Conspecifics as informers and competitors: an experimental study in foraging bumble-bees

Conspecifics are usually considered competitors negatively affecting food intake rates. However, their presence can also inform about resource quality by providing inadvertent social information. Few studies have investigated whether foragers perceive conspecifics as informers or competitors. Here, we experimentally tested whether variation in the density of demonstrators ('none', 'low' and 'high'), whose location indicated flower profitability, affected decision-making of bumble-bees Bombus terrestris. Bumble-bees foraged on either 'simple' (two colours) or 'complex' (four colours) artificial floral communities. We found that conspecifics at low density may be used as sources of information in first flower choices, whereas they appeared as competitors over the whole foraging sequence. Low conspecific densities improved foragers' first-visit success rate in the simple environment, and decreased time to first landing, especially in the complex environment. High conspecific densities did not affect these behavioural parameters, but reduced flower constancy in both floral communities, which may alter the efficiency of pollinating visits. These results suggest that the balance of the costs and benefits of conspecific presence varies with foraging experience, floral community and density. Spatio-temporal scales could thus be an important determinant of social information use. This behavioural flexibility should allow bumble-bees to better exploit their environment.     

Effects of sward structure on herbivore foraging behaviour in a South African savanna: An investigation of the forage maturation hypothesis

Abstract: The grass layer of savannas is characterized by strong temporal and spatial heterogeneity in the quantity and quality of forage. Besides this, there is strong variation in other aspects of sward structure, here defined as sward chemistry, morphology, architecture and species composition. The forage maturation hypothesis (FMH) is based on the temporal dynamics of forage quantity and quality of grasslands. Recently this hypothesis has been used to explain the foraging behaviour of large herbivores as an optimal solution for balancing forage ingestion and forage digestion, leading to a maximization of daily rates of forage intake in patches of intermediate forage quantity. However, so far studies using this hypothesis have been constrained to mono‐specific forage resources or have ignored inter‐patch variation in sward structure. We studied the foraging behaviour of cattle in a South African savanna. We explicitly addressed forage quantity and quality, and the structure of seven sward types in order to investigate (i) their effects on foraging behaviour; and (ii) the assumptions and predictions of the FMH for foraging behaviour in grasslands with pronounced variation in sward structure. The results indicated that the assumptions of the FMH were met and that forage quantity affected forage intake the most, with sward structure having little or no effect. The predicted maximum rate of daily forage intake agreed with the results of previous studies. However, the forage quantity at which this maximum intake rate was found, was larger than expected based on the results of some studies. It is likely that this discrepancy resulted from the use of artificial leaf‐only grass swards in these studies. The results suggest that the FMH can be used to explain the foraging behaviour of herbivores over a wide range of sward structures.     

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.     

Impacts of savanna trees on forage quality for a large African herbivore

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.     

Restricted habitat use by an African savanna herbivore through the seasonal cycle: key resources concept expanded

Areas within regional landscapes that make a disproportionate contribution to supporting large herbivore populations have been interpreted as key resource areas, hotspots, buffers, stepping stones or serving other functional roles. We investigated the role that the restricted extent of habitat types exploited at different stages of the seasonal cycle might play in limiting the abundance of a blue wildebeest subpopulation in the Kruger National Park, South Africa. GPS collars enabled the space use patterns of the animals to be related to available habitat types, and faecal nutrient concentrations to be related to the habitats exploited at that time. Wildebeest herds occupied primarily grazing lawn grasslands associated with gabbro uplands or sodic lowlands through the wet season into the early dry season. During the late dry season, they switched to seep‐zone grasslands in mid slope regions of granitic landscapes. Use of recently burned areas enhanced forage quality at the beginning of the wet season. The seasonal habitat shifts enabled wildebeest to obtain adequate nitrogen, phosphorus and sodium throughout the year. Lawn and seep‐zone grasslands combined constituted 10% of the available area. Grazing lawns, which encompassed only ? 3% of the study area, appeared to be the primary limitation on the abundance of wildebeest. However, the greater security from predation provided by the open vegetation cover in the grazing lawns is not easily disentangled from the resource benefits that they yield. Nevertheless, findings indicate how local abundance can be restricted by the extent of portions of the landscape providing crucial benefits during particular phases of the seasonal cycle. Hence the key resources concept needs to be expanded to accommodate the functionally distinct contributions made by different habitats towards supporting local herbivore populations.     

Large herbivore responses to surface water and land use in an East African savanna: implications for conservation and human-wildlife conflicts

Water, forage and predation constrain ungulate distributions in savannas. To understand these constraints, we characterized distributions of 15 herbivore species from water, locations of peak density and degree of clustering around the peaks using zero-inflated count data models and mapping census data collected in the Mara reserve and the adjoining pastoral ranches in Kenya during a wet and dry year. Herbivores followed a humped pattern (n = 46), suggesting constrained foraging in which they balance the benefits of proximity to water with the costs of foraging where food is depleted near water and travelling to more abundant food distant from water; an exponentially decreasing pattern (n = 11), indicating strong attraction to water or vegetation near water; or a uniform (n = 3) pattern. The details rather than the types of these patterns varied between years. Herbivores concentrated farther from water and more tightly around locations of their peak densities in the ranches than the reserve. Herbivores were more abundant and widely distributed from water in the wet than the dry year, and segregated along the distance-to-water gradient, presumably to minimize interspecific competition for food. Pastoralism compressed herbivore distributions and partially excluded some species (warthog, hartebeest, topi, wildebeest, zebra, eland, buffalo and elephant) from, while attracting others (Grant’s and Thomson’s gazelles, impala, giraffe) to the ranches, relative to the reserve. Regulating cultivation, fencing, settlements and livestock stocking levels in the ranches would allow continued wildlife access to water, reduce competition with, displacement or harassment of wildlife by people, livestock and dogs near water.     

Feeding while evading predators by a lotic mayfly: linking short-term foraging behaviours to long-term fitness consequences

Laboratory experiments were performed to quantify the combined effects of food abundance (low, high) and predation threat, imposed using a model fish (safe, risky) on fitness correlates (i.e. growth, time to emergence, adult body mass, fecundity, egg size) of the mayfly Baetis tricaudatus. These effects were determined by rearing larvae under different combinations of food abundance and predator threat. Fitness correlates were significantly affected by food abundance, predation threat or the interaction of these factors. High food abundance and the absence of predation threat significantly increased larval growth rates, adult body mass, fecundity, egg size, and decreased time to emergence. Long-term effects of predator threat and food abundance on fitness correlates of B. tricaudatus were compared to previously conducted short-term patch choice trials to test for concordance between short-term measures of patch choice and their potential longer-term fitness consequences. This comparison indicated that patches that were utilized the most were those that yielded the highest fitness benefits. These results suggest that behavioural costs of balancing mortality risk due to predation against food acquisition by B. tricaudatus can have strong fitness consequences.     

Effects of large herbivore browsing on the functional groups of woody plants in a southern African savanna

This study examined the effect of different large herbivore species and stocking rates in savanna ecosystems of Zimbabwe on the richness and abundances of woody plant functional groups and woody plant functional attributes. Seven fence-lines with different herbivore species and stocking rates on either side of the fence were sampled. Plots were placed on both sides of each fence at each of 18 randomly selected positions. The size and species of each woody plant was recorded for each plot. It was found that the number of species with different functional attributes of spinescence, leaf longevity, fruit type and dispersal mechanism and in the functional groups of palatability were not different on the different sides of the fence. However, there were differences in plant abundances for 26 out of the 35 tests carried out on plant abundances with different functional attributes and functional groups. It was hypothesised that the time needed to change woody plant species richness is hundreds of years in these systems, whereas the time needed to change woody plant abundances is decades.     

Fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an African savanna

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.     

Root foraging strategies and soil patchiness in a humid savanna

In Lamto (Côte d'Ivoire), the savanna is a patchy environment as far as soil is concerned: tree clumps and termite mounds lead to higher nutrient contents than in the surrounding savanna. Mature Borassus aethiopum (Mart.) specimens are tall palm trees dominating the community, with aerial parts located out of these nutrient-rich patches.Palm root densities were compared under tree clumps and in the surrounding savanna, and were also sampled along transects between palm trees and nutrient-rich patches (two clumps and one mound). Palm root densities were far higher (up to 10 times) in the nitrogen-rich soil of both clumps and termite mounds than in the surrounding savanna. Evidence is given that palm trees are able to extend their root system as far as 20 m towards these nutrient-rich patches where they proliferate. These results point out a particular root foraging strategy, which is one of the first known for a woody perennial. They also provide new insights for understanding nitrogen cycling and savannas high rate of primary production.     

Host plant phenology may determine the abundance of an ecosystem engineering herbivore in a tropical savanna

1. Variation in plant phenology allows plants to escape from herbivory. Insect herbivores manipulate their host plants by producing shelters, which they inhabit, and are protected against natural enemies and/or unfavourable environmental conditions. Environmental modifications induced by living organisms are characterised as ecosystem engineering. 2. In the Brazilian savanna, the interaction between the Malpighiaceae shrub, Byrsonima intermedia, and its main herbivore, the caterpillar Cerconota achatina, a shelter‐building organism, was studied. 3. It was focused on whether the phenological development of the host plant affects the caterpillars' infestation, and whether C. achatina acts as an ecosystem engineer by building shelters. All plant variables (number of leaves, flowers, buds, fruits and herbivores) were measured fortnightly. Phenological data were correlated with climatic information. The impact of the caterpillars acting as ecosystem engineers was measured experimentally. 4. Infestation levels of C. achatina covaried with the phenology of their host plant B. intermedia. It suggests that plant phenology might be an important factor in determining the abundances of this ecosystem engineer species.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

Dietary niche separation of rodents and shrews in an African savanna

While niche separation and relationships with environmental conditions of large mammals of the African savanna have been studied intensively, less conspicuous components have not received similar attention. This is the case of Kafue National Park (KNP), Zambia, where mechanisms supporting coexistence among rodent and shrew species remain unclear, much less the influence of fire on their dietary resource use. Here, we use stable carbon and nitrogen isotopes to assess dietary resource use and partitioning among rodents and shrews found in three vegetation formations of KNP. According to the nitrogen isotope signatures, rodents are one to two trophic levels above primary production, save for Mus triton that is above by two to three. Shrews are two trophic levels above primary production. Among shrews, factors allowing coexistence of similar sized species could not be resolved. Rodent species of the same assemblage either differ in body mass by a factor of two (following Hutchinson's rule) or similar sized species occupy different trophic levels or dietary guilds based on their isotopic nitrogen or carbon signatures. At sites with frequent fires, rodents have broader dietary niches than at sites with low fire frequencies. This could either indicate relaxed competition under high fire frequencies as rodent populations do not reach the carrying capacity of the habitat, or it could reflect reduced competition due to lower species numbers under high versus low fire recurrence regimes. The results indicate competition as an important component structuring rodent communities in Zambian savannas, thus suggesting limited resources.     

Evolution of nocturnality in bats: Potential competitors and predators during their early history

Despite their taxonomic and ecological diversity, modern bats (Order Chiroptera) are almost exclusively nocturnal. This behaviour is too ubiquitous to be explained by common patterns of temporal variation in availability of their diverse food sources or by the risk of hyperthermia when flying during the day. Other explanations for bat nocturnality include competition and increased predation risk from birds during the day. In the early and mid Eocene, the known bat fauna consisted of several insectivorous species of sizes similar to those of the modern European assemblage. This fauna was contemporaneous with several species of predatory birds, including owls (Strigiformes), hawks (Accipitridae), falcons (Falconidae) and rollers (Coraciiformes), which were the same size as modern predators on bats. Predation risk could therefore have been a significant factor preventing the early bats from becoming diurnal. Competition from aerial insectivorous birds, however, was less likely to have been significant for bats during the early Eocene, since very few such groups, mainly small Aegialornithidae, were present, with most of the major groups of aerial insectivores evolving later.     

Woody encroachment slows decomposition and termite activity in an African savanna

Woody encroachment can lead to a complete switch from open habitats to dense thickets, and has the potential to greatly alter the biodiversity and ecological functioning of grassy ecosystems across the globe. Plant litter decomposition is a critical ecosystem process fundamental to nutrient cycling and global carbon dynamics, yet little is known about how woody encroachment might alter this process. We compared grass decay rates of heavily encroached areas with adjacent nonencroached open areas in a semi‐arid South African savanna using litterbags that allowed or excluded invertebrates. We also assessed the effect of woody encroachment on the activity of termites— dominant decomposer organisms in savanna systems. We found a significant reduction in decomposition rates within encroached areas, with litter taking twice as long to decay compared with open savanna areas. Moreover, invertebrates were more influential on grass decomposition in open areas and termite activity was substantially lower in encroached areas, particularly during the dry season when activity levels were reduced to almost zero. Our results suggest that woody encroachment created an unfavourable environment for invertebrates, and termites in particular, leading to decreased decomposition rates in these areas. We provide the first quantification of woody encroachment altering the functioning of African savanna ecosystems through the slowing of aboveground plant decomposition. Woody encroachment is intensifying across the globe, and our results suggest that substantial changes to the carbon balance and biodiversity of grassy biomes could occur.     

Vegetation management shapes arthropod and bird communities in an African savanna

Habitat heterogeneity is a key driver of the diversity and distribution of species. African savannas are experiencing changes in their vegetation structure causing shifts towards increased woody plant cover, which results in vegetation structure homogenization. Given the impact that increasing woody plant cover has on patterns of animal use, resource managers across Africa are implementing habitat management practices that are intended to reduce woody plant cover. To understand the ecological implications of various habitat management practices on arthropod and bird communities, we leveraged large‐scale tree clearing and subsequent mowing in an African savanna to understand how changes in both the herbaceous layer and woody plant cover (i.e., structural heterogeneity) may shape arthropod and bird communities at the local scale. We focused on four replicated treatments: (1) annual summer mow, (2) annual winter mow, (3) >5 years since last mow (rest), and (4) an adjacent unmanipulated savanna to act as a control. We found that the mowing treatments significantly influenced vegetation structure both with respect to tree density and herbaceous layer. Both arthropod and bird community composition varied across treatments. Grass biomass was the best predictor of arthropod richness and abundance, with arthropods selecting for areas with high biomass. Insectivorous bird richness and abundance was driven by tree density (i.e., perching locations) and not arthropod abundance. Our results suggest that vegetation management practices contribute to habitat heterogeneity at the landscape scale and increase bird species richness through species turnover. However, we caution that if a single vegetation management practice dominates the landscape, it is plausible that it could lead to the simplification of the avian community.     

Climate‐driven fluctuations in surface‐water availability and the buffering role of artificial pumping in an African savanna: Potential implication for herbivore dynamics

Abstract In arid and semiarid environments surface‐water strongly constrains the distribution and abundance of large herbivores during the dry season. Surprisingly, we know very little about its variability in natural ecosystems. Here we used long‐term data on the dry‐season occurrence of water at individual waterholes to model the surface‐water availability across Hwange National Park, Zimbabwe, under contrasted climatic and management scenarios. Without artificial pumping only 19.6% of the park occurred within 5 km of water under average climatic conditions. However surface‐water availability was strongly influenced by annual rainfall, and over 20 years the variability of the surface area of the park occurring within 5 km of water was slightly larger than the variability of rainfall. This contrasts with the usual buffered response of vegetation production to rainfall fluctuations, and suggests that the variability in dry‐season foraging range determined by surface‐water availability could be the main mechanism regulating the population dynamics of large herbivores in this environment. Artificial pumping increased surface‐water availability and reduced its variability over time. Because changes in surface‐water availability could cause the greatest changes in forage availability for large herbivores, we urge ecologists to investigate and report on the variability of surface‐water in natural ecosystems, particularly where rapid climate changes are expected.