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Our study investigated the effects of fire frequency on the tree and grass components of Cumberland Plain Woodland, an endangered grassy eucalypt woodland in eastern Australia. We located three sites within each of three fire frequency classes (high, moderate, low) based on fire history and a similar time since last fire. For trees, we asked whether fire frequency affected density, eucalypt population structure, basal area and spatial patterning. For grasses, we tested for fire frequency effects on total abundance and the abundance of two common species. Density of trees was not significantly affected by fire frequency for juveniles, saplings or adult trees and neither was basal area per hectare. Features of a recruitment bottleneck model were present for eucalypts. There was a large pool of suppressed juveniles making up more than half the population at each fire frequency. Saplings were the smallest group and were susceptible to fire‐induced stem mortality, with particularly low numbers relative to juveniles and small trees where fire was frequent or intense. Despite this difference in sapling mortality, numbers of small trees did not differ with fire frequency. Saplings could transition to small trees in equal numbers if they did so more quickly at high than at lower fire frequencies, and if recruitment of saplings into the tree layer was controlled independently of fire frequency. The size hierarchy of small to medium eucalypt trees was homogeneous over eight of the nine sites, spatial patterning of adult trees was random tending to regular, and mean tree size decreased with density at all sites. These features of eucalypt population structure are indicative of possible resource competition which could regulate tree recruitment. Total grass cover index was high across all fire frequencies, with Themeda triandra dominating at high and moderate fire frequencies and Microlaena stipoides at low fire frequency.

For speciose, but poorly known groups, such as terrestrial arthropods, functional traits present a potential avenue to assist in predicting responses to environmental change. Species turnover is common along environmental gradients, but it is unclear how this is reflected in species traits. Community-level change in arthropod traits, other than body size, has rarely been explored across spatial scales comparable to those examined here. We hypothesized that the composition and morphological traits of spider assemblages would differ across a gradient of climate and habitat structure. We examined foliage-living spider assemblages associated with Themeda triandra grasslands along a 900 km climatic gradient in south-eastern Australia. We used sweep-netting to collect T. triandra-associated spiders and counted juveniles and identified adults. We also measured morphological traits of adult spiders and noted their hunting mode. Associations with measures of habitat structure were less consistent than relationships with climate. Both juvenile and adult spiders were more abundant in warmer sites, although species richness was not affected by temperature. We found distinct turnover in species composition along the climatic gradient, with hunting spiders, particularly crab spiders (Thomisidae), making up a greater proportion of assemblages in warmer climates. A range of traits of spiders correlated with the climatic gradient. For example, larger spider species and species that were active hunters were more common in warmer climates. Changes in morphological traits across species, rather than within species drove the morphology-climate relationship. Strong climate-trait correlations suggest that it may be possible to predict changes in functional traits of assemblages in response to anthropogenic disturbances such as climate change.

Species composition and productivity of natural grasslands are influenced by soil nutrient status. With high resource availability, productivity is expected to increase, and competition is assumed to gain prominence with predicted exclusion of species of lower competitive ability. During 2010 and 2011 we used the dry weight rank method to measure above-ground phytomass production of herbage in 96 plots (9 m × 2.7 m) fertilized for 60 years with two forms of nitrogen (N as limestone ammonium nitrate or ammonium sulphate at four levels: 0, 7.1, 14.1, 21.2 g m−2), phosphorus (P as superphosphate at two levels: 0, 33.6 g m−2), and lime (two levels: 0, 225 g m−2). Light attenuation was measured as the proportion of photosynthetically active radiation reaching the lower leaf layers of the grasses and the ground surface. Light conditions beneath the grass layer were reduced by nutrient addition to 30% of full sunlight but remained above 60% in non-fertilized plots. Grass total above-ground phytomass production increased with nutrient addition. The strongest yield responses were attained with N plus P addition. Species responses showed that Themeda triandra and Hyparrhenia hirta decreased in above-ground phytomass production with nutrient addition while Panicum maximum, Eragrostis curvula and E. plana increased. These findings are discussed in terms of competitive interactions among species, their position in the grass canopy and their physiological tolerances to high nitrogen environments.

Green leaves of six grass species in temperate Australia showed considerable circadian variation (especially in summer) in water content, which peaked late in the night and fell from sunrise to an afternoon minimum. In summer, water content of a set of ‘wetter’ species (Axonopus affinis, Pennisetum clandestinum and Paspalum dilatatum) was consistently higher and varied less profoundly through the 24 h than that of a ‘drier’ set (Imperata cylindrica, Poa labillardieri and Themeda triandra). Hour-to-hour and day-to-day variation in leaf water content of P. dilatatum in summer partly but imperfectly reflected ambient temperature. Recognizing the risks for herbivores of visiting water sources, we considered leaf water content as a source of free water for leaf-grazing herbivores, eastern grey kangaroos Macropus giganteus and common wombats Vombatus ursinus. Using known field metabolic rates, energy requirements and digestive efficiencies, we calculated these two species' free-water intakes when satisfying their energy needs with these grasses but distributing their grazing in different schedules. Both species would obtain more free water by their observed foraging schedules than by random or daytime-only schedules. We calculated that the measured grasses could satisfy the water requirements of wombats but not always those of kangaroos, who would need to drink when forage water content was less than about 70%. Water content of grass leaf could rise above and fall below this value within a day, creating some scope for kangaroos to manage their need to visit water by adjusting foraging schedules or choice of grass species. In future studies, circadian variation in leaf water content should be considered as a factor (together with their thermal physiology) driving herbivores' activity scheduling, and affecting their dietary selection.

Atmospheric CO2 has more than doubled since the last glacial maximum (LGM) and could double again within this century, largely due to anthropogenic activity. It has been suggested that low [CO2] contributed to reduced tree cover in savanna and grassland biomes at LGM, and that increasing [CO2] over the last century promoted increases in woody plants in these ecosystems over the past few decades. Despite the implications of this idea for understanding global carbon cycle dynamics and key global role of the savanna biome, there are still very few experimental studies quantifying the effects of CO2 on tree growth and demography in savannas and grasslands. In this paper we present photosynthetic, growth and carbon allocation responses of African savanna trees (Acacia karroo and Acacia nilotica) and a C4 grass, Themeda triandra, exposed to a gradient of CO2 concentrations from 180 (typical of LGM) to 1000 µmol mol−1 in open-top chambers in a glasshouse as a first empirical test of this idea. Photosynthesis, total stem length, total stem diameter, shoot dry weight and root dry weight of the acacias increased significantly across the CO2 gradient, saturating at higher CO2 concentrations. After clipping to simulate fire, plants showed an even greater response in total stem length, total stem diameter and shoot dry weight, signalling the importance of re-sprouting following disturbances such as fire or herbivory in savanna systems. Root starch (per unit root mass and total root starch per plant) increased steeply along the CO2 gradient, explaining the re-sprouting response. In contrast to the strong response of tree seedlings to the CO2 gradient, grass productivity showed little variation, even at low CO2 concentrations. These results suggest that CO2 has significant direct effects on tree recruitment in grassy ecosystems, influencing the ability of trees to recover from fire damage and herbivory. Fire and herbivore regimes that were effective in controlling tree increases in grassy ecosystems could thus be much less effective in a CO2-rich world, but field-based tests are needed to confirm this suggestion.

The importance of disturbance for regulating the structure and diversity of grassy ecosystems is widely recognized, but disturbance-mediated interactions between grassland composition and grassland resilience, and consequent implications for conservation management, are less well documented. We established replicated burning, mowing and (non-livestock) grazing regimes in two contrasting grassy woodland remnants in south-eastern Australia, and monitored the dynamics and resilience of the matrix-forming tussock grasses, Poa sieberiana (Poa) and Themeda australis (Themeda), over 12 years. Introduction of frequent burning to a Poa-dominated understorey in a rarely burnt woodland enhanced dominance by Themeda, and conversely, reduced fire frequency in a frequently burnt Themeda grassland substantially increased Poa abundance. Burning was potentially detrimental in the Poa-dominated woodland, but sward resilience (recovery after the 2002 burn) increased as Themeda increased with repeated burning. By contrast, the Themeda grassland was resilient to 4- and 8-yearly burning, but biennial burning led to poor resilience and high tussock mortality under drought conditions. Contrary to other mesic grasslands, cessation of burning had not caused sward collapse by 14 years post-fire despite high litter accumulation, potentially due to compensatory growth of Poa, lower site productivity and drought. Biennial mowing without slash removal was similar to 4-yearly burning in effects, while exclusion from kangaroo and rabbit grazing significantly increased sward biomass and contributed to increased Poa cover and inflorescence production. We conclude that functional complementarity associated with mixed dominants enhances resilience to variable disturbance regimes, and that below certain thresholds of abundance of each dominant, this resilience declines. Conservation management of Themeda-Poa ecosystems should thus aim to maintain an effective balance of these dominants.

Seven grass species were grown in monocultures and in multispecies mixtures along a gradient of total nutrient levels that ranged from 1/64 to 16 � the normal level of nutrient solution. The seven grasses represented three ecological groups: (i) three perennial species native to Australia ( Themeda triandra, Poa labillardieri and Danthonia carphoides ); (ii) two introduced annuals ( Vulpia bromoides and Hordeum leporinum ); and (iii) two introduced perennials ( Lolium perenne and Dactylis glomerata ). We hypothesized that the native grasses would prove less competitive when grown at increased nutrient levels than those introduced from Europe. Results supported the hypothesis. The native species were unable to compete in mixtures even at the lowest nutrient level, where T. triandra was the most productive species in monoculture. Lolium perenne and Dactylis glomerata dominated mixtures at intermediate nutrient levels. The responses of the annual introduced grasses differed in that Vulpia bromoides showed an optimum at intermediate nutrient levels in both monoculture and in mixtures, whereas Hordeum leporinum dominated at the highest nutrient levels in mixture but was suppressed by V. bromoides, L. perenne and D. glomerata at intermediate levels. The results are discussed in terms of predicting species responses in mixtures from their performance in monocultures as well as in terms of previous observations on the sequential changes in botanical composition of south-eastern Australian grasslands after 150 years of continuous grazing by sheep.

The effects of pasture management options (sowing introduced legumes and grasses, timber treatment, applying fertilizer, cultivation before sowing, and stocking rate) on species diversity were measured at two experimental sites (Hillgrove and Cardigan) near Charters Towers, northeast Queensland. Species were divided into three groups (sown, native and exotic) and diversity was measured as species density (number of species recorded in each plot and number of species/quadrat) annually from 1982 to 1992. The responses of individual native and naturalized species to treatment were also determined. All management options affected diversity but the responses varied with site and season, and with the different measurement scales. The density of sown species either increased or was unaffected by all the management options; there were no significant decreases. The density of native species showed both positive and negative responses; it increased at high stocking rates and with tree killing at Hillgrove, and decreased with pasture sowing and cultivation. The density of exotic species increased as stocking rate was increased and decreased when pastures were sown (although not at the quadrat scale at Hillgrove). Overall the most diverse vegetation was on plots grazed at high stocking rates; at the plot scale these were native pastures but at the quadrat scale the sown pastures had more species. Among the native and naturalized species, only Portulaca spp. were more frequent on the oversown plots than the native pasture plots; 48% (Hillgrove) and 68% (Cardigan) of the species were less frequent on the oversown plots. Fertilizer application had little effect on species frequencies, while timber treatment resulted in both increases and decreases in frequency of a small number of species. The species were divided into four groups on the basis of their responses to stocking rate: a grazing-sensitive group (e.g. Themeda triandra), two grazing-tolerant groups which either slightly decreased (e.g. Chrysopogon fallax) or slightly increased (e.g. Sida spinosa) in frequency as stocking rate increased, and a fourth group of species which were frequent only at high stocking rates (e.g. Bothriochloa pertusa). There were no close relationships between herbage yield and species density.

Using a randomized experimental design, plots of a savanna grassland were subjected to two levels of grass tuft removal (50% and 90%) in two ways; non-selective (all species removed in proportion to abundance) and selective (tufts of the most palatable species removed first, then the next most palatable, etc.). The plots were maintained in their cleared states for three years, then monitored for the next five. In general, the sward was resilient to the disturbance except for the 90% selectively cleared treatment, in which a dominant, palatable species (Themeda triandra) failed to recover (though die most palatable species, Sorghum plumosum, did recover). The recovery patterns were dependent on post-disturbance conditions, and markedly influenced by a particular rainy season and a fire during one of the dry seasons. In addition to species effects, the treatments induced changes in spatial patterning and associated micro-scale hydrology. These effects persisted in the 90% removal treatment. In this regard the results are scale-dependent, and the same percentage removals at different scales (e. g. 5 × 5 m patches rather than tuft × tuft scale) would lead to differences in ability to recover. In terms of value to livestock the selective 90% removal treatment was in a poor state at the end of die experiment. In all treatments die trajectory of species changes was back towards the controls, but the selective 90% plots were fully re-vegetated before this could be achieved. In these plots, the final steps to complete recovery will occur only after death of established new tufts.