Selective Herbivory Research Articles

A taste of space: Remote animal observations and discrete-choice models provide new insights into foraging and density dynamics for a large subarctic herbivore.

Competition for resources and space can drive forage selection of large herbivores from the bite through the landscape scale. Animal behaviour and foraging patterns are also influenced by abiotic and biotic factors. Fine-scale mechanisms of density-dependent foraging at the bite scale are likely consistent with density-dependent behavioural patterns observed at broader scales, but few studies have directly tested this assertion. Here, we tested if space use intensity, a proxy of spatiotemporal density, affects foraging mechanisms at fine spatial scales similarly to density-dependent effects observed at broader scales in caribou. We specifically assessed how behavioural choices are affected by space use intensity and environmental processes using behavioural state and forage selection data from caribou (Rangifer tarandus granti) observed from GPS video-camera collars using a multivariate discrete-choice modelling framework. We found that the probability of eating shrubs increased with increasing caribou space use intensity and cover of Salix spp. shrubs, whereas the probability of eating lichen decreased. Insects also affected fine-scale foraging behaviour by reducing the overall probability of eating. Strong eastward winds mitigated negative effects of insects and resulted in higher probabilities of eating lichen. At last, caribou exhibited foraging functional responses wherein their probability of selecting each food type increased as the availability (% cover) of that food increased. Space use intensity signals of fine-scale foraging were consistent with density-dependent responses observed at larger scales and with recent evidence suggesting declining reproductive rates in the same caribou population. Our results highlight potential risks of overgrazing on sensitive forage species such as lichen. Remote investigation of the functional responses of foraging behaviours provides exciting future applications where spatial models can identify high-quality habitats for conservation.

Herbivores can benefit both plants and their pathogens through selective herbivory on diseased tissue

Abstract Infectious disease can shape community structure, particularly when pathogens affect foundation species. Seagrasses are foundation species that form meadows along coasts worldwide, controlling sediment deposition and biogeochemical cycling while supporting a diverse community of fish and invertebrates. These plants are hosts to wasting diseases that cause necrotic tissue lesions, which could alter seagrass value as food, habitat and mediators of ecosystem processes. However, such biotic interactions and influences on disease dynamics are still not well understood. We investigated whether a common herbivore affected the development of wasting disease on eelgrass. We measured the severity and prevalence of eelgrass wasting disease in a meadow across two summers through repeated field surveys. We assessed the role of the eelgrass herbivore on disease spread and growth using microcosm and mesocosm experiments. We further explored herbivore feeding preferences in a choice trial, which was paired with chemical analysis of plant tissue and analysed using a structural equation model. While the herbivore facilitates the growth of new disease lesions among isolated leaves, on balance they reduce lesion severity by more than 50% in comparison with no‐herbivore controls in field‐realistic settings. This was likely because the herbivore strongly prefers to eat diseased rather than healthy tissue, consuming nearly twice as much lesion area in choice trials. This preference arises from pathogen‐driven changes in the host plant; lesioned tissue requires less force to penetrate than non‐lesioned tissue. Additionally, as lesions increase in size, their polyphenolic concentrations drop, which further increases the magnitude of preference for lesioned tissue. Synthesis: These results suggest that these herbivores could help maintain disease in this system at a high prevalence (by facilitating disease development) but low severity (through preferential consumption), which is consistent with our field observations of nearly 100% prevalence and low severity in a natural bed where herbivore density is high. Describing such multi‐species interactions in marine systems will advance our predictions of future disease states, as current understandings focus primarily on how environmental change contributes to pathogen outbreaks.

It takes two to Tango: Plant height and nutrient level determine the diet selection of wintering geese in Poyang Lake, a Ramsar wetland

In Poyang Lake, the largest and one of the most important wintering sites in the East Asian-Australasian Flyway, Carex (Carex cinerascens Kük) meadows provide the primary food source for the wintering geese. However, due to intensified river regulation and more frequent extreme climatic events such as drought, observational evidence suggests that the synchrony of geese migration and Carex phenology could not be maintained without human interventions, imposing a great risk of food shortage during the wintering period. Consequently, the current conservation priority in this Ramsar site is shifted to wet meadow improvement to ensure optimal food quality. Understanding the food preferences of wintering geese is the key for effective wet meadow management. As the growth stage and nutrient level of food plants are the decisive factors influencing the diet selection of herbivores, in this study, we sampled the preferred food items by tracking the foraging paths of the Greater White-fronted Goose (n = 84) and Bean Goose (n = 34) to quantify the “foraging window” in terms of plant height,protein level, and energy content. Further, we established relationships between the above three variables of Carex based on in-situ measurements. The results show that the geese prefer plants with height ranging from 2.4 to 25.0 cm, with protein content from 13.9 to 25.2 %, and energy content from 1440.0 to 1813.6 KJ/100 g. While plant energy content increases with height, the height-protein level relationship is negative. The opposite growth curves signify a conservation challenge to maintain the delicate balance between the quantity and quality requirements of wintering geese. Carex meadow management, such as mowing, should focus on optimizing the timing of action to maximize energy supply while maintaining the right protein level for the long-term fitness, reproduction and survival of the birds.

Native herbivore browsing alters plant physical and chemical traits in a eucalypt forest understory

AbstractMacropods exhibit selective herbivory on plant traits within and between species and hence can alter plant trait variation. In this study, we asked what are the effects of macropod herbivory on intraspecific variation in plant traits of forest understory species? We tested the effects of herbivory by macropods on eight plant traits, in nine understory species using a herbivore exclusion experiment. Generalized linear mixed models were used to test for differences between herbivory treatments in trait means as well as variation within species and at the community level. Macropod herbivory resulted in differences in either the mean trait value or trait variance within a species for seven of the eight tested plant traits. Macropod exclusion resulted in larger dimensions of size for most species, as well as changes in specific leaf area, moisture content, and carbon content of leaves. Further, community‐weighted means of specific leaf area, moisture content, and leaf nitrogen were lower in the presence of macropods. Plant trait variance was altered for three of the seven community‐weighted traits. This study demonstrates a mechanism through which macropods can drive changes in the traits of understory plant species. Plant traits have been linked to various larger scale ecological processes including ecosystem flammability, nutrient cycling, response to global change, and habitat suitability for other organisms. Therefore, herbivore‐induced changes in plant trait variation and community composition could have important flow‐on effects for other environmental processes and hence should be considered in management decisions regarding fire‐prone ecosystems.

Favorite Parts of a Single Leaf for Giant Flying Squirrels to Eat in Three Species of Food Trees.

To examine the effect of leaf chemical composition on selective herbivory by the Japanese giant flying squirrels (Petaurista leucogenys), we measured and compared the total phenolic, glucose, and water contents of leaves among their main food tree species, deciduous Quercus acutissima, and evergreen Q. sessilifolia and Phonitia serratifolia. Leaves of these three tree species were available in the warm season (April to October), but the flying squirrels mostly preferred the leaves of Q. acutissima, having higher glucose and water contents than those of the other two tree species. In the cold season (November to the next March), the two evergreen tree species were available, and the flying squirrels used both leaves without any apparent influence of the chemical compositions. On the other hand, the favorite parts of a single leaf differed among the three tree species. Flying squirrels dropped the individual leaves after partial consumption. Their feeding marks on the dropped leaves were distinguished into four types: apical, basal, central, and marginal parts of consumption. The basal parts of consumption were most frequent in Q. acutissima leaves in which more water was contained at the basal part, and the central part consumption followed, which may be related to a lower phenolic content and more glucose and water at the leaf center than its margin. In contrast, the apically consumed leaves dominated in Q. sessilifolia, with relatively homogeneous leaf chemical distribution except for more water at the center. In P. serratifolia, leaves consumed at the center were frequent, but those with marginal consumption were also observed, which may be related to its specific chemical distribution with less phenolics and more glucose at the leaf margin. Thus, the chemical distributions within the single leaf differ among tree species, and the flying squirrel's selectivity of the tree species and the part of each leaf depends partly on the relative compositions of preferable glucose and water and unpreferable phenolics.

Rotational grazing with cattle‐free zones supports the coexistence of cattle and wild herbivores in African rangelands

Abstract African wildlife populations are declining at an alarming rate. To stop further population declines and restore ecosystems, more areas for wildlife are needed. Community‐based conservation with wildlife‐livestock coexistence in the vast rangelands of Africa presents a major opportunity. However, the efficacy of wildlife conservation in mixed land‐use areas remains an outstanding question. To assess the ecological outcomes of land‐sharing between regulated livestock herds and wildlife populations in African savannas, we test how rotational cattle grazing affects spatiotemporal dynamics of 15 large herbivore species in the Maasai Mara, Kenya. First, we tested how wild herbivore distributions across the Greater Mara Ecosystem (the Mara, ~2600 km2) are related to cattle density and environmental variables using 584,561 observations of wild herbivores (ecosystem scale). In a second analysis, we tested how rotational cattle grazing affects wild herbivore distributions in a 300 km2 subsection of the Mara using 30,583 observations (landscape scale). Finally, we tested how functional traits of wild herbivores affect species‐level spatiotemporal responses to cattle grazing. At the ecosystem scale, the presence of five wild herbivore species was positively correlated with cattle density, while cattle effects on wild herbivore abundances were species‐dependent with both increases and decreases. At the landscape scale, rotational cattle grazing strongly impacted the spatiotemporal habitat selection of wild herbivores, resulting in distinct lag periods with which different species are attracted to areas previously grazed by cattle. These lag periods were linked to functional traits, with body mass and herd size explaining 35% of the interspecific differences. Small to medium‐sized herbivores with large herds select areas recently grazed by cattle, whereas large species with large herd sizes and small species with small herd sizes avoid recently grazed areas. Synthesis and applications: Our results revealed that the effect of cattle on wild herbivores varies considerably among species, suggesting that cattle‐wildlife interactions range from facilitation to competition. To maintain species that strongly avoid cattle, designated livestock‐free zones remain essential, also in rotational grazing systems. Rotational grazing systems with regulated livestock densities present an important opportunity to better manage wildlife‐livestock coexistence and thus improve wildlife conservation in African rangelands.

Differential accumulation of cardenolides from Asclepias curassavica by large milkweed bugs does not correspond to availability in seeds or biological activity on the bug Na+/K+-ATPase

Milkweed–herbivore systems are characterized by cardenolide chemical defenses and specialized herbivore adaptations such as physiological target site insensitivity. Cardenolide defenses in milkweeds can vary in terms of the total concentration, differences in the polarity of individual cardenolides, and the substitution of the steroidal structures that can contribute to the molecule's reactivity. The variability in cardenolide defenses could represent the plant's response to natural selection and adaptation of resistant herbivores and is a characteristic of phenotype-matching between defensive and offensive traits resulting from coevolution. Here, we test the phenotypic match of the cardenolide composition of seeds of Asclepias curassavica and those sequestered by nymphs and adults of the specialized seed herbivore Oncopeltus fasciatus, combined with tests of the inhibitory capacity of a subset of seed cardenolides against the Na+/K+-ATPase of O. fasciatus and a non-adapted insect (Drosophila melanogaster). We compare this with the inhibitory capacity against the highly sensitive porcine Na+/K+-ATPase. Among the five most abundant cardenolides present in milkweed seeds, glucopyranosyl frugoside, glucopyranosyl gofruside, and glucopyranosyl calotropin were significantly more abundant in the seeds than in the adults and nymphs; the bugs contained higher concentrations of the deglucosylated compounds. The most abundant compound, glucopyranosyl frugoside, was also the most inhibitory for O. fasciatus, but O. fasciatus was significantly more tolerant to all compounds compared to D. melanogaster and the highly sensitive porcine enzyme. Our results add to the evidence that O. fasciatus sequesters specific individual cardenolides from its Asclepias host plants that are not directly linked to the concentration and inhibitory potency.

Functional adaptive resources for large herbivores in African savannas: an ecological-gradient based framework

The viability of large herbivore populations in the face of climate change, environmental variability, disease and predation will be determined by their freedom to assess and respond to these factors through access to a range of functional seasonal resources and habitats. Their responses will be contingent upon various organismal traits, such as body size, mouth and digestive anatomy, which also facilitate coexistence among sympatric species. In this paper we develop a functional adaptive forage resource framework based on plant biomass, quality and phenology responses on ecological productivity gradients. We show how large herbivores coexist and respond to environmental variability, disease and predation by their foraging responses in relation to functional adaptive resources, as mediated by their anatomical traits. Below a critical body size, where predation limits population size, large herbivores adopt a variety of predation avoidance strategies, which are linked to their anatomical traits and foraging strategies. Mouth anatomy, and its interaction with body size, appears to be the major anatomical trait determining large herbivore selection for grass height. Body size is the major trait determining vulnerability to predators. Ecological productivity gradients underlain by variation in soil moisture availability over the annual cycle support high quality forage in the least productive (driest) regions, which promote growth and reproduction (a fecundity resource). Reserves of adequate quality forage in moderate productivity regions and buffers of low-quality forage in wetter and more productive regions of the gradient prevent loss of body stores over the dry season and starvation during droughts. Fire and grazing contribute towards providing high quality forage by removing old, low-quality material and preventing forage maturation. Consuming a high diversity of plant species distributed across ecological gradients promotes phytochemical diversity in the diet, which functions as medicinal resources to promote health while combating disease and parasites. Large herbivores are becoming increasingly restricted by ecosystem fragmentation in their access to the full range of these functional resource classes. The negative consequences for large herbivore populations of reduced access to these resource classes is compounded by climate change, where conditions are hotter and drought frequency and intensity is expected to be higher.

Individual‐plant selectivity by sheep in drylands scales‐up at plant population level and controls the forage supply and its accessibility

Abstract Diet selectivity by domestic herbivores controls plant community structure and dynamics and may induce rangeland degradation, particularly in drylands. However, management decisions frequently ignore herbivore selectivity. Here, we studied how grass morphology controls sheep selectivity for individual plants, and how this selectivity interacts with grazing intensity to determine population plant‐size distributions and the forage supply. In Patagonian steppes, we manipulated the plant morphology (size and standing‐dead proportion) of three dominant grass species differentially preferred by sheep for 4 years. Then, we evaluated how these morphological alterations affected intra‐ and inter‐specific preference patterns. We also evaluated how grazing intensity (ungrazed, moderate grazing and intensive grazing) affected the plant‐size distribution of the three species, the forage supply, and its accessibility. For the highly preferred species, herbivores selected plants that were either naturally or experimentally short, with low proportion of standing‐dead biomass. In contrast, morphological changes did not alter the within‐species selectivity of the least preferred species. Grazing intensity strongly changed the population plant‐size distribution of preferred species in ways that resembled the experimental manipulations of morphology. Moderate grazing showed the greatest morphological heterogeneity among individuals. When integrating the green biomass of forage species' individuals at ecosystem level, we found that the forage supply was the highest in ungrazed sites and decreased as grazing intensity increased. However, considering the dissuading effect of the standing‐dead proportion of plants, the accessible forage was the highest under moderate grazing. Synthesis and applications. Our findings (i) showed that, within preferred species, sheep selectivity at individual‐plant level is controlled by morphological characteristics that determine accessibility to green high‐quality biomass. This effect was as important as that of species identity; (ii) empirically proved plant–animal positive feedback at individual level; and (iii) revealed how the individual‐plant selectivity scales‐up at population level and controls the forage supply, but also its accessibility. Our complementary approach generates critical knowledge for developing management practices to control key forage species defoliation and to adjust the grazing pressure to the offer of accessible forage, avoiding the common carrying capacity overestimation. These aspects are essential for sustainable production in grazed drylands.

Interannual variations in diet and nutritional value of plants consumed by the Souss Valley tortoises, Testudo graeca graeca, in west-central Morocco: are they really selective foragers?

Abstract A previous study on the feeding habits of Moorish tortoises in west-central Morocco suggests that these animals are selective herbivores, but the nutritional consequences have not been examined. Because of the potassium excretion load, which requires water and/or nitrogen loss, we predicted that tortoises do not have salt glands. Moorish tortoises prefer plants rich in water and protein but low in potassium (PEP index). To this end, we studied the spring diet of adult tortoises in an arid steppe in west-central Morocco during two seasons of relatively dry years (2011-2012) using feces analyses. We also estimated the relative abundance of potential food plants by stratified sampling under the canopy of jujube bushes Ziziphus lotus. We statistically compared diet to plant abundance. Finally, we assessed plant species’ nutritional composition (water, crude protein, and potassium) available to tortoises. Results showed that species assemblages differed significantly between the two plant communities in both years. Nevertheless, tortoises consumed only about 5-6 and nearly the same species at the study site in 2011 and 2012, respectively. The plants consumed by the tortoises had the highest positive PEP index values indicating that there was more water and nitrogen in the food than is needed to excrete potassium.

Coexistence of grasses and shrubs in Patagonian steppes. Norm or exception?

AbstractQuestions(1) Is the co‐dominance of grasses and shrubs a general feature of Patagonian steppes? (2) At a local scale, does it change with domestic grazing intensity or decadal changes in precipitation?LocationsQuestion 1. Patagonia region in southern South America. Question 2. Shrub–grass steppes (45°24′S, 70°17′W) subjected to different grazing intensity for decades.MethodsWe sampled vegetation to estimate perennial grass and shrub cover at two scales: regional and local. At the regional (phytogeographical) scale, we sampled grazed steppes under different climates and soils (n = 324 sites over ca. 487,000 km2). At the local scale, we sampled paddocks subjected to known grazing management, from commercial intensive densities of sheep to total (experimental) exclusion. Also, in the same site, we sampled permanent plots (25 years old) in a grazing exclosure to study the effects of precipitation variation.ResultsMixed communities are the predominant form of community organization in the Patagonia region. At the regional scale, shrub cover was negatively related to grass cover and communities where shrubs had higher cover than grasses represent 54% of the data set. At the local scale for an arid steppe, perennial grasses had higher cover than shrubs, regardless of grazing pressure. The shrub‐to‐grass ratio was increased only by intensive grazing. Permanent plots in a grazing‐excluded paddock indicate that decadal xeric or humid climate periods decrease or increase total cover, respectively. Surprisingly, the relative shrub‐to‐grass ratio did not change significantly.ConclusionsAlthough perennial grasses and shrubs cover are negatively correlated, they coexist in the Patagonian steppes under a wide range of climate, soil and grazing conditions. At a local scale, selective herbivory promotes species replacement but did not substantially change the life‐form covers. Water shortages, as well as periods of water surplus, similarly affect both life forms without compromising their coexistence.

Changes in spinescence across leaf ontogeny support the optimal defence hypothesis in blackberries (Rubus adenotrichos)

Abstract Hypotheses based on allocation theory and herbivore selection offer opposite predictions about how defence levels against herbivores change as the plant tissue grows. The growth differentiation balance hypothesis (GDBH) assumes that defences will be resource-limited in immature tissues and predict that defence levels increase as the plant tissue grows. Conversely, the optimal defence hypothesis (ODH) proposes that plants would have the highest level of defences in the parts that have the highest value in terms of fitness and/or are more frequently attacked by herbivores, such as young tissues. We examine whether spinescence in the shrub Rubus adenotrichos (blackberry) change as the leaf grows, and if this change is consistent with the GDBH or the ODH. We compare the petiole area occupied by prickles, the prickles density and the individual prickle area in mature versus young petioles from Rubus adenotrichos. Our results show that, in R. adenotrichos, young tissues are more protected than mature tissues. Prickles density and the petiole area occupied by prickles were up to 25% higher in young petioles than in mature ones. These results support the ODH, reinforcing the idea that extrinsic factors such as herbivores pressure might drive the change of structural defences level across leaf ontogeny.

Effects of Inter- and Intra-Specific Interactions on Moose Habitat Selection Limited by Temperature

Habitat selection and daily activity patterns of large herbivores might be affected by inter- and intra-specific interaction, changes of spatial scale, and seasonal temperature. To reveal what factors were driving the habitat selection of moose, we collected moose (Alces alces) and roe deer (Capreolus pygargus bedfordi) occurrence data, analyzed the multi-scale habitat selection and daily activity patterns of moose, and quantified the effects of spatial heterogeneity distribution of temperature, as well as the occurrence of roe deer on these habitat selection processes. Our results suggested that moose and roe deer distribution spatially overlap and that moose habitat selection is especially sensitive to landscape variables at large scales. We also found that the activity patterns of both sexes of moose had a degree of temporal separation with roe deer. In the snow-free season, temperatures drove moose habitat selection to be limited by threshold temperatures of 17 °C; in the snowy season, there were no similar temperature driving patterns, due to the severe cold environment. The daily activity patterns of moose showed seasonal change, and were more active at dawn and nightfall to avoid heat pressure during the snow-free season, but more active in the daytime for cold adaptation to the snow season. Consequently, this study provides new insights on how the comprehensive effects of environmental change and inter- and intra- specific relationships influence the habitat selection and daily activity patterns of moose and other heat sensitive animals with global warming.

An underwater Serengeti: Seagrass‐mediated effects on intake and cultivation grazing behavior of a marine megaherbivore

AbstractPopulations of green turtles (Chelonia mydas), a megaherbivore that consumes seagrasses via cultivation grazing, are recovering worldwide. Information on plant‐mediated effects on herbivore foraging behavior is critical to understanding plant–herbivore interactions and sustainability of grazing as ecosystems continue to change. In a Caribbean seagrass ecosystem, we use stationary cameras and benthic surveys to evaluate the effects of seagrass morphology and leaf nitrogen content on green turtle grazing behavior.Thalassia testudinumleaf morphology has significant effects on forage intake (in milligrams of dry mass [DM] per minute) for green turtles, whereas leaf nitrogen content has no effect. Intake increases in grazed areas with shorter leaves and higher leaf biomass concentration (in milligrams of DM per cubic centimeter), indicating more efficient foraging under these conditions. Bite rate (in bites per minute) increases in grazed areas with short leaves, a result of reduced search time. Bite size (in milligrams of DM per bite) increases in grazed areas with short but dense canopies, because a turtle crops more shoots with each bite. Increased foraging efficiency and reduced search time in grazed areas with high biomass concentrations collectively maximize intake. Ingested leaves are shorter than the mean height of all available leaves in grazed areas, indicating herbivore selection for shorter leaves. Our estimate for daily intake is 86.1 g DM day−133‐kg turtle−1. Our study provides a novel contribution on the effects of plant‐level cues on the grazing behavior of a marine megaherbivore, and how cultivation grazing behavior optimizes the green turtle foraging strategy by maximizing foraging efficiency and intake.

Habitat selection of large herbivores evidenced as threats to alpine ecosystem

Across the globe, increasing populations of Cervidae have the potential to modify ecosystems. Long-term grazing pressure by deer not only alters vegetation but also affects ecosystem functions and interspecific interactions. It is recognized as a significant factor in ecological degradation. Alpine vegetation (i.e., alpine meadow) is an icon of alpine ecosystems and has thrived for centuries without the presence of large herbivores in Japan. However, in the alpine zone (>2700 m a.s.l.) of the South Alps National Park in central Honshu, Japan, sika deer (Cervus nippon) were first seen in the 1990s; thereafter, the vegetation noticeably changed starting in the 2000s. Nonetheless, virtually no information exists on habitat selection by these alpine sika deer. Here we analyzed (1) the altitudinal movement patterns of alpine sika deer, (2) the significant environmental factors affecting their habitat selection, and (3) estimated the relative probability of their habitat use via resource selection functions (RSF). The mean altitudinal difference between summer and winter home ranges was 1186 m, and the alpine sika deer arrived at the alpine ridgeline in June when the snow cover at >2400 m a.s.l. rapidly decreased. The mean altitude in the summer home range was 2635 ± 135 m. Alpine meadow, followed by Betula forests and tall grasslands, were preferred habitats by deer. The deer also preferred areas where snow melted in late spring and early summer. The deer's core habitat, indicated by higher RSF values (≥0.65), covered 45.2% of the study area, mostly distributed in the alpine zone. Our estimates clearly suggest the urgent need to prioritize specific areas for the conservation of the alpine vegetation in the South Alps National Park in Japan.

Soil cadmium stress reduced host plant odor selection and oviposition preference of leaf herbivores through the changes in leaf volatile emissions

The Elemental defense hypothesis suggested that metal accumulation in plant tissues could serve as direct defense to reduce herbivore feeding preference as metals are toxic to phytophagous insects. However, the indirectly defensive role of heavy metals on host plant odor selection and oviposition preference of leaf herbivores through the changes in leaf volatile organic compounds (VOCs) is still unknown. In this study, we used a local woody plant species, Populus yunnanensis, to investigate whether soil cadmium (Cd) stress could affect plant VOC production and whether Cd-mediated changes in leaf VOC emissions will further influence the host plant odor and oviposition preferences of female adults of a specialist and a generalist herbivore species. The results clearly showed that the soil Cd stress could prominently induce leaf total VOC emissions of P. yunnanensis and such induction was positively correlated with leaf Cd accumulation. Herbivore olfactometer bioassays further demonstrated that the VOCs released by P. yunnanensis under Cd exposure are far less attractive to both of the specialist and generalist female adults compared to control plants, leading to significant reduction in oviposition on Cd-treated plants. Moreover, the host plant odor selection and oviposition preference of the two herbivore species were all negatively correlated with leaf total VOC emissions, which confirmed the defensive role of Cd-induced VOCs for deterring the female insects. The result will extend the existing knowledge of the Elemental defense hypothesis and provide new insight into predicting the herbivore damage level of poplar species that naturally occurred in metal-polluted habitats.

Holling meets habitat selection: functional response of large herbivores revisited

BackgroundHolling (Can Entomol 91(5):293–320, 1959) was the first to describe a functional response between a predator’s consumption-rate and the density of its prey. The same concept can be applied to the habitat selection of herbivores, specifically, the change in relative habitat use with the change in habitat availability. Functional responses in habitat selection at a home-range scale have been reported for several large herbivores. However, a link to Holling’s original functional response types has never been drawn, although it could replace the current phenomenological view with a more mechanistically based understanding of functional responses.MethodsIn this study, discrete choice models were implemented as mixed-effects baseline-category logit models to analyze the variation in habitat selection of a large herbivore at seasonal and diurnal scales. Thus, changes in the use of land cover types with respect to their availability were investigated by monitoring 11 land cover types commonly used by roe deer (Capreolus capreolus) in the Bavarian Forest National Park, Germany. Functional response curves were then fitted using Holling’s formulas.ResultsStrong evidence of non-linear functional responses was obtained for almost all of the examined land cover types. The shape of the functional response curves varied depending on the season, the time of day, and in some cases between sexes. These responses could be referenced to Holling’s types, with a predominance of type II.ConclusionsOur results indicate that Holling’s types can be applied to describe general patterns of the habitat selection behavior of herbivores. Functional responses in habitat selection may occur in situations requiring a trade-off in the selection of land cover types offering different resources, such as due to the temporally varying physiological needs of herbivores. Moreover, two associated parameters defining the curves (prey density and predation rate) can aid in the identification of temporal variations and in determinations of the strength of the cost-benefit ratio for a specific land cover type. Application of our novel approach, using Holling’s equations to describe functional responses in the habitat selection of herbivores, will allow the assignment of general land cover attraction values, independent of availability, thus facilitating the identification of suitable habitats.

N deposition may accelerate grassland degradation succession from grasses- and sedges-dominated into forbs-dominated in overgrazed alpine grassland systems on Qinghai-Tibetan Plateau

Alpine grasslands are sensitive to grazing and atmospheric nitrogen (N) deposition. Despite increases in N deposition, few field studies have assessed the effects of grazing in conjunction with increased N deposition on alpine grassland vegetation.In this two-year field study, we examined the effects of overgrazing and overgrazing plus N deposition on eco-physiological processes of alpine grassland plants at the functional group level. We found that both overgrazing and overgrazing plus N deposition altered species composition and the dominance of three plant functional groups (grasses, sedges, and forbs) in an alpine meadow and alpine steppe.In the overgrazing and overgrazing plus N deposition treatments, forbs dominated the plant community. Grass and sedge dominance decreased substantially, relative to a ungrazed treatment. The underlying eco-physiological processes that led to the forb-dominated plant community differed between the overgrazing and overgrazing plus N deposition treatments. Overgrazing plus N deposition increased forb dominance both directly by selective herbivory and indirectly by enhancing forb photosynthetic rates.Our results suggest that overgrazing concomitant with increases in N deposition will likely shift the plant community composition of alpine grasslands on the Qinghai-Tibetan Plateau from grass/sedge communities to forb-dominated communities in the future.

Plant Volatiles Modulate Seasonal Dynamics between Hosts of the Polyphagous Mirid Bug Apolygus lucorum.

Plant-derived volatiles play a significant role in host selection of phytophagous insects, but their role in seasonal host shifts remain unclear. The polyphagous mirid bug Apolygus lucorum displays marked seasonal host alternation. During summer, volatiles from flowering plants play a key role in A. lucorum foraging. Though A. lucorum adults deposit overwintering eggs on jujube (Ziziphus jujuba) and grape (Vitis vinifera) during autumn, it is unclear whether plant volatiles equally mediate this host selection behavior. During 2015 and 2016, we found that population densities of A. lucorum adults on cotton (Gossypium hirsutum) during August were higher than those in September, whereas the opposite pattern was observed on fruit trees (i.e., jujube and grape). The dispersal factor of the adult population that dispersed from cotton fields during September was higher than in August, whereas opposite patterns were observed in the neighboring jujube/grape orchard. In Y-tube olfactometer trials, A. lucorum adults preferred cotton plant volatiles over fruit tree odors in August, whereas the opposite patterns were found in September. Three electro-physiologically active volatiles (butyl acrylate, butyl propionate and butyl butyrate) were identified from jujube and grape plants. During September, active volatiles are emitted in considerably greater amounts by jujube and grape than in August, while the amount of volatile emissions in cotton decreases in September. Temporal shifts in plant volatile emission thus may modulate host plant foraging of A. lucorum, and appear to guide its colonization of different host plants. Our findings help understand the role of plant volatiles in the host plant selection and seasonal dynamics of polyphagous herbivores.

Indirect Selection on Flower Color in Silene littorea.

Flower color, as other floral traits, may suffer conflicting selective pressures mediated by both mutualists and antagonists. The maintenance of intraspecific flower color variability has been usually explained as a result of direct selection by biotic agents. However, flower color might also be under indirect selection through correlated traits, since correlations among flower traits are frequent. In this study, we aimed to find out how flower color variability is maintained in two nearby populations of Silene littorea that consistently differ in the proportions of white-flowered plants. To do that, we assessed natural selection on floral color and correlated traits by means of phenotypic selection analysis and path analysis. Strong directional selection on floral display and flower production was found in both populations through either male or female fitness. Flower color had a negative indirect effect on the total male and female fitness in Melide population, as plants with lighter corollas produced more flowers. In contrast, in Barra population, plants with darker corollas produced more flowers and have darker calices, which in turn were selected. Our results suggest that the prevalence of white-flowered plants in Melide and pink-flowered plants in Barra is a result of indirect selection through correlated flower traits and not a result of direct selection of either pollinators or herbivores on color.