Leaf trait?palatability relationships differ between ungulate species: evidence from cafeteria experiments using na?ve tussock grasses

Leaf functional traits have been proposed as general indicators of plant palatability to ungulate herbivores, identifying which species are likely to be most at risk from ungulates, and how ungulate grazing may change ecosystem processes. However, few studies have tested whether leaf trait?palatability relationships are consistent across different ungulate species. The palatability of 44 native New?Zealand grass taxa (from the genera Festuca and Chionochloa) to two ungulate herbivores (sheep Ovis aries and red deer Cervus elaphus scoticus) was assessed in cafeteria experiments. There were significant differences between sheep and deer in the selection or avoidance of grass taxa, in part related to differences in response to variation in leaf functional traits. Deer had a greater tendency than sheep to select grasses with a higher specific leaf area (SLA) and to avoid taxa with a low SLA, suggesting that it is not possible to generalise leaf trait?palatability relationships across different ungulate species. Results suggest different ungulate species are likely to have additive effects on the biodiversity and ecosystem functioning of New?Zealand?s native grasslands. These findings indicate that the impacts of ungulate herbivory on ecosystem processes will depend on which grass species are present.     

Burning in a New Zealand snow-tussock grassland: Effects on soil microbial biomass and nitrogen and phosphorus availability

Fire has been an important management tool in the pastoral use of New Zealand tussock grasslands. The effects of a farm-scale pastoral fire and subsequent grazing by sheep on soil biochemical properties in tussock grasslands dominated by the narrow-leaved snow tussock (Chionochloa rigida ssp. rigida) were investigated, 1.5 and 2.5 years after the fire event, in 0-2 cm depth mineral soil at a site at 975 m altitude in Central Otago, New Zealand. The nitrogen (N) and phosphorus (P) concentrations of C. rigida leaves were also measured. Comparisons were made with soil and tussock leaves from an adjacent unburned site. At both samplings, values of total soil organic carbon (C), extractable C, microbial biomass C, and basal respiratory activity were, on average, 14%, 18%, 23%, and 40%, respectively, lower at the burned than at the unburned site. In contrast, microbial N values were roughly similar at both sites, while microbial P values were 42% higher at the burned site after 1.5 years. Phosphomonoesterase and phosphodiesterase activities were then also similar at both sites, whereas invertase activity was higher at the burned site. The greater availability of N and P at the burned site was confirmed by the higher concentrations of N and P in C. rigida leaves sampled 2 years after the fire. Ratios of microbial C:microbial N and microbial C:microbial P were significantly lower at both samplings at the burned site, and emphasise the importance of the soil microbial biomass in conserving N and P after pastoral burning in a grassland ecosystem.     

Leaf respiration and alternative oxidase in field-grown alpine grasses respond to natural changes in temperature and light

? We report the first investigation of changes in electron partitioning via the alternative respiratory pathway (AP) and alternative oxidase (AOX) protein abundance in field-grown plants and their role in seasonal acclimation of respiration. ? We sampled two alpine grasses native to New Zealand, Chionochloa rubra and Chionochloa pallens, from field sites of different altitudes, over 1 yr and also intensively over a 2-wk period. ? In both species, respiration acclimated to seasonal changes in temperature through changes in basal capacity (R??) but not temperature sensitivity (E?). In C. pallens, acclimation of respiration may be associated with a higher AOX : cytochrome c oxidase (COX) protein abundance ratio. Oxygen isotope discrimination (D), which reflects relative changes in AP electron partitioning, correlated positively with daily integrated photosynthetically active radiation (PAR) in both species over seasonal timescales. Respiratory parameters, the AOX : COX protein ratio and D were stable over a 2-wk period, during which significant temperature changes were experienced in the field. ? We conclude that respiration in Chionochloa spp. acclimates strongly to seasonal, but not to short-term, temperature variation. Alternative oxidase appears to be involved in the plant response to both seasonal changes in temperature and daily changes in light, highlighting the complexity of the function of AOX in the field.     

Little geographic or host plant genetic variation in a

The grass genus Chionochloa in New Zealand exhibits a high degree of mast seeding synchronised across species and habitats. Masting appears to be maintained by a predator satiation mechanism involving three pre-dispersal seed- and flower-feeding insects. It is not clear how important each of the three insects is in favouring the masting strategy. An undescribed cecidomyiid fly (Diptera: Cecidomyiidae) may be particularly important, since its conspicuous larvae are found throughout the South Island of New Zealand on many Chinochloa species. Despite the wide distribution of the larvae, it is not clear whether they are conspecific. Since the species is undescribed and adults are rarely seen, there may be different species on different host plants or in different geographic areas. We used Inter Simple Sequence Repeats (ISSRs) to determine whether cecidomyiid larvae found in four different areas in the South Island and on four species of Chionochloa exhibited molecular variation consistent with the presence of a single species of fly. Cluster analysis using Unweighted Pair-Group Method using Arithmetic averages (UPGMA) based on 38 ISSR fragments showed no clusters based on either host plant or geography. Analyis of Molecular Variance (AMOVA) analyses showed statistically significant differentiation among both host populations and geographic populations, but most of the molecular variation was explained by individual variation within geographic regions and host-plant populations. Thus, the molecular variation in the cecidomyiid larvae suggests the presence of a single species of cecidomyiid. Our data, combined with previous population surveys, suggest that the cecidomyiid is the most widespread of Chionochloa seed predators and may provide the selective benefit for the synchronous flowering observed among different Chionochloa populations in New Zealand.     

Adaptation of selected trees and grasses to low availability of phosphorus

High-nutrient-adapted and low-nutrient-adapted species of New Zealand tussock grasses (Chionochloa), barley (Hordeum), and several taiga trees were grown at three rates of phosphorus supply. Low-nutrient-adapted species in each group of species had similar (grasses) or lower (trees) capacities for phosphate absorption, were less efficient in producing biomass (i.e. had higher nutrient concentrations), and grew more slowly than high-nutrient-adapted species. I conclude that the major adaptation to low nutrient availability in each of these comparisons is a slow growth rate that reduces the annual nutrient requirement.     

Description of Eucalyptodiplosis chionochloae sp. nov., a cecidomyiid feeding on inflorescences of Chionochloa (Poaceae) in New Zealand

Abstract

A cecidomyiid that feeds on developing seeds in the inflorescences of the New Zealand tussock grasses Chionochloa australis, C. conspicua, C. crassiuscula, C.flavescens, C. macra, C. oreophila, C. pallens, C. rigida, C. rubra, C. spiralis and C. teretifolia is formally described from C. pallens. The new species, named Eucalyptodiplosis chionochloae Kolesik, is the most ubiquitous of flower feeders of Chionochloa. Its larvae do not form galls but feed on the developing seeds in autumn, overwinter as diapausing larvae inside the floret, and pupate then rapidly eclose in summer, sometimes after extended diapause. Methods for rearing adults are described. Based on its morphological characters this species is most closely related to two described congeners that form galls on buds of Eucalyptus trees in Australia. Seed predation by Eucalyptodiplosis chionochloae larvae appears to be the primary driver of the extreme mast seeding (variation among years in flower crops) seen in the host Chionochloa species.     

Nuclear DNA variation, chromosome numbers and polyploidy in the endemic and indigenous grass flora of New Zealand

BACKGROUND AND AIMS: Little information is available on DNA C-values for the New Zealand flora. Nearly 85 % of the named species of the native vascular flora are endemic, including 157 species of Poaceae, the second most species-rich plant family in New Zealand. Few C-values have been published for New Zealand native grasses, and chromosome numbers have previously been reported for fewer than half of the species. The aim of this research was to determine C-values and chromosome numbers for most of the endemic and indigenous Poaceae from New Zealand. SCOPE: To analyse DNA C-values from 155 species and chromosome numbers from 55 species of the endemic and indigenous grass flora of New Zealand. KEY RESULTS: The new C-values increase significantly the number of such measurements for Poaceae worldwide. New chromosome numbers were determined from 55 species. Variation in C-value and percentage polyploidy were analysed in relation to plant distribution. No clear relationship could be demonstrated between these variables. CONCLUSIONS: A wide range of C-values was found in the New Zealand endemic and indigenous grasses. This variation can be related to the phylogenetic position of the genera, plants in the BOP (Bambusoideae, Oryzoideae, Pooideae) clade in general having higher C-values than those in the PACC (Panicoideae, Arundinoideae, Chloridoideae + Centothecoideae) clade. Within genera, polyploids typically have smaller genome sizes (C-value divided by ploidy level) than diploids and there is commonly a progressive decrease with increasing ploidy level. The high frequency of polyploidy in the New Zealand grasses was confirmed by our additional counts, with only approximately 10 % being diploid. No clear relationship between C-value, polyploidy and rarity was evident.     

Interspecific and seasonal dietary differences of Himalayan thar, chamois and brushtail possums in the central Southern Alps, New Zealand

Himalayan thar or tahr (Hemitragus jemlahicus), Alpine chamois (Rupicapra rupicapra) and brushtail possums (Trichosurus vulpecula) are native to the Himalaya, Europe and Australia, respectively, but are now sympatric in parts of the central Southern Alps, New Zealand. All three species are managed as pests by the Department of Conservation. We analysed the diets of 246 thar, 78 chamois and 113 possums collected in the central Southern Alps during 1988–1996. The diets of thar and chamois strongly overlapped, but thar ate more grasses and less herbs and woody plants than chamois. The diet of possums differed from the diets of thar and chamois, containing a different suite of herbs and almost no grasses. The diets of thar, chamois and possums varied seasonally. Contrary to expectation, the diets of adult male and female thar were similar during the period when they are spatially segregated (October–May). Chamois sampled outside the thar range ate more species than those sampled inside the thar range, suggesting that the presence of thar modifies the diet of chamois. If managers wish to protect the dominant Chionochloa snow tussocks then thar should be controlled; chamois should be controlled to protect herbs such as Ranunculus spp.; and possums should be controlled if Podocarpus nivalis or Muehlenbeckia axillaris need to be protected. However, managers need to be aware of the potential for chamois to increase in abundance, at least in some sites, if thar are controlled to low abundance, and that this could lead to different impacts on plant populations.     

Palatability factors and nutritive values of the food of buffaloes (Syncerus caffer) in Uganda

The food preferences of tame buffaloes grazing different vegetation types were recorded. The effects of fire and rainfall were investigated. By initiating a new growth cycle, fire altered the food habits of large herbivores and also the distribution of small mammals. Seasonal variations in the effect of rainfall on plant growth explains why preferences did not change in proportion to precipitation. The more important grasses were collected and their parts analysed physically and chemically. Their stem to leaf ratio is considered to have influenced the species eaten by coarse grazers such as buffalo. A broad correlation was obtained between buffalo preferences and the percentage green leaf of Hyparrhenia filipendula and Themeda triandra. This correlation was improved when preferences were compared with the number of grams green leaf carbohydrate per 100 g dry grass. There was a negative response to the silica present in dead leaves, but stem fibre appeared to have little effect on preferences. A grazing animal experiences a combination of attractants and repellants, i.e. green leaf carbohydrate and protein less stem fibre and dead leaf silica. Other palatability factors were noted, in particular tannins and oils in grasses of the tribe Andropogoneae. These may act as repellants, possibly only when the grazing animal has alternative vegetation from which to choose. The mineral content of grasses appeared to be adequate and calcium and phosphorus were present in about the required proportions. The observations support the belief that selective grazing enables a herbivore to consume a diet of significantly higher nutritive value than that of the average sward. Browsing and new growth after fire help compensate for the low nutritive value of pasture during the dry season. The nutritive value of the diet is considered to be the proximal factor in the seasonal cycle of reproduction of the buffalo and probably the ultimate factor in the reproduction of certain other ungulates and rodents.     

A

A valley mire was sampled on the flanks of Swampy Hill, east Otago, New Zealand. It formed in a narrow valley, apparently originally comprising two basins. The end of the mire nearest the outlet contained species typical of fens (i.e., rheotrophic mires). At the head of the valley there was a section of the mire with mixed vegetation cover comprising the tussock grass Chionochloa rubra, Sphagnum species, and cushion/herb/shrub cover. Ombrotrophic status of this section was indicated by a slightly raised profile, greater acidity, lower exchangeable Na and K, and lower substrate cation exchange capacity, identifying it as a bog. Total Ca:Mg molar ratios were generally above 1.0, but this rule-of-thumb for ombrotrophic status may be inapplicable here. It is not known whether New Zealand Sphagnum species are as efficient at lowering the pH as those investigated elsewhere. Macrofosssil evidence indicates that some components of the bog, such as Sphagnum and epacridaceous subshrubs, have remained constant, almost since the inception of the bog. However, Empodisma minus, currently absent from the bog and rare in the region, was present at one stage. The change from cover with Empodisma and Dracophyllum as significant components, to the present Chionochloa/Sphagnum/cushion composition, occurred a few hundred years ago, probably initiated by fire. Comparison with preliminary information for other bogs suggests that those in the eastern part of the South Island vary considerably in species composition, with individualistic assemblages of species. The site is seen as having high conservation values. To protect these values the bog needs protection from invasive exotic weeds, and from damage by wild pigs.     

Plastic traits of an exotic grass contribute to its abundance but are not always favourable

In herbaceous ecosystems worldwide, biodiversity has been negatively impacted by changed grazing regimes and nutrient enrichment. Altered disturbance regimes are thought to favour invasive species that have a high phenotypic plasticity, although most studies measure plasticity under controlled conditions in the greenhouse and then assume plasticity is an advantage in the field. Here, we compare trait plasticity between three co-occurring, C(4) perennial grass species, an invader Eragrostis curvula, and natives Eragrostis sororia and Aristida personata to grazing and fertilizer in a three-year field trial. We measured abundances and several leaf traits known to correlate with strategies used by plants to fix carbon and acquire resources, i.e. specific leaf area (SLA), leaf dry matter content (LDMC), leaf nutrient concentrations (N, C:N, P), assimilation rates (Amax) and photosynthetic nitrogen use efficiency (PNUE). In the control treatment (grazed only), trait values for SLA, leaf C:N ratios, Amax and PNUE differed significantly between the three grass species. When trait values were compared across treatments, E. curvula showed higher trait plasticity than the native grasses, and this correlated with an increase in abundance across all but the grazed/fertilized treatment. The native grasses showed little trait plasticity in response to the treatments. Aristida personata decreased significantly in the treatments where E. curvula increased, and E. sororia abundance increased possibly due to increased rainfall and not in response to treatments or invader abundance. Overall, we found that plasticity did not favour an increase in abundance of E. curvula under the grazed/fertilized treatment likely because leaf nutrient contents increased and subsequently its' palatability to consumers. E. curvula also displayed a higher resource use efficiency than the native grasses. These findings suggest resource conditions and disturbance regimes can be manipulated to disadvantage the success of even plastic exotic species.     

Grazing-induced changes in plant composition affect litter quality and nutrient cycling in flooding Pampa grasslands

Changes in plant community composition induced by vertebrate grazers have been found to either accelerate or slow C and nutrient cycling in soil. This variation may reflect the differential effects of grazing-promoted (G+) plant species on overall litter quality and decomposition processes. Further, site conditions associated with prior grazing history are expected to influence litter decay and nutrient turnover. We studied how grazing-induced changes in plant life forms and species identity modified the quality of litter inputs to soil, decomposition rate and nutrient release in a flooding Pampa grassland, Argentina. Litter from G+ forbs and grasses (two species each) and grazing-reduced (G−) grasses (two species) was incubated in long-term grazed and ungrazed sites. G+ species, overall, showed higher rates of decomposition and N and P release from litter. However, this pattern was primarily driven by the low-growing, high litter-quality forbs included among G+ species. Forbs decomposed and released nutrients faster than either G+ or G− grasses. While no consistent differences between G+ and G− grasses were observed, patterns of grass litter decay and nutrient release corresponded with interspecific differences in phenology and photosynthetic pathway. Litter decomposition, N release and soil N availability were higher in the grazed site, irrespective of species litter type. Our results contradict the notion that grazing, by reducing more palatable species and promoting less palatable ones, should decrease nutrient cycling from litter. Plant tissue quality and palatability may not unequivocally link patterns of grazing resistance and litter decomposability within a community, especially where grazing causes major shifts in life form composition. Thus, plant functional groups defined by species’ “responses” to grazing may only partially overlap with functional groups based on species “effects” on C and nutrient cycling.     

A global comparison of the nutritive values of forage plants grown in contrasting environments

Forage plants are valuable because they maintain wild and domesticated herbivores, and sustain the delivery of meat, milk and other commodities. Forage plants contain different quantities of fibre, lignin, minerals and protein, and vary in the proportion of their tissue that can be digested by herbivores. These nutritive components are important determinants of consumer growth rates, reproductive success and behaviour. A dataset was compiled to quantify variation in forage plant nutritive values within- and between-plant species, and to assess variation between plant functional groups and bioclimatic zones. 1255 geo-located records containing 3774 measurements of nutritive values for 136 forage plant species grown in 30 countries were obtained from published articles. Spatial variability in forage nutritive values indicated that climate modified plant nutritive values. Forage plants grown in arid and equatorial regions generally contained less digestible material than those grown in temperate and tundra regions; containing more fibre and lignin, and less protein. These patterns may reveal why herbivore body sizes, digestion and migration strategies are different in warmer and drier regions. This dataset also revealed the capacity for variation in the nutrition provided by forage plants, which may drive consumer species coexistence. The proportion of the plant tissue that was digestible ranged between species from 2 to 91%. The amount of fibre contained within plant material ranged by 23–90%, protein by 2–36%, lignin by 1–21% and minerals by 2–22%. On average, grasses and tree foliage contained the most fibre, whilst herbaceous legumes contained the most protein and tree foliage contained the most lignin. However, there were individual species within each functional group that were highly nutritious. This dataset may be used to identify forage plant species or mixtures of species from different functional groups with useful nutritional traits which can be cultivated to enhance livestock productivity and inform wild herbivore conservation strategies.     

Leaf palatability and decomposability increase during a 200‐year‐old post‐cultural woody succession in New Zealand

Question: How do aggregate trait values and functional diversity of leaf traits linked to palatability and decomposability change during a woody post‐cultural succession spanning 200 years? Location: Coastal Marlborough, South Island, New Zealand. Methods: The biomass of all woody species was determined in 32 20‐m × 20‐m plots ranging from 10 to 200 years in time since last disturbance. Species abundances were combined with data on leaf nutrient, secondary metabolite and structural carbohydrate content to calculate biomass‐weighted trait means (i.e. aggregate trait values) and functional diversity index values for each plot. Aggregate trait values and functional diversity were regressed on successional age and total live above‐ground carbon content to examine functional shifts with succession and one consequence of succession – increasing above‐ground carbon. Results: Almost all significant regressions between aggregate trait values and both successional age and above‐ground carbon indicated a shift toward increased leaf palatability and decomposability during succession. The relationships were all non‐linear, with aggregate trait value shifts occurring relatively early in the successional sequence. There was weak evidence for an increase in functional richness with succession, but this was a secondary effect relative to the shifts in aggregate trait values. Conclusions: These results are in direct contrast with studies of the early stages of herbaceous post‐cultural successions from grasslands to shrublands, which have found a shift towards communities of decreasing palatability and decomposability, suggesting that functional shifts in woody succession may be fundamentally different.     

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.     

Clipping and irrigation enhance grass biomass and nutrients: Implications for rangeland management

Increasing frequency of drought and high herbivore pressure significantly affect individual grass functions in semiarid regions. Reseeding of degraded rangelands by native grass species has been recommended as a tool for restoration semiarid rangelands. However, how grass species used for reseeding respond to stressors has not been fully explored. We examined biomass allocation and nutrient contents of Cenchrus ciliaris and Chloris gayana in the semiarid Borana rangelands, Ethiopia. We tested clipped mature tufts of the same species for biomass allocation and nutritive values. Further, shifts in rainfall and herbivory were simulated by three irrigation and four clipping treatments, respectively, for newly established grasses in pot and field plot experiments. Aboveground biomass (AG_B) significantly declined by up to 75% under increased clipping in mature tufts. In contrast, clipping significantly stimulated up to 152% higher AG_B of newly established grasses. Lower irrigation reduced the AG_B by 24 and 42% in C. ciliaris and in C. gayana, respectively. Clipping, further, significantly enhanced grass nutrients in grass tufts by up to 82 and 105% in C. ciliaris and C. gayana, respectively. Hence, management should focus on balancing this trade-off in mature grasses for nutritious rangeland production by clipping and storing for later supplemental feeding when grass nutrients drop. Further, young pastures should be moderately clipped/grazed for better establishment and biomass allocation. Additionally, our experiments established the first interactive effect of clipping and irrigation frequencies on the biomass allocation of native grasses in the semiarid Borana rangelands, Ethiopia. Knowledge of these interacting factors is deemed essential for policy makers to enhance productivity of degraded rangelands such as the Borana rangelands.     

Kinetics of nitrate uptake by different species from nutrient-rich and nutrient-poor habitats as affected by the nutrient supply

The species Urtica dioica L., Plantago major ssp. major L., Plantago lanceolata L., Hypochaeris radicata L. ssp. radicata and Hypochaeris radicata ssp. ericetorum Van Soest were grown under high and low nutrient conditions (1/4 Hoagland and 2% of 1/4 Hoagland further called the 100% and 2% treatment, containing 3.75 mM NO^-₃ and 0.075 mM NO^-₃, respectively). After a certain period half of the plants were transferred from low to high or high to low nutrients, yielding the 100%/2% and the 2%/100% treatments. The kinetics of nitrate uptake in the range of system I of the five species grown under the different nutrient conditions were measured during a three week experimental period. The nitrate uptake of all the species showed the characteristic features of Michaelis-Menten kinetics. Under low nutrient conditions the apparent V_max of U. dioica expressed per g dry root was lower than under high nutrient conditions. For H. radicata ssp. radicata and for H. radicata ssp. ericetorum the reverse was found. The V_max values of P. major ssp. major were almost the same for the two treatments. The apparent V_max in young plants of P. lanceolata was higher in the 100% treatment than in 2%; whereas the reverse was found in mature plants. The results are explained in relation to the relative growth rate, the shoot to root ratio and the natural environment of the species. The apparent K_m values were not influenced by the different treatments. Differences in K_m between the species, if any, were very small. It is suggested that the V_max is a more important parameter for the distribution of plant species in the field than the K_m. The rate of nitrogen accumulation was calculated from growth data and the contents of nitrate and reduced nitrogen. It is concluded that the V_max of system I for nitrate uptake in most cases was sufficient to explain the observed growth rates.     

New high latitude Capnuchosphaera species (Triassic Radiolaria) from Waipapa Terrane,New Zealand

Two new species and one new subspecies of genus Capnuchosphaera, (Capnuchosphaera tumida nov. sp., C. waihekeensis nov. sp. and C. texensis australis nov. ssp.) are described herein from phosphatic nodules included in mudstone and sandstone beds of the Waipapa Terrane, Waiheke Island, New Zealand. The phosphatic nodules yielded a rich Late Triassic (Carnian-Norian) radiolarian fauna, with a high abundance of spumellarian taxa including numerous species of the genera Capnuchosphaera, Vinassaspongus, Kahlerosphaera, Sarla and Dumitricasphaera. Waiheke Island Capnuchosphaera are characterized by a large cortical shell and a distinctively low ratio of spine length to cortical shell diameter. These features differ significantly from those of Capnuchosphaera in the Tethyan Realm and are considered to be the result of adaptation to an Austral-New Zealand peripheral ocean of Gondwanaland in the Mesozoic Southern Hemisphere.     

Grazing effects on herbage nutritive values depend on precipitation and growing season in Inner Mongolian grassland

Aims Grasslands are widely used for production of livestock, which depend on the nutritive value of herbage species. However, there are still large uncertainties as to how grazing, precipitation and growing season interactively affect herbage nutritive value.Methods Here, based on a grazing experiment with seven grazing intensities in an Inner Mongolian grassland since the year 2005, we analysed nutritive value of four dominant grass species from June to September in both a relatively wetter year (2008) and a drier year (2010). Herbage with high nutritive value is characterized by high concentration of crude protein, high concentration of cellulase digestible organic matter and low concentration of neutral detergent fibre.Important findings We found that (i) grazing significantly increased the nutritive value of Leymus chinensis, Agropyron michnoi and Cleistogenes squarrosa but had minor effects on the nutritive value of Stipa grandis. (ii) For all species, nutritive values were greater in the wetter year than in the drier year and were greatest in the early growing season (June) and lowest at the end of the growing season (September). Inter-annual and inter-seasonal variations in nutritive value were much greater for L. chinensis and A. michnoi than for C. squarrosa and S. grandis, suggesting higher water use efficiency for the latter two species. (iii) Grazing significantly decreased the drought resistance of three species, but not of S. grandis. (iv) Grazing significantly increased inter-month variation in nutritive value for L. chinensis and A. michnoi but had relatively minor or no effects on that of C. squarrosa and S. grandis. Therefore, grazing effects on species nutritive values showed strong inter-annual and seasonal patterns, and species-specific responses might be related to species traits (i.e. water use efficiency, digestibility). Our results have important implications for ecosystem management of arid and semi-arid grasslands under intense grazing and global climatic change.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.