Low nutritive quality as a plant defence: Effects of herbivore-mediated interactions

Summary A plant may lower its nutritive quality, for herbivores, by using secondary compounds, morphological characters and/or having a lowered nutrient content. If such traits decrease the amount of resources lost through herbivory, then they act as antiherbivore defences. However, if herbivores compensate for the lowered nutrient availability, by increasing their intake rates or by prolonging their feeding periods, then this may render the defence useless. I analyse the conditions for evolution of this type of plant defences in a game theoretical model. The predictions of the model depend on the amount of compensatory feeding performed by the herbivores and on the herbivores' mobility in relation to the spatial structure of the plant population. When herbivores cannot compensate for a lowered nutritive quality, the defence can evolve irrespective of the type of herbivore. When herbivores can compensate for such defences, the outcome depends on how the herbivores compensate. In situations where herbivores compensate only on defended plants, which could correspond to immobile herbivores, this type of defence can evolve only if the level of compensation is lower than a certain critical value. When herbivores compensate more on defended than on undefended plants, e.g. because of low mobility, the outcome depends on the level of compensation performed on defended plants. If this level of compensation is high, then the model predicts a stable coexistence of defended and undefended plants and, if it is low, then the populations can consist of only defended plants. When herbivores compensate more on undefended plants than on defended ones, e.g. highly mobile herbivores, the result is populations consisting of either only defended plants, or only undefended plants. Consequently, the fact that herbivores may compensate for lowered nutrient quality does not, as such, nullify the notion of low nutrient quality as a plant defence. However, compensatory feeding may restrict the conditions for the evolution of such defences.     

Spatial Coupling of Plant and Herbivore Dynamics: The Contribution of Herbivore Dispersal to Transient and Persistent "Waves" of Damage

Spatial variations in the abundance of insect herbivores and in herbivore damage are both striking an commonplace. The standard explanations for heterogeneity in herbivore attack emphasize spatial variations in plant genetype, soils, or physical environment. Here I examine an alternative hypothesis-that heterogeneity arises in plant-herbivore systems, even in homogeneous environments, as a result of the direct coupling of herbivore movement to herbivore density and plant quality. Using a mathematical model for plant quality and herbivore growth and dispersal, I demonstrate how spatial instabilities about homogeneous steady state values result in both transient and stationary waves of damage to the plant. Key herbivore movement behaviors include the tendendy for herbivores to aggregate over a range of spatial scales for increased feeding efficiency and the tendency for herbivores to move up gradients in plant quality (herbivory-taxis). My approach translates the biased "random walk" behavior of individual herbivores into a continuum partial differential equation model. Analytical and numerical methods are used to demonstrate the nature of the spatio-temporal variations in plant quality and herbivore density.     

GENOTYPIC VARIATION IN LEAF DAMAGE IN PIPER ARIEIANUM (PIPERACEAE) BY A MULTISPECIES ASSEMBLAGE OF HERBIVORES

The shrub Piper arieianum (Piperaceae) has a diverse herbivore fauna (95 species total) in Costa Rican rain forest. The effect of plant genotype on leaf damage by individual herbivore species and total leaf area removed was studied in P. arieianum through a cloning experiment. Damage patterns were measured over 3.5 years for two plots, four genotypes per plot, in the understory of lowland rain forest. In both plots, there were significant differences among genotypes in total leaf area missing throughout the study period. Rankings of genotypes based on overall damage remained constant over time in plot 1 but changed in plot 2. Certain individual herbivore species caused significantly higher damage in some genotypes than in others; the change in genotype rankings in plot 2 was associated with increased damage to particular genotypes by specific herbivore groups. The genotype most heavily damaged by a given insect species varied depending on the herbivore species; thus, resistance to one herbivore species did not necessarily confer resistance against all species. Those herbivore species causing the greatest proportion of damage for a given plant changed over time. Because total damage resulted from the summation of losses to individual herbivore species, whether an individual plant lost more leaf area than its neighbors depended on the relative abundance of the herbivore species at any one time. Finally, for a portion of the study period in each plot, more heavily damaged clones grew less than lesser damaged clones. Together with previous reports that naturally growing plants differ significantly in damage and that these differences are sufficient to cause fitness differences, the results presented here suggest that the herbivores of P. arieianum represent a selective force for changes in resistance but that this selective force changes both in intensity and quality over time.     

Density dependence in insect performance within individual plants: induced resistance to Spodoptera exigua in tomato

Net intraspecific density dependence experienced by insect herbivores at the scale of single plants can be a function both of induced resistance in the plant and other interactions among individual herbivores. Theory suggests that non‐linearity in the form of this density dependence can influence the effects of plants on herbivore population dynamics. This study examined both net density dependence at the scale of single plants, and changes in plant quality with herbivore density for Spodoptera exigua caterpillars on tomato plants. One experiment measured the growth of caterpillars moving freely about the plant at different densities, the distribution of damage by these caterpillars, and the quality of the plant as food for caterpillars (growth of caterpillars on undamaged leaf tissue excised from the plant). A second experiment measured plant quality for plants with different amounts of damage by caterpillars confined to particular leaves in mesh bags. Growth of S. exigua caterpillars was found to be negatively density dependent, and this was in part due to decreases in plant quality both as herbivore density increased and as the amount of damage increased. The response of plant quality to herbivores was found to have non‐linear features; there was both a threshold below which no significant decreases in quality (as measured by herbivore growth) occurred, and the decrease in herbivore performance saturated at the highest damage levels. In addition, it was found that caterpillar damage was significantly more aggregated than expected when multiple caterpillars occupy a single plant. This study confirms that host plants have the potential to be a source of density dependence that affects herbivore performance.     

Consequences of plant–chemical diversity for domestic goat food preference in Mediterranean forests

The domestic goat, a major herbivore in the Mediterranean basin, has demonstrated a strong ability to adapt its feeding behaviour to the chemical characteristics of food, selecting plants according to their nutritive quality. In this study, we determine some chemical characteristics related to plant nutritional quality and its variability among and within five tree species, these being the main components of the mountain forests of SE Spain, with the aim of determining their influence on food selection by this generalist herbivore. We analyse nitrogen, total phenols, condensed tannins and fibre concentration as an indicator of the nutritive value of the different trees. To determine the preference by the domestic goat, we performed two types of feeding-choice assays, where goats had to select between different species or between branches of the same species but from trees of different nutritional quality. The analysis of the plant nutritional quality showed significant differences in the chemical characteristics between species, and a high variability within species. However, when faced with different tree species, the domestic goat selected some of them but showed striking individual differences between goats. When selecting between trees of the same species, the goats showed no differential selection. This limited effect of chemical plant characteristics, together with the variability in foraging behaviour, resulted in a widespread consumption of diverse plant species, which can potentially modulate the effect of the goat on vegetation composition, and open the way for the conservation of traditional livestock grazing on natural protected areas.     

Disentangling effects of induced plant defenses and food quantity on herbivores by fitting nonlinear models

Abstract Plants can respond to herbivore damage through both broad-scale (systemic) and localized induced responses. While many studies have quantified the impact of systemic responses on herbivores, measuring the impact of localized changes is difficult because plant tissues that have suffered direct damage may represent both a lower quality and a lower quantity of food. This article uses nonlinear models to disentangle the confounding effects of prior herbivory on food quantity and quality. The first (null) model assumes that herbivore performance is determined only by the quantity of food available to an average herbivore. Modified models allow two distinct effects of damage-induced defenses: an increase in the amount of food each herbivore is required to consume in order to achieve maximum performance and a reduction in the maximum performance even when herbivores are fed ad lib. Maximum likelihood methods were used to fit the models to data from field experiments in which Colorado potato beetle (Leptinotarsa decemlineata) larvae were reared on three varieties of potatoes that had been damaged to varying degrees by adult beetles. Prior damage reduced the mean mass of beetles at pupation, and this effect was due to both a decrease in food quantity and induced changes in food quality. In contrast, beetle survival was affected in some cases by reduced food quantity but showed no responses that could be attributed to induced defenses. I discuss this result in the context of previous studies of induced (mostly systemic) responses in the potato-potato beetle system, and I suggest that detailed studies of particular chemical responses and the proposed method of combining bioassays with quantitative models should be used as complementary approaches in future studies of herbivore-induced defenses in plants.     

Variations in leaf traits and susceptibility to insect herbivory within a Salix miyabeana population under field conditions

Variations in leaf traits (toughness, total nitrogen and total phenolic concentrations) and susceptibility to herbivory in Salix miyabeana were studied among individual trees within a population under field conditions. Leaf quality clearly decreased as season progressed, i.e. increases in leaf toughness and total phenolics and decrease in leaf nitrogen. Seasonal pattern and extent of herbivore attack were similar between years. Significant correlation between leaf traits and susceptibility to herbivore attack was detected, while effects of sex and plant size on leaf traits and herbivory were less clear. There was a negative correlation between total nitrogen and total phenolics, and a positive correlation between leaf toughness and total phenolics among trees. Trees with high quality leaves tended to suffer from frequent herbivore attack and leaf damage. Such a clear relationship between leaf traits and susceptibility to herbivory may be related with a life-history strategy of willows, which are rapid-growing pioneer species and generally respond to herbivorous damage not by induced resistance but by compensative growth. This revised version was published online in August 2006 with corrections to the Cover Date.     

Insect herbivores and their frass affect Quercus rubra leaf quality and initial stages of subsequent litter decomposition

Defoliation‐induced changes in plant foliage are ubiquitous, though factors mediating induction and the extent of their influence on ecosystem processes such as leaf litter decomposition are poorly understood. Soil nitrogen (N) availability, which can be affected by insect herbivore frass (feces), influences phytochemical induction. We conducted experiments to test the hypotheses that insect frass deposition would (1) reduce phytochemical induction following herbivory and (2) increase the decomposition and nutrient release of the subsequent leaf litter. During the 2002 growing season, 80 Quercus rubra saplings were subjected to a factorial experiment with herbivore and frass manipulations. Leaf samples were collected throughout the growing season to measure the effects of frass deposition on phytochemical induction. In live foliage, herbivore damage increased tannin concentrations early, reduced foliar N concentrations throughout the growing season, and lowered lignin concentrations in the late season. Frass deposition apparently reduced leaf lignin concentrations, but otherwise did not influence leaf chemistry. Following natural senescence, litter samples from the treatment groups were decomposed in replicated litterbags for 18 months at the Coweeta Hydrologic Laboratory, NC. In the dead litter samples, initial tannin concentrations were lower in the herbivore damage group and higher in the frass addition group relative to their respective controls. Tannin and N release rates in the first nine months of decomposition were also affected by both damage and frass. However, decomposition rates did not differ among treatment groups. Thus, nutrient dynamics important for some ecosystem processes may be independent from the physical loss of litter mass. Overall, while lingering effects of damage and even frass deposition can therefore carry over and affect ecosystem processes during decomposition, their effects appear short lived relative to abiotic forces that tend to homogenize the decomposition process.     

Model analysis for plant disease dynamics co-mediated by herbivory and herbivore-borne phytopathogens

Plants are subject to diseases caused by pathogens, many of which are transmitted by herbivorous arthropod vectors. To understand plant disease dynamics, we studied a minimum hybrid model combining consumer-resource (herbivore-plant) and susceptible-infected models, in which the disease is transmitted bi-directionally between the consumer and the resource from the infected to susceptible classes. Model analysis showed that: (i) the disease is more likely to persist when the herbivore feeds on the susceptible plants rather than the infected plants, and (ii) alternative stable states can exist in which the system converges to either a disease-free or an endemic state, depending on the initial conditions. The second finding is particularly important because it suggests that the disease may persist once established, even though the initial prevalence is low (i.e. the R(0) rule does not always hold). This situation is likely to occur when the infection improves the plant nutritive quality, and the herbivore preferentially feeds on the infected resource (i.e. indirect vector-pathogen mutualism). Our results highlight the importance of the eco-epidemiological perspective that integration of tripartite interactions among host plant, plant pathogen and herbivore vector is crucial for the successful control of plant diseases.     

Thresholds in plant–herbivore interactions: predicting plant mortality due to herbivore browse damage

Patterns of herbivore browse at small scales, such as the rate of leaf consumption or plant preferences, drive the impact of herbivores on whole-plant processes, such as growth or survival, and subsequent changes in plant population structure. However, herbivore impacts are often non-linear, highly variable and context-dependent. Understanding the effect of herbivores on plant populations therefore requires a detailed understanding of the relationships that drive small-scale processes, and how these interact to generate dynamics at larger scales. We derive a mathematical model to predict annual rates of browse-induced tree mortality. We model individual plant mortality as a result of rates of foliage production, turnover and herbivore intake, and extend the model to the population scale by allowing for between-tree variation in levels of herbivore browse. The model is configurable for any broadleaved tree species subject to vertebrate or invertebrate browse, and is designed to be parameterized from field data typically collected as part of browse damage assessments. We parameterized and tested the model using data on foliage cover and browse damage recorded on kamahi trees (Weinmannia racemosa) browsed by possums (Trichosurus vulpecula) in New Zealand forests. The model replicated observed patterns of tree mortality at 12 independent validation sites with a wide range of herbivore densities and browse damage. The model reveals two key thresholds; in plant foliar cover, indicating when individual trees may be at high risk from browse-induced mortality, and in herbivore intake, leading to high rates of mortality across the whole population.     

The effects of herbivory and soil fertility on the growth patterns of <Emphasis Type="Italic">Quercus serrata</Emphasis> and <Emphasis Type="Italic">Q. crispula</Emphasis> saplings at the shoot and individual levels

We investigated the combined effects of herbivore damage and soil fertility on shoot growth patterns of Quercus serrata and Q. crispula saplings at both the shoot and individual levels. Saplings were grown in herbivore-damaged or undamaged areas of the greenhouse with the two fertilization treatment levels, low or high. We measured the leaf area loss, number of flushes, length of extension units (EUs; the first vs the higher), number of leaves on each individual, and number of EUs. At the shoot level, the leaf area loss at high soil fertility was significantly greater than that at low soil fertility among the highest EUs of Q. serrata, while this difference was not significant in Q. crispula, suggesting that effect of soil fertility on leaf area loss is species-specific. Furthermore, herbivore damage was associated with a significant increase in the number of EUs and a reduction in the length of the higher EUs under both soil fertility treatments, although saplings had a tendency to produce significantly more flushes and longer individual EUs under the high soil fertility. At the individual level, herbivore-damaged saplings exhibited a significant increase in leaf numbers; however, the total length of the EUs in Q. serrata or Q. crispula was not significantly affected by herbivore damage, regardless of soil fertility. These results suggest that Q. serrata and Q. crispula saplings produce shorter EUs in response to herbivore damage in order to reduce the cost of mechanical support and spread the risk for any subsequent herbivore damage.     

Sex-biased herbivory: a meta-analysis of the effects of gender on plant-herbivore interactions

We used sets of meta-analyses to review the evidence of sex-biased herbivory in dioecious plants as well as intersexual differences in plant characteristics that might affect herbivores. Thirtythree studies were reviewed to assess the effects of plant sex on herbivore abundance, survivorship and damage imposed to host plants, whereas 54 studies were reviewed for the effects of plant sex on plant morphological, physiological, and nutritional characteristics. The standardized mean difference between males and females was chosen as the measure to calculate effect sizes. Male plants exhibited significantly higher numbers of herbivores (d++=1.074) and significantly higher herbivory measured in terms of plant damage (d++=0.577) compared to female plants. Effects of plant sex on herbivore abundance were stronger for folivores and gallformers compared to the other guilds, whereas effects of plant sex on herbivore damage were stronger for flower predators compared to browsers and folivores. No difference in herbivore survivorship was observed between sexes. Male plants exhibited significantly more leaves (d++=0.202), larger leaves (d++=0.91), fewer flowers (d++=−0.89) and longer stems (d++=0.614) than female plants. Although male plants exhibited significantly lower concentrations of secondary compounds (d++=−0.209) and other defenses (d++=−0.53) than female plants, no difference in nutrient concentration, such as foliar nitrogen, was observed between sexes.     

Geographic patterns of plant–herbivore interactions are driven by soil fertility

Aims Geographic patterns of the intensity of plant herbivory in relation to climate factors have garnered little general support and appear to be species specific. However, plant–herbivore interactions are also driven by resource availability, such as soil nutrient content, and it remains unclear whether broad-scale variation in soil factors is reflected in herbivore consumption rates across species’ ranges. Additionally, we know little of how intraspecific variation in tissue quality associates with edaphic and climatic factors, and how this variation controls herbivore consumption. The resource availability hypothesis (RAH) predicts that plant individuals growing in low-resource environments will have lower leaf nutritional quality and more constitutive defenses, which will result in lower rates of leaf consumption.     

Plant-herbivore interactions along elevational gradient: Comparison of field and common garden data

In response to climate change, various organisms tend to migrate to higher elevations and latitudes. Unequal migration rates of plants and animals are expected to result in changes in the type and intensity of their interactions such as plant-herbivore interactions. In the present study, we studied the extent of herbivore damage in Salvia nubicola along an elevational gradient in Manang, central Nepal. A common garden experiment was also carried out by sowing seeds collected from different populations along the elevational gradient. As expected, the extent of herbivore damage in the field was significantly lower at higher elevations, and it increased with the population size and at sites without shrubs. In the common garden experiment, herbivore damage was higher in plants originating from lower elevations and from more open habitats. While higher herbivore pressure in the field at lower elevations may suggest that plants will be better protected against herbivores at lower elevations, the common garden study demonstrated the opposite. A possible explanation could be that plants from higher elevations have to adapt to extreme conditions, and lower palatability is a side effect of these adaptations. Thus, S. nubicola in the Himalayan region is likely to survive the expected higher herbivore pressure caused by an upward shift of herbivores under future climate change. Future studies should attempt to elucidate generality of such a conclusion by studying multiple species along similar gradients. Our results from comparison of the field and common garden study suggest that future experiments need to include comparisons in common environments to understand the expected response of plants to changes in herbivore pressure.     

Variation in plant quality and the population dynamics of herbivores: there is nothing average about aphids

In the attempt to use results from small-scale studies to make large-scale predictions, it is critical that we take into account the greater spatial heterogeneity encountered at larger spatial scales. An important component of this heterogeneity is variation in plant quality, which can have a profound influence on herbivore population dynamics. This influence is particularly relevant when we consider that the strength of density dependence can vary among host plants and that the strength of density dependence determines the difference between exponential and density- dependent growth. Here, we present some simple models and analyses designed to examine the impact of variable plant quality on the dynamics of insect herbivore populations, and specifically the consequences of variation in the strength of density dependence among host plants. We show that average values of herbivore population growth parameters, calculated from plants that vary in quality, do not predict overall population growth. Furthermore, we illustrate that the quality of a few individual plants within a larger plant population can dominate herbivore population growth. Our results demonstrate that ignoring spatial heterogeneity that exists in herbivore population growth on plants that differ in quality can lead to a misunderstanding of the mechanisms that underlie population dynamics.     

CO2‐mediated changes of plant traits and their effects on herbivores are determined by leaf age

1. Concentration of atmospheric CO₂ is predicted to double during the 21st century. However, quantitative effects of increased CO₂ levels on natural herbivore–plant interactions are still little understood. 2. In this study, we assess whether increased CO₂ quantitatively affects multiple defensive and nutritive traits in different leaf stages of cyanogenic wildtype lima bean plants (Phaseolus lunatus), and whether plant responses influence performance and choice behaviour of a natural insect herbivore, the Mexican bean beetle (Epilachna varivestis). 3. We cultivated lima bean plants in climate chambers at ambient, 500, 700, and 1000 ppm CO₂ and analysed cyanogenic precursor concentration (nitrogen‐based defence), total phenolics (carbon‐based defence), leaf mass per area (LMA; physical defence), and soluble proteins (nutritive parameter) of three defined leaf age groups. 4. In young leaves, cyanide concentration was the only parameter that quantitatively decreased in response to CO₂ treatments. In intermediate and mature leaves, cyanide and protein concentrations decreased while total phenolics and LMA increased. 5. Depending on leaf stage, CO₂‐mediated changes in leaf traits significantly affected larval performance and choice behaviour of adult beetles. We observed a complete shift from highest herbivore damage in mature leaves under natural CO₂ to highest damage of young leaves under elevated CO_2. Our study shows that leaf stage is an essential factor when considering CO₂‐mediated changes of plant defences against herbivores. Since in the long run preferred consumption of young leaves can strongly affect plant fitness, variable effects of elevated CO₂ on different leaf stages should receive highlighted attention in future research.     

Fluctuations in food supply in an insularized and heavily grazed savanna ecosystem in Kenya

The herbaceous plant standing crop differed significantly between seasons and vegetation types. It showed similar seasonal fluctuations in all habitat types although herbivore offtake rates differed considerably among habitats. Nutritive value of the pasture also differed between seasons, sometimes falling below minimum levels for herbivore maintenance. These conditions created considerable nutritive stress for the high-density large herbivore community. This study indicates the need for active management of plant and animal communities to maintain species and habitat diversity.     

A short-lived herbivore on a long-lived host: tree resistance to herbivory depends on leaf age

Short-lived insect herbivores should be able to adapt to the resistance mechanisms of their long-lived woody hosts because the life span of a single host will encompass numerous generations of herbivores. However, adaptation may be slowed down if host genotypes can create, in a single genotype, such large phenotypic variation in traits relevant for the herbivore that it matches variance among host genotypes. We tested this hypothesis by measuring leaf consumption by, and growth of, half-sibs of the geometrid moth Epirrita autumnata on individual birch trees, during three instars. The instar×tree interaction, rather than tree identity alone, was a significant variance component for both consumption and growth, indicating that different larval instars ranked individual trees differently. Both consumption and growth varied most between the 3rd and the later (4th and 5th) instars, coinciding with rapid seasonal changes in numerous nutritive and phenolic traits of maturing leaves. Thus, developmental variance in the leaf quality of individual trees may reduce the likelihood of E. autumnata genotypes adapting to the defenses of their host trees. We did not find evidence of in the ability of different half-sibs to utilize individual trees or leaf stages, indicating that E. autumnata larvae are generalists over a wide variety of host traits.     

Variation in insect herbivory across a salt marsh tidal gradient influences plant survival and distribution

Herbivore damage and impact on plants often varies spatially across environmental gradients. Although such variation has been hypothesized to influence plant distribution, few quantitative evaluations exist. In this study I evaluated patterns of insect herbivory on an annual forb, Atriplex patula var. hastata, across a salt marsh tidal gradient, and performed experiments to examine potential causes and consequences of variation in herbivory. Damage to plants was generally twice as great at mid-tidal elevations, which are more frequently inundated, than at higher, less stressful, elevations at five of six surveyed sites. Field herbivore assays and herbivore preference experiments eliminated the hypothesis that plant damage was mediated by herbivore response to differences in host plants across the gradient. Alternately, greater herbivore densities in the mid-marsh, where densities of an alternate host plant (Salicornia europaea) were high, were associated with greater levels of herbivory on Atriplex, suggesting spillover effects. The effect of insect herbivores on host plant performance varied between the two sites studied more intensively. Where overall herbivore damage to plants was low, herbivory had no detectable effect on plant survival or seed production, and plant performance did not significantly differ between zones. However, where herbivore damage was high, herbivores dramatically reduced both plant survival (>50%) and fruit production (40-70%), and their effects were stronger in the harsher mid-marsh than the high marsh. Thus herbivores likely play a role in maintaining lower Atriplex densities in mid-marsh. Overall, these results suggest that variation in herbivore pressure can be an important determinant of patterns of plant abundance across environmental gradients.     

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

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