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.     

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.     

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.     

Diet overlap between small ruminants and the European hare in a Mediterranean shrubland

Seasonal diets of goats, sheep and European hares (Lepus europaeus) were examined using microhistological analysis of feces collected when these herbivores grazed together in a typical Mediterranean shrubland. Approximately half of the total diet content of goats was shrubs (mainly kermes oak, Quercus coccifera), while that of hares was grasses (mostly brush grass, Chrysopogon gryllus). Sheep had a more balanced diet consisting mainly of grasses, forbs, and shrubs. Dietary overlap between goats and sheep was high throughout the year. In contrast, there was very low dietary overlap between small ruminants and hares. Dietary diversity was high in spring and low in winter across all species, with sheep in general displaying higher dietary diversity across all seasons than goats and hares. Goats had intermediate and hares had low dietary diversity across all seasons. Communal grazing by small ruminants and hares ensures that there is a more uniform use of the available forage resources than if a single herbivore is left to graze an area.     

Silica in grasses as a defence against insect herbivores: contrasting effects on folivores and a phloem feeder

1. Silica, deposited as opaline phytoliths in the leaves of grasses, constitutes 2-5% of dry leaf mass, yet its function remains unclear. It has been proposed that silica may act as an antiherbivore defence by increasing the abrasiveness and reducing the digestibility of grass leaves, although there is little direct experimental evidence to support this. 2. We investigated the effects of manipulated silica levels on the abrasiveness of the leaves of five grass species. We also examined the effects of silica levels on the feeding preferences, growth performance and digestion efficiency of two folivorous insects and one phloem-feeding insect. 3. Silica addition resulted in increases to leaf abrasiveness in four of the five grass species studied. Silica addition also deterred feeding by both folivores and reduced their growth rates and digestion efficiency. 4. These effects resulted in lower pupal mass of the lepidopteron larvae Spodoptera exempta and compensatory feeding by the orthopteran, Schistocerca gregaria. In contrast, silica had no effects on the feeding preference or the population growth of the phloem feeder, Sitobion avenae. 5. Our results demonstrate that silica is an effective defence against folivorous insects, both as a feeding deterrent, possibly mediated by increased abrasiveness, and as a digestibility reducer. The effects of silica on pupal mass and development time may impact on herbivore fitness and exposure to natural enemies. 6. These results are the first demonstration of a direct effect of silica on the abrasiveness of grasses and the adverse impact of silica on herbivore preference and performance.     

More than herbivory: levels of silica‐based defences in grasses vary with plant species,genotype and location

Silica defences in grasses have recently been suggested to be a potential driver of vole population dynamics. However, the ability of grasses to induce silica in response to herbivory has not been tested in northern ecosystems where small rodents are important herbivores. We conducted a large‐scale field experiment in subarctic tundra using three river catchments differing in herbivore densities, and examined the effects of small rodent and/or reindeer exclusion on leaf silica levels in five grass species (Avenella flexuosa, Anthoxanthum nipponicum, Calamagrostis phragmitoides, Deschampsia cespitosa and Phleum alpinum). We also conducted a greenhouse experiment using three of these species (A. flexuosa, A. nipponicum and D. cespitosa) and Festuca ovina to determine whether intraspecific genotypic variation affects baseline silica concentrations and the capacity to induce silica in response to simulated grazing. Baseline leaf silica concentrations and silica induction varied with plant species in both experiments, with catchment in the field experiment and with genotype in the greenhouse experiment. These findings show that the allocation to silica defences in grasses is highly variable, and suggest that the combined effects of grazing pressure, plant species and intraspecific genotypic differences are likely to determine the circumstances under which silica induction may be an optimal defence strategy. A better understanding of the interplay between grazing and other factors influencing silica induction is necessary to interpret the role of silica in plant–herbivore interactions.     

Grazing and productivity alter individual grass size dynamics in semi-arid woodlands

The spatial arrangement of perennial vegetation is critical for ecosystem function in drylands. While much is known about how vegetation patches respond to grazing and abiotic conditions, the size dynamics of individual plants is mostly limited to theoretical studies. We measured the size distribution (mean, variance, skewness) and density of individual grasses, and grass species composition at 451 sites spanning a range of grazing intensities across three broad vegetation communities in semi-arid eastern Australia. We assessed the relative role of grazing by livestock (cattle and sheep), native (kangaroos) and introduced (rabbits) free ranging herbivores, and several environmental measures (productivity, diversity, composition and groundstorey plant cover) on the size distribution and density of individual grasses. We found mean grass size and density were more sensitive to shifts in grazing intensity and environmental conditions than size variance or the frequency of the smallest individuals (skewness), and shifts were mostly driven by site productivity and cattle and kangaroo grazing. Sheep grazing only reduced mean grass size, and rabbit grazing had no consistent effects. Importantly, we found that site productivity and species composition altered the impacts of grazing on grass density and size distribution. For example, increasing cattle grazing led to larger grasses in low productivity sites. It also led to larger, denser, more variable-sized grasses among grass species from sites with finer soil texture. Increasing kangaroo grazing led to smaller, denser individuals among grass species from sites with coarse soil texture. At high diversity sites kangaroo grazing led to denser, more homogenised grass sizes with a lower frequency of small individuals. Understanding the in situ response of individual plant sizes gives us insights into the processes driving shifts in perennial vegetation patchiness, improving our ability to predict how the spatial arrangement of ecosystems might change under global change scenarios.     

Preference for C4 shade grasses increases hatchling performance in the butterfly,Bicyclus safitza

The Miocene radiation of C₄ grasses under high‐temperature and low ambient CO₂ levels occurred alongside the transformation of a largely forested landscape into savanna. This inevitably changed the host plant regime of herbivores, and the simultaneous diversification of many consumer lineages, including Bicyclus butterflies in Africa, suggests that the radiations of grasses and grazers may be evolutionary linked. We examined mechanisms for this plant–herbivore interaction with the grass‐feeding Bicyclus safitza in South Africa. In a controlled environment, we tested oviposition preference and hatchling performance on local grasses with C₃ or C₄ photosynthetic pathways that grow either in open or shaded habitats. We predicted preference for C₃ plants due to a hypothesized lower processing cost and higher palatability to herbivores. In contrast, we found that females preferred C₄ shade grasses rather than either C₄ grasses from open habitats or C₃ grasses. The oviposition preference broadly followed hatchling performance, although hatchling survival was equally good on C₄ or C₃ shade grasses. This finding was explained by leaf toughness; shade grasses were softer than grasses from open habitats. Field monitoring revealed a preference of adults for shaded habitats, and stable isotope analysis of field‐sampled individuals confirmed their preference for C₄ grasses as host plants. Our findings suggest that plant–herbivore interactions can influence the direction of selection in a grass‐feeding butterfly. Based on this work, we postulate future research to test whether these interactions more generally contribute to radiations in herbivorous insects via expansions into new, unexploited ecological niches.     

Plant sex and hare feeding preferences

Summary To evaluate the general extent to which sex-related differences in palatability occur in boreal dioecious woody plants, males and females of five dioecious woody plant species were presented to free-ranging mountain hares (Lepus timidus) during winter. Hares strongly preferred branches from male plants when feeding on Populus tremula and Salix caprea and weakly preferred male S. pentandra. However, they did not show any sex-related preference when feeding on the other two species studied (Myrica gale and Juniperus communis). Nitrogen concentration and, to some degree, digestibility showed strong relationships with hare food preferences. By contrast, the concentration of phenolics was only weakly related to feeding preference. Phenolics could, nevertheless, still be important if only one or a few specific compounds deter hare feeding. These results indicate that sex-related differences in plant palatability in the boreal forest might be more widespread than previously believed, particularly for species of the family Salicaceae. Thus, herbivores might be responsible for the female-biased sex ratios found in willow populations in northern Scandinavia (e.g. Elmqvist et al. 1988).     

A Comparative Test of Adaptive Explanations for Hypsodonty in Ungulates and Rodents

Hypsodonty has long been recognized as an adaptation for grazing: grazing is suggested to increase tooth wear due to endogenous (e.g., fiber, silica) and/or exogenous (e.g., dust, grit) properties of ingested food. However, it is unknown whether tooth crown height is correlated with the mastication of high fiber or silica in grasses, the ingestion of external abrasives, or both. Furthermore, comparative studies of hypsodonty have not explicitly taken into account phylogenetic biases due to shared ancestry in tooth morphology and/or feeding behavior. This study highlights the relationship between molar crown height and feeding habits in African ungulates and South American rodents when phylogenetic effects are controlled. Among ungulates, high hypsodonty indices are significantly associated with specific plant and foraging height preferences, while habitat and climate show no correlation with tooth crown height. For rodents, grass-eating species are significantly more hypsodont than frugivorous or folivorous species, and arboreal rodents are less hypsodont than terrestrial species. These results as well as those of a posteriori analyses controlling for aspects of the behavioral ecology (e.g., grass-eating, substrate preference) of the sample species confirm the role of both diet and grit in shaping the evolution of cheek tooth crown height in herbivorous mammals.     

Vegetation changes following sheep grazing in abandoned mountain meadows

Abstract. Sheep grazing was investigated as an alternative to traditional management of meadows in the Krkono?e Mts. Until the second World War these meadows were mown in mid‐summer and grazed by cattle for the rest of the season. Subsequent abandonment of the meadows has resulted in decreasing species richness. Degradation phases of the former communities have been replacing the original species‐rich vegetation. Significant changes were apparent six years after the introduction of sheep grazing. In grazed plots the proportion of dominant herbs (Polygonum bistorta and Hypericum maculatum) decreased and grasses (Deschampsia cespitosa, Festuca rubra, Agrostis capillaris, Anthoxanthum alpinum) increased. The increase in grasses was positively correlated with an increase in several herbs. The proportion of some herbs increased despite being selectively grazed (Adenostyles alliariae, Melandrium rubrum, Veratrum lobelianum). Any losses caused by grazing of mature plants were probably compensated by successful seedling establishment. Cessation of grazing resulted in significant changes in vegetation within three years. The cover of nitrophilous tall herbs and grasses (e.g. Rumex alpestris, Holcus mollis, Deschampsia cespitosa, Geranium sylvaticum) increased in the abandoned plots. In the plots grazed for nine years cover of species‐rich mountain meadow species increased (e.g. fine‐leaved grasses, Campanula bohemica, Potentilla aurea, Viola lutea, Silene vulgaris). The main conservation risk is the expansion of a competitive species with low palatability, Deschampsia cespitosa. This species can be suppressed by a combination of grazing and mowing. In order for grazing to be effective, the number of sheep should be proportional to meadow production. This may be difficult to maintain as production is variable and is impossible to predict at the beginning of a growing season. A large part of the biomass may thus remain intact in some years. Negative effects of grazing may be, at least partly, eliminated by a combination of cutting and grazing.     

A fungal endophyte of a palatable grass affects preference of large herbivores

Temperate grasses frequently acquire resistance to herbivores through a symbiosis with epichloid fungi that produces alkaloids of variable deterrent effects. However, in those cases without apparent endophyte negative effects on domestic herbivores, it is not clear whether plant consumption or preference is affected or not. We performed three experiments with 1‐year‐old steers (Bos taurus, Aberdeen Angus) and the annual grass Lolium multiflorum, infected or not by Epichloë occultans to evaluate preference and to identify the underlying tolerance mechanisms. The first experiment evaluated steer preference for L. multiflorum cultivated in plots with three endophyte infection frequencies (low, medium and high), and investigated the role of canopy structure and plant nutritional traits on preference. The second experiment evaluated preference for chopped grass, offered in individual trays with contrasting infection frequencies (low and high), to discard possible effects associated with canopy structure and to focus on nutritional traits. The third experiment was performed with a tray + basket design that separated visual and olfactory stimuli from nutritional traits. High endophyte infection frequencies reduced consistently animal preference, even after short (~10 min) feeding events. However, we did not find significant evidence of plant structural, nutritional, visual or olfactory traits. Our results discarded several potential mechanisms; therefore, the dissuasive effect of fungal endophytes on animal consumption might be related to other mechanisms, including, likely, alkaloids and changes on grass metabolome.     

Experimental demonstration of the antiherbivore effects of silica in grasses: impacts on foliage digestibility and vole growth rates

The impact of plant-based factors on the population dynamics of mammalian herbivores has been the subject of much debate in ecology, but the role of antiherbivore defences in grasses has received relatively little attention. Silica has been proposed as the primary defence in grasses and is thought to lead to increased abrasiveness of foliage so deterring feeding, as well as reducing foliage digestibility and herbivore performance. However, at present there is little direct experimental evidence to support these ideas. In this study, we tested the effects of manipulating silica levels on the abrasiveness of grasses and on the feeding preference and growth performance of field voles, specialist grass-feeding herbivores. Elevated silica levels did increase the abrasiveness of grasses and deterred feeding by voles. We also demonstrated, for the first time, that silica reduced the growth rates of both juvenile and mature female voles by reducing the nitrogen they could absorb from the foliage. Furthermore, we found that vole feeding leads to increased levels of silica in leaves, suggesting a dynamic feedback between grasses and their herbivores. We propose that silica induction due to vole grazing reduces vole performance and hence could contribute to cyclic dynamics in vole populations.     

Convergent grazing responses of different‐sized ungulates to low forage quality in a wet savanna

We studied how grazing intensity by small and mid‐sized ungulate grazers varied with nutritional quality and grass species composition in wet oligotrophic tallgrass savanna of coastal northern Tanzania. Average grazing intensity was low (3–15% by cover), and most grass species were scarcely used by herbivores. Only two grasses, Panicum infestum and Digitaria milanjiana, had nitrogen and phosphorus concentrations that were consistently above the minimum levels (e.g. nitrogen concentrations <7 mg g^?1) required by the three commonest grazers, Bohor reedbuck, waterbuck and wildebeest. The best predictors of grazing intensity were cover of P. infestum (the most abundant grass, with a mean cover of 15%) and canopy height of ungrazed vegetation. Models did not contain separate predictors for nutritional quality, presumably because quality varied mainly at the grass species level and therefore was fully represented by the variable ‘cover of P. infestum’. Given that the three grazers differed greatly in body size and muzzle width (parameters known to influence nutrient requirements and the ability of grazers to feed selectively at the smallest spatial scale), we expected there to be strong resource partitioning that would be detectable in terms of grazing strategies and feeding sites. However, apart from minor differences in canopy height, greenness and diameter of grazed patches (albeit consistent with our expectations), feeding stations of the three grazers were similar and strongly dominated by P. infestum. We conclude that the low quality of herbage in wet oligotrophic savannas restricts foraging choices, which produces a characteristic yet impoverished grazing community that exhibits only limited resource partitioning.     

Ecosystem consequences of selective feeding of an insect herbivore: palatability–decomposability relationship revisited

1. The relationship between leaf palatability and litter decomposability is critical to understanding the effects of selective feeding by herbivores on decomposition processes, and several studies have reported that there is a positive relationship between them. 2. However, palatability is not always positively correlated with decomposability, because of species‐specific feeding adaptation of herbivores to host plants. Moreover, the effects of selective feeding by herbivores on soil decomposition processes should be understood in terms of the inputs of leaf litter and excrement. 3. The present study examined the relationships between leaf palatability and the decomposability of litter and frass, using Lymantria dispar Linnaeus and 15 temperate deciduous tree species. 4. Larvae of L. dispar exhibited a clear feeding preference, and subsequently the excreted frass mass differed among tree species. Litter and frass decomposability also differed among tree species, and frass was more rapidly decomposed than litter. There were no positive or negative correlations between palatability and decomposability of litter and frass. 5. These results indicate that L. dispar larvae may accelerate the decomposition process in temperate deciduous forests through selective feeding on plants with relatively low litter decomposability and the production of frass with higher decomposability than the litter.     

Diet Switching by Mammalian Herbivores in Response to Exotic Grass Invasion

Invasion by exotic grasses is a severe threat to the integrity of grassland ecosystems all over the world. Because grasslands are typically grazed by livestock and wildlife, the invasion is a community process modulated by herbivory. We hypothesized that the invasion of native South American grasslands by Eragrostis plana Nees, an exotic tussock-forming grass from Africa, could be deterred by grazing if grazers switched dietary preferences and included the invasive grass as a large proportion of their diets. Bos taurus (heifers) and Ovis aries (ewes) grazed plots with varying degrees of invasion by E. plana in a replicated manipulative experiment. Animal positions and species grazed were observed every minute in 45-min grazing session. Proportion of bites and steps in and out of E. plana tussocks were measured and used to calculate several indices of selectivity. Both heifers and ewes exhibited increasing probability of grazing E. plana as the proportion of area covered by tussocks increased, but they behaved differently. In agreement with expectations based on the allometry of dietary preferences and morphology, ewes consumed a low proportion of E. plana, except in areas that had more than 90% E. plana cover. Heifers consumed proportionally more E. plana than ewes. Contrary to our hypothesis, herbivores did not exhibit dietary switching towards the invasive grass. Moreover, they exhibited avoidance of the invasive grass and preference for short-statured native species, both of which should tend to enhance invasion. Unless invasive plants are highly palatable to livestock, the effect of grazing to deter the invasion is limited, due to the inherent avoidance of the invasive grass by the main grazers in the ecosystem, particularly sheep.     

Effects of large herbivore grazing on grasshopper behaviour and abundance in a meadow steppe

1. Adaptive phenotypic plasticity has been a major subject in evolutionary ecology, but how a species' behaviour may respond to certain environmental change is still not clear. In grasslands worldwide, large herbivores are increasingly used as a tool for grazing management, and occur to interact with grasshoppers that dominate grassland insect communities. Previous studies have been well-documented about grazing effects on diversity and abundance of grasshoppers. Yet, how grazing may alter grasshopper behaviour, and potential effects on their abundance remains elusive. 2. We conducted a field experiment by manipulating grazing using sheep, cattle, and their mix to examine the behavioural responses and abundance of the grasshoppers (Euchorthippus unicolor) to grazing in a Leymus chinensis-dominated grassland. 3. Results showed that the grasshoppers spent less time on feeding and resting on grasses, but more time on switching and resting on forbs under cattle grazing and mixed grazing with cattle and sheep. In contrast, the grasshoppers spent more time on feeding but less time on switching and resting on forbs under sheep grazing. The behavioural changes were also potentially linked to grasshopper abundance in the context of grazing management. 4. The responses of grasshopper behaviour and abundance to grazing may be largely triggered by altered vegetation and microclimates. Such behavioural flexibility of grasshoppers must be considered when large herbivores are recognised as a management tool for influencing grasshopper abundance, and grazer species should be paid more attention both individually and jointly for better grassland conservation.     

Consumption rates and food preferences of slugs in a calcareous grassland under current and future CO<Subscript>2</Subscript> conditions

This study explored consumption of a generalist herbivore feeding on leaf tissue of various plant species of a calcareous grassland, and tested whether consumption levels and preferences changed when plants were exposed to 5 years of in situ CO₂ enrichment. The first part of this experiment tested whether the consumption patterns of slugs (Deroceras reticulatum) observed in single-species feeding tests were altered when slugs were given a choice of food sources. Overall consumption increased 270% when slugs were given a choice, and they preferred having a choice of food sources more than they preferred having any one food source. Surprisingly, slugs consumed fewer legumes and grasses and more non-leguminous forbs when given a choice. In the second part of this experiment, feeding behaviors of slugs in response to elevated CO₂ were investigated by feeding them leaves of two legumes, one grass, and a non-leguminous forb (Trifolium medium, Lotus corniculatus, Bromus erectus, and Sanguisorba minor, respectively) in two or four species combinations. In the leguminous species mix, the non-leguminous species mix, and the combined mix (legumes and non-legumes), neither overall consumption by herbivores nor species preference was significantly altered by long-term CO₂ enrichment. In the combined species mix, slugs preferred legumes to non-legumes (P=0.012) and exhibited a weak functional group preference shift from non-legumes to legumes (P=0.089) in response to CO₂ enrichment. This is the first time such a shift has been observed, and provides evidence that there may be multiple herbivore responses to rising atmospheric CO₂ concentrations. Numerous single-species feeding tests using insects have shown that consumption by herbivores may increase when herbivores are fed plants grown in enriched CO₂ atmospheres. This study clearly demonstrates the limited applicability of non-choice feeding trials to generalist herbivores in species-rich communities. Received: 19 August 1999 / Accepted: 20 April 2000     

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.     

Seasonal variation in secondary plant compounds in relation to the palatability of Pteridium aquilinum

A correlation was found between the levels of cyanide production and of tannins and the palatability of bracken Pteridium aquilinum L. Kuhn to sheep and deer and to the phytophagous locust Schistocerca gregaria. In the case of the mammals, cyanogenesis, which is genetically polymorphic, appeared to be a complete deterrent to feeding, but tannins probably also play a part. For the insect there were two times during the season when feeding was strongly inhibited and these corresponded to peaks in cyanogenesis in the young developing fronds late May/early June and in tannins which reached the highest level in late August/early September. Changes in total flavonoid levels did not appear to correlate with such feeding preferences.