Root trait variation in African savannas

Plant and Soil - While patterns of variation in woody plant aboveground traits related to disturbance and resource availability in savanna ecosystems are fairly well understood, dimensions of...     

Biogeographical patterns in Artomyces pyxidatus

Artomyces pyxidatus (Auriscalpiaceae) is a lignicolous, coralloid basidiomycete found throughout temperate regions of the Northern Hemisphere. Previous studies established that populations from the eastern United States, Sweden, and China were conspecific based on mating compatibility and enzyme profiles. In this study, mating compatibility was extended to include collections from Russia, Costa Rica, Mexico, and Utah. The molecular diversity of A. pyxidatus was examined by DNA sequence and restriction site analyses of the nuclear ribosomal internally transcribed spacer region (ITS1-5.8S-ITS2). A phylogenetic analysis of twelve isolates based on ITS sequences revealed a broad geographical pattern in which Eurasian isolates comprise a sister clade to North American isolates. North American isolates appear to be further subdivided into northeastern and southwestern clades. A survey of 255 A. pyxidatus isolates using restriction enzymes revealed variable RFLP patterns that follow similar geographical patterns.     

Local versus landscape-scale effects of anthropogenic land-use on forest species richness

The study investigated the effects of human-induced landscape patterns on species richness in forests. For 80 plots of fixed size, we measured human disturbance (categorized as urban/industrial and agricultural land areas), at ‘local’ and ‘landscape’ scale (500 m and 2500 m radius from each plot, respectively), the distance from the forest edge, and the size and shape of the woody patch. By using GLM, we analyzed the effects of disturbance and patch-based measures on both total species richness and the richness of a group of specialist species (i.e. the ‘ancient forest species’), representing more specific forest features. Patterns of local species richness were sensitive to the structure and composition of the surrounding landscape. Among the landscape components taken into account, urban/industrial land areas turned out as the most threatening factor for both total species richness and the richness of the ancient forest species. However, the best models evidenced a different intensity of the response to the same disturbance category as well as a different pool of significant variables for the two groups of species. The use of groups of species, such as the ancient forest species pool, that are functionally related and have similar ecological requirements, may represent an effective solution for monitoring forest dynamics under the effects of external factors. The approach of relating local assessment of species richness, and in particular of the ancient forest species pool, to land-use patterns may play an important role for the science-policy interface by supporting and strengthening conservation and regional planning decision making.     

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

Diversity and occurrence of herbaceous communities in West African savannas in relation to climate,land use and habitat

In the face of the current changes in land use and climate as well as habitat destruction, it is important to study herbaceous vegetation as an indicator of changes occurring in savanna ecosystems. We investigated the effects of climate, land use and habitat, both alone and in combination, on the diversity and occurrence of West African savanna herbaceous plant communities. Floristic data and environmental variables were sampled in Burkina Faso and subjected to ordination and indicator species analysis to explore the variation in nine vegetation types. Regression analyses showed that climate, land use, humidity gradient, soil fraction and vegetation structure discriminate herbaceous plant communities. Climate, habitat and their interaction had the greatest effect on the occurrence of these communities. Changes in species richness of the studied communities were mainly due to climate, land use and their interaction, which were more important for increasing rather than decreasing diversity. In all cases, climate conditions remained the most important environmental factor driving vegetation variation in West African savannas. Beside this, the effects of habitat degradation in interaction with land use and climatic conditions indicate land use to be a threat for the diversity of the herbaceous vegetation.     

Superpredation patterns in four large European raptors

Predatory interactions among top predators, like superpredation or intraguild predation (IGP), can influence community structure. Diurnal raptors occupy high trophic levels in terrestrial food webs, and thus can regulate the presence of mesopredators. We studied superpredation (the killing and eating of another predator) in four large European raptors. We gathered 121 dietary studies, totalling 161,456 prey for the Goshawk Accipiter gentilis L., Golden Eagle Aquila chrysaetos L., Bonelli’s Eagle Aquila fasciata Vieillot, and Eagle Owl Bubo bubo L. Results showed that superpredation: (1) is a widespread interaction in large raptors, but it can vary according to the top predator species; (2) is not an important energetic resource for large raptors, but rather seems mostly related to diet diversification when the main prey decreases; (3) is spatially clustered reflecting habitat heterogeneity, but shows no temporal or large-scale spatial trends; and (4) it is associated with lower breeding success of the top predator species. These findings support the food stress hypothesis as the main driving force behind increases in superpredation and IGP in raptors, with the decrease in breeding performance as a side effect. Superpredation by large raptors deserves future research to understand its effects on mesopredators, because on one hand it might contribute to promote biodiversity, while on the other hand, it can sometimes represent an additional risk for small populations of endangered mesopredators.     

Humans and elephants as treefall drivers in African savannas

Humans have played a major role in altering savanna structure and function, and growing land‐use pressure will only increase their influence on woody cover. Yet humans are often overlooked as ecological components. Both humans and the African elephant Loxodonta africana alter woody vegetation in savannas through removal of large trees and activities that may increase shrub cover. Interactive effects of both humans and elephants with fire may also alter vegetation structure and composition. Here we capitalize on a macroscale experimental opportunity – brought about by the juxtaposition of an elephant‐mediated landscape, human‐utilized communal harvesting lands and a nature reserve fenced off from both humans and elephants – to investigate the influence of humans and elephants on height‐specific treefall dynamics. We surveyed 6812 ha using repeat, airborne high resolution Light Detection and Ranging (LiDAR) to track the fate of 453 685 tree canopies over two years. Human‐mediated biennial treefall rates were 2–3.5 fold higher than the background treefall rate of 1.5% treefall ha^–1, while elephant‐mediated treefall rates were 5 times higher at 7.6% treefall ha^–1 than the control site. Model predictors of treefall revealed that human or elephant presence was the most important variable, followed by the interaction between geology and fire frequency. Treefall patterns were spatially heterogeneous with elephant‐driven treefall associated with geology and surface water, while human patterns were related to perceived ease of access to wood harvesting areas and settlement expansion. Our results show humans and elephants utilize all height classes of woody vegetation, and that large tree shortages in a heavily utilized communal land has transferred treefall occurrence to shorter vegetation. Elephant‐ and human‐dominated landscapes are tied to interactive effects that may hinder tree seedling survival which, combined with tree loss in the landscape, may compromise woodland sustainability.     

Hunting flight speeds of five southern African raptors

The flight speeds of hunting falconry birds were determined using global positioning system data loggers. Until now, the hunting flight speed of African raptors has not been directly measured. We predicted that hunting flight speeds would differ between species and that flight dynamics, such as altitude, and bird morphology, particularly wing surface area, would influence maximum and mean flight speeds. This study considered five African raptor species, which included two long-wing species, Lanner Falcon Falco biarmicus and Peregrine Falcon F. peregrinus, one short-wing species, Black Sparrowhawk Accipiter melanoleucus, and two broad-wing species, African Hawk-eagle Aquila spilogaster and Jackal Buzzard Buteo rufofuscus. Maximum and mean hunt speeds differed significantly between the long- and short-wing species. There was no difference in acceleration or deceleration rates between these species, but this could be due to small sample sizes. There was a significant positive correlation between maximum hunt speed and maximum flight height for the long-wing species. Maximum and mean flight speeds were significantly negatively correlated with wing area for all five species in this study. However, following phylogenetic correction, no significant relationship between wing area and maximum hunt speeds was found. This study presents baseline data of hunting speeds in African raptors and further highlights the importance of inter-species variation, which can provide accuracy to flight speed models and the understanding of hunting strategies.     

Elephants limit aboveground carbon gains in African savannas

Understanding the drivers of vegetation carbon dynamics is essential for climate change mitigation and effective policy formulation. However, most efforts focus on abiotic drivers of plant biomass change, with little consideration for functional roles performed by animals, particularly at landscape scales. We combined repeat airborne Light Detection and Ranging with measurements of elephant densities, abiotic factors, and exclusion experiments to determine the relative importance of drivers of change in aboveground woody vegetation carbon stocks in Kruger National Park, South Africa. Despite a growing elephant population, aboveground carbon density (ACD) increased across most of the landscape over the 6‐year study period, but at fine scales, bull elephant density was the most important factor determining carbon stock change, with ACD losses recorded only where bull densities exceeded 0.5 bulls/km². Effects of bull elephants were, however, spatially restricted and landscape dependent, being especially pronounced along rivers, at mid‐elevations, and on steeper slopes. In contrast, elephant herds and abiotic drivers had a comparatively small influence on the direction or magnitude of carbon stock change. Our findings demonstrate that animals can have a substantive influence on regional‐scale carbon dynamics and warrant consideration in carbon cycling models and policy formulation aimed at carbon management and climate change mitigation.     

A continental-scale analysis of tree cover in African savannas

Aim We present a continental‐scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. Location African savannas. Methods Tree cover is represented as a resource‐driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi‐empirical modelling and information theory to identify the best models. Results Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. Main conclusions The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These ‘perturbation’ effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi‐arid and mesic savannas (400–1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600–2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24–34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.     

Biogeographical patterns of endemic terrestrial Afrotropical birds

Biogeographical zones are described for terrestrial bird species endemic to the Afrotropics using up‐to‐date distributional data and multivariate statistical techniques. This provides an objective basis for a hierarchy of subregions, provinces and districts, based on a set of rules. Results are compared to previous studies at continental and regional scales. Biogeographical zones for passerines and non‐passerines are compared and found to be similar. Peaks of species richness and narrow endemism are described for the six major subdivisions (subregions) identified by the cluster analysis. Coincidence of peaks of species richness and narrow endemism is found to be low, such that areas selected to represent high species richness tallies will often fail to represent narrow endemics. Strong regionalization of Afrotropical birds indicates the need to use a biogeographical framework in conservation priority setting exercises to ensure that unique, but species‐poor, avifaunas are not neglected.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Influential environmental gradients and patterns of fish assemblages in a West African basin

Twenty-two sites were monitored in the Lower Sassandra River Basin (Côte d’Ivoire, West Africa) to examine patterns in fish species assemblage structure along environmental gradients. Hierarchical clustering of the species presence–absence data identified four types of assemblages corresponding to the man-made Lake Buyo, the main channel, and the two major tributaries. Canonical Correspondence Analysis (CCA) indicated that canopy closure, leaves-wood, aquatic plants, temperature, width, total dissolved solids and depth were the environmental factors most strongly correlated with variation in the fish assemblages. Five species: Papyrocranus afer, Micralestes elongatus, Parailia pellucida, Tilapia busumana and T. mariae were reported from the basin for the first time. The investigation of the variation in species richness led to the assessment of a major anthropogenic disturbance induced by a palm oil factory. In conjunction with the fact that the Sassandra River hosts an endemic species, Synodontis koensis, it was concluded that this basin is of high conservation priority.     

Comparative foraging adaptations of small raptors in a dense African savanna

Our aim was to investigate how ecologically similar species with different evolutionary adaptations may coexist in a moist Guinean savanna, which is the optimal habitat of the Lizard Buzzard Kaupifalco monogrammicus whereas the Grey Kestrel Falco ardosiaceus and the Grasshopper Buzzard Butastur rufipennis are of more Sudanian origin. Activity budgets, foraging behaviour and success were assessed by focal sampling and were related to habitat parameters and prey availability. Each species had its own daily foraging pattern which seemed to be related to the activity patterns of their main prey, grasshoppers or lizards. Interspecific habitat segregation reflected their morphological and behavioural adaptations and their respective tolerance to grass height, the most important factor in habitat selection. The resident Lizard Buzzard selected the densest woodlands, whereas the Grey Kestrel was restricted to the most open areas and the dry season migrant Grasshopper Buzzard to intermediate, recently burnt, savanna types. The main foraging variables (perch height, inter-perch distance and frequency of moves) differed significantly between species. Maximum detection distances increased with perch height but decreased with increasing grass cover. Perch height was lower when the grass became too high. When attack rates increased, birds tended to move more often, but for shorter distances, and stayed longer within restricted patches. The highly insectivorous Grasshopper Buzzard had the highest and least variable attack rates, negatively correlated with grass height. It exhibited a behaviour typical of a migrant—superficial and opportunistic exploitation of superabundant, easily accessible, and seasonal food sources. Conversely, the residents were associated with denser, more stable habitats and larger, less abundant, but more predictable prey.     

Ecosystem‐scale effects of megafauna in African savannas

Natural protected areas are critically important in the effort to prevent large‐scale megafaunal extinctions caused by hunting and habitat degradation. Yet such protection can lead to rapid increases in megafauna populations. Understanding ecosystem‐scale responses of vegetation to changing megafaunal populations, such as the case of the African elephant Loxodonta africana in savannas, requires large‐scale, high‐resolution monitoring over time. From 2008 to 2014, we repeatedly surveyed the fate of more than 10.4 million woody plant canopies throughout the Kruger National Park, South Africa using airborne Light Detection and Ranging (LiDAR), to determine the relative importance of multiple environmental, biotic and management factors affecting treefall rates and patterns. We report a mean biennial treefall rate of 8 trees or 12% ha^?1, but with heterogeneous patterns of loss in both space and time. Throughout Kruger, the influence of elephant density on treefall was matched only by spatial variation in soils and elevation, and all three factors co‐dominated park‐wide treefall patterns. Elephant density was up to two times more influential than fire frequency in determining treefall rates, and this pattern was most pronounced for taller trees (> 2 m in height). Our results suggest that confining megafauna populations to protected areas, or reintroducing them into former or new habitat, can greatly alter the structure and functioning of the host ecosystem. Conservation strategies will need to accommodate and manage these massive ecological changes in the effort to save megafauna from extinction, without compromising system functionality.     

Biogeographical determinants for total and endemic species richness in a continental archipelago

We examined the relationship between plant species richness and biogeographical variables (island area, island maximum elevation, distance from nearest inhabited island, distance from nearest mainland) using a data set comprising 201 islands of the Aegean archipelago. We found that endemic species richness was strongly correlated to total species richness. Single-island endemic species richness was most strongly correlated to island maximum elevation, and then to island area, with an apparent small island effect for islands smaller than 47 km². Total species richness was most strongly correlated to island area (with no apparent small island effect), and less strongly correlated to island maximum elevation. Distance from the mainland or other inhabited islands displayed limited predictive value in our data set. The slope of the relationship between species richness and geographical factors (island area, elevation, distance from island/mainland) was steeper for endemic species richness than for total richness. Finally, the different scales of endemicity (single-island endemics, island group endemics and Aegean regional endemics) displayed similar qualitative trends and only differed quantitatively. Thus, we conclude that different biogeographical factors act as drivers for total species richness than for endemic species richness.