Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Assessing the Jarman–Bell Principle: Scaling of intake,digestibility, retention time and gut fill with body mass in mammalian herbivores

Differences in allometric scaling of physiological characters have the appeal to explain species diversification and niche differentiation along a body mass (BM) gradient — because they lead to different combinations of physiological properties, and thus may facilitate different adaptive strategies. An important argument in physiological ecology is built on the allometries of gut fill (assumed to scale to BM^1.0) and energy requirements/intake (assumed to scale to BM^0.75) in mammalian herbivores. From the difference in exponents, it has been postulated that the mean retention time (MRT) of digesta should scale to BM^1.0–0.75 = BM^0.25. This has been used to argue that larger animals have an advantage in digestive efficiency and hence can tolerate lower-quality diets. However, empirical data does not support the BM^0.25 scaling of MRT, and the deduction of MRT scaling implies, according to physical principles, no scaling of digestibility; basing assumptions on digestive efficiency on the thus-derived MRT scaling amounts to circular reasoning. An alternative explanation considers a higher scaling exponent for food intake than for metabolism, allowing larger animals to eat more of a lower quality food without having to increase digestive efficiency; to date, this concept has only been explored in ruminants. Here, using data for 77 species in which intake, digestibility and MRT were measured (allowing the calculation of the dry matter gut contents (DMC)), we show that the unexpected shallow scaling of MRT is common in herbivores and may result from deviations of other scaling exponents from expectations. Notably, DMC have a lower scaling exponent than 1.0, and the 95% confidence intervals of the scaling exponents for intake and DMC generally overlap. Differences in the scaling of wet gut contents and dry matter gut contents confirm a previous finding that the dry matter concentration of gut contents decreases with body mass, possibly compensating for the less favorable volume–surface ratio in the guts of larger organisms. These findings suggest that traditional explanations for herbivore niche differentiation along a BM gradient should not be based on allometries of digestive physiology. In contrast, they support the recent interpretation that larger species can tolerate lower-quality diets because their intake has a higher allometric scaling than their basal metabolism, allowing them to eat relatively more of a lower quality food without having to increase digestive efficiency.     

A review of feeding and nutrition of herbivorous land crabs: adaptations to low quality plant diets

This paper reviews the nutritional ecology, the digestive physiology, and biochemistry of herbivorous land crabs and the adaptations that they possess towards a diet of plant material. Land crab species that breathe air and forage out of water can be divided into three feeding specialisations: primarily carnivorous, deposit feeders feeding on micro-organisms and organic matter in the sediment, and herbivores consuming mainly plant material and its detritus. The last forms the focus of this review. The diets of the herbivores are low in nitrogen and high in carbon, are difficult to digest since they contain cellulose and hemicellulose, and may disrupt digestion due to the presence of tannins. Herbivorous crustaceans are able to efficiently utilise plant material as their primary nutrient source and are indeed able to meet their nitrogen requirements from it. Herbivorous land crabs display a range of adaptations towards a low nitrogen intake and these are discussed in this review. They also appear to endogenously produce cellulase and hemicellulase enzymes for the digestion of cellulose and hemicellulose. Generalised and specific adaptations allow them to inhibit the potentially negative digestive effects of tannins. To digest plant material, they possess a plastic digestive strategy of high food intake, short retention time, high assimilation of cell contents, and substantial digestion of cellulose and hemicellulose.     

Flexibility of digestive responses in two generalist herbivores,the tortoises Geochelone carbonaria and Geochelone denticulata

Summary To test the prediction that digestive responses digestibility, intake and passage time-of generalist herbivores vary with different diets, feeding trials were conducted in Venezuela with two sympatric tortoise species, Geochelone carbonaria and G. denticulata. Three single-species diets (two fruit, one foliage) were fed to both species. For a given diet, digestibility, mass-specific intake and passage time did not differ between the two tortoise species, nor did they vary by sex or body mass within each species. However, the digestive parameters varied for tortoises feeding on the different diets. The responses ranged from nearly abandoning cell wall fermentation and depending entirely on extraction of cell contents to relying heavily on cell wall fermentation. Therefore, these generalist herbivores have flexible digestive responses that are influenced by diet, not fixed digestive responses that limit the diet, as previously observed in other generalist herbivores. A three-part classification of herbivores (specialist, specialized mixed feeder and opportunistic mixed feeder) is suggested as an approach to understanding flexible and inflexible digestive strategies in herbivores.     

Multiple pathways from three types of sugar receptor sites to metabotropic transduction pathways of the blowfly: study by the whole cell-clamp experiments

Differences in the allometric scaling between gut capacity (with body mass, BM^1.00) and food intake (with BM^0.75) should theoretically result in a scaling of digesta retention time with BM^0.25 and therefore a higher digestive efficiency in larger herbivores. This concept is an important part of the so-called ‘Jarman–Bell principle’ (JBP) that explains niche differentiation along a body size gradient in terms of digestive physiology. Empirical data in herbivorous mammals, however, do not confirm the scaling of retention time, or of digestive efficiency, with body mass. Here, we test these concepts in herbivorous reptiles, adding data of an experiment that measured food intake, digesta retention, digestibility and gut capacity in 23 tortoises (Testudo graeca, T. hermanni , Geochelone nigra, G. sulcata, Dipsochelys dussumieri) across a large BM range (0.5–180 kg) to a literature data collection. While dry matter gut fill scaled to BM^1.07 and dry matter intake to BM^0.76, digesta mean retention time (MRT) scaled to BM^0.17; the scaling exponent was not significantly different from zero for species > 1 kg. Food intake level was a major determinant of MRT across reptiles and mammals. In contrast to dietary fibre level, BM was not a significant contributor to dry matter digestibility in a General Linear Model. Digestibility coefficients in reptiles depended on diet nutrient composition in a similar way as described in mammals. Although food intake is generally lower and digesta retention longer in reptiles than in mammals, digestive functions scale in a similar way in both clades, indicating universal principles in herbivore digestive physiology. The reasons why the theoretically derived JBP has little empirical support remain to be investigated. Until then, the JBP should not be evoked to explain niche differentiation along a body size axis in terms of digestive physiology.     

The relationship between forage cell wall content and voluntary food intake in mammalian herbivores

• 1 It is generally assumed that animals compensate for a declining diet quality with increasing food intake. Differences in the response to decreasing forage quality in herbivores have been postulated particularly between cattle (ruminants) and horses (hindgut fermenters). However, empirical tests for both assumptions in herbivorous mammals are rare.
• 2 We collected data on voluntary food intake in mammals on forage‐only diets and related this to dietary neutral detergent fibre (NDF) content, assuming a nonlinear correlation between these measurements. Generally, the paucity of corresponding data is striking.
• 3 Elephants and pandas showed very high food intakes that appeared unrelated to dietary fibre content. Only in small rodents, and possibly in rabbits, was an increase in food intake on forages of higher NDF content evident. In particular, other large herbivores, including horses, followed patterns of decreasing intake with increasing forage NDF, also observed in domestic cattle or sheep.
• 4 For large herbivores, empirical data therefore do not – so far – support the notion that intake is increased in response to declining diet quality. However, data are in accord with the assumption that most large herbivores have an anticipatory strategy of acquiring body reserves when high‐quality forage is available, and reducing food intake (and potentially metabolic losses) when only low‐quality forage is available.
• 5 Intake studies in which the influence of digestive strategy on food intake capacity is tested should be designed as long‐term studies that outlast an anticipatory strategy and force animals to ingest as much as possible.
• 6 We suggest that a colonic separation mechanism coupled with coprophagy, in order to minimize metabolic faecal losses, is necessary below a body size threshold where an anticipatory strategy (living off body reserves, migration) is not feasible. Future studies aimed at investigating fine‐scale differences, for example between equids and bovids, should focus on non‐domesticated species.

     

Physical defences wear you down: progressive and irreversible impacts of silica on insect herbivores

1. Silica in the leaves of grasses can act as a defence against both vertebrate and invertebrate herbivores. The mechanisms by which silica affects herbivore performance are not well characterized. Here we expose an insect herbivore Spodoptera exempta to high-silica diets and test two mechanisms by which silica has been proposed to act as a defence. First, that silica reduces the digestibility of leaves and second, that silica causes wear to insect mandibles, both of which could potentially impact on herbivore performance. 2. Silica reduced the efficiency with which S. exempta converted ingested food to body mass and the amount of nitrogen absorbed from their food, leading to reduced insect growth rates. The measure of how efficiently herbivores utilize digested food (ECD) was unaffected by silica. 3. These effects occurred even with short-term exposure to silica-rich diets, but they also increased markedly with the duration of exposure and affected late instars more than early instar larvae. This appears to be due to the progressive impacts of silica with longer exposure times and suggests that herbivores cannot adapt to silica defences, nor do they develop a tolerance for silica with age. 4. Exposure to silica-rich diets caused increased mandible wear in S. exempta. This effect was extremely rapid, occurring within a single instar, further reducing feeding efficiency and growth rates. These effects on insect growth and feeding efficiency are nonreversible, persisting after the herbivore has switched diets. Up to a third of this residual impact can be explained by the degree of mandible wear caused by previous silica-rich diets. 5. The impacts of silica on S. exempta larvae were progressive with exposure time and could not be compensated for, even by switching to a different diet. Thus, herbivores cannot easily adapt to physical defences such as silica, suggesting this defence will have major implications for herbivore fitness.     

Evolution of specialization and ecological character displacement of herbivores along a gradient of plant quality

We study the combined evolutionary dynamics of herbivore specialization and ecological character displacement, taking into account foraging behavior of the herbivores, and a quality gradient of plant types. Herbivores can adapt by changing two adaptive traits: their level of specialization in feeding efficiency and their point of maximum feeding efficiency along the plant gradient. The number of herbivore phenotypes, their levels of specialization, and the amount of character displacement among them are the result of the evolutionary dynamics, which is driven by the underlying population dynamics, which in turn is driven by the underlying foraging behavior. Our analysis demonstrates broad conditions for the diversification of a herbivore population into many specialized phenotypes, for basically any foraging behavior focusing use on highest gains while also including errors. Our model predicts two characteristic phases in the adaptation of herbivore phenotypes: a fast character-displacement phase and a slow coevolutionary niche-shift phase. This two-phase pattern is expected to be of wide relevance in various consumer-resource systems. Bringing together ecological character displacement and the evolution of specialization in a single model, our study suggests that the foraging behavior of herbivorous arthropods is a key factor promoting specialist radiation.     

Optimal foraging and community structure: The allometry of herbivore food selection and competition

I address the selection of plants with different characteristics by herbivores of different body sizes by incorporating allometric relationships for herbivore foraging into optimal foraging models developed for herbivores. Herbivores may use two criteria in maximizing their nutritional intake when confronted with a range of food resources: a minimum digestibility and a minimum cropping rate. Minimum digestibility should depend on plant chemical characteristics and minimum cropping rate should depend on the density of plant items and their size (mass). If herbivores do select for these plant characteristics, then herbivores of different body sizes should select different ranges of these characteristics due to allometric relationships in digestive physiology, cropping ability and nutritional demands. This selectivity follows a regular pattern such that a herbivore of each body size can exclusively utilize some plants, while it must share other plants with herbivores of other body sizes. I empirically test this hypothesis of herbivore diet selectivity and the pattern of resource use that it produces in the field and experimentally. The findings have important implications for competition among herbivores and their population and community ecology. Furthermore, the results may have general applicability to other types of foragers, with general implications for how biodiversity is influenced.     

Resource partitioning along multiple niche dimensions in differently sized African savanna grazers

Resource partitioning among mammalian savanna herbivores is thought to be predominantly driven by differences in body size. In general, large herbivore species utilize abundant low quality forage while small herbivores focus on scarcer high quality food items. However, in a natural system other factors such as digestive strategy, season and the presence of megaherbivores (body size > 1000 kg) are likely to complicate allometric predictions. Non‐ruminants are probably better able to cope with abundant low quality food than ruminants of the same size causing a non‐ruminant to act ‘larger’ than allometrically predicted. Also, the effect of alternating seasons with high and low food availability on diet choice and hence the competitive interactions between co‐occurring herbivores is still poorly understood. Lastly, how megaherbivores deviate from allometric predictions (based on smaller species) is still not well quantified. In this study we examine resource partitioning among three ruminant and three non‐ruminant grazers: impala, wildebeest, buffalo, warthog, zebra and white rhinoceros (megaherbivore) in the savanna of Hluhluwe iMfolozi Park, South Africa. We analysed habitat and diet overlap, specifically grass species (something not commonly investigated) and grass height eaten, in both the wet and the dry seasons. We found that habitat utilization differences among the species were generally small and did not vary between seasons. Diets within feeding patches overlapped during the wet season but highly diverged during the dry season. Body mass differences among species explained their dry season resource partitioning for all species except for comparisons with the megaherbivore (white rhino), while differences in digestive strategy were not related to niche overlap in either season. We conclude that savanna herbivores in this system coexist mostly through body size‐driven resource partitioning in the dry‐season, with the exception of the white rhino (megaherbivore).     

Intraspecific difference among herbivore lineages and their host‐plant specialization drive the strength of trophic cascades

Trophic cascades – the indirect effect of predators on non‐adjacent lower trophic levels – are important drivers of the structure and dynamics of ecological communities. However, the influence of intraspecific trait variation on the strength of trophic cascade remains largely unexplored, which limits our understanding of the mechanisms underlying ecological networks. Here we experimentally investigated how intraspecific difference among herbivore lineages specialized on different host plants influences trophic cascade strength in a terrestrial tri‐trophic system. We found that the occurrence and strength of the trophic cascade are strongly influenced by herbivores’ lineage and host‐plant specialization but are not associated with density‐dependent effects mediated by the growth rate of herbivore populations. Our findings stress the importance of intraspecific heterogeneities and evolutionary specialization as drivers of trophic cascade strength and underline that intraspecific variation should not be overlooked to decipher the joint influence of evolutionary and ecological factors on the functioning of multi‐trophic interactions.     

Pyric Herbivory and the Nexus Between Forage,Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

     

Responses of belowground communities to large aboveground herbivores: Meta‐analysis reveals biome‐dependent patterns and critical research gaps

The importance of herbivore–plant and soil biota–plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta‐analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root‐feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.     

Legacy of avian-dominated plant?herbivore systems in New Zealand

Avian herbivores dominated New Zealand?s pre-settlement terrestrial ecosystems to an unparalleled extent, in the absence of a terrestrial mammal fauna. Approximately 50% (88 taxa) of terrestrial bird species consumed plant foliage, shoots, buds and flowers to some degree, but fewer than half these species were major herbivores. Moa (Dinornithiformes) represent the greatest autochthonous radiation of avian herbivores in New Zealand. They were the largest browsers and grazers within both forest and scrubland ecosystems. Diverse waterfowl (Anatidae) and rail (Rallidae) faunas occupied forests, wetlands and grasslands. Parrots (Psittacidae) and wattlebirds (Callaeidae) occupied a range of woody vegetation types, feeding on fruits/seeds and foliage/ fruits/nectar, respectively. Other important herbivores were the kereru (Columbidae), stitchbird (Notiomystidae) and two honeyeaters (Meliphagidae). Cryptic colouration, nocturnal foraging and fossil evidence suggest that avian populations were strongly constrained by predation. With the absence of migratory avian herbivores, plant structural, constitutive defences prevailed, with the unusual ?wire syndrome? representing an adaptation to limit plant offtake by major terrestrial avian browsers. Inducible plant defences are rare, perhaps reflecting longstanding nutrient-limitations in New Zealand ecosystems. Evidence from coprolites suggests moa were important dispersers of now rare, annual, disturbance-tolerant herb species, and their grazing may have maintained diverse prostrate herbs in different vegetation types. The impact of moa on forest structure and composition remains speculative, but many broadleaved woody species would likely have experienced markedly reduced niches in pre-settlement time. Several distinctive avian-mediated vegetation types are proposed: dryland woodlands, diverse turf swards, coastal herb-rich low-forest-scrubland, and conifer-rich forests. Since human settlement (c. 750 yrs ago), c. 50% of endemic avian herbivore species or c. 40% overall have become extinct, including all moa, 60% of waterfowl and 33% of rail species. Numerically, avian herbivore introductions (c. 24 taxa) since European settlement have compensated for extinctions (c. 27 taxa), but the naturalised birds are mostly small, seed-eating species restricted to human-modified landscapes. Several naturalised species (e.g. Canada goose, Branta canadensis; brown quail, Coturnix ypsilophorus) may provide modes and levels of herbivory comparable with extinct species. The original avian and current introduced mammal herbivore regimes were separated by several centuries when New Zealand lacked megaherbivores. This ?herbivory hiatus? complicates comparisons between pre-settlement and current herbivore systems in New Zealand. However, predation, animal mobility, feeding mode, nutrient transfer patterns and soil impacts were different under an avian regime compared with current mammalian herbivore systems. Levels of ecological surrogacy between avifauna and introduced mammals are less evident. Ungulates generally appear to have impacts qualitatively different from those of the extinct moa. Because of New Zealand?s peculiar evolutionary history, avian herbivores will generally favour the persistence of indigenous vegetation, while mammalian herbivores continue to induce population declines of select plant species, change vegetation regeneration patterns, and generally favour the spread and consolidation of introduced plant species with which they share an evolutionary history.     

Evolution of nutrient uptake reveals a trade-off in the ecological stoichiometry of plant-herbivore interactions

Nutrient limitation determines the primary production and species composition of many ecosystems. Here we apply an adaptive dynamics approach to investigate evolution of the ecological stoichiometry of primary producers and its implications for plant-herbivore interactions. The model predicts a trade-off between the competitive ability and grazing susceptibility of primary producers, driven by changes in their nutrient uptake rates. High nutrient uptake rates enhance the competitiveness of primary producers but also increase their nutritional quality for herbivores. This trade-off enables coexistence of nutrient exploiters and grazing avoiders. If herbivores are not selective, evolution favors runaway selection toward high nutrient uptake rates of the primary producers. However, if herbivores select nutritious food, the model predicts an evolutionarily stable strategy with lower nutrient uptake rates. When the model is parameterized for phytoplankton and zooplankton, the evolutionary dynamics result in plant-herbivore oscillations at ecological timescales, especially in environments with high nutrient availability and low selectivity of the herbivores. High herbivore selectivity stabilizes the community dynamics. These model predictions show that evolution permits nonequilibrium dynamics in plant-herbivore communities and shed new light on the evolutionary forces that shape the ecological stoichiometry of primary producers.     

Modeling herbivore competition mediated by inducible changes in plant quality

Competition between herbivorous insects often occurs as a trait mediated indirect effect mediated by inducible changes in plant quality rather than a direct effect mediated by plant biomass. While plant-mediated competition likely influences many herbivores, progress linking studies of plant-mediated competition in terrestrial phytophagous insects to longer-term consequences for herbivore communities has been elusive, and there is little relevant theory to guide this effort. We present simple models describing plant-mediated interactions between two herbivorous insects or other functionally equivalent organisms. These models consider general features of plant-mediated competition including specificity of elicitation by and effects on herbivores, positive and negative interactions among herbivores, competition independent of changes in plant biomass, and the existence of multiple relevant plant traits. Our analyses generate four important conclusions. First, herbivores competing strongly via only one plant quality phenotype exhibit a limited range of outcomes. These include coexistence and competitive exclusion of either herbivore, but do not include initial condition dependence. Second, when the outcome of competition is competitive exclusion, the herbivore that persists is the one that can do so under the highest inducible reductions in plant quality. Third, competition via more than one inducible phenotype can exhibit a wider range of outcomes including multiple equilibria and initial condition dependence. Finally, transient dynamics may not predict the eventual outcome of competition when changes in plant quality are slow relative to herbivore population growth, especially when herbivores compete through multiple phenotypes. We interpret our results in terms of competition outcomes reported in the literature, and suggest directions for the future empirical study of herbivore competition mediated by inducible changes in plant quality.     

A conserved pattern in plant‐mediated interactions between herbivores

Plant‐mediated interactions between herbivores are important determinants of community structure and plant performance in natural and agricultural systems. Current research suggests that the outcome of the interactions is determined by herbivore and plant identity, which may result in stochastic patterns that impede adaptive evolution and agricultural exploitation. However, few studies have systemically investigated specificity versus general patterns in a given plant system by varying the identity of all involved players. We investigated the influence of herbivore identity and plant genotype on the interaction between leaf‐chewing and root‐feeding herbivores in maize using a partial factorial design. We assessed the influence of leaf induction by oral secretions of six different chewing herbivores on the response of nine different maize genotypes and three different root feeders. Contrary to our expectations, we found a highly conserved pattern across all three dimensions of specificity: The majority of leaf herbivores elicited a negative behavioral response from the different root feeders in the large majority of tested plant genotypes. No facilitation was observed in any of the treatment combinations. However, the oral secretions of one leaf feeder and the responses of two maize genotypes did not elicit a response from a root‐feeding herbivore. Together, these results suggest that plant‐mediated interactions in the investigated system follow a general pattern, but that a degree of specificity is nevertheless present. Our study shows that within a given plant species, plant‐mediated interactions between herbivores of the same feeding guild can be stable. This stability opens up the possibility of adaptations by associated organisms and suggests that plant‐mediated interactions may contribute more strongly to evolutionary dynamics in terrestrial (agro)ecosystems than previously assumed.     

Review: Feeding conserved forage to horses: recent advances and recommendations

The horse is a non-ruminant herbivore adapted to eating plant-fibre or forage-based diets. Some horses are stabled for most or the majority of the day with limited or no access to fresh pasture and are fed preserved forage typically as hay or haylage and sometimes silage. This raises questions with respect to the quality and suitability of these preserved forages (considering production, nutritional content, digestibility as well as hygiene) and required quantities. Especially for performance horses, forage is often replaced with energy dense feedstuffs which can result in a reduction in the proportion of the diet that is forage based. This may adversely affect the health, welfare, behaviour and even performance of the horse. In the past 20 years a large body of research work has contributed to a better and deeper understanding of equine forage needs and the physiological and behavioural consequences if these are not met. Recent nutrient requirement systems have incorporated some, but not all, of this new knowledge into their recommendations. This review paper amalgamates recommendations based on the latest understanding in forage feeding for horses, defining forage types and preservation methods, hygienic quality, feed intake behaviour, typical nutrient composition, digestion and digestibility as well as health and performance implications. Based on this, consensual applied recommendations for feeding preserved forages are provided.     

Environmental gradients and herbivore feeding preferences in coastal salt marshes

Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.     

Phylogenetic and geographic variation in host breadth and composition by herbivorous amphipods in the family Ampithoidae

Predicting the host range for herbivores has been a major aim of research into plant-herbivore interactions and an important model system for understanding the evolution of feeding specialization. Among many terrestrial insects, host range is strongly affected by herbivore phylogeny and long historical associations between particular herbivore and plant taxa. For small herbivores in marine environments, it is known that the evolution of host use is sculpted by several ecological factors (e.g., food quality, value as a refuge from predators, and abiotic forces), but the potential for phylogenetic constraints on host use remains largely unexplored. Here, we analyze reports of host use of herbivorous amphipods from the family Ampithoidae (102 amphipod species from 12 genera) to test the hypotheses that host breadth and composition vary among herbivore lineages, and to quantify the extent to which nonpolar secondary metabolites mediate these patterns. The family as a whole, and most individual species, are found on a wide variety of macroalgae and seagrasses. Despite this polyphagous host use, amphipod genera consistently differed in host range and composition. As an example, the genus Peramphithoe rarely use available macrophytes in the order Dictyotales (e.g., Dictyota) and as a consequence, display a more restricted host range than do other genera (e.g., Ampithoe, Cymadusa, or Exampithoe). The strong phylogenetic effect on host use was independent of the uneven distribution of host taxa among geographic regions. Algae that produced nonpolar secondary metabolites were colonized by higher numbers of amphipod species relative to chemically poor genera, consistent with the notion that secondary metabolites do not provide algae an escape from amphipod herbivory. In contrast to patterns described for some groups of phytophagous insects, marine amphipods that use chemically rich algae tended to have broader, not narrower, host ranges. This result suggests that an evolutionary advantage to metabolite tolerance in marine amphipods may be that it increases the availability of appropriate algal hosts (i.e., enlarges the resource base).