Rainfall and topo-edaphic influences on woody community phenology in South African savannas

Woody community phenology was studied in the central lowveld, South Africa, over a twelve month period at three sites along a rainfall gradient, with both toplands and bottomlands sampled at each site. Each month, individual plants, in replicated samples, were scored into a number of categories describing their phenological state. Position on the rainfall gradient influenced: (1) onset and magnitude of leaf emergence, (2) onset and duration of mature leaves, and (3) the proportion of leafless trees. Generally, the moist site demonstrated earlier leaf growth than the intermediate or arid sites. Emergent and mature leaves were recorded earlier, and in the case of mature leaves, retained longer. Overall, there was a lower proportion of leafless trees during the dry season at the moist site, followed by the semi-arid site, followed by the arid site. Differences with respect to catenal position were evident for the proportion of trees in winter with mature leaves, and the proportion of trees with senescent leaves. Bottomlands had a greater proportion of trees with leaves during winter, but a lower proportion of trees recorded with senescent leaves. Both of these findings were a result of the greater proportion of evergreen species in bottomlands, as well as increased leaf retention by the deciduous species. Phenological activity of leaves was related to plant stem size. In particular, there was greater leaf retention during the dry period by small stems, relative to large stems.     

Zooplankton body size and community structure: Effects of thermal and toxicant stress

Episodic heat waves and an increase in pesticide use are widely cited as consequences of climatic warming. Recent studies show that these stressors often cause declines in the mean body size of zooplankton. Results from laboratory and field studies, as well as observations from both thermal- or toxicant-stressed natural systems, show (1) reductions in mean body size within stressed populations, or (2) changes in community composition that favor small-bodied over large-bodied species. During the past decade, it has become widely accepted that a shift in zooplankton body size can dramatically affect water clarity, rates of nutrient regeneration and fish abundances. Thus, climatic warming and associated change in pesticide use has the potential to cause striking change in the structure and functioning of temperate-zone lakes.     

Fire season affects size and architecture of Colophospermum mopane in southern African savannas

Wildfires may be started naturally by lightning or artificially by humans. In the savanna regions of southern Africa, lightning fires tend to occur at the start of the wet season, during October and November, while anthropogenic fires are usually started during the dry season, between July and August. A long-term field manipulation experiment initiated in the Kruger National Park in 1952 was used to explore whether this seasonal divergence affects tree abundance, spatial pattern, size and architecture. After 44 years of prescribed burning treatments that simulated the seasonal incidence of lightning and anthropogenic fires, mean densities of the locally-dominant shrub, Colophospermum mopane, were 638 and 500 trees ha^–1 respectively. Trees in burnt plots had aggregated distributions while trees in unburnt plots had random distributions. Significant differences (p < 0.001) were recorded in a range of morphological parameters including tree height, canopy diameter, mean stem circumference and number of stems. The incidence of resprouting also differed significantly between treatments, with burnt trees containing a high proportion of coppiced stems. The differences in tree size and architecture between the mid-dry season and early-wet season burning plots suggest that anthropogenic fires applied during July and August cannot substitute for a natural lightning fire regime. Anthropogenic fire yields a landscape that is shorter, more scrubby and populated by numerous coppiced shrubs than the landscape generated by natural lightning fire conditions.     

Integrating environmental and spatial processes in ecological community dynamics

The processes controlling the abundances of species across multiple sites form the cornerstone of modern ecology. In these metacommunities, the relative importance of local environmental and regional spatial processes is currently hotly debated, especially in terms of the validity of neutral model. I collected 158 published data sets with information on community structure, environmental and spatial variables. I showed that approximately 50% of the variation in community composition is explained by both environmental and spatial variables. The majority of the data sets were structured by species-sorting dynamics (SS), followed by a combination of SS and mass-effect dynamics. While neutral processes were the only structuring process in 8% of the collected natural communities, disregarding neutral dispersal processes would result in missing important patterns in 37% of the studied communities. Moreover, metacommunity characteristics such as dispersal type, habitat type and spatial scale predicted part of the detected variation in metacommunity structure.     

Changes in otolith area: sensory area ratio with body size and depth

Synopsis The ratio between the sensory areas of the sulcus acusticus and the sagittal otolith (S : O ratio) in the species Merluccius capensis and M. paradoxus was analyzed using a digital image processing system. Sagittal growth in both species is negatively allometric with respect to total fish length but is more negative in the deeper-dwelling species, Merluccius paradoxus. In contrast, the sulcus acusticus undergoes a relative increase in size, that is, the S : O ratio increases with fish length. There was a clear relationship between the increase in the S : 0 ratio and depth.     

Abundance, diversity and body size: patterns from a grassland arthropod community

1. The empirical relationships among body size, species richness and number of individuals may give insight into the factors controlling species diversity and the relative abundances of species. To determine these relationships, we sampled the arthropods of grasslands and savannahs at Cedar Creek, MN using sweep nets (90 525 individuals of 1225 species) and pitfall traps (12 721 individuals of 92 species). Specimens were identified, enumerated and measured to determine body size.
2. Both overall and within abundant taxonomic orders, species richness and numbers of individuals peaked at body sizes intermediate for each group. Evolution could create unimodal diversity patterns by random diversification around an ancestral body size or from size-dependent fitness differences. Local processes such as competition or predation could also create unimodal diversity distributions.
3. The average body size of a species depended significantly on its taxonomic order, but on contemporary trophic role only within the context of taxonomic order.
4. Species richness ( S _i) within size classes was related to the number of individuals ( I _i) as S _i =  I i^0·5. This relationship held across a 100 000-fold range of body sizes. Within size classes, abundance distributions of size classes were all similar power functions. A general rule of resource division, together with similar minimum population sizes, is sufficient to generate the relationship between species richness and number of individuals.
5. Smaller bodied species had slightly shallower abundance distributions and may, in general, persist at lower densities than larger species.
6. Our results suggest there may be fewer undescribed small arthropod species than previously thought and that most undescribed species will be smaller than arthropods.     

Alien plant invasions in tropical and sub-tropical savannas: patterns,processes and prospects

Biological invasions affect virtually all ecosystems on earth, but the degree to which different regions and biomes are invaded, and the quality of information from different regions, varies greatly. A large body of literature exists on the invasion of savannas in the Neotropics and northern Australia where invasive plants, especially African grasses, have had major impacts. Less has been published on plant invasions in African savannas, except for those in South Africa. Negative impacts due to plant invasions in African savannas appear to be less severe than in other regions at present. As savannas cover about 60% of the continent, with tens of millions of people relying on the services they provide, it is timely to assess the current status of invasions as a threat to these ecosystems. We reviewed the literature, contrasting the African situation with that of Neotropical and Australian savannas. A number of drivers and explanatory factors of plant invasions in savannas have been described, mostly from the Neotropics and Australia. These include herbivore presence, residence time, intentional introductions for pasture improvements, fire regimes, the physiology of the introduced species, and anthropogenic disturbance. After comparing these drivers across the three regions, we suggest that the lower extent of alien plant invasions in African savannas is largely attributable to: (1) significantly lower rates of intentional plant introductions and widespread plantings (until recently); (2) the role of large mammalian herbivores in these ecosystems; (3) historical and biogeographical issues relating to the regions of origin of introduced species; and (4) the adaptation of African systems to fire. We discuss how changing conditions in the three regions are likely to affect plant invasions in the future.     

Functional significance of ungulate diversity in African savannas and the ecological implications of the spread of pastoralism

The African savanna biome supports a higher diversity of ungulate species than is found in any other biome or continent. This exceptional faunal diversity and herbivore biomass density is directly linked to the high spatial heterogeneity of African savanna ecosystems. The dependence of herbivore dietary tolerance on body size translates into important size-related differences between savanna ungulate species in terms of habitat specificity, geographical range, and the share of community resources exploited. Intact savanna ungulate communities, with species distributed across body size classes and feeding guilds (grazer/browser), have strong regulatory influences on savanna ecosystem structure and function. Replacement with livestock systems of low diversity and high biomass density within a narrow body size range has occurred through the removal of competitors, pathogens, and predators, and the widespread provisioning of water. Overgrazing by livestock, coupled with episodic droughts, has caused widespread rangeland degradation and loss of floristic and faunal diversity which, by current models, is unlikely to recover to 'climax conditions even with destocking. In selected regions where potential still exists, African savanna biodiversity and human economic development will both be best served by the integration of sustainable wildlife utilization into multispecies animal production systems.     

Local community size mediates ecological drift and competition in metacommunities

The outcome of competitive interactions is likely to be influenced by both competitive dominance (i.e. niche-based dynamics) and ecological drift (i.e. neutral dynamics governed by demographic stochasticity). However, spatial models of competition rarely consider the joint operation of these two processes. We develop a model based on the original competition-colonization trade-off model that incorporates niche and neutral processes and several realistic facets of ecological dynamics: it allows local competition (i.e. competition within a patch) to occur within communities of a finite size, it allows competitors to vary in the degree of competitive asymmetry, and it includes the role of local migration (i.e. propagule pressure). The model highlights the role of community size, i.e. the number of competitors in the local community, in mediating the relative importance of stochastic and deterministic forces. In metacommunities where local communities are small, ecological drift is substantial enough that strong competitors become effectively neutral, creating abrupt changes in the outcome of competition not predicted by the standard competition-colonization trade-off. Importantly, the model illustrates that, even when other aspects of species interactions (e.g. migration ability, competitive ability) are unchanged, local community size can alter the dynamics of metacommunity persistence. Our work demonstrates that activities which reduce the size of local communities, such as habitat destruction and degradation, effectively compound the extinction debt.     

On size and area: Patterns of mammalian body size extremes across landmasses

We describe a biogeographic pattern in which mammalian body size extremes scale with landmass area. The relationship between the largest and the smallest mammal species found on different landbridge islands, mountaintops and continents shows that the size of the largest species increases, while that of the smallest species decreases, with increase in the area of the landmass. We offer two possible explanations: (1) that the pattern is the result of sampling artefacts, which we call the ‘statistical artefact hypothesis’, or (2) that the pattern is the result of processes related to the way body size affects the number of individuals that a particular species can pack in a given area, which we call the ‘area-scaling hypothesis’. Our results point out that the pattern is not a statistical artefact resulting from random sampling, but can be explained by considering the scaling of individual space requirements and its effect on population survival on landmasses of different area. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

The 'island rule' in birds: medium body size and its ecological explanation

Do birds show a different pattern of insular evolution from mammals? Mammals follow the ''island rule'', with large-bodied species getting smaller on islands and small-bodied species getting bigger. By contrast, the traditional view on birds is that they follow no general island rule for body size, but that there is an insular trend for large bills. Insular shifts in feeding ecology are, therefore, widely assumed to be the primary cause of divergence in island birds. We use a comparative approach to test these ideas. Contrary to the traditional view, we find no evidence for increased bill size in insular populations. Instead, changes in both bill size and body size obey the ''island rule''. The differences between our results and the traditional view arise because previous analyses were based largely on passerines. We also investigate some ecological factors that are thought to influence island evolution. As predicted by the traditional view, shifts in bill size are associated with feeding ecology. By contrast, shifts in body size are associated with the potential for intraspecific competition and thermal ecology. All these results remain qualitatively unchanged when we use different methods to score the ecological factors and restrict our analyses to taxa showing pronounced morphological divergence. Because of strong covariation between ecological factors, however, we cannot estimate the relative importance of each ecological factor. Overall, our results show that the island rule is valid for both body size and bill length in birds and that, in addition to feeding ecology, insular shifts in the level of intraspecific competition and the abiotic environment also have a role.     

Relationship of sexual dimorphism in canine size and body size to social,behavioral, and ecological correlates in anthropoid primates

Among anthropoid primates there are interspecific differences in the degree of sexual dimorphism in both body size and canine size. Within the suborder body size dimorphism and canine size dimorphism are positively correlated,r=0.76. This correlation suggests that the two dimorphisms are equally developed in some species, while in other species there is a differential degree of sexual dimorphism. An analysis of these results and their relation to social organization and other ecological variables reveals: (1) the degree of canine size dimorphism is closely related to the amount of male intrasexual selection in a given mating system; and (2) the degree of body size dimorphism is also related to male intrasexual selection, but may be modified (either enhanced or diminished) by selection pressure from factors such as habitat, diet, foraging behavior, antipredator behavior, locomotory behavior, and female preference.     

Phylogenetic analysis of coadaptation in behavior, diet, and body size in the African antelope

Several authors have suggested that African antelope (familyBovidae) exemplify coadaptation of ecological, behavioral,and morphological traits. We tested four hypotheses relatedto the ecology and behavior of 75 species of African antelopeusing both conventional statistical techniques and techniques that account for the nonindependence of species by consideringtheir phylogenetic relationships. Specifically, we tested thehypotheses that (1) dietary selectivity is correlated negativelywith body mass, (2) dietary selectivity is correlated negativelywith group size, (3) gregarious species either flee or counterattackwhen approached by predators, but solitary and pair-livingspecies seek cover to hide, and (4) body mass and group sizeare correlated positively. Each of these hypotheses was examinedfor the global data set (family Bovidae) and, when possible,within the two antelope subfamilies (Antilopinae and Bovinae)and within 7 of the 10 antelope tribes. The results of ourconventional and phylogenetically corrected analyses supportedthe hypotheses that group and body size vary predictably with feeding style and that antipredator behavior varies with groupsize. The hypothesis that body mass and group size are correlatedpositively was supported by conventional statistics, but thesetwo traits were only weakly related using a phylogeneticallycorrected analysis. Moreover, qualitative and quantitativecomparisons within each of the eight major African antelope tribes generally gave little support for the four hypothesestested. Thus, although our analyses at the subfamily levelprovided results that were consistent with prior hypotheses,our analyses at the level of tribes were equivocal. We discussseveral possible explanations for these differences.     

Biodiversity of galling insects: historical, community and habitat effects in four neotropical savannas

Five hypotheses were tested to explain the pattern of galling insect species richness in four neotropical savanna physiognomies, 'canga ', 'campo sujo', 'cerrado' s. st., and 'cerradão', that occur in Minas Gerais, southeastern Brazil. We found 125 species of galling insects on 80 host plant species. The increase of plant species richness explained 35% of the variation in galling insect richness, corroborating the plant species richness hypothesis. Most of the galling species occurred on trees, followed by shrubs, and herbs. However, the difference in mean number of galls was only statistically significant between herbs and trees, corroborating partially the plant structural complexity hypothesis. A significant relationship was observed between galling species richness and density of herbs, and shrubs, corroborating partially the resource concentration hypothesis. Galling insect richness showed a negative correlation with magnesium, potassium, and zinc on soil, corroborating the soil fertility hypothesis. The content of magnesium, potassium, iron and CTC (T) explained 72% of the variation in galling insect richness. Plant family size positively influenced galling insect richness, corroborating the plant family size hypothesis. Overall, the results corroborate the hypothesis that predicts that habitat stress is the main factor generating the patterns of galling insect richness in Brazilian savannas.     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

Connecting landscape structure and patterns in body size distributions

Understanding the interaction between community structure and landscape structure represents a pressing theoretical challenge of great applied importance considering the increasing structural modification of ecosystems through habitat loss and fragmentation. Dispersal ability and energetic demands coupled to body size determine the landscape structure experienced by an organism, which could essentially be fragmented for small individuals but continuous for large ones. Although discontinuities in species assemblages have been predicted and detected, no explicit association between habitat structure and body size distributions has been demonstrated. In this contribution, we propose that body size structure in local communities should reflect such different perceptions of landscape structure. To this end, we explore this association in a simple metacommunity located in the Atacama Desert, in northern Chile. Using graph theory we found that species of different size and trophic position (carnivores and herbivores) perceive the landscape at contrasting spatial scales. In each community (n = 31) we determined the observed and the expected body size distributions – in a random sample from the metacommunity of 18 727 individuals –, which allowed us to identify the body sizes at which an overrepresentation or underrepresentation of individuals occur. Such aggregations and discontinuities in body sizes were related, for carnivores, to patch location within the landscape, and to the internal banded vegetation pattern within patches for herbivores. Our study shows, for the first time, an empirical connection between the spatial distribution of communities, their local attributes, and the existence and locations of discontinuities and aggregations in body size distributions.     

Conflicting processes in the evolution of body size and development time

Body size and development time of Manduca sexta are both determined by the same set of three developmental–physiological factors. These define a parameter space within which it is possible to analyse and explain how phenotypic change is associated with changes in the underlying factors. Body size and development time are determined by the identical set of underlying factors, so they are not independent, but because the mechanisms by which these factors produce each phenotype are different, the two phenotypes are only weakly correlated, and the correlation is context dependent. We use a mathematical model of this mechanism to explore the association between body size and development time and show that the correlation between these two life-history traits can be positive, zero or negative, depending entirely on where in parameter space a population is located, and on which of the underlying factors has a greater variation. The gradient within this parameter space predicts the unconstrained evolutionary trajectory under directional selection on each trait. Calculations of the gradients for body size and development time revealed that these are nearly orthogonal through much of the parameter space. Therefore, simultaneous directional selection on body size and development time can be neither synergistic nor antagonistic but leads to conflicting selection on the underlying developmental parameters.