Spinescence and Total Phenolic Content Do Not Influence Diet Preference of a Critically Endangered Megaherbivore, but the Mix of Compounds Does.

In contrast to understanding spinescence in savanna woody species, little is known about the functions of plant secondary metabolites (PSM). Negative effects of PSMs on individual animal performance potentially translate into negative effects on herbivore population growth. Hence, understanding PSM functions is important for the conservation of savanna megafauna. We tested the view that black rhinoceros (Diceros bicornis) diet preference is not affected by spinescence or total phenolic abundance. We hypothesized that the composition of phenolic mixtures, however, would affect preference. Furthermore, we tested our data from 71 woody species for a trade-off between structural and chemical defenses. Spinescence type, and spinescence generally, did not deter black rhino feeding. Using eco-metabolomic data, we found that total abundance of phenolics did not affect preference, but mixture composition did and that the probability of spinescence trading off against phenolics depended on the mixture. We note that our study was restricted to black rhino and that diet preferences of other mammal herbivores might be influenced by subtle differences in phenolic mixtures. However, our results did support a previous, more detailed study of phenolic profiles of six species showing the same patterns in relation to preference generalised across mammal herbivore species in savannas. Our results represent substantial advancement in the understanding of the roles of PSMs, especially flavonoid compounds, in the functioning of savanna ecosystems, and highlight the need to dig deeper into broad groups of traits such as spinescence or total phenolics to improve understanding of woody plant defenses in savannas.

Does the Growth Differentiation Balance Hypothesis Explain Allocation to Secondary Metabolites in Combretum apiculatum , an African Savanna Woody Species?

The growth differentiation balance hypothesis (GDBH) provides a framework that predicts a trade-off between costs of secondary metabolites (SMs) relative to the demand for photosynthate by growth. However, this hypothesis was developed using empirical evidence from plant species in northern boreal and temperate systems, leaving its applicability to species under different abiotic and biotic conditions questionable and generalizations problematic. The objective of this study was to investigate whether the GDBH explains allocation to SMs in the deciduous African savanna woody species C. apiculatum along a 6-point N gradient. The cornerstone prediction of the GDBH, i.e., the parabolic response in SMs along the N gradient, was not observed, with secondary metabolism showing compound-specific responses. Quercetin, myricetin, and kaempferol glycoside concentrations, all produced via the same pathway, responded differently across the N gradient. Flavonol glycoside, cinnamic acid, and quercetin glycoside concentrations decreased as N increased, which provides partial support for the carbon nutrient balance hypothesis. Simulated herbivory had no effect on photosynthesis, decreased foliar N and consequently increased C:N ratio, but did not induce an increase in SMs, with condensed tannins and flavonol glycosides being unaffected. Defoliated plants at low N concentration compensated for lost biomass, which suggests a tolerance response, but as predicted by the limiting resource model, plants at higher N concentration were evidently C limited and thus unable to compensate. Our results show that the GDBH does not explain allocation to SMs in C. apiculatum, and suggest that mechanistic explanations of plant allocation should consider the integrative defensive effect of changed SMs.

Effects of season, browse species and polyethylene glycol addition on gas production kinetics of forages in the subhumid subtropical savannah, South Africa.

BACKGROUND: This study was conducted to investigate the effects of season, species and polyethylene glycol addition on gas production (GP) and GP kinetic parameters by in vitro incubation (72 h) of five plant species from the subhumid subtropical savannah, South Africa. Plant species used were Acacia natalitia, Acacia nilotica, Dichrostachys cinerea, Scutia myrtina and Chromolaena odorata, leaves of which were harvested during the dry (June/July), early wet (November/December) and late wet (February/March) seasons. An automated in vitro gas production technique was used in two experiments carried out with nine replicates. The first experiment was to test the effect of season and species, while the second experiment tested the effect of tannins using polyethylene glycol 4000 (PEG). The PEG treatment was applied to samples in the early wet and late wet seasons. RESULTS: There were wide variations among seasons and species in crude protein (CP), neutral detergent fibre (NDF), acid detergent fibre (ADF) and condensed tannin (CT). Season and species affected the maximum GP and GP kinetic parameters. During the three seasons, C. odorata had the highest CP (186-226 g kg(-1) dry matter (DM)) and GP (87-104 mL g(-1) DM) and S. myrtina had the lowest CP (105-129 g kg(-1) DM), while A. nilotica, A. natalitia, D. cinerea and S. myrtina had similar and low GP (23-50 mL g(-1) DM). The maximum GP, its degradation rate and GP from the soluble fraction were positively correlated with CP both without and with PEG. With PEG, GP from the soluble fraction was negatively correlated with NDF, ADL and CT; without PEG, it was negatively correlated with CT. CONCLUSION: Both season and species affected the GP parameters. The addition of PEG emphasises that the inhibitory effect of tannins on rumen microbes was greater for all but C. odorata, confirming that these browse species can be used as feed supplements.

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.

Severe drought limits trees in a semi-arid savanna.

Increasingly frequent and severe droughts under climate change are expected to have major impacts on vegetation worldwide. However, research to date has focused on tree vulnerability to drought in forests. Less is known about trees and drought in savannas, where a sparse tree layer coexists with grass. These tree-grass interactions (often mediated by fire and herbivory) shape savanna tree ecology, and confound predictions of how strongly drought might affect trees. On the one hand, drought is physiologically stressful, which could harm trees and be exacerbated by herbivore impacts; on the other hand, trees adapted to semiarid savannas might be relatively drought tolerant, and the considerable impacts of drought on grass could even benefit trees via reduced grass competition and fire risk, especially in the year following a drought. Here, we sought to understand the net effects of severe drought on the savanna tree layer, and how fire and herbivory mediate these effects. We monitored tree growth, mortality, and community structure for 2 yr within existing long-term fire and herbivory experiments across a drought-severity contrast, following a major drought in Kruger National Park, South Africa. Overall, severe drought was a major stressor for trees. Tree mortality rates in most species increased by an order of magnitude in the year following drought, and slower growth rates for some persisted for 2 yr. At the community level, this translated into substantial decreases in tree densities. Herbivory and fire did little either to mitigate or exacerbate drought effects on trees, and overall, drought swamped effects of herbivory and fire that have otherwise been observed. However, species differed in their responses to drought, with some dominant encroaching species especially vulnerable. We suggest that increasing drought frequency and severity could drastically alter savanna vegetation by repeatedly killing off trees.

Seasonal regulation of condensed tannin consumption by free-ranging goats in a semi-arid savanna.

Although condensed tannins (CTs) are known to reduce forage intake by mammalian herbivores in controlled experiments, few studies have tested these effects in the field. Thus the role of CTs on foraging ecology of free-ranging herbivores is inadequately understood. To investigate the effects of CTs under natural savanna conditions, we pre-dosed groups of goats with polyethylene glycol (PEG, a CT-neutralising chemical), CT powder or water before observing their foraging behaviour. While accounting for the effects of season and time of the day, we tested the hypothesis that herbivores forage in ways that reduce the intake rate (g DM per minute) of CTs. We expected pre-dosing goats with CTs to reduce CT intake rates by (1) consuming diets low in CTs, (2) reducing bite rates, (3) increasing the number of foraging bouts, or (4) reducing the length of foraging bouts. Lastly, (5) expected CT to have no influence the number of dietary forage species. In both wet and dry seasons, pre-dosing goats with CTs resulted in lower CT consumption rates compared to PEG goats which seemed relieved from the stress associated with CT consumption. During dry season, the number of dietary forage species was similar across treatments, although goats that were dosed with PEG significantly increased this number in the wet season. Dosing goats with PEG increased the number and length of browsing bouts compared to goats from the other treatments. Pre-loading goats with PEG also tended to increase bite rates on browse forages, which contributed to increased consumption rates of CTs. Based on the behavioural adjustments made by goats in this study and within the constraints imposed by chemical complexity in savanna systems, we concluded that herbivores under natural conditions foraged in ways that minimised CTs consumption. More research should further elucidate the mechanism through which CTs regulated feeding behaviour.

Differential phenolic profiles in six African savanna woody species in relation to antiherbivore defense.

Low molecular weight phenolics are suggested to have a role in mediating diet selection in mammalian herbivores. However, very little is known about low molecular weight phenolic profiles of African savanna woody species. We determined low molecular weight phenolic profiles of six woody species with different life history, morphological and functional traits. We investigated interspecific phytochemical variation between species and found that: (1) related Acacia species were chemically dissimilar; (2) similarity percentage analysis revealed that Acacia grandicornuta was most dissimilar from other species and that the evergreen and unpalatable Euclea divinorum had a qualitatively similar chemical profile to the deciduous and palatable Acacia exuvialis and Combretum apiculatum; (3) C. apiculatum had the highest chemical diversity; (4) relative to spineless plants, spinescent plants contained significantly less HPLC phenolics and condensed tannins; and (5) the major quantitative difference between the evergreen and unpalatable E. divinorum and other species was its high myricitrin concentration.