Sources of variation in plant responses to belowground insect herbivory: a meta-analysis

Growing interest in belowground herbivory and the remarkable diversity of the accumulated information on this topic inspired us to quantitatively explore the variation in the outcomes of individual studies. We conducted a meta-analysis of 85 experimental studies reporting the effects of root-feeding insect herbivores (36 species) on plants (75 species). On average, belowground herbivory led to a 36.3% loss of root biomass, which was accompanied by a reduction in aboveground growth (-16.3%), photosynthesis (-11.7%) and reproduction (-15.5%). The effects of root herbivory on aboveground plant characteristics were significant in agricultural and biological control studies, but not in studies of natural systems. Experiments conducted in controlled environments yielded larger effects on plants than field experiments, and infestation experiments resulted in more severe effects than removal studies employing natural levels of herbivory. Simulated root herbivory led to greater aboveground growth reductions than similar root loss imposed by insect feeding. External root chewers caused stronger detrimental effects than sap feeders or root borers; specialist herbivores imposed milder adverse effects on plants than generalists. Woody plants suffered from root herbivory more than herbaceous plants, although root loss was similar in these two groups. Evergreen woody plants responded to root herbivory more strongly than deciduous woody plants, and grasses suffered from root herbivory more than herbs. Environmental factors such as drought, poor nutrient supply, among-plant competition, and aboveground herbivory increased the adverse effects of root damage on plants in an additive manner. In general, plant tolerance to root herbivores is lower than tolerance to defoliating aboveground herbivores.     

Evolutionary lability of a complex life cycle in the aphid genus <Emphasis Type="Italic">Brachycaudus</Emphasis>

Background  

Most aphid species complete their life cycle on the same set of host-plant species, but some (heteroecious species) alternate between different hosts, migrating from primary (woody) to secondary (herbaceous) host plants. The evolutionary processes behind the evolution of this complex life cycle have often been debated. One widely accepted scenario is that heteroecy evolved from monoecy on woody host plants. Several shifts towards monoecy on herbaceous plants have subsequently occurred and resulted in the radiation of aphids. Host alternation would have persisted in some cases due to developmental constraints preventing aphids from shifting their entire life cycle to herbaceous hosts (which are thought to be more favourable). According to this scenario, if aphids lose their primary host during evolution they should not regain it. The genus Brachycaudus includes species with all the types of life cycle (monoecy on woody plants, heteroecy, monoecy on herbs). We used this genus to test hypotheses concerning the evolution of life cycles in aphids.     

Little strokes fell great oaks: minor but chronic herbivory substantially reduces birch growth

Modern concepts of plant tolerance to herbivory are primarily based on studies of short‐term severe damage, whereas the effects of minor chronic damage to long‐lived woody plants, corresponding to background herbivory (2–15% annual loss of foliar biomass in boreal and temperate forests), remain poorly understood. In our experiment, the annual removal of 2, 4, 8 and 16% of the leaf area from naturally growing mountain birch Betula pubescens subsp. czerepanovii saplings during a seven‐year period resulted in a pronounced reduction of plant vertical growth (–30, –34, –45 and –78%, respectively). Leaf size decreased first (already after one year of the 16% treatment), resulting in the reduction of the total leaf area. This effect was followed by a considerable decrease in the length of long shoots in all treatments. Leaf number on the plant was maintained for a longer time, being reduced by the end of the experiment in 16% treatment only; no changes in specific leaf area or chlorophyll fluorescence were observed in either of the treatments. This pattern may indicate that the plant reallocates resources from the growth of the woody parts to the maintenance of the photosynthetic area, and can be seen as a strategy of tolerance to minor herbivory, whereas compensatory responses typical of severe herbivory (increased photosynthesis rates and shoot regrowth) have not been detected. The predicted 2–5% increase in background herbivory due to climate warming can potentially produce previously unrecognised negative impacts on tree growth. We conclude that in the long term, background herbivory is likely to impose stronger effects on the growth of woody plants than short‐term devastating outbreaks of defoliators, thus contributing more to the development of plant evolutionary adaptations to herbivory than severe but episodic bouts of damage.     

Species richness in plant genera disjunct between temperate eastern Asia and North America

The species richness of 109 amphi-Pacific disjunct genera was examined in eastern Asia and North America. Although the entire flora of eastern Asia contains approximately one-third more species than that of North America, the difference in species richness among disjunct taxa is less. When woody and herbaceous genera are considered separately, the former exhibit a strong diversity bias favouring eastern Asia whereas there is no significant difference in diversity between continents among herbaceous genera. This result is not due to habitat differences between woody and herbaceous genera, because the disjunct herbs inhabit primarily moist forests and woodlands. This result is also not related to relative phylogenetic advancement, even though older major lineages of plants tend to have a predominance of woody taxa. Woody genera are distributed in lower latitudes than herbaceous genera on both continents, and both woody and herbaceous genera are distributed in lower latitudes in eastern Asia than in North America. The North American temperate flora is primarily a relict of a flora form 7 more widespread throughout the Northern Hemisphere. Contemporary patterns of diversity suggest that the effects of climate changes in the late Tertiary were less severe in eastern Asia and promoted diversification, but were more severe in North America and may have caused widespread extinction. The difference in the effect of climate change on diversity in herbaceous and woody lineages reflects the different ecological relationships of species having these contrasting life forms. Clearly, the contemporary floras of eastern Asia and North America bear the imprint of history and emphasize the important interface between ecological relationships and evolutionary responses.     

Effects of mycorrhizal symbiosis on tallgrass prairie plant–herbivore interactions

Complex relationships occur among plants, mycorrhizal fungi, and herbivores. By altering plant nutrient status, mycorrhizas may alter herbivory or plant tolerance to herbivory via compensatory regrowth. We examined these interactions by assessing grasshopper preference and plant growth and fungal colonization responses to herbivory under mycorrhizal and non‐mycorrhizal conditions within tallgrass prairie microcosms. Mycorrhizal symbiosis increased plant regrowth following defoliation, and some strongly mycotrophic plant species showed overcompensation in response to herbivory when they were mycorrhizal. Although grasshoppers spent more time on mycorrhizal plants, herbivory intensity did not differ between mycorrhizal and non‐mycorrhizal plants. Aboveground herbivory by grasshoppers significantly increased mycorrhizal fungal colonization of plant roots. Thus mycorrhizas may greatly benefit plants subjected to herbivory by stimulating compensatory growth, and herbivores, in turn, may increase the development of the symbiosis. Our results also indicate strong interspecific differences among tallgrass prairie plant species in their responses to the interaction of aboveground herbivores and mycorrhizal symbionts.     

Climbing strategy in herbs does not necessarily lead to lower investments into stem biomass

Herbaceous climbers (vines) represent a growth strategy in which the stem lacks most of its supporting function. This has led to the hypothesis that herbaceous climbers are structural parasites that invest less into stems than self-supporting plants. So far, the support for this idea has been ambiguous, as woody and herbaceous plants have been discussed jointly and evidence is often based on young plants in pot experiments. We collected in wild fully grown temperate herbaceous climbers and self-supporting herbs to examine the idea. We made a phylogenetically informed comparison of biomass allocation into stems and leaves of 16 climber species and 74 self-supporting herbs. Furthermore, we compared our results with those published for woody climbers to gain insight into different biomass allocation between herbaceous and woody growth forms. We found that herbaceous climbers and self-supporting herbs do not differ in their proportion of stem biomass to leaf biomass. Herbaceous climbers reach much higher in the canopy thanks to their climbing habit and in average more than seven times longer stems, but contrary to the expectation and unlike their woody counterparts, they do not save on investment into the stem. Herbaceous climbers and self-supporting herbs represent a study system which provides insight into biomass scaling with versus without supporting function where both stems as well as leaves are seasonal.

     

Foliar herbivory and its effects on plant growth in native and exotic species in the Patagonian steppe

Studies of herbivory and its consequences on the growth of native and exotic plants could help elucidate some processes involved in plant invasions. Introduced species are likely to experience reduced herbivory in their new range due to the absence of specialist enemies and, thus, may have higher benefits if they reduce the investment in resistance and increase their compensatory capacity. In order to evaluate the role of herbivory in disturbed areas within the Patagonian steppe, we quantified and compared the leaf levels of herbivory of four native and five exotic species and recorded the associated insect fauna. We also performed greenhouse experiments in which we simulated herbivory in order to evaluate the compensatory capacity of native and exotic species under different herbivory levels that resembled naturally occurring damage. Natural herbivory levels in the field were similar between the studied exotic and native plants. Field observations confirmed that they both shared some herbivore insects, most of which are generalists. In the greenhouse experiments, both exotic and native plants fully compensated for herbivory. Our results suggest that the studied exotic plants are not released from herbivory in the Patagonian steppe but are able to fully compensate for it. The capacity to recover from herbivory coupled with other potential adaptations, such as a better performance under disturbance and greater competitive ability than that of the native species, may represent some of the mechanisms responsible for the success of plant invasion in the Patagonian steppe.     

Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management

Plant species richness and range-size rarity in Africa south of the Sahara is concentrated in centres of plant diversity and endemism. Distribution patterns of plants mapped in the Distributiones Plantarum Africanum series and selected taxonomic monographs are analysed using the computer programme WORLDMAP. The plants are divided into four groups: herbaceous geophytes, mesophytic herbs, light-demanding shrubs and woody genera. Each group has peaks of species richness and range-size rarity at locations different to the other groups. Herbaceous geophytes and mesophytic herbs have their peaks of species richness and range-size rarity in the same location, the western Cape for geophytes and the Crystal Mountain for mesophytic herbs, whereas light-demanding shrubs and woody genera have peaks in different places. The results are discussed in relation to possible factors determining species richness and endemism and their likely conservation significance.     

Loss of specificity: native but not invasive populations of Triadica sebifera vary in tolerance to different herbivores

During introduction, invasive plants can be released from specialist herbivores, but may retain generalist herbivores and encounter novel enemies. For fast-growing invasive plants, tolerance of herbivory via compensatory regrowth may be an important defense against generalist herbivory, but it is unclear whether tolerance responses are specifically induced by different herbivores and whether specificity differs among native and invasive plant populations. We conducted a greenhouse experiment to examine the variation among native and invasive populations of Chinese tallow tree, Triadica sebifera, in their specificity of tolerance responses to herbivores by exposing plants to herbivory from either one of two generalist caterpillars occurring in the introduced range of Triadica. Simultaneously, we measured the specificity of another defensive trait, extrafloral nectar (EFN) production, to detect potential tradeoffs between resistance and tolerance of herbivores. Invasive populations had higher aboveground biomass tolerance than native populations, and responded non-specifically to either herbivore, while native populations had significantly different and specific aboveground biomass responses to the two herbivores. Both caterpillar species similarly induced EFN in native and invasive populations. Plant tolerance and EFN were positively correlated or had no relationship and biomass in control and herbivore-damaged plants was positively correlated, suggesting little costs of tolerance. Relationships among these vegetative traits depended on herbivore type, suggesting that some defense traits may have positive associations with growth-related processes that are differently induced by herbivores. Importantly, loss of specificity in invasive populations indicates subtle evolutionary changes in defenses in invasive plants that may relate to and enhance their invasive success.     

Warming the tundra: reciprocal responses of invertebrate herbivores and plants

Rapid warming in northern ecosystems is simultaneously influencing plants, herbivores and the interactions among them. Recent studies suggest that herbivory could buffer plant responses to environmental change, but this has only been shown for vertebrate herbivores so far. The role of invertebrate herbivory in tundra ecosystems is often overlooked, but can be relevant in determining the structure and dynamics of tundra plant communities and may also affect how plants respond to warming. Invertebrate herbivores are also likely to respond more rapidly to warming than vertebrates because their behaviour and life cycles strongly depend on temperature. We investigated the effects of current season warming on Arctic moth caterpillars, their herbivory rates, and the subsequent responses of two common tundra plants, Salix arctica and Dryas octopetala. We manipulated both herbivore presence and temperature in a full‐factorial field experiment at two elevations, using enclosures and passive warming chambers. Changes in temperature achieved through elevation and/or experimental warming directly affected caterpillars, herbivory and the responses of plants. Caterpillars performed worse (higher respiration rates and lower growth rates) in warmer, lower elevation plots and shifted their diets towards more nutritious foods, such that the relative intensity of herbivory changed for the two studied plants. Within‐season responses of both forage plant species were weak, but invertebrate herbivores affected the responses of plants to elevation or experimental warming. Our results suggest that increased temperatures can reduce the performance of cold‐adapted invertebrate herbivores, with potential consequences to the longer term responses of tundra plants to warming due to changes in herbivory rates and selective foraging.     

Differential effects of goat browsing on herbaceous plant community in a two-phase mosaic

The impact of herbivores on herbaceous plant communities is usually attributed to direct consumption of plants. We hypothesized that goats affect herbaceous plants both directly (consumption by foraging) and indirectly, by changing environmental conditions through modification of woody plant structure. We assessed the effects of goats browsing on environmental conditions, landscape structure, and herbaceous plants to link the direct and indirect effects of goats on herbaceous communities. Our model system was the Mediterranean woodland in Mt. Carmel, Israel. This is a two-phase mosaic landscape, composed of herbaceous (open) and woody patches. We delineated 10 plots of 1000 m², goats were introduced to five plots and five plots remained without goats. We monitored plant species richness and composition in two adjacent patch types (woody and open) in each plot. For each patch type, in all plots, we collected data on environmental conditions. We analyzed landscape structure using landscape metrics derived from a high-resolution vegetation map. We found that goats modified the structure of woody plants and hence the landscape mosaic. This alteration was associated with changes in environmental conditions, with more light penetration and higher temperatures. The impact of goats on the herbaceous plant community depended on patch type. In open patches, goats affected the herbaceous community mostly by direct consumption, whereas in woody patches they affected the herbaceous community mainly by modification of abiotic conditions. Our results stress the importance of considering landscape and patch structure in analyzing the effect of herbivory on plant communities.     

Vegetation development on deposit soils starting at different seasons

Permanent plots were created in different seasons (autumn and spring) and filled with two substrates: nutrient-rich topsoil and nutrient-poor ruderal soil (n = 5 for each treatment). My objectives were to assess the influence of starting season on initial species composition, whether differences at the start cause divergent or convergent pathways of succession and which mechanisms are operating during vegetation development. Mean species richness (number of species per plot) and mean total cover of herb layer differed significantly between substrates and changed significantly during 10 year succession, but there were no significant differences with respect to starting season. However, seasonal as well as substrate effects were evident for particular dominant species and for the pattern of successional sequences. When succession on topsoil plots started in spring, first summer annuals dominated, then monocarpic and polycarpic perennial herbs, then herbaceous perennials together with woody perennials, and at the end of the decade woody perennials. When succession started in autumn, polycarpic perennial herbs dominated from the beginning, and then were replaced by woody perennials in the second half of the decade. On ruderal soil, there was a less rapid but continuous increase of polycarpic perennial herbs and woody species, both on spring and on autumn plots, whereas short-lived plants were more abundant in the first years and then decreased. Species turnover was very high from the first to the second year for all treatments (except topsoil plots starting in autumn), but slowed down during succession. Priority effects due to starting season caused high dissimilarity at the start on the nutrient-rich substrate, but convergent succession towards the end of the first decade. The main mechanisms during early succession on the nutrient-rich topsoil were tolerance based on different life-history traits and inhibition due to reduced light availability. There was no evidence for obligate facilitation. However, an indirect facilitative effect by annuals, which slowed the development of herbaceous perennials down, and thus facilitated growth of woody species, could be seen on topsoil when succession started in spring.     

Plant trait responses to grazing – a global synthesis

Herbivory by domestic and wild ungulates is a major driver of global vegetation dynamics. However, grazing is not considered in dynamic global vegetation models, or more generally in studies of the effects of environmental change on ecosystems at regional to global scale. An obstacle to this is a lack of empirical tests of several hypotheses linking plant traits with grazing. We, therefore, set out to test whether some widely recognized trait responses to grazing are consistent at the global level. We conducted a meta‐analysis of plant trait responses to grazing, based on 197 studies from all major regions of the world, and using six major conceptual models of trait response to grazing as a framework. Data were available for seven plant traits: life history, canopy height, habit, architecture, growth form (forb, graminoid, herbaceous legume, woody), palatability, and geographic origin. Covariates were precipitation and evolutionary history of herbivory. Overall, grazing favoured annual over perennial plants, short plants over tall plants, prostrate over erect plants, and stoloniferous and rosette architecture over tussock architecture. There was no consistent effect of grazing on growth form. Some response patterns were modified by particular combinations of precipitation and history of herbivory. Climatic and historical contexts are therefore essential for understanding plant trait responses to grazing. Our study identifies some key traits to be incorporated into plant functional classifications for the explicit consideration of grazing into global vegetation models used in global change research. Importantly, our results suggest that plant functional type classifications and response rules need to be specific to regions with different climate and herbivory history.     

Beaver herbivory on aquatic plants

Herbivores have strong impacts on marine and terrestrial plant communities, but their impact is less well studied in benthic freshwater systems. For example, North American beavers (Castor canadensis) eat both woody and non-woody plants and focus almost exclusively on the latter in summer months, yet their impacts on non-woody plants are generally attributed to ecosystem engineering rather than herbivory. Here, we excluded beavers from areas of two beaver wetlands for over 2 years and demonstrated that beaver herbivory reduced aquatic plant biomass by 60%, plant litter by 75%, and dramatically shifted plant species composition. The perennial forb lizard’s tail (Saururus cernuus) comprised less than 5% of plant biomass in areas open to beaver grazing but greater than 50% of plant biomass in beaver exclusions. This shift was likely due to direct herbivory, as beavers preferentially consumed lizard’s tail over other plants in a field feeding assay. Beaver herbivory also reduced the abundance of the invasive aquatic plant Myriophyllum aquaticum by nearly 90%, consistent with recent evidence that native generalist herbivores provide biotic resistance against exotic plant invasions. Beaver herbivory also had indirect effects on plant interactions in this community. The palatable plant lizard’s tail was 3 times more frequent and 10 times more abundant inside woolgrass (Scirpus cyperinus) tussocks than in spatially paired locations lacking tussocks. When the protective foliage of the woolgrass was removed without exclusion cages, beavers consumed nearly half of the lizard’s tail leaves within 2 weeks. In contrast, leaf abundance increased by 73–93% in the treatments retaining woolgrass or protected by a cage. Thus, woolgrass tussocks were as effective as cages at excluding beaver foraging and provided lizard’s tail plants an associational refuge from beaver herbivory. These results suggest that beaver herbivory has strong direct and indirect impacts on populations and communities of herbaceous aquatic plants and extends the consequences of beaver activities beyond ecosystem engineering.     

The evolution of plant life span: facts and hypotheses

There are two different views on the evolution of life forms in Cormophyta: from woody plants to herbaceous ones or in opposite direction - from herbs to trees. In accordance with these views it is supposed that life span in plants changed in the course of evolution from many years (perennials) to few years (annuals, biennials), or went in reverse - from few years to many years. The author discusses the problems of senescence and longevity in Cormophyta in the context of various hypotheses of ageing (programmed death theory, mutation accumulation, antagonistic pleiotropy, disposable soma, genes of ageing, genes of longevity). Special attention is given to bio-morphological aspects of longevity and cases of non-ageing plants ("negative senescence", "potential immortality"). It is proposed to distinguish seven models of simple ontogenesis in Cormophyta that can exemplify the diversity of mechanisms of ageing and longevity. The evolution of life span in plants is considered as an indirect result of natural selection of other characteristics of organisms or as a consequence of fixation of modifications (episelectional evolution). It seems that short life span could emerge several times during evolution of one group of plants, thus favoring its adaptive radiation.     

Trade-offs between flowering time,plant height,and seed size within and across 11 communities of a QingHai-Tibetan flora

Flowering timing is of fundamental biological importance for its tight association with pre-flower growth states and subsequent reproduction success. Here, we selected plant height and seed size to represent plant growth and reproduction states, respectively, and analyzed their associations with flowering time in 11 communities together representing a QingHai-Tibetan flora. Trait associations were examined using Pearson correlation analyses (TIPs) and phylogenetically independent contrasts (PICs) within individual communities and meta-analyses across all communities. The results of TIPs-based and PICs-based analyses were generally congruent, although fewer contrasts were significant with PICs, probably because of low statistical power. Overall, flowering time was negatively correlated with seed size and plant height (i.e., plants with larger seeds and stature started flowering earlier) in various woody communities, but correlations were neutral or positive in herbaceous communities. The seed size–flowering time relationship was negative for woody and herbaceous perennials but not for annual herbs in most communities. The relationship between plant height and flowering time was negative for woody but positive for herbaceous plants. Moreover, the lack of difference in time–size relationships between anemophilous and entomophilous plants suggests that pollination type may only be a secondary force in controlling flowering phenology. Our studies demonstrate that environmental conditions, community structure, and plant life history strategies may affect community flowering time singly or in combination.     

Tolerating herbivory: does the plant care if the herbivore has a backbone?

Differences in size-related ecology and behaviour between vertebrate and invertebrate herbivores lead to differences in the rates, tissue specificity, and spatial distribution of their damage, as well as in their indirect effects. As a result, many features of tolerance to herbivory by these groups also may differ. Tolerating vertebrate herbivory may demand the ability to tolerate sporadic non-specific impacts; this may be achieved by broad responses promoting regrowth and resource acquisition. In contrast, the diversity of different types of invertebrate damage seems likely to demand a correspondingly great variety of responses. These conclusions suggest that tolerance to invertebrates may involve a broader set of responses than tolerance to vertebrates; conversely, the greater specificity of these responses may make it more difficult for arthropod-tolerant plants to achieve cross-tolerance to other types of damage. This revised version was published online in July 2006 with corrections to the Cover Date.     

Exotic herbivores and fire energy drive standing herbaceous biomass but do not alter compositional patterns in a semiarid savanna ecosystem

Questions

Fire regime alterations are pushing open ecosystems worldwide past tipping points where alternative steady states characterized by woody dominance prevail. This reduces the frequency and intensity of surface fires, further limiting their effectiveness for controlling cover of woody plants. In addition, grazing pressure (exotic or native grazers) can reinforce woody encroachment by potentially reducing fine-fuel loads. We investigated the effects of different fire energies on the herbaceous plant community, together with mammalian wildlife herbivory (exotic and native combined) exclusion, to inform best management practices.

Location

Texas semi-arid savanna, southern Great Plains, USA.

Methods

We conducted an experiment in which we manipulated fire intensity and herbivore access to herbaceous biomass in a split-plot design. We altered fire energy via fuel addition rather than applying fire under different environmental conditions to control for differences in standing biomass and composition attributable to differential plant physiological status and fire season.

Results

High-energy fire did not reduce herbaceous biomass or alter plant community composition, although it did increase among-plot variability in composition and forb biomass relative to low-energy fire and non-burned controls. Grazing pressure from native and non-native mammalian herbivores reduced above-ground herbaceous biomass regardless of fire treatments, but did not alter community composition.

Conclusions

Managers seeking to apply high-intensity prescribed fire to reduce woody encroachment will not negatively impact herbaceous plant productivity or alter community composition. However, they should be cognizant that repeated fires necessary for greatly reducing woody plants in heavily invaded areas might be difficult to accomplish due to fine-fuel reduction from wild herbivores. High fencing to restrict access by wildlife herbivores or culling might be necessary to build fuels sufficient to conduct high-intensity burns for woody-plant reduction.     

Flowering phenology and compensation for herbivory in<Emphasis Type="Italic"> Ipomopsis aggregata</Emphasis>

The mechanisms and circumstances that affect a plant's ability to tolerate herbivory are subjects of ongoing interest and investigation. Phenological differences, and the timing of flowering with respect to pollinators and pre-dispersal seed predators, may provide one mechanism underlying variable responses of plants to herbivore damage. The subalpine wildflower, Ipomopsis aggregata, grows across a wide range of elevations and, because phenology varies with elevation, phenological delays associated with elevation may affect the ability of I. aggregata to compensate for or tolerate browsing. Thus, we examined the response of I. aggregata to herbivory across an elevation gradient and addressed the interactions among phenological delays imposed by damage, elevation, pre-dispersal seed predation and pollination, on I. aggregata's compensatory response. Among high and low elevation populations in areas near the Rocky Mountain Biological Laboratory (RMBL) in Gothic, Colorado, we compared the responses of naturally browsed, artificially browsed (clipped), and unbrowsed (control) plants of I. aggregata. We compared responses in the date of initiation of flowering, timing of peak bloom, floral display, nectar production and sugar concentration, oviposition and fruit destruction by the pre-dispersal seed predator Hylemya sp. (Anthomyiidae), fruit production, and aboveground biomass production. Clipping had the greatest effect on reproductive success and clipped plants at high elevation exhibited the lowest tolerance for herbivory. The effects of browsing appear to be mediated by flowering phenology, and both browsing and elevation delayed flowering phenology. Time needed for regrowth delays flowering, and thus affects the overlap with seed predators and pollinators. As a result of delayed flowering, naturally browsed and clipped plants incurred lower rates of seed predation. In the absence of seed predation, plants would exhibit a lower tolerance to herbivory since naturally and artificially browsed plants had fewer fruits destroyed by Hylemya larvae. We provide additional evidence that, for populations near the RMBL, clipping and natural browsing do not have the same effect on I. aggregata plants. This may be due to the selection of larger plants by herbivores. Although under some conditions plants may tolerate browsing, in areas where the growing season is short a phenological delay imposed by damage is likely to significantly reduce plant fitness. Identifying the mechanisms that allow plants to tolerate herbivore damage will help to develop a general framework for understanding the role of tolerance in plant population and community dynamics, as well as plant-herbivore interactions.     

Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.