Girdling behavior of the longhorn beetle modulates the host plant to enhance larval performance.

BACKGROUND: Preingestive behavioral modulations of herbivorous insects on the host plant are abundant over insect taxa. Those behaviors are suspected to have functions such as deactivation of host plant defenses, nutrient accumulation, or modulating plant-mediated herbivore interactions. To understand the functional consequence of behavioral modulation of insect herbivore, we studied the girdling behavior of Phytoecia rufiventris Gautier (Lamiinae; Cerambycidae) on its host plant Erigeron annuus L. (Asteraceae) that is performed before endophytic oviposition in the stem. RESULTS: The girdling behavior significantly increased the larval performance in both field monitoring and lab experiment. The upper part of the girdled stem exhibited lack of jasmonic acid induction upon larval attack, lowered protease inhibitor activity, and accumulated sugars and amino acids in compared to non-girdled stem. The girdling behavior had no effect on the larval performance of a non-girdling longhorn beetle Agapanthia amurensis, which also feeds on the stem of E. annuus during larval phase. However, the girdling behavior decreased the preference of A. amurensis females for oviposition, which enabled P. rufiventris larvae to avoid competition with A. amurensis larvae. CONCLUSIONS: In conclusion, the girdling behavior modulates plant physiology and morphology to provide a modulated food source for larva and hide it from the competitor. Our study implies that the insect behavior modulations can have multiple functions, providing insights into adaptation of insect behavior in context of plant-herbivore interaction.

Symbiosis: Did bacteria bias the beetle big bang?

The massive species richness of certain taxonomic groups has long enchanted evolutionary biologists, but even within such groups there are biases in cladogenesis. A study of Metazoa's greatest radiation - the beetles - points to metabolic symbioses with bacteria as a possible driver of enhanced diversification in herbivorous clades.

Functional traits-not nativeness-shape the effects of large mammalian herbivores on plant communities.

Large mammalian herbivores (megafauna) have experienced extinctions and declines since prehistory. Introduced megafauna have partly counteracted these losses yet are thought to have unusually negative effects on plants compared with native megafauna. Using a meta-analysis of 3995 plot-scale plant abundance and diversity responses from 221 studies, we found no evidence that megafauna impacts were shaped by nativeness, "invasiveness," "feralness," coevolutionary history, or functional and phylogenetic novelty. Nor was there evidence that introduced megafauna facilitate introduced plants more than native megafauna. Instead, we found strong evidence that functional traits shaped megafauna impacts, with larger-bodied and bulk-feeding megafauna promoting plant diversity. Our work suggests that trait-based ecology provides better insight into interactions between megafauna and plants than do concepts of nativeness.

Contrasting plant transcriptome responses between a pierce-sucking and a chewing herbivore go beyond the infestation site.

BACKGROUND: Plants have acquired a repertoire of mechanisms to combat biotic stressors, which may vary depending on the feeding strategies of herbivores and the plant species. Hormonal regulation crucially modulates this malleable defense response. Jasmonic acid (JA) and salicylic acid (SA) stand out as pivotal regulators of defense, while other hormones like abscisic acid (ABA), ethylene (ET), gibberellic acid (GA) or auxin also play a role in modulating plant-pest interactions. The plant defense response has been described to elicit effects in distal tissues, whereby aboveground herbivory can influence belowground response, and vice versa. This impact on distal tissues may be contingent upon the feeding guild, even affecting both the recovery of infested tissues and those that have not suffered active infestation. RESULTS: To study how phytophagous with distinct feeding strategies may differently trigger the plant defense response during and after infestation in both infested and distal tissues, Arabidopsis thaliana L. rosettes were infested separately with the chewing herbivore Pieris brassicae L. and the piercing-sucker Tetranychus urticae Koch. Moderate infestation conditions were selected for both pests, though no quantitative control of damage levels was carried out. Feeding mode did distinctly influence the transcriptomic response of the plant under these conditions. Though overall affected processes were similar under either infestation, their magnitude differed significantly. Plants infested with P. brassicae exhibited a short-term response, involving stress-related genes, JA and ABA regulation and suppressing growth-related genes. In contrast, T. urticae elicited a longer transcriptomic response in plants, albeit with a lower degree of differential expression, in particular influencing SA regulation. These distinct defense responses transcended beyond infestation and through the roots, where hormonal response, flavonoid regulation or cell wall reorganization were differentially affected. CONCLUSION: These outcomes confirm that the existent divergent transcriptomic responses elicited by herbivores employing distinct feeding strategies possess the capacity to extend beyond infestation and even affect tissues that have not been directly infested. This remarks the importance of considering the entire plant's response to localized biotic stresses.

The first intron of EIJ1 confers a specific response to wounding and herbivore stresses.

Plants are constantly exposed to different kinds of biotic stress, such as herbivore attack and wounding. To deal with these stresses, plants have evolved sophisticated defence mechanisms to protect themselves. Previously, we found that EIJ1 (EDS1-interacting J protein 1) plays a negative regulatory role in plant disease resistance in Arabidopsis thaliana. Follow-up studies revealed that EIJ1 specifically responds to wounding and herbivore stresses. The expression of EIJ1 was specifically induced by wounding or herbivore stress, as demonstrated by similar results in EIJ1 protein assay. Interestingly, GUS staining found that the promoter of EIJ1 is not involved in the induction of expression under wounding stress. Instead, we identified the first intron of EIJ1 as a key factor in response to wounding stress. Deleting the first intron of EIJ1 resulted in a loss of response to wounding stress in plants. Our results broaden the role of EIJ1 in plant resistance to biotic stress and provide new insights into plant responses to biotic stress.

Predator mass mortality events restructure food webs through trophic decoupling.

Predators have a key role in structuring ecosystems1-4. However, predator loss is accelerating globally4-6, and predator mass-mortality events7 (MMEs)-rapid large-scale die-offs-are now emblematic of the Anthropocene epoch6. Owing to their rare and unpredictable nature7, we lack an understanding of how MMEs immediately impact ecosystems. Past predator-removal studies2,3 may be insufficient to understand the ecological consequences of MMEs because, in nature, dead predators decompose in situ and generate a resource pulse8, which could alter ensuing ecosystem dynamics by temporarily enhancing productivity. Here we experimentally induce MMEs in tritrophic, freshwater lake food webs and report ecological dynamics that are distinct from predator losses2,3 or resource pulses9 alone, but that can be predicted from theory8. MMEs led to the proliferation of diverse consumer and producer communities resulting from weakened top-down predator control1-3 and stronger bottom-up effects through predator decomposition8. In contrast to predator removals alone, enhanced primary production after MMEs dampened the consumer community response. As a consequence, MMEs generated biomass dynamics that were most similar to those of undisturbed systems, indicating that they may be cryptic disturbances in nature. These biomass dynamics led to trophic decoupling, whereby the indirect beneficial effects of predators on primary producers are lost and later materialize as direct bottom-up effects that stimulate primary production amid intensified herbivory. These results reveal ecological signatures of MMEs and demonstrate the feasibility of forecasting novel ecological dynamics arising with intensifying global change.

Fatty alcohols, a minor component of the tree tobacco surface wax, are associated with defence against caterpillar herbivory.

Despite decades of research resulting in a comprehensive understanding of epicuticular wax metabolism, the function of these almost ubiquitous metabolites in plant-herbivore interactions remains unresolved. In this study, we examined the effects of CRISPR-induced knockout mutations in four Nicotiana glauca (tree tobacco) wax metabolism genes. These mutations cause a wide range of changes in epicuticular wax composition, leading to altered interactions with insects and snails. Three interaction classes were examined: chewing herbivory by seven caterpillars and one snail species, phloem feeding by Myzus persicae (green peach aphid) and oviposition by Bemisia tabaci (whitefly). Although total wax load and alkane abundance did not affect caterpillar growth, a correlation across species, showed that fatty alcohols, a minor component of N. glauca surface waxes, negatively affected the growth of both a generalist caterpillar (Spodoptera littoralis) and a tobacco-feeding specialist (Manduca sexta). This negative correlation was overshadowed by the stronger effect of anabasine, a nicotine isomer, and was apparent when fatty alcohols were added to an artificial lepidopteran diet. By contrast, snails fed more on waxy leaves. Aphid reproduction and feeding activity were unaffected by wax composition but were potentially affected by altered cutin composition. Wax crystal morphology could explain the preference of B. tabaci to lay eggs on waxy wild-type plants relative to both alkane and fatty alcohol-deficient mutants. Together, our results suggest that the varied responses among herbivore classes and species are likely to be a consequence of the co-evolution that shaped the specific effects of different surface wax components in plant-herbivore interactions.

How plant traits respond to and affect vertebrate and invertebrate herbivores-Are measurements comparable across herbivore types?

Despite plants realistically being affected by vertebrate and invertebrate herbivores simultaneously, fundamental differences in the ecology and evolution of these two herbivore guilds often means their impacts on plants are studied separately. A synthesis of the literature is needed to understand the types of plant traits examined and their response to, and effect on (in terms of forage selection) vertebrate and invertebrate herbivory, and to identify associated knowledge gaps. Focusing on grassland systems and species, we found 138 articles that met our criteria: 39 invertebrate, 97 vertebrate and 2 focussed on both vertebrate and invertebrate herbivores. Our study identified invertebrate focussed research, research conducted in the Southern Hemisphere and research on nondomesticated herbivores was significantly underrepresented based on our search and should be a focus of future research. Differences in study focus (trait response or trait effect), along with differences in the types of traits examined, led to limited opportunity for comparison between the two herbivore guilds. This review therefore predominantly discusses the response and effect of plant traits to each herbivore guild separately. In future studies, we suggest this review be used as a guide for trait selection, to improve comparability and the broader significance of results.

Electrical signalling and plant response to herbivory: A short review.

For a long time, electrical signaling was neglected at the expense of signaling studies in plants being concentrated with chemical and hydraulic signals. Studies conducted in recent years have revealed that plants are capable of emitting, processing, and transmitting bioelectrical signals to regulate a wide variety of physiological functions. Many important biological and physiological phenomena are accompanied by these cellular electrical manifestations, which supports the hypothesis about the importance of bioelectricity as a fundamental 'model' for response the stresses environmental and for activities regeneration of these organisms. Electrical signals have also been characterized and discriminated against in genetically modified plants under stress mediated by sucking insects and/or by the application of systemic insecticides. Such results can guide future studies that aim to elucidate the factors involved in the processes of resistance to stress and plant defense, thus aiding in the development of successful strategies in integrated pest management. Therefore, this mini review includes the results of studies aimed at electrical signaling in response to biotic stress. We also demonstrated how the generation and propagation of electrical signals takes place and included a description of how these electrical potentials are measured.

The Capsicum terpenoid biosynthetic module is affected by spider-mite herbivory.

In response to herbivory, Capsicum annuum leaves adapt their specialized metabolome that may protect the plant against herbivore feeding either directly or indirectly through volatile metabolites acting as cues for natural enemies of the herbivore. The volatile blend of spider-mite infested leaves differs from non-challenged leaves predominantly by a higher contribution of mono- and sesquiterpenes. In addition to these terpenoids released into the headspace, the terpenoid composition of the leaves alters upon herbivory. All this suggests an important role for terpenoids and their biosynthetic machinery in the defence against herbivory. Here, we show that the C. annuum genome contains a terpene synthase (TPS) gene family of 103 putative members of which structural analysis revealed that 27 encode functional enzymes. Transcriptome analysis showed that several TPS loci were differentially expressed upon herbivory in leaves of two C. annuum genotypes, that differ in susceptibility towards spider mites. The relative expression of upstream biosynthetic genes from the mevalonate and the methylerythritol phosphate pathway also altered upon herbivory, revealing a shift in the metabolic flux through the terpene biosynthetic module. The expression of multiple genes potentially acting downstream of the TPSs, including cytochrome P450 monooxygenases, UDP-glucosyl transferases, and transcription factors strongly correlated with the herbivory-induced TPS genes. A selection of herbivory-induced TPS genes was functionally characterized through heterologous expression and the products that these enzymes catalysed matched with the volatile and non-volatile terpenoids induced in response to herbivory.

Soil salinization and chemically mediated plant-insect interactions in a changing climate.

Increase in soil salinization due to climate change is a global phenomenon that can induce significant changes in plant growth, physiology, and chemistry, exacerbating growing threats to insect biodiversity. Insects that rely on plants are likely to be indirectly impacted by changes in soil salt content through changes in plant chemistry, yet few studies link changes in plant metabolism to impacts on higher trophic levels. Some salinity-mediated changes in specialized metabolites may be predictable due to highly conserved metabolic pathways shared between herbivore defense and stress resistance, but recent studies also suggest substantial variation across plant species and habitats. To date, most of the research on salinity and chemically mediated plant-insect interactions has focused on herbivores, particularly in agricultural systems. Published effects of salinity on pollinators and parasitoids are scarce. Future research will need to focus more on the role of plant chemistry to bridge the divide between studies of plant and insect responses to salinization.

Microbial degradation of plant toxins.

Plants produce a variety of secondary metabolites in response to biotic and abiotic stresses. Although they have many functions, a subclass of toxic secondary metabolites mainly serve plants as deterring agents against herbivores, insects, or pathogens. Microorganisms present in divergent ecological niches, such as soil, water, or insect and rumen gut systems have been found capable of detoxifying these metabolites. As a result of detoxification, microbes gain growth nutrients and benefit their herbivory host via detoxifying symbiosis. Here, we review current knowledge on microbial degradation of toxic alkaloids, glucosinolates, terpenes, and polyphenols with an emphasis on the genes and enzymes involved in breakdown pathways. We highlight that the insect-associated microbes might find application in biotechnology and become targets for an alternative microbial pest control strategy.

Direct and indirect impacts of ocean acidification and warming on algae-herbivore interactions in intertidal habitats.

Anthropogenically induced global climate change has caused profound impacts in the world ocean. Climate change related stressors, like ocean acidification (OA) and warming (OW) can affect physiological performance of marine species. However, studies evaluating the impacts of these stressors on algae-herbivore interactions have been much more scarce. We approached this issue by assessing the combined impacts of OA and OW on the physiological energetics of the herbivorous snail Tegula atra, and whether this snail is affected indirectly by changes in biochemical composition of the kelp Lessonia spicata, in response to OA and OW. Our results show that OA and OW induce changes in kelp biochemical composition and palatability (organic matter, phenolic content), which in turn affect snails' feeding behaviour and energy balance. Nutritional quality of food plays a key role on grazers' physiological energetics and can define the stability of trophic interactions in rapidly changing environments such as intertidal communities.

Long-term livestock exclusion increases plant richness and reproductive capacity in arid woodlands.

Herbivore exclusion is implemented globally to recover ecosystems from grazing by introduced and native herbivores, but evidence for large-scale biodiversity benefits is inconsistent in arid ecosystems. We examined the effects of livestock exclusion on dryland plant richness and reproductive capacity. We collected data on plant species richness and seeding (reproductive capacity), rainfall, vegetation productivity and cover, soil strength and herbivore grazing intensity from 68 sites across 6500 km2 of arid Georgina gidgee (Acacia georginae) woodlands in central Australia between 2018 and 2020. Sites were on an actively grazed cattle station and two destocked conservation reserves. We used structural equation modeling to examine indirect (via soil or vegetation modification) versus direct (herbivory) effects of grazing intensity by two introduced herbivores (cattle, camels) and a native herbivore (red kangaroo), on seasonal plant species richness and seeding of all plants, and the richness and seeding of four plant groups (native grasses, forbs, annual chenopod shrubs, and palatable perennial shrubs). Non-native herbivores had a strong indirect effect on plant richness and seeding by reducing vegetative ground cover, resulting in decreased richness and seeding of native grasses and forbs. Herbivores also had small but negative direct impacts on plant richness and seeding. This direct effect was explained by reductions in annual chenopod and palatable perennial shrub richness under grazing activity. Responses to grazing were herbivore-dependent; introduced herbivore grazing reduced native plant richness and seeding, while native herbivore grazing had no significant effect on richness or seeding of different plant functional groups. Soil strength decreased under grazing by cattle but not camels or kangaroos. Cattle had direct effects on palatable perennial shrub richness and seeding, whereas camels had indirect effects, reducing richness and seeding by reducing the abundance of shrubs. We show that considering indirect pathways improves evaluations of the effects of disturbances on biodiversity, as focusing only on direct effects can mask critical mechanisms of change. Our results indicate substantial biodiversity benefits from excluding livestock and controlling camels in drylands. Reducing introduced herbivore impacts will improve soil and vegetation condition, ensure reproduction and seasonal persistence of species, and protect native plant diversity.

Role of herbivory in shaping the dryland vegetation ecosystem: Linking spiral vegetation patterns and nonlinear, nonlocal grazing.

Self-organized vegetation patterns are an amazing aspect of dryland ecosystems; in addition to being visually appealing, patterns control how these water-deprived systems react to escalating environmental stress. Although there is a wide variety of vegetation patterns, little is known about the mechanisms behind spiral patterns. The well-known models that explain other vegetation patterns such stripes, rings, and fairy circles cannot account for these spirals. Here we have adopted a modeling approach in which the interplay between herbivore grazing and vegetation is found to be the reason why spirals form. To comprehend the nonlinear dependence of grazing on the availability vegetation, we have introduced a grazing term that gets saturated when forage is abundant. To account for the impact of the spatial nonhomogeneity in vegetation layout, it is thought that grazing is dependent on mean vegetation density rather than density at a single site. Results show how the system dynamics is changed fundamentally depending on the different types of grazing response. Incorporation of nonlocality into the herbivore grazing leads to spiral-shaped vegetation patterns only in natural grazing scenarios; however, no patterning is seen in human controlled herbivory. Overall, our research points to the nonlocal, nonlinear grazing behavior of herbivores as one of the major driving forces for the development of spiral patterns.

Impacts of large herbivores on terrestrial ecosystems.

Large herbivores play unique ecological roles and are disproportionately imperiled by human activity. As many wild populations dwindle towards extinction, and as interest grows in restoring lost biodiversity, research on large herbivores and their ecological impacts has intensified. Yet, results are often conflicting or contingent on local conditions, and new findings have challenged conventional wisdom, making it hard to discern general principles. Here, we review what is known about the ecosystem impacts of large herbivores globally, identify key uncertainties, and suggest priorities to guide research. Many findings are generalizable across ecosystems: large herbivores consistently exert top-down control of plant demography, species composition, and biomass, thereby suppressing fires and the abundance of smaller animals. Other general patterns do not have clearly defined impacts: large herbivores respond to predation risk but the strength of trophic cascades is variable; large herbivores move vast quantities of seeds and nutrients but with poorly understood effects on vegetation and biogeochemistry. Questions of the greatest relevance for conservation and management are among the least certain, including effects on carbon storage and other ecosystem functions and the ability to predict outcomes of extinctions and reintroductions. A unifying theme is the role of body size in regulating ecological impact. Small herbivores cannot fully substitute for large ones, and large-herbivore species are not functionally redundant - losing any, especially the largest, will alter net impact, helping to explain why livestock are poor surrogates for wild species. We advocate leveraging a broad spectrum of techniques to mechanistically explain how large-herbivore traits and environmental context interactively govern the ecological impacts of these animals.

BSR1, a Rice Receptor-like Cytoplasmic Kinase, Positively Regulates Defense Responses to Herbivory.

Crops experience herbivory by arthropods and microbial infections. In the interaction between plants and chewing herbivores, lepidopteran larval oral secretions (OS) and plant-derived damage-associated molecular patterns (DAMPs) trigger plant defense responses. However, the mechanisms underlying anti-herbivore defense, especially in monocots, have not been elucidated. The receptor-like cytoplasmic kinase Broad-Spectrum Resistance 1 (BSR1) of Oryza sativa L. (rice) mediates cytoplasmic defense signaling in response to microbial pathogens and enhances disease resistance when overexpressed. Here, we investigated whether BSR1 contributes to anti-herbivore defense responses. BSR1 knockout suppressed rice responses triggered by OS from the chewing herbivore Mythimna loreyi Duponchel (Lepidoptera: Noctuidae) and peptidic DAMPs OsPeps, including the activation of genes required for biosynthesis of diterpenoid phytoalexins (DPs). BSR1-overexpressing rice plants exhibited hyperactivation of DP accumulation and ethylene signaling after treatment with simulated herbivory and acquired enhanced resistance to larval feeding. As the biological significance of herbivory-induced accumulation of rice DPs remains unexplained, their physiological activities in M. loreyi were analyzed. The addition of momilactone B, a rice DP, to the artificial diet suppressed the growth of M. loreyi larvae. Altogether, this study revealed that BSR1 and herbivory-induced rice DPs are involved in the defense against chewing insects, in addition to pathogens.

Movement drives population dynamics of one of the most mobile ungulates on Earth: Insights from a mechanistic model.

Long-distance movements are hypothesized to positively influence population size and stability of mobile species. We tested this hypothesis with a novel modeling approach in which moving herbivores interact with the environment created by a dynamic global vegetation model using highly mobile Mongolian gazelles in the eastern Mongolian grasslands as a case study. Gazelle population dynamics were modeled from 1901 to 2018 under two scenarios, one allowing free movement and one restricting movement. Gazelles were 2.2 times more abundant when they could move freely and were extirpated in 71% of the study area when mobility was restricted. Mobility resulted in greater population increases during times of abundant forage and smaller population decreases during drought. Reduced thermoregulatory costs associated with climate change, combined with an increase in vegetation biomass, increased gazelle abundance. Since high abundances often resulted in overgrazing and, thus, extirpation when movement was restricted, mobility had an important role in maintaining higher densities. The novel modeling approach shows how accounting for not just herbivore but also plant ecophysiology can improve our understanding of the population dynamics of highly mobile herbivores, in particular when examining the effects of habitat and climate change. Since the model simulates herbivores based on general physiological mechanisms that apply across large herbivores and the vegetation model can be applied globally, it is possible to adapt the model to other large-herbivore systems.

Bringing Fundamental Insights of Induced Resistance to Agricultural Management of Herbivore Pests.

In response to herbivory, most plant species adjust their chemical and morphological phenotype to acquire induced resistance to the attacking herbivore. Induced resistance may be an optimal defence strategy that allows plants to reduce metabolic costs of resistance in the absence of herbivores, allocate resistance to the most valuable plant tissues and tailor its response to the pattern of attack by multiple herbivore species. Moreover, plasticity in resistance decreases the potential that herbivores adapt to specific plant resistance traits and need to deal with a moving target of variable plant quality. Induced resistance additionally allows plants to provide information to other community members to attract natural enemies of its herbivore attacker or inform related neighbouring plants of pending herbivore attack. Despite the clear evolutionary benefits of induced resistance in plants, crop protection strategies to herbivore pests have not exploited the full potential of induced resistance for agriculture. Here, we present evidence that induced resistance offers strong potential to enhance resistance and resilience of crops to (multi-) herbivore attack. Specifically, induced resistance promotes plant plasticity to cope with multiple herbivore species by plasticity in growth and resistance, maximizes biological control by attracting natural enemies and, enhances associational resistance of the plant stand in favour of yield. Induced resistance may be further harnessed by soil quality, microbial communities and associational resistance offered by crop mixtures. In the transition to more sustainable ecology-based cropping systems that have strongly reduced pesticide and fertilizer input, induced resistance may prove to be an invaluable trait in breeding for crop resilience.

Grazing-induced cattle behaviour modulates the secondary production in a Eurasian steppe ecosystem.

Livestock-grassland interactions are among the most important relationships in grazed grassland ecosystems, where herbivores play a crucial role in plant community and ecosystem functions. However, previous studies primarily have focused on the responses of grasslands to grazing, with few focussing on the effects of livestock behaviour that in turn would influence livestock intake and primary and secondary productivity. Through a 2-year grazing intensity experiment with cattle in Eurasian steppe ecosystem, global positioning system (GPS) collars were used to monitor animal movements, where animal locations were recorded at 10-min intervals during the growing season. We used a random forest model and the K-means method to classify animal behaviour and quantified the spatiotemporal movements of the animals. Grazing intensity appeared to be the predominant driver for cattle behaviour. Foraging time, distance travelled, and utilization area ratio (UAR) all increased with grazing intensity. The distance travelled was positively correlated with foraging time, yielding a decreased daily liveweight gain (LWG) except at light grazing. Cattle UAR showed a seasonal pattern and reached the maximum value in August. In addition, the canopy height, above-ground biomass, carbon content, crude protein, and energy content of plants all affected cattle behaviour. Grazing intensity and the resulting change in above-ground biomass and forage quality jointly determined the spatiotemporal characteristics of livestock behaviour. Increased grazing intensity limited forage resources and promoted intraspecific competition of livestock, which induced longer travelling distance and foraging time, and more even spatial distribution when seeking habitat, which ultimately led to a reduction in LWG. In contrast, under light grazing where there were sufficient forage resources, livestock exhibited higher LWG with less foraging time, shorter travelling distance, and more specialized habitat occupation. These findings support the Optimal Foraging Theory and the Ideal Free Distribution model, which may have important implications for grassland ecosystem management and sustainability.