The induction of spinescence in European holly leaves by browsing ungulates

Evidence is presented which suggests that the spinescence of leaves of European holly, Ilex aquifolium, deters feeding by ungulates and is induced by browsing. Spinescence decreased as leaf size increased; hence, spinescence may be achieved by reducing adult leaf size. Holly shrubs with very spiny leaves were browsed less often than less spiny shrubs. In the absence of browsing ungulates during a one year period, the spinescence of leaves of holly shrubs significantly decreased. Browsed shrubs exhibited reduced annual shoot growth, increased branching, and produced smaller leaves with high spinescence. The regrowth on browsed branches of holly trees was characterized by increased leaf spinescence relative to unbrowsed branches. Hence, the induced response was localized, thereby reducing the ability of browsing ungulates to exert selective pressures on holly trees.     

The input of terrestrial invertebrates from tree canopies to a stream

SUMMARY. The input of terrestrial invertebrates from different tree canopies to a trout stream was determined for a 28-week period from April to October, 1980. Sycamore produced the greatest number of animals, followed by oak and alder. Ash was not significantly different from the controls. Coleoptera, Diptera, Homoptera and Arachnida made up the greatest number of animals caught, with Lepidoptera larvae important beneath oak. The input of biomass (g m^-2 dry wt) was also greatest beneath sycamore (35.80), followed by oak (27.76), alder (20.39), ash (11.15) and control (9.92). The input of biomass was bimodal. The significance of terrestrial invertebrates as food for salmonids is discussed.     

Food–niche relationships within a guild of alpine ungulates including an introduced species

To assess competition for trophic resources within a guild of ungulates, we studied the food niches of sympatric ungulates in an alpine area of the Italian Western Alps and evaluated the extent of their dietary overlap, especially with regard to the presence of the introduced mouflon Ovis musimon . The roe deer Capreolus capreolus had the most diverse diet including dicotyledons, young sprouts of trees and an unusually high percentage of graminoids. The red deer Cervus elaphus and chamois Rupicapra rupricapra mainly consumed graminoids, supplemented with dicotyledonous herbs and woody plants. The mouflon was a grass feeder, with graminoids representing the majority of items ingested in all seasons. Among the native species, we recorded a high overlap of feeding choices between red deer and chamois, which frequently used the same areas and are both intermediate feeders. The mouflon showed considerable dietary overlap with the chamois and red deer in all seasons, while the similarity with roe deer was low. High food–niche overlap can imply competition but only if resources are limited and the species use the same habitats.     

Significance of diet type and diet quality for ecological diversity of African ungulates

1. We test two nutritional hypotheses for the ecological diversity of ungulates, the browser/grazer (diet type) and diet quality models, among free-ranging herbivores in a South African savanna, the Kruger National Park. Tests are based on assessment of relationships between diet type and diet quality with body mass and hypsodonty, two morphological features that have been associated with both elements. 2. We use stable carbon isotope ratios of faeces to reconstruct diet in terms of proportions of C(3) plants (browse) and C(4) plants (grass) consumed by different species in different seasons. These data are combined with proxies for diet quality (per cent nitrogen, neutral detergent fibre, acid detergent fibre, and acid detergent lignin) from faeces to track changes in diet quality. 3. Two statistical approaches are used in model selection, i.e. tests of significant correlations based on linear regression analyses, and an information-theory approach (Akaike's Information Criterion) providing insight into strength of evidence for models. 4. Results of both methods show that, contrary to many predictions, body mass and diet type are not related, but these data confirm predictions that diet quality decreases with increasing body size, especially during the dry season. Hypsodonty, as expected, varies with diet type, increasing with increased grass intake. 5. These findings support both a diet type and diet quality model, implying some degree of exclusivity. We propose that congruence between models may be achieved through addition of diet quality proxies not included here, because hypsodonty is more likely a reflection of the abrasive properties of consumed foods, i.e. related to food quality, rather than food type. This implies that adaptation to diets of varying quality, through changes in body size and dental features, has been the primary mechanism for diversification in ungulates. 6. Our interpretation contrasts with several recent studies advocating diet type as the primary factor, exemplifying that further reconciliation between the two models is needed. We discuss the implications of this study for future approaches to achieve a more cohesive understanding of the evolutionary outcomes of herbivore nutrition.     

Condensed tannins deter feeding by browsing ruminants in a South African savanna

Summary The palatability of 14 species of woody plant was assessed for three species of browsing ruminant, namely kudus, impalas and goats. Results show that palatability was most clearly related to leaf contents of condensed tannins. The effect was a threshold one, with all plants containing more than 5% condensed tannins being rejected as food during the wet season period. In contrast palatability was not influenced by concentrations of protein-precipitating polyphenols, and only weakly related to contents of nitrogen, phosphorus, cations, fibre components and other secondary metabolites. Insect herbivory shows a different pattern. These findings support the hypotheses that (i) condensed tannins function to protect plant cell walls against microbial attack; (ii) hydrolyzable tannins function to inactivate the digestive enzymes of insect herbivores. Large mammalian herbivores are influenced by condensed tannins due to their dependance upon microbial fermentation of plant cell walls for part of their energy needs.     

TrackBeam, a simple tool for estimating potential exposure to the solar beam underneath tree canopies

A simple tool, TrackBeam, for rapid in situ estimates of potential exposure to the solar beam of objects underneath tree canopies is presented, tested and demonstrated to be useful in insect conservation studies. The tool can be used whenever accurate data on sun exposure is of use e.g., in ecological studies of saproxylic insects. TrackBeam draws upon the principles behind the analysis of hemispherical photographs but its use is much less dependent on the weather at the time of sampling. It may be used for detecting canopy openings in directions corresponding to the solar path. Based on this, periods of time with potential beam exposure and/or the proportion of sky that is not obscured by canopy for objects such as dead wood may be estimated. The results of using TrackBeam compares well between operators and with the corresponding results of analysed hemispherical photographs. Results are presented which show that TrackBeam was successfully used to characterise the habitat light requirements of the saproxylic beetle Melanotus castanipes (Coleoptera: Elateridae).     

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.     

Selectivity in large generalist herbivores: feeding patterns of African ungulates in a semi-arid habitat

Feeding habits of free-ranging wildebeest and zebra were monitored in a semi-arid nature reserve, bordering the southwestern part of Kruger National Park, South Africa. The purpose of study was to distinguish and define the feeding niches of two roughage grazers that occur in similar habitat types. The monthly compositions of diets were evaluated by direct observations of feeding bouts over a period of two years when rainfall patterns were average and animal populations were stable. Other analyses evaluated the standing biomass of grass species in the reserve during the wet summer and dry winter seasons. A considerable overlap of grass species composition was found in the diets of wildebeest and zebra. Ordination of bi-monthly records of the diet composition showed greater variations in scores of grasses in zebra diet in comparison to wildebeest. Seasonal patterns were more apparent in the wildebeest diet. Preference ranking of grass species indicated that zebra diet remained constant in winter and summer. Wildebeest diet however, alternated with seasons, showing high preferences during the winter months for grass species which were rejected during summer. The combined assessment of results from three separate statistical methods analysing temporal patterns and preferences in diet composition revealed contradictory trends. The solution, however, relied on the initial assumptions posed. Hence, wildebeest and zebra are essentially generalist feeders which show a limited amount of preference in their choice of diet.     

Vertical zonation of coral reef fishes in the Sudanese Red Sea

The vertical zonation of 38 species of fish inhabiting coral reefs in the Sudanese Red Sea is described. A marked degree of vertical zonation of the fishes is shown to exist. The usefulness of a depth oriented survey method for studying reef fish is discussed in relation to these results.General relationships between vertical zonation and species diversity are noted.     

The role of photoperiodism in the seasonal breeding of tropical African ungulates

Records of ungulate calving peaks, from different latitudes within the African tropics, are reviewed. Converting these peaks to conception dates, and plotting them against latitude, reveals whether the initiation of conception may be connected with a photoperiodic response. The north-south plots of a short-day species should lie in diagonally-opposed quadrants, and those of long-day species in the opposite quadrants. The analysis suggests that a photoperiodic stimulus could be a factor initiating conception in some species, but others lack any apparent relationship.