Seasonal variation in patch use in a tropical African environment

We used giving-up densities (GUD) to study patch use decisions of small granivorous passerines throughout the year. We measured GUDs continuously in four sites for a period of 9–10 months per year during 2004 and 2005 in a savannah area in Jos, central Nigeria. The study thus covered a period from the middle of the dry season, through the wet season to the beginning of the next dry season in each year. We placed experimental food patches in both open areas and in cover to investigate possible effects of predation risk and thermal hazard on the foraging behavior of the birds. We found a difference in GUDs between the microhabitats, with a consistently lower GUD in cover throughout the year and for the two years. During both years GUDs followed a pattern coinciding with the seasonal change in local seed availability. An initial decline in GUDs late in the dry season was followed by a steady increase during and after the rains. A similar trend in GUDs observed for both years supports the conclusion that GUDs measure the feeding birds' assessment of environmental quality, possibly in combination with other factors changing predictably during the year. We conclude that food abundance may act with other environmental and ecological factors to affect foraging decisions throughout the year.     

Herbaceous Forage and Selection Patterns by Ungulates across Varying Herbivore Assemblages in a South African Savanna

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.     

Land management implications for ecosystem services in a South African rangeland

In South Africa, restoration and sustainable management of historically overgrazed and degraded rangelands are promoted to increase biodiversity and ecosystem service provision. This study evaluates different land management scenarios in terms of ecosystem services in a South African rangeland, the Baviaanskloof catchment. As measured data were limited, we used simple models to quantify and map the effect of the different combination of agricultural, nature conservation and restoration practices on multiple ecosystem services. The land management scenarios were evaluated against management targets set for individual ecosystem services. Results highlight how the provision of ecosystem services is related to land management as unmanaged, pristine ecosystems provide a different mix of ecosystem services than ecosystems recently restored or managed as grazing lands. Results also indicate that historically overgrazed lands provide no forage, may retain 40% less sediment and have 38% lower biodiversity, while providing 60% more fuel wood and supplying two and half times more water (i.e. retaining less water), than pristine or restored lands. We conclude that a combination of light grazing, low input agriculture, nature conservation and restoration is the best for the sufficient provision of multiple ecosystem services. Applying such mixed management would improve biodiversity, ecotourism and maintain forage production and regulating services on farmers’ land. This management option also fits into and further optimizes local decision-makers’ vision regarding the future management of the area.     

Denitrification in a semi-arid grazing ecosystem

The effect of large herbivores on gaseous N loss from grasslands, particularly via denitrification, is poorly understood. In this study, we examined the influence of native migratory ungulates on denitrification in grasslands of Yellowstone National Park in two ways, by (1) examining the effect of artificial urine application on denitrification, and (2) comparing rates inside and outside long-term exclosures at topographically diverse locations. Artificial urine did not influence denitrification 3 and 12 days after application at hilltop, mid-slope, and slope-bottom sites. Likewise, grazers had no effect on community-level denitrification at dry exclosure sites, where rates were low. At mesic sites, however, ungulates enhanced denitrification by as much as 4 kg N ha⁻¹ year⁻¹, which was double atmospheric N inputs to this ecosystem. Denitrification enzyme activity (DEA, a measure of denitrification potential) was positively associated with soil moisture at exclosure sites, and herbivores stimulated DEA when accounting for the soil moisture effect. Glucose additons to soils increased denitrification and nitrate additions had no influence, suggesting that denitrification was limited by the amount of labile soil carbon, which previously has been shown to be enhanced by ungulates in Yellowstone. These results indicate that denitrification can be an ecologically important flux in portions of semi-arid landscapes, and that there is a previously unsuspected regulation of this process by herbivores. Received: 6 March 1998 / Accepted: 28 August 1998     

Interactive response of herbivores,soils and vegetation to annual burning in a South African savanna

Abstract Despite decades of research, the primary factors determining savanna structure remain elusive – a conundrum termed ‘the savanna problem’. After 47 years of annual burning in Terminalia woodland and Acacia/Combretum savanna on sandy, granite‐derived soils in the southern Kruger National Park, South Africa, a dense cover of trees and shrubs persists on some burnt plots and is largely absent from others. We postulated that intense browsing pressure by antelope and other herbivores prevents recruitment of trees in burnt plots and that herbivores concentrate on plots that are richest in nutrients. Herbivore abundance did not show a relationship with soil macronutrients and we consequently investigated micronutrient status. The reduction in tree cover as a result of annual burning was positively correlated with mass of herbivores (15–1500 kg) (r ² = 0.61, n = 8). This index of herbivore abundance was in turn positively correlated with total Zn (r ² = 0.64, n = 8). Other indices of herbivore abundance showed significant relationships with total clay content and total Mn. We suggest that herbivores concentrate on sites with greater clay content (possibly due to a greater availability of micronutrients), and that tree cover can remain relatively dense under a regime of annual burning if browsing pressure is not intense. The long‐term burn experiments in the Kruger National Park savanna provide a platform for unravelling the savanna problem. Determining possible interactions between soil properties, herbivory and fire is a step in this direction.     

放牧对草地昆虫多样性的影响研究进展

在草地生态系统中,大型草食动物放牧是重要的管理方式之一,对草地生物多样性起着关键的驱动作用。昆虫是草地生态系统中生物多样性的重要组成成分,对生态系统的食物网结构以及其功能与稳定性起着关键作用。已有研究结果表明,大型草食动物与昆虫存在密切联系,放牧对草地昆虫多样性或有正向、或负向、或无明显作用,这依赖于放牧管理方式、昆虫类群以及草地类型。放牧必然通过直接(采食、践踏或粪尿)或间接(植物群落组成或植被结构)作用对昆虫多样性产生显著的影响。当前,关于大型草食动物放牧对草地昆虫多样性影响研究较多,但是,从研究系统性、深入性和延续性来说还存在一定问题。本文在综述国内外对放牧对草地昆虫多样性的影响研究基础上,提出了今后的研究方向,对于理解放牧管理的草地昆虫多样性变化规律,以及为积极探索维持草地昆虫多样性的长期有效的科学管理措施提供理论指导。     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

Climbing asparagus,Asparagus scandens Thunb.: a South African in your forest patch

Climbing asparagus, Asparagus scandens Thunb., currently has a patchy distribution in New Zealand, mainly in disturbed lowland forest remnants near towns and the coast, predominantly in the North Island but found as far south as Banks Peninsula. Here it prevents regeneration of native seedlings and smothers or strangles understorey shrubs and saplings. Based on its current distribution, it tolerates open sunny sites to deep shade and damp or dry forest. Left unchecked and aided by birds and humans, climbing asparagus is likely to spread considerably in the next decade, consolidating its current distribution in New Zealand. Several characteristics of the species make it difficult to control: the small, fine seedlings are hard to find, adult plants entwine with native plants, much of the plant’s biomass is underground and the apical meristem will resprout if left in the ground after control.     

放牧家畜食性选择机制研究进展

动物的食性选择是一个十分复杂的主题.本文从不同角度,结合多学科的研究成果对放牧家畜食性选择机制问题进行了阐述：1）食性选择行为发生的机制,包括基于遗传和学习两方面的解释,其中动物具有的学习机制主要有来自动物个体的学习（采食后反馈、试错学习）、向其他社会群体的学习,以及在进行斑块、景观水平选择时,所采用的空间记忆、视觉线索和皮肤防御系统机制等;2）放牧家畜食性选择策略的4个综合性假说：经验法则、边际值法则、优化采食理论、最小总不适感概念;3）食性选择机制研究存在的问题与展望,诸如空间尺度问题、动物消化生理因素以及多因素互作研究的不足等.     

Plant community response to loss of large herbivores: comparing consequences in a South African and a North American grassland

Loss of biodiversity poses one of the greatest threats to natural ecosystems throughout the world. However, a comprehensive understanding of the impacts of species losses from upper trophic levels is still emerging. Here we compare the impacts of large mammalian herbivore species loss on grassland plant community structure and composition in a South African and North American grassland. Herbaceous plant communities were surveyed at sites without large mammalian herbivores present and at sites with a single species of herbivore present in both locations, and additionally at one site in South Africa with multiple herbivore species. At both the North American and South African locations, plant communities on sites with a single herbivore species were more diverse and species rich than on sites with no herbivores. At the multi-herbivore site in South Africa, plant diversity and richness were comparable to that of the single herbivore site early in the growing season and to the no herbivore site late in the growing season. Analyses of plant community composition, however, indicated strong differences between the multi-herbivore site and the single and no herbivore sites, which were more similar to each other. In moderate to high-productivity ecosystems with one or a few species of large herbivores, loss of herbivores can cause a significant decrease in plant diversity and richness, and can have pronounced impacts on grassland plant community composition. In ecosystems with higher herbivore richness, species loss may also significantly alter plant community structure and composition, although standard metrics of community structure may obscure these differences.     

Seasonal patterns of habitat use by insectivorous bats in a subtropical African agro‐ecosystem dominated by macadamia orchards

We report on acoustic surveys of insectivorous bats conducted during seven months of the year using ANABAT recordings in two habitats (macadamia orchards and adjacent riparian bush) in a subtropical agro‐ecosystem in northern South Africa. We defined two functional foraging groups of bats based on their echolocation calls: (i) open‐air foragers (family Molossidae) having narrow‐band, low‐frequency, low duty cycle calls; and (ii) clutter‐edge foragers (families Miniopteridae and Vespertilionidae), having broad‐band, higher frequency, low duty cycle calls. Bat activity (number of bat passes) was not significantly influenced by habitat. Total bat activity and activity of both functional groups varied significantly between seasons, being highest in summer and autumn (coinciding with annual peaks in numbers of Twin spotted (Bathycoelia natalicola) and Green (Nezara spp) Stinkbugs, order Heteroptera, family Pentatomidae, and Macadamia Nut Borer moths, Cryptophlebia ombrodelta) and lower in winter and spring. No significant effect of moon phase was detected, either on total activity or activity of the two functional groups. We postulate that the significant pattern of seasonality of commuting and/or foraging activity of bats in macadamia orchards (which is more marked in open‐air foragers) may be driven by the seasonal abundance of pest insects such as stinkbugs and Macadamia Nut Borer moths.     

Comparative habitat use by grazing fishes in a Bornean stream

Habitat use by four grazing fishes in a rainforest stream was determined by direct observation (snorkeling) and microhabitat measurements for individual fish. Significant species heterogeneity (P?<?0.004) occurred along two principal component gradients of velocity, depth and substrate and most pairwise species contrasts were significant. Abundant Paracrossochilus acerus (Cyprinidae) occupied the slower, deeper end of the gradient and Gastromyzon punctulatus (Balitoridae) the fast, shallow end with common G. cranbrooki and rare G. aeroides intermediate. However, overlap was substantial and as many as three species grazed on a single rock with no apparent interaction. All species were primarily day-active. Incomplete experiments suggest the fishes rapidly abandon rocks with reduced algal cover. Frequent spates with high discharge, turbidity and bedload movement disturbed the river. Disturbance, rather than biotic interactions, may be the dominant factor in the ecology of these fishes.     

Forage quality and patch choice by wapiti (Cervus elaphus)

Recent models suggest that herbivores might optimize energygain by selecting patches of intermediate vegetation biomass.We tested this hypothesis in wapiti (Cervus elaphus) by estimatingdaily rates of energy gain in relation to grass biomass andby measuring patch choice in experimental pastures in whichgrass biomass was manipulated by mowing. The digestible energycontent of grasses declined with increasing biomass due to maturationalchanges in fiber and lignin content. Daily rates of dry matterintake by wapiti increased with grass biomass at a deceleratingrate, implying a Type II functional response. Linking thesevalues to published ad libitum energy intake and energy expenditureparameters, Fryxell's (1991) model predicted that the dailyrate of energy gain should be highest when wapiti feed in grasslandswith 1000–1100 kg/ha. In trials in which grass biomasswithin a mosaic of patches was manipulated experimentally between800–2900 kg/ha, wapiti preferred patches of 1200 kg/ha,close to the value predicted by the energy gain model. Our resultssuggest that the rate of energy gain by wapiti is constrainedby both grass biomass and grass fiber content, the latter ofwhich varies inversely with grass biomass. Behavioral preferencefor grass patches of intermediate biomass and fiber contentcould help explain patterns of aggregation and seasonal migrationreported previously for wapiti.     

Large herbivores affect the stoichiometry of nutrients in a regenerating woodland ecosystem

Previous studies on the effects of herbivores on nutrient cycling have given little consideration to the relationship between soil nitrogen (N) and phosphorus (P) availability. Here we examined how browsing by red deer influences the relative availability of N and P in a regenerating woodland ecosystem. We found that removal of browsing by fencing for 14 years led to a shift from N toward P limitation of the dominant tree species Betula pubescens . This was evidenced by a significant increase in foliar N:P ratio of B. pubescens as a result of removal of browsing; mean N:P ratio of foliage from browsed areas was 13.2 suggesting that trees growing in browsed areas were N limited, whereas foliage from unbrowsed areas had a mean N:P ratio of 15.8, suggesting that these areas were more P limited. Further evidence of a shift toward P limitation in unbrowsed areas came from the finding that root uptake of labelled ³²P was significantly greater in roots collected from unbrowsed than browsed trees. Soil phosphatase activity did not significantly differ between browsed and unbrowsed areas. Our data indicate therefore that herbivores have the potential to significantly affect the stoichiometry of N and P in forest ecosystems.     

Soil and vegetation differences across ecological boundaries in an arid South African ecosystem

Boundaries are the most reactive nodes in landscapes and may be hypersensitive to global change in climate and land use. Investigations on how soils govern vegetation boundaries are scant, particularly in arid and semiarid ecosystems. The Tankwa Karoo National Park (TKNP) is a unique arid biodiversity hotspot with an unrivalled aridity gradient from < 100 mm MAP to about 700 mm in < 10 km. We investigated the abruptness of four soil‐vegetation boundaries separating eight communities. Two 50 m transects were established across four boundaries for 24 descending point transects, in which the cover‐abundance of each plant encounter at 1 m intervals was recorded. In addition, three soil samples were collected from the top 5 cm in each of the four boundaries and twelve patches. Soil and vegetation parameters altogether indicated three boundary syndromes that were context dependent: (a) a sharp boundary, (b) a gradual boundary or (c) no boundary exists. Soil respiration recorded here, and perhaps other ecosystem processes, was mediated by the soil‐vegetation boundaries. These nodes should be the focus of ecological studies since they reveal much more than the constituent patches themselves.     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

The importance of scale of patchiness for selectivity in grazing herbivores

The notion that spatial scale is an important determinant of foraging selectivity and habitat utilization has only recently been recognized. We predicted and tested the effects of scale of patchiness on movements and selectivity of a large grazer in a controlled field experiment. We created random mosaics of short/high-quality and tall/low-quality grass patches in equal proportion at grid sizes of 2×2 m and 5×5 m. Subsequently, we monitored the foraging behaviour of four steers in 16 20×40 m plots over 30-min periods. As predicted on the basis of nutrient intake maximization, the animals selected the short patches, both by walking in a non-random manner and by additional selectivity for feeding stations. The tortuosity of foraging paths was similar at both scales of patchiness but selectivity was more pronounced in large patches than in small ones. In contrast, the number of bites per feeding station was not affected by patch size, suggesting that selection between and within feeding stations are essentially different processes. Mean residence time at individual feeding stations could not be successfully predicted on the basis of the marginal-value theorem: the animals stayed longer than expected, especially in the less profitable patch type. The distribution of the number of bites per feeding station suggests a constant probability to stay to feed or to move on to the next feeding station. This implies that the animals do not treat larger patches as discrete feeding stations but rather as a continuous resource. Our results have important implications for the application of optimal foraging theory in patchy environments. We conclude that selectivity in grazers is facilitated by large-scale heterogeneity, particularly by enhancing discrimination between feeding stations and larger selection units. Received: 1 March 1999 / Accepted: 14 July 1999