterium (δD) isotopes, bulk carbon isotopes (δ¹³Corg), grain size, geochemical properties) from Lake Otjikoto at high taxonomical and temporal resolution.
3. We provide evidence that state changes in semiarid environments may occur on a scale of one century and that transitions between stable states can span around 80 years and are characterized by a unique vegetation composition. We demonstrate that the current grass/woody ratio is exceptional for the last 170 years, as supported by n‐alkane distributions and the δ¹³C and δ¹³Corg records. Comparing vegetation records to environmental proxy data and census data, we infer a complex network of global and local drivers of vegetation change. While our δD record suggests physiological adaptations of woody species to higher atmospheric pCO₂ concentration and drought, our vegetation records reflect the impact of broad‐scale logging for the mining industry, and the macrocharcoal record suggests a decrease in fire activity associated with the intensification of farming. Impact of selective grazing is reflected by changes in abundance and taxonomical composition of grasses and by an increase of nonpalatable and trampling‐resistant taxa. In addition, grain‐size and spore records suggest changes in the erodibility of soils because of reduced grass cover.
4. Synthesis. We conclude that transitions to an encroached savanna state are supported by gradual environmental changes induced by management strategies, which affected the resilience of savanna ecosystems. In addition, feedback mechanisms that reflect the interplay between management legacies and climate change maintain the encroached state.

     

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.     

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 environmental variables on vesicular‐arbuscular mycorrhizal abundance in wild populations of Vangueria infausta

Abstract. Vesicular‐arbuscular endomycorrhizal fungi include among its members some of the most widespread root symbiont species. It is not known whether these fungal species show environmental or host preferences. In 13 semi‐arid savanna sites in Botswana, we found positive correlations between individual environmental factors and the abundance of VAM associations in the roots of an important host, the indigenous fruit tree Vangueria infausta (Rubiaceae). The concentration of phosphorus in the leaves of the host was positively correlated with both phosphorus in the soil and the abundance of VAM associations in its roots, indicating direct benefits to the host of this association. Abundance of VAM associations was significantly different between the studied sites and between seasons and was positively correlated with the mean annual rainfall. In addition, there was a negative correlation with phosphorus concentration in the soil. The differences between the summer and winter abundance of VAM associations were positively correlated with the density of bush cover and the amount of grazing. The apparent correlation found between the abundance of VAM in V. infausta and bush and tree community composition is presumably related to correlations between these two parameters and environmental conditions.     

Fire resilience of ant assemblages in long‐unburnt savanna of northern Australia

Abstract Tropical savannas and rainforests contrast in their flammability and the fire resilience of their associated species. While savanna species generally exhibit high resilience to burning, there is much debate about the fire resilience of forest‐associated species, and the persistence of forest patches in a flammable savanna matrix. Where fire has been excluded, savanna tends on a trajectory towards forest, with an increase in forest‐associated plants and animal species. This study tested the idea that given the high proportion of forest‐associated taxa in long‐unburnt savanna, the fauna of these areas would be expected to exhibit less resilience to fire than the fauna in frequently burnt savannas. The study investigated the immediate and short‐term effects on ant assemblages of re‐introducing fire into long‐unburnt savanna in northern Australia. The ant fauna exhibited high resistance to fires, with no significant short‐term change in mean abundance or species richness; instead, seasonality had a far stronger influence on overall ant activity. Fire caused dramatic declines in dominance of the patchily distributed forest‐associated species Oecophylla smaragdina and Papyrius sp., but had no effect on overall dominance by open savanna species of Iridomyrmex. Dominance by Iridomyrmex pallidus declined, but this was compensated for by increases in I. reburrus, while two other species of Iridomyrmex showed no change. This indicates a high level of functional redundancy among dominant species of Iridomyrmex, which universally dominate open savanna communities, but not of dominant forest‐associated species. Overall, our findings demonstrate a high degree of fire‐resilience of the long‐unburnt savanna ant fauna. Despite the occurrence of forest‐associated species, the high proportion of savanna species persisting in this habitat means that long‐unburnt savanna retains the general response characteristics of frequently burnt savanna.     

Characterization of a stretch‐activated potassium channel in chondrocytes

Chondrocytes possess the capacity to transduce load‐induced mechanical stimuli into electrochemical signals. The aim of this study was to functionally characterize an ion channel activated in response to membrane stretch in isolated primary equine chondrocytes. We used patch‐clamp electrophysiology to functionally characterize this channel and immunohistochemistry to examine its distribution in articular cartilage. In cell‐attached patch experiments, the application of negative pressures to the patch pipette (in the range of 20–200 mmHg) activated ion channel currents in six of seven patches. The mean activated current was 45.9 ± 1.1 pA (n = 4) at a membrane potential of 33 mV (cell surface area approximately 240 µm²). The mean slope conductance of the principal single channels resolved within the total stretch‐activated current was 118 ± 19 pS (n = 6), and reversed near the theoretical potassium equilibrium potential, E_K+, suggesting it was a high‐conductance potassium channel. Activation of these high‐conductance potassium channels was inhibited by extracellular TEA (K_d approx. 900 µM) and iberiotoxin (K_d approx. 40 nM). This suggests that the current was largely carried by BK‐like potassium (MaxiK) channels. To further characterize these BK‐like channels, we used inside‐out patches of chondrocyte membrane: we found these channels to be activated by elevation in bath calcium concentration. Immunohistochemical staining of equine cartilage samples with polyclonal antibodies to the α1‐ and β1‐subunits of the BK channel revealed positive immunoreactivity for both subunits in superficial zone chondrocytes. These experiments support the hypothesis that functional BK channels are present in chondrocytes and may be involved in mechanotransduction and chemotransduction. J. Cell. Physiol. 223: 511–518, 2010. © 2010 Wiley‐Liss, Inc.     

灌丛化对黄土高原草地植物群落结构和地上生物量的影响

中国北方草地普遍出现灌丛化现象,灌丛化改变植物群落结构、植物多样性和生产力,直接影响着草地生态保护与可持续利用。该研究以黄土高原灌丛化草地为研究对象,通过植被调查,分析比较不同坡向的灌丛斑块与禾草斑块植物群落结构(物种组成、优势种及物种多样性)和地上生物量的差异。结果发现:(1)灌丛化草地不同坡向对物种多样性及地上生物量均无显著影响(P 0.1),但不同斑块植物群落结构(P=0.001)及地上生物量(P0.001)存在显著差异。(2)灌丛化草地共出现植物29种,其中禾草斑块有27种,灌丛斑块有18种;灌丛化显著改变了植物群落的物种组成,优势种由长芒草(Stipa bungeana)更替为矮脚锦鸡儿(Caragana brachypoda),且灌丛化降低了草地物种丰富度,增加了群落均匀度。(3)灌丛化显著改变了草地地上生物量,其中灌丛斑块地上生物量较禾草斑块地上生物量增加251.2 g·m~(-2),灌丛斑块中灌木/半灌木地上生物量提高了452.1 g·m~(-2),多年生丛生禾草减少了176.5 g·m~(-2),其余功能群植物的地上生物量减少了24.4 g·m~(-2)。(4)灌丛化过程(从禾草斑块—灌丛斑块)中,植物种丢失对地上生物量减少的影响较小,新增物种和群落优势种更替促进了灌木斑块地上生物量增加;虽然灌丛化导致草地地上生物量增加,但植物物种丰富度降低和优势种更替很有可能改变草地多样性和稳定性维持机制。     

Environmental and species‐specific controls on δ13C and δ15N in dominant woody plants from central‐western Argentinian drylands

Spatial variation in mean annual precipitation is the principal driver of plant water and nitrogen status in drylands. The natural abundance of carbon stable isotopes (δ¹³C) in photosynthetic tissues of C3 plants is an indicator of time‐integrated behaviour of stomatal conductance; while that of nitrogen stable isotopes (δ¹⁵N) is an indicator of the main source of plant N (soil N vs. atmospheric N₂). Previous studies in drylands have documented that plant δ¹³C and δ¹⁵N values increase with decreasing mean annual precipitation due to reductions in stomatal conductance, and soil enriched in ¹⁵N, respectively. However, evidence for this comes from studies focused on stable isotopes measurements integrated at the plant community level or on dominant plants at the site level, but little effort has been made to study C and N isotope variations within a species growing along rainfall gradients. We analysed plant δ¹³C, δ¹⁵N and C/N values of three woody species having different phenological leaf traits (deciduous, perennial and aphyllous) along a regional mean annual precipitation gradient from the central‐western Argentinian drylands. Noticeably, plant δ¹³C and δ¹⁵N values in the three woody species did not increase towards sites with low precipitation or at the start of the growing season (drier period), as we expected. These results suggest that environmental factors other than mean annual precipitation may be affecting plant δ¹³C and δ¹⁵N. The short‐term environmental conditions may interact with species‐specific plant traits related to water and nitrogen use strategies and override the predictive influence of the mean annual precipitation on plant δ¹³C and δ¹⁵N widely reported in drylands.     

Assessing the frequency and drivers of early‐greening in broad‐leaved woodlands along a latitudinal gradient in southern Africa

Savannas are the only deciduous system where new leaf flush pre‐empts the onset of suitable conditions for growth, a phenological phenomenon known as early‐greening. Limited understanding of the frequency and drivers of the occurrence of early‐greening in southern African savanna trees exists. We aimed to estimate the frequency of early‐greening events across southern Africa and investigated potential environmental drivers of green‐up. We selected and compared seven broad‐leaved woodland sites where Burkea africana was a dominant species using remotely sensed data along a latitudinal gradient from South Africa to Zambia. Normalized difference vegetation index (NDVI) values were extracted from the Moderate Resolution Imaging Spectroradiometer (MODIS) satellite imagery at each site from January 2002 to June 2014. Using an austral year (July 1st–June 30th), early‐greening was recorded if the green‐up start date occurred prior to the onset date of seasonal rainfall. A latitudinal gradient of early‐green‐up was detected across southern Africa (R² = 0.74) with the two most northerly (Zambian) sites showing the earliest and most consistent green‐up start dates (3 October ± 5.34 days). A strong latitudinal gradient was observed between the variability in the amount of rainfall in the first 6 months of green‐up and the green‐up start dates across southern Africa (R² = 0.92). Photoperiod appeared to play a role in areas where the onset of rainfall commenced late into the austral year. Mean maximum temperatures recorded 10 days prior to green‐up start dates suggested a potential threshold of about 35°C, which could drive early‐greening in the absence of rainfall. Correlations between the proportion of early‐greening years and the above mentioned environmental factors indicated that rainfall variability had the strongest influence over the observed phenological gradient (R² = 0.96). Understanding early‐greening in complex savanna systems is a vital step in furthering predictive phenological models under changing climatic conditions.     

Long‐term livestock exclusion facilitates native woody plant encroachment in a sandy semiarid rangeland

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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.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

Regional patch dynamics of Cirsium arvense and possible implications for plant‐animal interactions

Abstract. Plant population biology considers the dynamics of plant modules within stands. However, stands themselves may have considerable regional turnover in space and time. These changes in the number, distribution and size of plant stands generate a dynamic spatial pattern with important implications for the spatial and temporal dynamics of phytophagous insects using these plants as a host. During five successive years we studied the regional distribution and patch dynamics of the creeping thistle Cirsium arvense and the distribution of associated populations of the herbivore Urophora cardui (Diptera: Tephritidae), a specialist stem gall former. The study conducted was in a 15 km² heterogeneous, agricultural area in northeastern Bavaria. The distribution of the number of plants per patch was skewed with many more small C. arvense patches than large ones. During the five years of study, there was a 50% increase in the number of C. arvense patches, and a decrease in the mean number of plants per patch (= patch size) to less than half the patch size of the first year. Whilst patch size was randomly distributed in space, patch density showed a consistent, non‐random spatial pattern. Patch density was spatially auto‐correlated, with areas of high or low patch density having a characteristic dimension of ca. 1 km. Patch size was predictable in time and appeared to be regulated by size dependent processes, with the extinction probability of a patch being negatively correlated with its size. Correlated with the decline of C. arvense patch size during the study, the occupancy and total numbers of the herbivore U. cardui had a marked decrease, suggesting that the regional distribution of the stem gall former is not only influenced by patch number but more importantly by the mean patch size. With decreasing patch sizes, U. cardui was faced with an increasingly dynamic landscape due to higher extinction rates of small patches, although the mean distance between host plant patches decreased.     

Savanna woody encroachment is widespread across three continents

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta‐analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO₂, changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N‐fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.     

Carbon and nitrogen dynamics across a natural precipitation gradient in Patagonia,Argentina

Abstract. Both ecosystem carbon gain and nutrient availability are largely constrained by the magnitude and seasonality of precipitation in arid and semi‐arid ecosystems. We investigated the role of precipitation on ecosystem processes along an International Geosphere Biosphere Programme (IGBP) transect in temperate South America. The transect consists of a contiguous precipitation gradient in the southern region of Argentinean Patagonia (44–45° S), from 100 mm to 800 mm mean annual precipitation (MAP) and vegetation ranging from desert scrub to closed canopy forest. Gravimetric soil water content tracked changes in seasonal and annual precipitation, with a linear increase in soil water content with increasing MAP. Above‐ground net primary production (ANPP) increased linearly along the gradient of precipitation (ANPP =– 31.2 + 0.52 MAP, r²= 0.84, p= 0.028), supporting the relationship that carbon assimilation is largely controlled by available water in these sites, and was in general agreement with regional models of ANPP and rainfall. However, inorganic soil nitrogen was also highly linearly correlated with both MAP ([N] = 0.19 MAP – 32, r²= 0.96, p= 0.003) and ANPP (ANPP = 2.6 [N_inorganic]+59.4, r²= 0.79, p= 0.042), suggesting a direct control of precipitation on nitrogen turnover and an interaction with nitrogen availability in controlling carbon gain. The asynchrony of precipitation and changes in dominant vegetation may play important roles in determining the carbon‐nitrogen interactions along this rainfall gradient.