Leaf‐cutting ants,seasonal burning and nutrient distribution in Cerrado vegetation

Abstract Fire and herbivory are known to modify plant community structure. Many studies have suggested that fire ashes may increase soil nutrients in dystrophic soils. Herbivores may also change plant community structure through direct effects of herbivory and affecting nutrient cycling. Leaf‐cutting ants were traditionally viewed as herbivores, although their role may be more complex, because their nests affect both chemical and physical soil properties, thus affecting plants indirectly. We investigated the effects of frequent burning and of leaf‐cutting ants on the nutrient status of an herbaceous and a shrub species occurring in the Brazilian Cerrado, a habitat that is characterized by natural burnings. The proximity of ant nests resulted in an increase of nutrients in the leaves of both vegetation strata, whereas burning sometimes resulted in a decrease of nutrients. Our results do not lead to a possible positive effect of fire on plant nutrient content. On the other hand, ant nests may represent an important source of nutrients for plants on the nutrient‐depleted Cerrado soils and may accelerate vegetation recovery after burning.     

Effectiveness of protein-rich pea flour for the control of stored-product beetles

Abstract The association between visiting ants and the extrafloral nectaries (EFN)‐bearing shrub Hibiscus pernambucensis Arruda (Malvaceae) was investigated in two different coastal habitats – a permanently dry sandy forest and a regularly inundated mangrove forest. In both habitats the frequency of plants with ants and the mean number of ants per plant were much higher on H. pernambucensis than on non‐nectariferous neighbouring plants. In the sandy forest the proportion of live termite baits attacked by ants on H. pernambucensis was much higher than on plants lacking EFNs. In the mangrove, however, ants attacked equal numbers of termites on either plant class. Ant attendance to tuna/honey baits revealed that overall ant activity in the sandy forest is higher than in the mangrove area. The vertical distribution (ground vs. foliage) of ant activity also differed between habitats. While in the mangrove foraging ants were more frequent at baits placed on foliage, in the sandy forest ant attendance was higher at ground baits. Plants housing ant colonies were more common in the mangrove than in the sandy forest. Frequent flooding in the mangrove may have resulted in increased numbers of ant nests on vegetation and scattered ant activity across plant foliage, irrespective of possession of EFNs. Thus plants with EFNs in the mangrove may not experience increased ant aggression towards potential herbivores relative to plants lacking EFNs. The study suggests that the vertical distribution of ant activity, as related to different nest site distribution (ground vs. foliage) through a spatial scale, can mediate ant foraging patterns on plant foliage and probably affect the ants’ potential for herbivore deterrence on an EFN‐bearing plant species.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.     

Nutrient stress: an explanation for plant anti-herbivore responses to defoliation

Summary A hypothesis is put forward that the long-lasting inducible responses of trees to herbivores, particularly lepidopteran defoliators, may not be active defensive responses, but a by-product of mechanisms which rearrange the plant carbon/nutrient balance in response to nutrient stress caused by defoliation. When defoliation removes the foliage nutrients of trees growing in nutrient-poor soils, it increases nutrient stress wich in turn results in a high production of carbon-based allelochemicals. The excess of carbon that cannot be diverted to growth due to nutrient stress is diverted to the production of plant secondary metabolites. The level of carbon-based secondary substances decays gradually depending on the rate at which nutrient stress is relaxed after defoliation. In nutrient-poor soils and in plant species with slow compensatory nutrient uptake rates the responses induced by defoliation can have relaxation times of several years. The changes in leaf nitrogen and phenolic content of mountain birch support this nutrient stress hypothesis. Defoliation reduces leaf nitrogen content while phenolic content increases. These responses of mountain birch to defoliation are relaxed within 3–4 years.     

More salt,please: global patterns,responses and impacts of foliar sodium in grasslands

Sodium is unique among abundant elemental nutrients, because most plant species do not require it for growth or development, whereas animals physiologically require sodium. Foliar sodium influences consumption rates by animals and can structure herbivores across landscapes. We quantified foliar sodium in 201 locally abundant, herbaceous species representing 32 families and, at 26 sites on four continents, experimentally manipulated vertebrate herbivores and elemental nutrients to determine their effect on foliar sodium. Foliar sodium varied taxonomically and geographically, spanning five orders of magnitude. Site‐level foliar sodium increased most strongly with site aridity and soil sodium; nutrient addition weakened the relationship between aridity and mean foliar sodium. Within sites, high sodium plants declined in abundance with fertilisation, whereas low sodium plants increased. Herbivory provided an explanation: herbivores selectively reduced high nutrient, high sodium plants. Thus, interactions among climate, nutrients and the resulting nutritional value for herbivores determine foliar sodium biogeography in herbaceous‐dominated systems.     

Convergence and non-convergence of plant traits in climatically and edaphically matched sites in Mediterranean Australia and South Africa

Abstract This paper compares the occurrence of plant traits in five edaphically matched sites at the Barrens, southwestern Australia and the Agulhas Plain, southwestern South Africa. The two regions are very closely matched in terms of their Mediterranean-type climates, landforms, soil types and disturbance regimes. On both continents, matched sites on all substrata (siliceous sand, quartzite, laterite, limestone and calcareous sand) support sclerophyllous shrublands with a similar mix of growth forms. Soils from all substrata in both Australia and South Africa are extremely nutrient-poor except for the calcareous sands where high levels of phosphorus were recorded. Contrary to expectations, Australian soils are not generally less fertile than their South African counterparts. The frequency of species in different leaf consistence categories was similar on the two continents, as was the leaf specific mass of overstorey shrubs from all substrata. Woody plants with leaf spines are significantly more frequent on Australian nutrient-poor substrata. Among woody plants, species with canopy-stored seed are significantly more frequent on Australian nutrient-poor sites, whereas species with bird-dispersed fruits and inter-fire germination are significantly more frequent on South African limestone and calcareous sand. There was good evidence for convergence between the two continents in the frequency of other seed biological traits. The study indicates strong convergence between Australian and South African shrublands in the frequency of a wide range of traits relating to plant form and function. Examples of non-convergence are probably due to regional and historical processes rather than differences in the contemporary physical environments of the two study areas.     

Interplant variation in creosotebush foliage characteristics and canopy arthropods

Summary We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.     

Effects of plant species on nutrient cycling

Plant species create positive feedbacks to patterns of nutrient cycling in natural ecosystems. For example, in nutrient-poor ecosystems, plants grow slowly, use nutrients efficiently and produce poor-quality litter that decomposes slowly and deters herbivores. /n contrast, plant species from nutrient-rich ecosystems grow rapidly, produce readily degradable litter and sustain high rates of herbivory, further enhancing rates of nutrient cycling. Plants may also create positive feedbacks to nutrient cycling because of species' differences in carbon deposition and competition with microbes for nutrients in the rhizosphere. New research is showing that species' effects can be as or more important than abiotic factors, such as climate, in controlling ecosystem fertility.     

Nutrient and secondary metabolite concentrations in a savanna are independently affected by large herbivores and shoot growth rate

Carbon-based secondary metabolites (CBSMs) such as tannins are assumed to function as plant defences against herbivores. CBSMs are thought to be inversely related to growth rate and nutrient concentrations because a physiological trade-off exists between cellular growth and differentiation, but CBSM concentrations can be altered by herbivory-induced changes in the trade-off. We predicted that a significant interaction exists between herbivory and growth phase, such that the effects of large herbivores (or their exclusion) on nutrient or CBSM concentrations are greatest during phases of rapid shoot or leaf growth. Leaf samples were collected during phases of different growth rate from six woody species 4 years after establishment of a large-scale long-term herbivore exclusion experiment in Kruger National Park, South Africa. Samples were analysed for N, P, condensed tannins and total phenolics. Interactions between growth phase and herbivores were rare. However, the assumption that elevated nutrients and reduced CBSMs occurs during fast phases of growth was supported by four species (consistent with the growth-differentiation balance hypothesis), but not the other two. Large herbivores generally did not affect nutrients, but CBSMs in four species were reduced by large herbivores other than elephants, while CBSMs in two species were reduced by elephants. Carbon limitation ultimately prevailed among woody plants taller than 2 m under long-term browsing. Large herbivores and plant growth phase are independent and important determinants of nutrients or CBSMs in African savannas, but the effects depend on the interacting assemblages of species, which poses challenges to the application of current general hypotheses of plant defence.     

Incidence of the introduced parasitoids Cotesia kazak and Microplitis croceipes (Hymenoptera: Braconidae) from Helicoverpa armigera (Lepidoptera: Noctuidae) in tomatoes, sweet corn, and lucerne in New Zealand

Morphological defense traits of plants such as trichomes potentially compromise biological control in agroecosystems because they may hinder predation by natural enemies. To investigate whether plant trichomes hinder red imported fire ants, Solenopsis invicta Buren (Hymenoptera: Formicidae), as biological control agents in soybean, field and greenhouse experiments were conducted in which we manipulated fire ant density in plots of three soybean isolines varying in trichome density. Resulting treatment effects on the abundance of herbivores, other natural enemies, plant herbivory, and yield were assessed. Trichomes did not inhibit fire ants from foraging on plants in the field or in the greenhouse, and fire ant predation of herbivores in the field was actually greater on pubescent plants relative to glabrous plants. Consequently, fire ants more strongly reduced plant damage by herbivores on pubescent plants. This effect, however, did not translate into greater yield from pubescent plants at high fire ant densities. Intraguild predation by fire ants, in contrast, was weak, inconsistent, and did not vary with trichome density. Rather than hindering fire ant predation, therefore, soybean trichomes instead increased fire ant predation of herbivores resulting in enhanced tritrophic effects of fire ants on pubescent plants. This effect was likely the result of a functional response by fire ants to the greater abundance of caterpillar prey on pubescent plants. Given the ubiquity of lepidopteran herbivores and the functional response to prey shown by many generalist arthropod predators, a positive indirect effect of trichomes on predation by natural enemies might be more far more common than is currently appreciated.     

Do nutrient-poor soils inhibit development of forests? A nutrient stock analysis

Nutrient-poor soils often support low-stature grasslands, savannas and shrublands where the climate is warm enough and wet enough for closed forests. Though this pattern has long been recognised, the causes are debated and poorly explored. I tested the hypothesis that forest fails to develop where the total nutrient pool is too small to construct both the foliage and the wood. I estimated potential woody biomass from the difference between soil nutrient stocks and forest foliage stocks. Nutrient stocks required for foliage were estimated from leaf tissue concentrations and foliage biomass typical of Amazon forests. Potential wood biomass was estimated from wood nutrient concentrations typical of Amazon forests. Data on soil nutrient stocks were assembled from studies from South American and African forests and savannas and from south-western Australian and south-west African heathlands. According to these calculations, estimated nutrient stocks (kg ha^?1) to build a forest would need to be > = P: 20–30, K 200–350, Ca 300–600 and Mg 55–65. Many surface soil horizons from both savanna and heathland sites were below these thresholds. However when deeper soil layers were included, most soils had adequate nutrient stocks. The nutrients in shortest supply were Ca and K and not P. This study suggests that nutrient stocks are usually adequate for constructing the wood needed to build a forest, except where soils are highly leached and very shallow. The implication is that, at steady state, low nutrient stocks seldom constrain forest development. The apparent failure of low nutrient stocks to explain the missing forests on nutrient-poor soils emphasises the need for new ideas on how nutrients, alone or in combination with other factors such as fire, influence vegetation structure.     

Small mammalian herbivore determines vegetation response to patchy nutrient inputs

Nutrient inputs to plant communities are often spatially heterogeneous, for example those deriving from the dung and urine of large grazing animals. The effect of such localised elevation of nutrients on plant growth and composition has been shown to be modified by the grazing of large herbivores. However, there has been little work on interactions between small mammalian herbivores and such patchy nutrient inputs, even though these interactions are potentially of major significance for plant performance and community structure.
We examined the effect of simulated cattle urine deposition on the vegetation structure, above-ground biomass and species composition of chalk grassland within enriched patches. Short-term exclosures were used to determine whether a small herbivore (rabbit) would preferentially graze the vegetation in enriched patches and what impact this interaction would have on the performance of plants in such patches. Rabbit grazing pressure determined whether nutrient inputs had a negative or positive effect on plant biomass. Nutrients increased plant biomass in the absence of grazing, but when exposed to grazing, plants in nutrient-rich patches had more biomass consumed by herbivores than neighbouring plants. Further, nutrients increased the relative palatability of a less preferred forage species ( Brachypodium pinnatum ), contributing to changes in plant community composition. We conclude that a small herbivore can drive plant responses to patchily distributed nutrients.     

Post‐fire pickings: Large herbivores alter understory vegetation communities in a coastal eucalypt forest

Fire and herbivores alter vegetation structure and function. Future fire activity is predicted to increase, and quantifying changes in vegetation communities arising from post‐fire herbivory is needed to better manage natural environments. We investigated the effects of post‐fire herbivory on understory plant communities in a coastal eucalypt forest in southeastern Australia. We quantified herbivore activity, understory plant diversity, and dominant plant morphology following a wildfire in 2017 using two sizes of exclosures. Statistical analysis incorporated the effect of exclusion treatments, time since fire, and the effect of a previous prescribed burn. Exclusion treatments altered herbivore activity, but time since fire did not. Herbivory reduced plant species richness, diversity, and evenness and promoted the dominance of the most abundant plants within the understory. Increasing time since fire reduced community diversity and evenness and influenced morphological changes to the dominant understory plant species, increasing size and dead material while decreasing abundance. We found the legacy effects of a previous prescribed burn had no effect on herbivores or vegetation within our study. Foraging by large herbivores resulted in a depauperate vegetation community. As post‐fire herbivory can alter vegetation communities, we postulate that management burning practices may exacerbate herbivore impacts. Future fire management strategies to minimize herbivore‐mediated alterations to understory vegetation could include aggregating management burns into larger fire sizes or linking fire management with herbivore management. Restricting herbivore access following fire (planned or otherwise) can encourage a more diverse and species‐rich understory plant community. Future research should aim to determine how vegetation change from post‐fire herbivory contributes to future fire risk.     

Drainage, soil fertility and fire frequency determine composition and structure of gumland heaths in northern New?Zealand

Vegetation and soils were sampled at remaining gumland heath ecosystems in northern New?Zealand to determine vegetation patterns, environmental controls and major threats to long-term persistence. Classification and ordination techniques identified six vegetation types reflecting differences in drainage, rainfall, altitude, nutrients, and time since fire. Two modal types reflected opposite ends of the main environmental spectra. Leptospermum scoparium (Myrtaceae) shrubland occurred on relatively better drained sites with lower rainfall, altitude, and soil nutrient levels, whereas Gleichenia dicarpa (Gleicheniaceae) fernland typically occurred on more poorly drained sites with higher rainfall, altitude, and nutrient levels. Another widespread vegetation type dominated by both Leptospermum scoparium and Gleichenia dicarpa occupied plots of intermediate drainage, rainfall, altitude and nutrients. The three remaining types were of limited distribution and reflected uncommon combinations of environmental conditions or recent fire. Low soil nutrients in gumlands (mean total N = 0.182%, total P = 0.004%, oven-dry weight) are reflected in low Leptospermum scoparium foliage nutrients (mean total N = 0.858%, total P = 0.034%, δ¹⁵N = δ6.06‰, oven-dry weight) and slow growth rates (mean annual height growth rate = 11.90 cm year^?1), as in heathlands in Australia and South Africa. Gumlands are threatened by non-native plant species invasion, especially Hakea sericea (Proteaceae); habitat destruction for agricultural, industrial, and suburban development; and nutrient enrichment from adjacent agricultural land. Currently, fire is much less common in gumlands (mean time since fire = 18.4 years) than during early European settlement and some communities are apparently reverting to forest. Research to investigate the use of fire as a management tool is recommended for long-term conservation of New?Zealand gumlands.     

Ant Species Identity has a Greater Effect than Fire on the Outcome of an Ant Protection System in Brazilian Cerrado

Although fire‐ and ant–plant interactions influence the community structure and dynamics of Neotropical savannas, no previous studies have considered their simultaneous effects on target host plants. We monitored the effect of ant exclusion for 3 years on leaf area loss to leaf chewing insects, thrips abundance, and reproductive output of the extrafloral nectary‐bearing shrub, Peixotoa tomentosa (Malpighiaceae). We predicted that the impact of ants on herbivores and plants would depend on the ant species, and that fire would reduce the effect of ants. We deliberately chose control plants that differed in their occupant ant species. Fire occurred in the second year of the study, allowing us to determine its effect on the benefit afforded by ants. Ants reduced leaf area loss and thrips abundance, and increased fruit and seed production in all 3 years. Some ant species were more effective than others, while plants with multiple ant species suffered higher leaf area loss than plants with a single ant species. In the year following the fire, leaf damage was greater than in the other years, regardless of the ant species, and the proportional effect of ants in reducing damage was less. Interactions affecting thrips abundance did not change following fire, nor was the benefit to the plant proportionally reduced. Overall, the identity of the ant species had a greater effect than did the occurrence of fire on the ant–herbivore–plant interaction: the identity of the ant species influenced leaf area loss, thrips numbers, and bud and seed production, while fire only modified the impact of ants on the amount of leaf area consumed by insect herbivores.     

Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system

In the Brazilian savanna many plant species bear regular associations with patrolling ants that are aggressive towards insect herbivores. However, not only ants but also several species of predatory wasps are attracted to plants due to the extrafloral nectaries (EFNs). Such wasps feed on both herbivores and plant exudates. In this study we describe the foraging behavior of the social Polistinae wasp Brachygastra lecheguana in the extrafloral nectaried shrub Banisteriopsis malifolia, and investigated the influence of patrolling ants Camponotus blandus on the activity of the wasp. Brachygastra lecheguana fed on the endophytic larvae of Anthonomus (Curculionidae) beetles that developed inside flower buds. The wasp lacerated the bud layers to reach the beetle larvae located at the bud core. The wasp visits to Ba. malifolia were statistically related to the abundance of flower buds and beetles. Ant exclusion experiments revealed that the hunting behavior of B. lecheguana on beetles was not related to the absence of C. blandus. However we found that wasps spent more time consuming extrafloral nectar on branches where ants were excluded. This is the first study reporting extrafloral nectar consumption by B. lecheguana, as well as the predation on herbivores in natural areas. In cerrado vegetation, ants benefit the plant by reducing insect herbivores, and our study provides evidence that the B. lecheguana – Ba. malifolia system represents a potential interaction where the wasp may also benefit the host plant. The value of this wasp species as a plant‐guard is discussed.     

Effects of experimental season of prescribed fire and nutrient addition on structure and function of previously grazed grassland

Aims Understanding the drivers of grassland structure and function following livestock removal will inform grassland restoration and management. Here, we investigated the effects of fire and nutrient addition on structure and function in a subtropical semi-native grassland recently released from grazing in south-central Florida. We examined responses of soil nutrients, plant tissue nutrients, biomass of live, standing dead and litter, and plant species composition to experimental annual prescribed fire applied during different seasons (wet season vs. dry season), and nutrient additions (N, P and N + P) over 9 years.Methods Experimental plots were set up in a randomized block split-plot design, with season of prescribed fire as the main treatment and nutrient addition as the subplot treatment. Species cover data were collected annually from 2002 to 2011 and plant tissue and plant biomass data were collected in 2002–2006 and 2011. Soil nutrients were analyzed in 2004, 2006 and 2011.Important findings Soil total phosphorus (P) levels increased substantially with P addition but were not influenced by prescribed fire. Addition of P and N led to increased P and N concentrations in live plant tissues, but prescribed fire reduced N in live tissue. Levels of tissue N were higher in all plots at the beginning of the experiment, an effect that was likely due to grazing activity prior to removal of livestock. Plant tissue N steadily declined over time in all plots, with annually burned plots declining faster than unburned plots. Prescribed fire was an important driver of standing dead and litter biomass and was important for maintaining grass biomass and percent cover. Nutrient addition was also important: the addition of both N and P was associated with greater live biomass and woody forbs. Removal of grazing, lack of prescribed fire, and addition of N + P led to a reduction of grass biomass and a large increase in biomass of a woody forb. Annual prescribed fire promoted N loss from the system by reducing standing dead and litter, but maintained desirable biomass of grasses.     

Constraint on woody cover in relation to nutrient content of soils in western southern Africa

The influence of soil nutrients on woody plants is poorly understood. Are trees – fire and other disturbance factors being equal – generally promoted by nutrient‐rich or by nutrient‐poor soils? To determine the edaphic parameters controlling woody cover, we sampled soils and summed the extent of the crowns of trees and tall shrubs on 364 plots at 20 sites in Namibia and adjacent South Africa, ranging from desert lichen‐fields to caesalpiniaceous woodland with associated mean annual rainfall of 11 mm to 535 mm. Our analysis included the macro‐nutrients N, P, Mg, K and Ca and the trace elements Mn, Fe, Cu and Zn. A boundary line analysis showed that woody cover was densest, with the greatest large‐scale heterogeneity, at intermediate nutrient contents, but consistently constrained at extreme nutrient richness as well as poverty. If aridity exerted the ultimate constraint at extreme nutrient richness, no such correlation with climate apparently applies at extreme nutrient poverty, where our graphs show an ‘oligotrophic decline’. Notwithstanding the importance of water, we suggest that extreme nutrient richness and poverty both favour grasses over tree seedlings. This is because catabolic dystrophy – a regime in which the supply of catabolic nutrients shortfalls their demand – is unlikely in environments where nutrient richness allows catabolic rates to match anabolic rates or where nutrient poverty constrains anabolic rates. We also reason that surpluses of photosynthate resulting from dystrophy can be allocated to lignin and that the potential for woody growth thus corresponds to soils of intermediate nutrient content. This explains why woody cover is consistently but not homogeneously densest in nutritionally intermediate plots in our dataset. Hence, the abundance of woody plants in various biomes may be determined partly by soil nutrient content, particularly of Cu, Zn, and other elements indispensible for catabolism.     

Fire and Grazing Change Herbaceous Species Composition and Reduce Beta Diversity in the Kalahari Sand System

After rainfall and soils, fire and herbivory are two of the main determinants of savanna ecosystems. Although the interactive effects of fire and herbivores on soil and vegetation are widely acknowledged few studies have addressed these two factors in concert, and none of the studies has focused on the Kalahari sand system. We experimentally studied how annual late dry season fires and grazing affect herbaceous plant species composition, above- and belowground biomass, and soil and grass nutrient concentrations in the nutrient-poor semi-arid Kalahari system in northern Botswana. Four treatments (fire, grazing, fire + grazing, and no-fire–no-grazing) were applied for two consecutive years in the late dry season. Plant species composition was affected by treatment and year. The no-fire–no-grazing treatment was distinctly different from all the other treatments in terms of species composition. Beta diversity was lower on the fire treatment and grazing treatment, but not where fire and grazing were combined. Fire and grazing alone or in combination did not have a substantial effect on biomass, soil and plant nutrients or plant species alpha diversity. Plant nitrogen was the only element that differed between treatments, with high concentrations on all the grazed treatments in the first year and low levels on the fire-alone treatment during the second year. The results show that fire and grazing mainly affect species composition and large-scale biodiversity patterns as indicated by the no-fire–no-grazing treatment being distinctly different from other treatments, suggesting the evolutionary adaptation of this dystrophic Kalahari sand system to herbivory and fire.