Undertaking large‐scale forest restoration to generate ecosystem services

The global community is seeking to substantially restore the world's forest cover to improve the supply of ecosystem services. However, it is not clear what type of reforestation must be used and there is a risk that the techniques used in industrial timber plantations will become the default methodology. This is unlikely to be sufficient because of the well‐known relationship between biodiversity and ecological functioning. Restoration may be achieved through natural regeneration but this may not always occur at critical locations. Ecological restoration involving species‐rich plantings might also be used but can be difficult to implement at landscape scales. I review here the consequence of planting more limited numbers of species and the effects of this on the delivery of ecosystem services. Evidence suggests many commonly sought ecosystem services—though not all—may be generated by the modest levels of species richness provided these species have appropriate traits. The literature also shows that the alpha diversity of restored forests is not the only driver of functionality and that the location and extent of any reforestation are significant as well; beta and gamma diversity may also affect functionality but these relationships remain unclear. Encouraging the adoption of even moderately diverse plantings at landscape scales and at key locations will require policies and institutions to balance the type, location, and scale of restoration and make the necessary trade‐offs between national and local aspirations. New approaches and metrics will have to be developed to monitor and assess restoration success at these larger scales.     

Biocomplexity in large tree mortality: interactions between elephant,fire and landscape in an African savanna

The vegetation dynamics of the savanna ecosystem are driven by complex interactions between biotic and abiotic factors, and thus are expected to exhibit emergent properties of biocomplexity. We explore the relative importance of static and dynamic drivers in explaining the patterns of mortality of large trees in the Kruger National Park, South Africa. Data on large trees were collected from 22 transects in April 2006, and these transects were re‐sampled in November 2008. Of the 2546 individually‐identified trees that were re‐sampled, 290 (11.4%) died in the interim. We tested several competing hypotheses with varying levels of complexity, and found that mortality of large trees was affected mainly by both static (geophysical and landscape characteristics) and dynamic (elephant damage and fire) factors that were either additive or interactive in their effects. Elephant damage was the main predictor of tree mortality, but fire also played an important role depending on the landscape type. Other static variables such as position‐on‐slope, height below canopy, and altitude had weak effects in explaining tree mortality. These results indicate that keystone features such as large trees, show differential vulnerability to mortality that is landscape‐specific. For conservation managers, this implies that the dynamic drivers (elephant and fire) of tree mortality have to be managed at the specific landscape‐level. We suggest that this emergent biocomplexity in the spatial and temporal patterns of large tree mortality is not unique to the African savannas, but is likely widespread across heterogeneous landscapes.     

Resources driving landscape‐scale distribution patterns of grazers in an African savanna

In order to effectively manage and conserve indigenous herbivores, a good understanding is needed of how resources drive their distribution patterns. This study employed a unique dataset to test a range of ecological theories and hypotheses on free‐ranging grazers. Using aerial census data collected over 14 yr across the 2 million ha Kruger National Park (South Africa), this study employs spatial autologistic regression models to explore the spatial relationships that exist between the distribution of eight indigenous grazer species and a set of resource variables. It was found that ecological theories relating to feeding guild, water‐dependence, allometric scaling, gut‐morphology and vulnerability to predation could explain most of the grazer distribution patterns observed in relation to surface‐water, forage quality, forage quantity and habitat openness. All the grazers studied were water‐dependent and occurred close to a permanent source of water in the dry season. This was ascribed to the lack of moisture in the diet of grazers during the dry season. Most ruminants’ distribution patterns were significantly associated with areas of high forage quality whereas hind‐gut fermentors were neutral towards forage quality. Average forage quantity was not a significant predictor of long‐term, landscape‐scale distribution patterns for any of the grazer species studied. Most small‐ and medium‐bodied grazers preferred open habitats above closed habitats, probably due to higher visibility and a lower predation risk. Large‐bodied grazers did not bias their distribution patterns towards open habitats. The way in which grazers distribute themselves with respect to different resources can potentially inform management actions on appropriate scales.     

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.     

Natural regeneration in the context of large‐scale forest and landscape restoration in the tropics

Large‐scale and long‐term restoration efforts are urgently needed to reverse historical global trends of deforestation and forest degradation in the tropics. Restoration of forests within landscapes offers multiple social, economic, and environmental benefits that enhance lives of local people, mitigate effects of climate change, increase food security, and safeguard soil and water resources. Despite rapidly growing knowledge regarding the extent and feasibility of natural regeneration and the environmental and economic benefits of naturally regenerating forests in the tropics, tree planting remains the major focus of restoration programs. Natural regeneration is often ignored as a viable land‐use option. Here, we assemble a set of 16 original papers that provide an overview of the ecological, economic, and social dimensions of forest and landscape restoration (FLR), a relatively new approach to forest restoration that aims to regain ecological integrity and enhance human well‐being in deforested or degraded forest landscapes. The papers describe how spontaneous (passive) and assisted natural regeneration can contribute to achieving multiple social and ecological benefits. Forest and landscape restoration is centered on the people who live and work in the landscape and whose livelihoods will benefit and diversify through restoration activities inside and outside of farms. Given the scale of degraded forestland and the need to mitigate climate change and meet human development needs in the tropics, harnessing the potential of natural regeneration will play an essential role in achieving the ambitious goals that motivate global restoration initiatives.     

A large‐scale latitudinal pattern of life‐history traits in a strictly univoltine damselfly

1. Variation in thermal conditions and season length along latitudinal gradients affect body size‐related traits over different life stages. Selection is expected to optimise these size traits in response to the costs and benefits. 2. Egg, hatchling, larval and adult size in males and females were estimated along a latitudinal gradient of 2730 km across Europe in the univoltine damselfly Lestes sponsa, using a combination of field‐collection and laboratory‐rearing experiments. In the laboratory, individuals were grown in temperatures and photoperiod simulating those at the latitude of origin, and in common‐garden conditions. 3. The size of adults sampled in nature was negatively correlated with latitude. In all populations the females were larger than the males. Results from simulated and common‐garden rearing experiments supported this pattern of size difference across latitudes and between sexes, suggesting a genetic component for the latitudinal size trend and female‐biased size dimorphism. In contrast, hatchling size showed a positive relationship with latitude, but egg size, although differing between latitudes, showed no such relationship. 4. The results support a converse Bergmann cline, i.e. a negative body size cline towards the north. This negative cline in body size is probably driven by progressively stronger seasonal time and temperature constraints towards the higher latitudes and by the obligate univoltine life cycle of L. sponsa. As egg size showed no relationship with latitude, other environmental factors besides temperature, such as desiccation risk, probably affect this trait.     

Effects of fire on bird diversity and abundance in an East African savanna

Fire is an important determinant of many aspects of savanna ecosystem structure and function. However, relatively little is known about the effects of fire on faunal biodiversity in savannas. We conducted a short‐term study to examine the effects of a replicated experimental burn on bird diversity and abundance in savanna habitat of central Kenya. Twenty‐two months after the burn, Shannon diversity of birds was 32% higher on plots that had been burned compared with paired control plots. We observed no significant effects of burning on total bird abundance or species richness. Several families of birds were found only on plots that had been burned; one species, the rattling cisticola (Cisticola chiniana), was found only on unburned plots. Shrub canopy area was negatively correlated with bird diversity on each plot, and highly correlated with grass height and the abundance of orthopterans. Our results suggest that the highest landscape‐level bird diversity might be obtained through a mosaic of burned and unburned patches. This is also most likely to approximate the historical state of bird diversity in this habitat, because patchy fires have been an important natural disturbance in tropical ecosystems for millennia.     

Do rabbits eat voles? Apparent competition,habitat heterogeneity and large‐scale coexistence under mink predation

Habitat heterogeneity is predicted to profoundly influence the dynamics of indirect interspecific interactions; however, despite potentially significant consequences for multi-species persistence, this remains almost completely unexplored in large-scale natural landscapes. Moreover, how spatial habitat heterogeneity affects the persistence of interacting invasive and native species is also poorly understood. Here we show how the persistence of a native prey (water vole, Arvicola terrestris ) is determined by the spatial distribution of an invasive prey (European rabbit, Oryctolagus cuniculus ) and directly infer how this is defined by the mobility of a shared invasive predator (American mink, Neovison vison ). This study uniquely demonstrates that variation in habitat connectivity in large-scale natural landscapes creates spatial asynchrony, enabling coexistence between apparent competitive native and invasive species. These findings highlight that unexpected interactions may be involved in species declines, and also that in such cases habitat heterogeneity should be considered in wildlife management decisions.     

The importance of post‐fire regrowth for sable antelope in a Southern African savanna

Burning is commonly used in savannas to stimulate grass regrowth for grazing ungulates. We recorded the relative use of burns occurring at different stages in the seasonal cycle, as well as in different regions of the landscape by two herds of sable. We also recorded behavioural measures of foraging efficiency and faecal nutrient contents as an indication of nutrient gains. Sable consistently concentrated their grazing on burned areas provided there was sufficient green regrowth during the dry season. In these circumstances they grazed for longer per feeding station, showed a slower step rate while foraging, and shorter between‐patch moves, and a higher probability of encountering acceptable food per step taken while foraging than on unburnt areas. In the year when only a burn with insufficient regrowth was available, sable continued to forage in the area that had been burned during the previous year. Faecal crude protein was substantially higher at the end of the dry season in the year when burned areas were utilized. Accordingly early dry season fires can be important in helping sable bridge the nutritional limitations posed by the dry season, provided sufficient soil moisture remains to promote adequate grass regrowth.     

Multi‐scale ant diversity in savanna woodlands: an intercontinental comparison

Ecological patterns and processes are highly scale‐dependent, but few studies have used standardized methodology to examine how scale dependency varies across continents. This paper examines scale dependency in comparative ant species richness and turnover in savannas of Australia and Brazil, which are well‐matched climatically but whose ant faunas have contrasting biogeographic origins. The study was conducted in savanna woodland near Darwin in northern Australia and Uberlândia in central Brazil. The sampling design consisted of eight 400‐m line transects, four in each continent, with eight pitfall traps located on and around each of 20 trees evenly spaced along each transect. Ant richness and species turnover were compared at three spatial scales: pitfalls associated with a tree, trees within a transect and transects within a savanna. The composition of the Australian and Brazilian savanna ant faunas was broadly similar at the subfamily level, despite the very low proportion of shared genera and species. The ground and arboreal ant faunas were very distinct from each other in both savannas, but especially in Brazil. Overall ant abundance was almost three times higher in Australia than in Brazil, both on the ground and on vegetation, but overall species richness was higher in Brazil (150 species) than in Australia (93). There was no significant difference in the mean number of species per pitfall trap, but the mean species richness was significantly higher in Brazil than in Australia at both the tree and transect scales. We attribute these scale‐dependent intercontinental differences to biogeographical and historical factors in Brazil that have led to a large regional pool of arboreal species of rainforest origin. Our study underlines the importance of biogeographical context when conducting comparative analyses of community structure across biogeographical scales, and highlights the importance of process acting at regional scales in determining species richness in ant communities.     

Responses of vertebrates to pastoralism,military land use and landscape position in an Australian tropical savanna

Despite the hegemony of pastoralism over most of Australia’s tropical savannas, its impacts upon biodiversity are poorly known. There is even less knowledge about the impacts of military training, a recent, but rapidly expanding, alternative land use. We compare impacts of these land uses upon mammals, birds, reptiles and frogs at a site in north‐eastern Australia, with sampling from 24 quadrats stratified by four landscape positions (upper slope to riparian) and three current land‐use types (pastoralism, military training and undisturbed). Prior to exclusion in 1967, the whole study area had been subjected to grazing over the course of approximately 100 years, so differences observed strictly reflect responses to changed land use (largely cessation from grazing) over the period of 32 years subsequent to the imposition of the present regime. The four classes of vertebrates showed contrasting responses. Frog distribution was unrelated to land use, but strongly associated with landscape position. Reptiles showed a very strong response to land‐use type but not to landscape position. The total abundance and richness of reptiles was greater in ungrazed (i.e. military and undisturbed) than in grazed quadrats. The total abundance and species richness of birds varied strongly with landscape position but was unrelated to land use. However, many individual bird species showed significant responses to land‐use type, and bird species composition was significantly related to both land‐use type and landscape position. The richness of the mammal fauna was weakly related to landscape position and not related to land‐use type. A few individual mammal species showed significant responses to either or both factors, but mammal species composition was significantly (albeit weakly) related only to land‐use type. With due regard to some interpretative constraints in the study design, and the history of the site prior to this study, these results suggest that pastoralism leads to a substantial rearrangement of the vertebrate fauna, and particularly so for reptiles and those mammals and birds associated with the ground and understorey layers. Given the extent of pastoralism across the tropical savannas, these results suggest that this industry has contributed to major and widespread change in the savanna fauna. In contrast to pastoralism, military land use (at least at the relatively low intensity examined here) produced little change in vertebrate assemblages.     

Weak impact of fine‐scale landscape heterogeneity on evolutionary potential in Arabidopsis lyrata

Microhabitat heterogeneity can lead to fine‐scale local adaptation when gene flow is restricted, which may be important for the maintenance of genetic variation within populations. This study tested whether microhabitat heterogeneity was associated with trait differences in a population of Arabidopsis lyrata and studied its impact on the genetic variance–covariance (G) matrix. Maternal seed families were collected from dune tops and bottoms, two microhabitats known to vary significantly in water availability. In a common garden experiment, replicate individuals per family were raised under wet and dry conditions, and physiological, morphological and life‐history traits were assessed. Plants from the two microenvironments differed in their response to treatment in two performance components, in stomata density and most strongly in flowering time. Under wet conditions, plants originating from dune bottoms flowered 4 weeks earlier than those from dune tops. Only one of three G‐matrix comparisons revealed that habitat heterogeneity and evolutionary potential were positively linked. The number of independent trait dimensions was larger in the entire population than within subpopulations separated by microhabitat under wet conditions. However, the size of the G‐matrix was no larger in the entire population than within subpopulations separated by microhabitat, and trait correlation structure between microhabitats and treatments was not significantly different. These results indicate that fine‐scale habitat heterogeneity likely led to local adaptation, which weakly affected levels of across‐trait genetic variation.     

Fine‐scale group structure and demography of African savanna elephants recolonizing lands outside protected areas

Aim Dispersal is a critical component of animal ecology that is poorly understood for most species. In particular, savanna elephants (Loxodonta africana) have been studied for decades in national parks across Africa, but little is known about their dispersal into new or unused habitats or their population dynamics in human‐dominated landscapes. We capitalized on a natural dispersal event of savanna elephants recolonizing communal land in southern Kenya to document their demographic characteristics and genetic relationships. Location Rift Valley province of Kenya. Methods We collected faecal samples and used genetic methods to identify individuals, estimate the sex ratio and evaluate the patterns of relatedness within the female groups and male aggregations. We also measured dung bolus circumference to assign age classes to individuals and estimate the age structure. Results We identified 112 individuals with a sex ratio not different from one (1.32:1.00). The age structure was skewed towards younger elephants (71%), suggesting the potential for rapid growth from reproduction. We detected significantly higher kinship levels within female groups (R = 0.124 ± 0.023), suggesting that family groups colonized the site, but found little support for higher‐order genetic relationships among female groups. Males detected together were unrelated (R = 0.003 ± 0.030). Main conclusions Our results suggest that highly social mammals, such as savanna elephants, disperse into unoccupied habitat as family groups and that a young demographic structure and a large number of males might be expected in establishing populations. These findings highlight the potential value of indirect, non‐invasive methods for assessing elephant herd and demographic characteristics when direct observations are difficult.     

Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover

When tropical and sub-tropical ecosystems burn, considerable amounts of N present in the biomass fuel may be released. This ultimately results in a loss of fixed N to the atmosphere. It is often assumed that this volatilization loss of N with frequent fire will result in a reduction of plant-available N and total system N. By changing the amount of woody biomass fire may, however, also have indirect effects on N and C dynamics. Here we consider the effects of 50 years of frequent fire on total soil N and soil organic C (SOC) and total soil N in a mesic savanna in the Kruger National Park, South Africa. We also determine how changes in woody biomass may affect total soil N and SOC. We measured soil and fine root N and C concentrations as well as total soil N and SOC pools in four burning treatments, including fire exclusion, of a long-term fire experiment. Our results show that regardless of soil depth, fire treatment had no significant effect on total soil N and SOC. Our results also show that under trees total soil N and SOC concentrations of the surface soil increase, and pools of N and SOC increase to a depth of 7 cm. However, the extent to which soil N and C dynamics differed under canopies and away from canopies was dependent on fire treatment. Our results show that the effect of fire on soil N and C is mediated both through the indirect effect of changes in woody cover and the direct effects of fire (volatilization losses of nutrients). We suggest that woody thickening in this mesic savanna will have pronounced effects on long-term N and C dynamics.     

Stream order controls geomorphic heterogeneity and plant distribution in a savanna landscape

We posed the question: does viewing a savanna as a network of streams linked to a matrix of terrestrial hillslopes provide a useful framework to research and understand plant distribution in these landscapes? Our study area, the Phugwane River network, lies in the semi‐arid savanna of Kruger National Park, South Africa. We examined changes in hillslope geomorphology from first‐, third‐ and fifth‐order hillslopes with regression equations. The distribution of geomorphic boundaries was enumerated by moving window analysis and the relationship between geomorphology and plant distribution was explored through ordination. First‐order hillslopes had a simple geomorphology, fewer geomorphic boundaries and a relatively homogeneous plant assemblage. By contrast, fifth‐order hillslopes were more complex in geomorphology, with more boundaries and a relatively heterogeneous vegetation pattern. Stream order classification of a savanna drainage network resulted in landscape units distinguishable by geomorphology, geomorphic boundaries and vegetation pattern. Therefore, the drainage network is a useful template to expose and organize the complexity in savanna landscapes into easily managed and researched units. This perspective should inform a shift from single‐scale phytosociological views of homogeneous vegetation units towards multi‐scale conceptualizations of savannas as water dependent ecosystems.     

Cooperation,collective action,and the archeology of large‐scale societies

Archeologists investigating the emergence of large‐scale societies in the past have renewed interest in examining the dynamics of cooperation as a means of understanding societal change and organizational variability within human groups over time. Unlike earlier approaches to these issues, which used models designated voluntaristic or managerial, contemporary research articulates more explicitly with frameworks for cooperation and collective action used in other fields, thereby facilitating empirical testing through better definition of the costs, benefits, and social mechanisms associated with success or failure in coordinated group action. Current scholarship is nevertheless bifurcated along lines of epistemology and scale, which is understandable but problematic for forging a broader, more transdisciplinary field of cooperation studies. Here, we point to some areas of potential overlap by reviewing archeological research that places the dynamics of social cooperation and competition in the foreground of the emergence of large‐scale societies, which we define as those having larger populations, greater concentrations of political power, and higher degrees of social inequality. We focus on key issues involving the communal‐resource management of subsistence and other economic goods, as well as the revenue flows that undergird political institutions. Drawing on archeological cases from across the globe, with greater detail from our area of expertise in Mesoamerica, we offer suggestions for strengthening analytical methods and generating more transdisciplinary research programs that address human societies across scalar and temporal spectra.     

The co‐evolution of social institutions,demography, and large‐scale human cooperation

Human cooperation is typically coordinated by institutions, which determine the outcome structure of the social interactions individuals engage in. Explaining the Neolithic transition from small‐ to large‐scale societies involves understanding how these institutions co‐evolve with demography. We study this using a demographically explicit model of institution formation in a patch‐structured population. Each patch supports both social and asocial niches. Social individuals create an institution, at a cost to themselves, by negotiating how much of the costly public good provided by cooperators is invested into sanctioning defectors. The remainder of their public good is invested in technology that increases carrying capacity, such as irrigation systems. We show that social individuals can invade a population of asocials, and form institutions that support high levels of cooperation. We then demonstrate conditions where the co‐evolution of cooperation, institutions, and demographic carrying capacity creates a transition from small‐ to large‐scale social groups.