Seasonality and edge effect determine herbivory risk according to different plant association models

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.     

The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions

Diet selection by mammalian herbivores is often influenced by plant community composition, and numerous studies have focused on the relationships between herbivore foraging decisions and food/plant species abundance. However, few have examined the role of neighbour palatability in affecting foraging of a target plant by large mammalian herbivores. We used a large-scale field dataset on diet selection by red deer Cervus elaphus in Fiordland National Park, New Zealand to: (1) estimate the palatability of native forest plant species to introduced deer from observed patterns of browse damage; and (2) examine whether intraspecific variation in browsing of plants can be related to variation in the local abundance of alternative forage species. Overall, 21 of the 53 forest species in our dataset were never browsed by deer. At a community level, plants were more likely to be browsed if they were in a patch of vegetation of high forage quality, containing high abundances of highly palatable species and/or low abundances of less-palatable species. Our findings suggest that deer make foraging decisions at both a coarse-grain level, selecting vegetation patches within a landscape based on the overall patch quality, and at a fine-grain level by choosing among individual plants of different species.     

Conditional outcomes in plant–herbivore interactions: neighbours matter

Spatial distribution of palatable and unpalatable plants can influence the foraging behaviour of herbivores, thereby changing plant‐damage probabilities. Moreover, the immediate proximity to certain plants can benefit other plants that grow below them, where toxicity or spines act as a physical barrier or concealment against herbivores. This paper presents the results of a multi‐scale experiment performed to test the effect of shrubs as protectors of tree saplings against herbivores and the mechanism involved in Mediterranean ecosystems. We performed a factorial design in two mountain ranges, similar in physiognomy and vegetation, planting saplings of a palatable tree, the maple (Acer opalus subsp. granatense), and an unpalatable tree, the black pine (Pinus nigra), under three different types of shrubs. We considered four experimental microhabitats: highly palatable shrub (Amelanchier ovalis), palatable but spiny shrub (Crataegus monogyna or Prunus ramburii), unpalatable spiny shrub (Berberis vulgaris subsp. australis) and control (gaps of bare soil without shrubs). Three main factors were found to determine the probability of sapling attack: sapling palatability, experimental microhabitat and plot. Palatable saplings (maples) were browsed much more than unpalatable ones (pines). The degree of protection provided by the shrub proved greater as its palatability decreased with respect to sapling palatability, the unpalatable spiny shrub being the safest microhabitat for palatable saplings and bare soil for unpalatable ones. The differences found in number of attacked saplings between plots may be attributable to differences in herbivore pressure. The community context in which interaction takes place, namely the characteristics of the neighbours and the intensity of herbivore pressure, are determining factors for understanding and predicting the damage undergone by a target plant species. The mechanism that best explains these results is associational avoidance of saplings that grow near to unpalatable shrubs. It is necessary to introduce this neighbour effect in theoretical models and food‐web approaches that analyse the plant–herbivore relationships, since it can strongly determine not only the intensity of the interaction, but also the spatial distribution and diversity of the plant community.     

Effects of spatial distribution on plant associational defense against herbivory

Several studies have shown that consumption of a focal plant by herbivores depends not only on its own defense traits but also on the characteristics of the neighboring plants. A number of studies have reported on plant associational defense in relation to neighboring plant palatability but the effect of the spatial distribution of the focal plant within patches of different neighboring plants has received less attention. We conducted a manipulative experiment to determine whether and how spatial distribution of focal plants affects the associational defense between plant species. In our experimental setup sheep encountered two patches varying in spatial distribution of the focal plant within patches (dispersed or clumped) and patch quality, good patch and bad patch, where the focal plant, Lathyrus quinquenervius, was neighbored to high- (Chloris virgata) or low-palatable (Kalimeris integrifolia) species, respectively. Results showed that, when focal plants were dispersed within both patches, the risk of attack was significantly lower for focal plants in the patches with low- than high-palatable neighbors, indicating associational defense. Alternatively, when focal plants were clumped within both patches, they were consumed in bad-patch as much as in good-patch plots, which indicates the absence of associational defense. However, if the focal plants have different spatial distributions in the two patches (dispersed in good-patch and clumped in bad-patch or vice versa), sheep foraging success for focal plants was greatly reduced in dispersed spatial pattern irrespective of the palatability of neighboring plants. Therefore, we concluded that spatial distribution is as important as traits of neighboring plants in predicting vulnerability of the focal plant to grazing by generalist herbivores. The outcome of plant associational defense for different types of neighborhood strongly depends on the magnitude of herbivore foraging selectivity between and within patches, which further depended on the contrasts between plant species or between patches.     

Scale-dependent neighborhood effects: shared doom and associational refuge

A resource’s susceptibility to predation may be influenced by its own palatability and the palatability of its neighbors. We tested for effects of plant chemical defenses on seed survival by manipulating the frequency of palatable and less palatable sunflower seeds in food patches subject to harvest by fox squirrels (Sciurus niger) and gray squirrels (Sciurus carolinensis). We varied resource distributions at three scales: among stations (aggregates of patches ca. 50 m apart), among patches immediately adjacent to each other, and within patches. When food patches were segregated into high-palatability and low-palatability stations (Experiment 1), seeds suffered greater mortality at stations with high levels of palatable seeds. In the same experiment, within patches, squirrels selected strongly for palatable seeds over less palatable seeds. When high- and low-palatability food patches were placed together at the same stations (Experiment 2), increasing densities of co-occurring palatable seeds amplified the mortality of less palatable seeds, indicating “shared doom.” When palatable and less palatable seeds were partitioned into micropatches (Experiment 3), associational effects disappeared, as predicted. Furthermore, selectivity in less palatable patches increased as the initial densities of palatable seeds increased, and selectivity in palatable patches decreased as the initial densities of less palatable seeds increased. Foraging theory predicts associational effects among prey that vary in palatability. Our results show how the type and magnitude of associational effects emerge from the interplay among the spatial scale of prey heterogeneity, the diet selection strategy, and the scale-dependent foraging responses of the consumer.     

Associational resistance and shared doom: effects of epibiosis on herbivory

The potential for spatial associations between palatable and unpalatable plant species to reduce herbivore pressure on the palatable species has been described as associational resistance, associational refuge or associational defense for numerous terrestrial and marine communities. One of the closest associations between species-epibiosis-has not been thoroughly investigated in this regard. In this study we evaluated how different associations between host seaweeds and epibiotic plants and animals influenced the movement of an omnivorous sea urchin (Arbacia punctulata) to the host and subsequent feeding on the host. A. punctulata showed clear preferences when given pairwise choices between 12 prey species (3 animals, 9 algae). These preferences were consistent and allowed us to rank the six epibiont species and six host species linearly from least to most preferred by A. punculata. Most host-epibiont associations dramatically changed urchin preference, increasing or decreasing urchin grazing on fouled hosts as compared to clean conspecifics. Herbivory on the host increased when the epibiont was more preferred, and decreased when it was less preferred than the unfouled host alga. Taking the host species as a point of reference, we classified epibiosis-caused decrease in herbivory as associational resistance, while epibiont-caused increases in herbivory were defined as shared doom. These epibiont-host-herbivore interactions could select for hosts that facilitate the growth of certain low preference epibionts on their surfaces in situations where the resulting decreases in herbivory would offset the various negative effects of being fouled. In contrast, in situations where herbivores are common, the negative effects of being fouled by palatable epibionts may be much greater than is generally assumed. In our assays, unpalatable hosts fouled by palatable epibionts became much more attractive to urchins and rose several ranks on the urchins' preference hierarchy.     

Spatially complex neighboring relationships among grassland plant species as an effective mechanism of defense against herbivory

Close spatial relationships between plant species are often important for defense against herbivory. The associational plant defense may have important implications for plant community structure, species diversity, and species coexistence. An increasing number of studies have focused on associational plant defense against herbivory at the scale of the individual plant and its nearest neighbors. However, the average neighborhood effects between plant species at the scale of whole plant communities have received almost no attention. The aims of this study were to determine patterns of spatial relationship between different plant species that can provide effective defense against herbivory. We conducted a manipulative experiment using sheep and three native plant species with different palatability. Consumption of palatable plants by herbivores was largest when the three plant species were isolated in three patches and independent of each other. A homogenous and spatially equal neighbor relationship between the three species did not reduce the risk of herbivory of palatable species compared to isolation of these species, but it reduced the total intake of all plant species. The palatable species was subject to less herbivory in a complex spatial neighborhood of several plant species. High complexity of spatial neighborhood resulted in herbivores passively reducing selectivity, thereby reducing the probability of damage to palatable species in the community, or making inaccurate judgments in foraging selectivity between and within patches, thereby reducing the vulnerability of palatable plants and even the whole plant community. We conclude that compelling herbivores to passively reduce the magnitude of foraging selectivity by establishing spatially complex neighborhoods between plant species is a compromise and optimal spatial strategy by plants to defend themselves again herbivory. This may contribute not only to maintenance of plant species diversity but also to a stable coexistence between herbivores and plants in grassland ecosystems.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.     

The larval ecology of the butterfly Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant quantity and quality as main predictors of habitat quality

Corresponding to theory, the persistence of metapopulations in fragmented landscapes depends on the area of suitable habitat patches and their degree of isolation, mediating the individual exchange between habitats. More recently, habitat quality has been highlighted as being equally important. We therefore assess the role of habitat area, isolation and quality for the occupancy of larval stages of the regionally threatened butterfly Euphydryas desfontainii occurring in grassland habitats comprising the host plant Dipsascus comosus. We put a special focus on habitat quality which was determined on two spatial scales: the landscape (among patches) and the within-patch level. On the landscape level, occupancy of caterpillars was determined by a presence-absence analysis at 28 host plant patches. On the within-patch level, oviposition site selection was studied by comparing 159 host plants with egg clutches to a random sample of 253 unoccupied host plants within six habitat patches. The occupancy of caterpillars and presence of egg clutches on host plants was then related to several predictors such as patch size and isolation on the landscape level and host plant characteristics and immediate surroundings on the within patch level. On the landscape level, only host plant abundance was related to the presence of caterpillars, while size and isolation did not differ between occupied and unoccupied patches. However, the weak discrimination of larval stages among patches changed on the within-patch level: here, several microclimatic predictors such as sunshine hours and topography, host plant morphology and phenology as well as further potential host plants in the immediate surroundings of the plant chosen for oviposition strongly determined the presence of egg clutches. We strongly suggest promoting the presence of the host plant in topographically and structurally rich habitat patches to offer potential for microclimatic compensation for a species considered threatened by climate change.     

Associational susceptibility in broccoli: mediated by plant volatiles,impeded by ozone

Plant‐emitted volatile organic compounds (VOCs) mediate interactions within a plant community. Typically, receiving a signal from a damaged neighbour enhances the defensive attributes of a receiver plant. The mechanisms underlying plant–plant interactions may be divided into active and passive processes, both of which involve transit of VOCs between plants and are vulnerable to environmental perturbation. Numerous studies have documented between‐plant interactions, but the specific effects on a receiver plant's interactions with herbivores have received little attention. Moreover, the relative contributions of active and passive processes to plant defence and the effects of environmental pollutants on the processes have been largely unexplored. We used a system comprising Brassica oleracea var. italica (broccoli) and the specialist herbivore Plutella xylostella to test whether plants previously exposed to herbivore‐damaged neighbours differed from nonexposed plants in their susceptibility to oviposition. We then investigated the roles of active and passive mechanisms in our observations and whether differences in susceptibility remained under elevated ozone concentrations. Plants exposed to herbivore‐damaged neighbours were more susceptible to oviposition than plants exposed to undamaged neighbours, which indicates associational susceptibility. Mechanistically, active and passive volatile‐mediated processes occurred in tandem with the passive process – involving adsorption of sesquiterpenes to receiver plants – appearing to structure the oviposition response. Exposure to ozone rapidly degraded the sesquiterpenes and eliminated the associational susceptibility. Plant volatiles have typically been thought to play roles in between‐plant interactions and to promote receiver plant defence. Here, we show that receiver plants may also become more susceptible to oviposition and thus more likely to be damaged. Extensive disruption of volatile‐mediated interactions by an atmospheric pollutant highlights the need to consider the pervading environment and changes therein when assessing their ecological significance.     

Herbivore-induced coexistence of competing plant species

We study a series of spatially implicit lottery models in which two competing plant species, with and without defensive traits, are grazed by a herbivore in a homogeneous habitat. One species (palatable) has no defensive traits, while the other (defended) has defensive traits but suffers reduced reproduction as the result of an assumed trade-off. Not surprisingly, coexistence of these plants cannot occur when the herbivore density is very low (the palatable plant always wins) or very high (the defended plant wins). At intermediate densities, however, herbivory can mediate plant coexistence, even in a homogeneous environment. If the herbivore eats several plants per bite, and its forage-selection depends on the average palatability of the plants it eats, then palatable species in the immediate neighbourhood of defended plants may be more likely to persist (associational resistance) even at higher grazing pressure. If the herbivore shows a positive numerical response to the average palatability of the habitat as a whole, then both plant populations are stabilized and coexistence is promoted, because both species obtain a minority advantage through the negative feedback caused by herbivory. If the herbivore exhibits both of these traits, the system may have at most two non-trivial equilibria, one of which is stable and the other unstable. This means that coexistence in such a system is vulnerable to large fluctuations in herbivore density and identity, and this has implications for conservation in systems where large herbivores are managed to promote plant diversity.     

Associational refuge in practice: can existing vegetation facilitate woodland restoration?

Herbivores can dramatically diminish revegetation success, but associational refuge theory predicts that neighbouring plants could hinder browsing of planted seedlings. The key to strategic restoration using associational refuge is to define which patch variables are effective against the appropriate herbivores, at multiple scales, and to understand which stages of the foraging process these variables disrupt. Our study aimed to test the capacity of existing vegetation to act as associational refuge for planted seedlings by affecting search, detection and consumption decisions, and more generally influence herbivore foraging patterns. We conducted a field trial with free‐ranging, mammalian herbivores and nursery‐raised, native tree seedlings. We quantified seedling browsing damage over time in relation to a suite of existing patch variables at two spatial scales (100 m² and 4 m²). After two months, 78% of seedlings were browsed, suffering mean foliage loss of 90.5%. Focal seedlings were almost exclusively consumed by swamp wallabies Wallabia bicolor, an abundant generalist browser. Once a swamp wallaby investigated a seedling, the probability of consumption was high (86%). At the large scale, browsing of seedlings was delayed in patches with lower canopy cover and fewer browsed plant species. Seedlings in fern‐dominated patches escaped browsing for longer than those in grass‐dominated patches. At the small scale, browsing was delayed with higher cover of understorey vegetation. Associational refuge was provided by vegetation with characteristics, and at spatial scales, consistent with disrupted search and detection of focal seedlings by herbivores. Thus strategic placement of seedlings in existing vegetation – based on understanding which herbivore species is responsible and how it responds to vegetation – can take advantage of associational refuge during restoration. However, given rapid seedling detection by herbivores, associational refuge may be inadequate in the long‐term under high browsing pressure unless high absolute numbers of seedlings are planted among refuge.     

Feeding patterns by roe deer and rabbits on pine, willow and birch in relation to spatial arrangement

The location of a plant and its association with the surrounding vegetation may be a strong determinant of herbivore foraging decisions. The attractiveness of a food plant may be reduced if associated with a less preferred one (repellent–plant hypothesis) or, conversely, it may escape herbivory via association with a more preferred species (attractant–decoy hypothesis). In this study we tested the hypothesis that selection of the same food plants by two herbivore species with different body mass, i.e. roe deer and rabbits, is affected by the spatial disposition of preferred plants in relation to less preferred ones. We planted willows, birches and pines, representing food species of higher, intermediate and lower preference, respectively, in different spatial arrangements, to manipulate patch quality and accessibility to herbivores. Contrasting patches were constructed by planting willows or pines in the midst of birches and single-species arrays of all three species, in a blocked design, replicated six times in an area occupied by both roe deer and rabbits. Preference patterns were studied by recording browsing on current years' shoots and older plant parts. Across all trees in the experiment, we confirmed that roe deer clearly preferred willow over birch, pine was not browsed at all. There was greater herbivory by roe deer on birches occurring peripherally in an array, when the associated central species was willow or pine, as compared to birches in a single-species birch patch. Therefore food choices by roe deer appear to be determined by both patch and individual level selection. The rabbits only showed preference at a species level and did not respond to spatial arrangement of trees. The results from this study indicate that the location and association of trees can strongly affect foraging patterns of roe deer, but not in a manner wholly consistent with the repellent–plant or the attractant–decoy hypotheses.     

Herbivore-induced responses and patch heterogeneity affect abundance of arthropods on plants

Abstract.  1. Plants respond to herbivore damage by inducing defences that can affect the abundance of herbivores and predators. These tritrophic interactions may be influenced by heterogeneity in plant neighbourhood.
2. In the present study, the effects of induced responses on the abundance of herbivores (flea beetles and aphids), omnivores (pirate bugs and thrips), and predators (lady beetles and spiders) on individual plants and their neighbours between and within patches composed of three tomato plants was investigated.
3. Herbivore damage was manipulated to create homogeneous patches where either all or none of the plants had defences induced by herbivore damage, and heterogeneous patches where only one of the plants was induced.
4. Arthropod abundance on plants at different scales was compared by testing between patch effects (patch level), for neighbourhood effects at the plant phenotype level (neighbourhood level), and between near and far plants (within patch position).
5. At the patch level , plants in homogeneously induced patches contained fewer flea beetles and pirate bugs, but more lady beetles, compared with homogeneously non-induced patches. There was no effect of patch type on the abundance of aphids, thrips, and spiders on plants.
6. At the neighbourhood level , induced plants in heterogeneous patches contained more flea beetles and pirate bugs compared with induced plants in homogeneous patches, indicating that the abundance of some herbivores and omnivores on induced plants varied depending on the phenotype of the other plants within the patch. Within patch position, there was no evidence that the abundance of herbivores or predators on non-induced plants was affected by proximity to an induced plant.
7. Therefore, variation in plant neighbourhood generated by induced plant responses affected the abundance of three arthropods from three feeding guilds.     

Unpreferred plants affect patch choice and spatial distribution of European brown hares

Many herbivore species prefer to forage on patches of intermediate biomass. Plant quality and forage efficiency are predicted to decrease with increasing plant standing crop which explains the lower preference of the herbivore. However, often is ignored that on the long-term, plant species composition is predicted to change with increasing plant standing crop. The amount of low-quality, unpreferred food plants increases with increasing plant standing crop. In the present study the effects of unpreferred plants on patch choice and distribution of European brown hare in a salt-marsh system were studied. In one experiment, unpreferred plants were removed from plots. In the second experiment, plots were planted with different densities of an unpreferred artificial plant. Removal of unpreferred plants increased hare-grazing pressure more than fivefold compared to unmanipulated plots. Planting of unpreferred plants reduced hare-grazing pressure, with a significant reduction of grazing already occurring at low unpreferred plant density. Spatial distribution of hares within this salt-marsh system was related to spatial arrangement of unpreferred plants. Hare-grazing intensity decreased strongly with increasing abundance of unpreferred plants despite a high abundance of principal food plants. The results of this study indicate that plant species replacement is an important factor determining patch choice and spatial distribution of hares next to changing plant quality. Increasing abundance of unpreferred plant species can strengthen the decreasing patch quality with increasing standing crop and can decrease grazing intensity when preferred food plants are still abundantly present.     

Mass emergence of a specialist sawfly species on unpalatable herbs under severe feeding pressure by sika deer

In recent years, the increase in sika deer populations has had a significant impact on forest ecosystems. Many studies have reported that this increase has led to the dominance of plants unpalatable to the deer; however, few studies have investigated the effects of this dominance on the specialist sawfly species that feed on the unpalatable plants. We observed the mass emergence of Aglaostigma amoorensis (Hymenoptera: Tenthredinidae), a rare specialist sawfly of the false helleborine Veratrum plants unpalatable to sika deer, in areas under severe feeding pressure by sika deer. An exponential increase in the number of emergent A. amoorensis adults was observed to occur with an increase in Veratrum album patch sizes. In addition, the increase in the A. amoorensis density had a negative effect on reproductive success of V. album. Our findings suggest that the relative abundance of specialist sawfly to the host‐plant abundance has changed for successive years after sika deer irruption.     

Habitat use at fine spatial scale: how does patch clustering criteria explain the use of meadows by red deer?

Large mammalian herbivores are keystone species in different ecosystems. To mediate the effects of large mammalian herbivores on ecosystems, it is crucial to understand their habitat selection pattern. At finer scales, herbivore patch selection depends strongly on plant community traits and therefore its understanding is constrained by patch definition criteria. Our aim was to assess which criteria for patch definition best explained use of meadows by wild, free-ranging, red deer (Cervus elaphus) in a study area in Northeast Portugal. We used two clustering criteria types based on floristic composition and gross forage classes, respectively. For the floristic criteria, phytosociological approach was used to classify plant communities, and its objectivity evaluated with a mathematical clustering of the floristic relevés. Cover of dominant plant species was tested as a proxy for the phytosociological method. For the gross forage classes, the graminoids/forbs ratio and the percentage cover of legumes were used. For assessing deer relative use of meadows we used faecal accumulation rates. Patches clustered according to floristic classification better explained selection of patches by deer. Plant community classifications based on phytosociology, or proxies of this, used for characterizing meadow patches resulted useful to understand herbivore selection pattern at fine scales and thus potentially suitable to assist wildlife management decisions.     

Refuge effect of an unpalatable forb on community structure and grass morphology in a temperate grassland

The role of unpalatable plant species as biological antiherbivore refuges for palatable species is well-documented at community level particularly in harsh environments. In productive sub-humid temperate grassland subjected to domestic grazing, we examined the protective effect of Eryngium horridum on plant community structure and floristic composition, and evaluated whether these changes impacted on a number of morphological traits of grasses, related to grazing resistance. We also investigated, for a palatable grass species (Stipa neesiana) the existence of morphological differences between protected and unprotected plants and if this eventual variation was either plastic or genetic. The study consisted of a field survey where we compared paired patches, with and without E. horridum, and a greenhouse experiment where we evaluated individuals of S. neesiana coming from both patch types over a 11 months period. Patches dominated by E. horridum had lower richness and cover of forbs than patches without the forb, and similar richness but greater cover of cool-season tussock palatable grasses, which suggests a protective role on the latter. Grasses in these patches also had longer blades and sheaths and lower specific leaf area. The morphological differences of S. neesiana individuals collected from both patch types disappeared after 11 months growth in a common environment which revealed significant phenotypic plasticity in this species. These results suggest the existence of plant-to-plant facilitation in a productive ecosystem not only at community level, through changes in species richness and the promotion of palatable grasses, but also at population level, through plastic changes in aboveground morphological traits. Both facilitation and plasticity, would contribute to the persistence of threatened palatable grasses in the heavy grazed productive ecosystems.     

Distribution and colonisation ability of three parasitoids and their herbivorous host in a fragmented landscape

Habitat fragmentation can disrupt communities of interacting species even if only some of the species are directly affected by fragmentation. For instance, if parasitoids disperse less well than their herbivorous hosts, habitat fragmentation may lead to higher herbivory in isolated plant patches due to the absence of the third trophic level. Community-level studies suggest that parasitoids tend to have limited dispersal abilities, on the order of tens of metres, much smaller than that of their hosts, while species-oriented studies document dispersal by parasitoids on the scale of kilometres. In this study the distribution patterns of three parasitoid species with different life histories and their moth host, Hadena bicruris, a specialist herbivore of Silene latifolia, were compared in a large-scale network of natural fragmented plant patches along the rivers Rhine and Waal in the Netherlands. We examined how patch size and isolation affect the presence of each species. Additionally, experimental plots were used to study the colonisation abilities of the species at different distances from source populations.In the natural plant patches the presence of the herbivore and two of the parasitoids, the gregarious specialist Microplitis tristis and the gregarious generalist Bracon variator were not affected by patch isolation at the scale of the study, while the solitary specialist Eurylabus tristis was. In contrast to the herbivore, the presence of all parasitoid species declined with plant patch size. The colonisation experiment confirmed that the herbivore and M. tristis are good dispersers, able to travel at least 2 km within a season. B. variator showed intermediate colonisation ability and E. tristis showed very limited colonisation ability at this spatial scale. Characteristics of parasitoid species that may contribute to differences in their dispersal abilities are discussed.     

Soil nutrients indirectly influence intraspecific plant selection in white-tailed deer

Few attempts have been made to determine how soil productivity influences diet selection in herbivores, likely because environmental characteristics known to influence diet selection such as plant community structure and herbivore nutritional demands are often confounded with changes in soil productivity. We designed a soil-amendment experiment to isolate the effects of soil productivity on diet selection by manipulating soil productivity and quantifying intraspecific plant selection within a population of white-tailed deer (Odocoileus virginianus). We hypothesized soil productivity would indirectly influence deer plant selection by directly affecting plant tissue chemistry. Soil productivity indeed influenced diet selection indirectly because soil amendments only affected deer plant selection when palatable plants were present. Soil amendments increased plant phosphorus concentrations, and plant phosphorus concentrations explained 47% of the variation in diet selection. Thus, our data indicate plant nutritional quality mediates the indirect effects of soil productivity on herbivore diet selection. Previous research demonstrating differential influences of herbivory on plant communities across a soil productivity gradient may in part be explained by indirect effects of soil productivity on diet selection.