Being a generalist herbivore in a diverse world: how do diets from different grasslands influence food plant selection and fitness of the grasshopper Chorthippus parallelus?

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing.     

Diet switching in a generalist mammalian folivore: fundamental to maximising intake

Generalist mammalian herbivores exploit a diverse diet. A generalised feeding strategy utilises a mixed diet to obtain a range of nutrients and to reduce the detoxication load of similar groups of plant secondary metabolites (PSMs). There is limited research investigating how mammalian herbivores achieve this dietary mixing in their daily foraging activities. We investigated the patterns of, and behaviours associated with, dietary mixing in a generalist mammalian folivore, the common brushtail possum (Trichosurus vulpecula). Possums were offered foliage of two eucalypt species (Eucalyptus globulus and E. regnans) as either (a) Full choice: both species offered for 8 h; (b) Restricted choice: both species offered for 2×2 h blocks; (c) G–R no choice: E. globulus offered for the first 4 h, E. regnans offered for next 4 h; and (d) R–G no choice: E. regnans offered for first 4 h, E. globulus offered for next 4 h. We hypothesised that possums would maximise intake on the Full choice diet, where time availability was greatest in combination with a choice of foliage. We also hypothesised that diet switching, defined as the frequency of changing between food types while feeding, would play a fundamental role in maximising intake. Possums achieved maximum intake on the Full choice diet and minimum intake on the No choice diets. Although intake was similar between the Restricted choice and No choice diets, possums achieved this intake in half the amount of time when able to switch regularly between foliage on the Restricted choice diet. We conclude that a generalist herbivore’s ability to effectively switch diets when foraging is fundamental to maximising intake. Hence, the degree of plant heterogeneity in an environment, and the spatial scale at which it occurs, may affect an herbivore’s foraging decisions and, ultimately, influence its foraging efficiency.     

Food choice causes interrupted feeding in the generalist grasshopper Schistocerca americana: further evidence for inefficient decision-making

Individual grasshoppers were observed foraging on either single food plant species or binary mixtures of them. Individuals ate slightly less on the mixtures, but grew faster on them. In the mixtures, individuals fed regularly from both plant species, typically eating both within a single meal. Feeding bouts were longer on the preferred plant species in the mixture. The pattern of foraging behavior was markedly different in mixtures and single food treatments. In mixtures, although the total length of time engaged in ingestion over the day was slightly less than in single food treatments, there were relatively more short feeding bouts and more gaps between feeding bouts within meals, so that the time spent in activities associated with the food was markedly longer on mixtures. This is discussed in terms of efficiency and the problem of choice that faces individual generalists. The efficiency factor and its impact on vigilance is added to the other costs of being a polyphagous herbivore.     

Feeding ecology of the Iberian cyprinid Barbus bocagei Steindachner, 1865 in a lowland river

Patterns of food resource utilization by Barbus bocagei in a lowland river in the central part of the Iberian Peninsula are presented and discussed. Focal points include feeding activity, diet composition and seasonal and ontogenetic diet shifts. Striking features of overall feeding pattern of barbel are its continuous feeding activity, bottom habits, high consumption of detritus and dipteran larvae and narrow diet breadth. Seasonal changes in food resource availability elicited changes in food utilization patterns, revealing a trend to more generalist feeding during spring. Barbel increase consumption of plant material and large prey during ontogeny, but overlap in prey resource utilization between size classes is high. The versatility of barbel foraging behaviour may be an important feature in its exploitation of Iberian lotic systems.     

Dietary mixing in three generalist herbivores: nutrient complementation or toxin dilution?

We reared larvae of three generalist insect species on plants occurring in their habitats. Individuals of each species were kept either on mixed diets, or on each plant species separately. We measured food plant preference in the mixed-diet group and compared insect performance on single plants to the performance on the mixed diet. For all three insect species, food choice within the mixed-diet groups was non-random and delivered the best overall performance, thus fulfilling the criteria for self-selected diets. When a single diet was as good as the mixed diet for one particular aspect of performance (Adenostyles alliariae and Petasites albus for Miramella alpina; A. alliariae for Callimorpha dominula), it was never the most preferred food plant in the mixed-diet treatment. Whether the benefit achieved by mixing diets is due to nutrient complementation or toxin dilution, we argue that there is no easy way to distinguish between the two hypotheses on the basis of consumption and performance measurements, as has previously been proposed. From the interpretation of utilisation plots, the ANCOVA equivalent of nutritional indices, we were able to gain insight into where in the sequence from ingestion to growth (preingestive, predigestive or postdigestive) single diets caused differences from mixed diets. The elements of this control system which were influenced by single diets varied considerably, both within and between insect species. No food plant was toxic or deterrent to all experimental insect species; a food plant that caused consumption effects (preingestive) for one insect species could be dealt with metabolically (postdigestive) by another; different food plants could cause behavioural effects (preingestive), metabolic effects (postdigestive), or a combination of both effects, all within the same insect species. However, one generality did emerge: once a food was ingested, further growth-relevant effects occurred metabolically (postdigestive) rather than via differential egestion (digestibility). Received: 5 October 1998 / Accepted: 1 March 1999     

Testing the efficiency of three 15N-labeled nitrogen compounds for indirect labeling of grasshoppers via plants in the field

In field studies of plant–insect herbivore interactions it is often difficult to establish which herbivore has fed on a particular plant. We investigated the suitability of three different ¹⁵N‐labeled nitrogen compounds (ammonium, nitrate, and glycine) for indirect marking of three grasshopper species [Omocestus viridulus (L.), Chorthippus parallelus (Zett.), and Chorthippus biguttulus (L.) (Orthoptera: Acrididae)] through labeling their food plants in the field. In two short‐term experiments grassland plots of 1 m² were separately labeled with either one of the different nitrogen compounds. Grasshoppers were caged on three food‐plant species [Dactylis glomerata L., Holcus lanatus L. (Poaceae), and Trifolium repens L. (Fabaceae)] present in these plots for 72 h. Significantly enriched δ¹⁵N values in grasshoppers were found in all plant/grasshopper combinations. Enrichment in grasshoppers was positively correlated with the enrichment of plants and labeling with nitrate resulted in highest ¹⁵N enrichment. In a long‐term experiment, individuals of C. biguttulus were placed in a cage covering an area of 1 m² for 37 days, with sampling of grasshoppers at regular intervals. δ¹⁵N values of the grasshopper and a common food plant, D. glomerata, increased steadily over time, up to 40‐fold by the end of the experiment. Our results demonstrate that ¹⁵N‐labeling of plants is an appropriate tool for the investigation of insect–plant interactions under natural conditions.     

The impact of plant diversity and fertilization on fitness of a generalist grasshopper

In many environments land use intensification is likely to result in a decrease in species richness and in an increase in eutrophication. Although the importance of both factors for higher trophic levels such as insect herbivores is well documented, their impact has rarely been studied in combination. Herbivorous insects have a strong impact on the functioning of ecosystems and it is therefore important to understand how they are affected by eutrophication in high or low diversity environments.We used a grassland biodiversity experiment to investigate the combined effect of fertilization and plant diversity loss on the fitness of the generalist grasshopper Chorthippus parallelus by rearing grasshopper nymphs for four weeks in cages on unfertilized or fertilized (NPK) subplots across a species richness gradient from 1 to 60 plant species. Survival, the number of oothecae, body mass and the number of hatchlings were measured separately for each cage. Plant diversity had no effect on any of the grasshopper fitness measures, neither in unfertilized nor in fertilized plots. NPK-fertilization reduced grasshopper survival but increased body mass of males and reproductive success of the surviving females. Fertilization effects were not mediated by plant community structure, productivity or composition, suggesting that higher food plant quality was one of the main drivers. There was no interaction between plant diversity and fertilization on any of the measures. In conclusion, an increase in eutrophication, in both species-rich and species-poor grasslands, could lead to higher reproductive success and therefore higher abundances of herbivorous insects including insect pests, with fertilization effects dominating plant diversity effects.     

Herbivore phenology can predict response to changes in plant quality by livestock grazing

Livestock grazing can have a strong impact on herbivore abundance, distribution and community. However, not all species of herbivores respond the same way to livestock grazing, and we still have a poor understanding of the underlying mechanisms driving these differential responses. Here, we investigate the effect of light intensity cattle grazing on the abundance of two grasshoppers (Euchorthippus cheui and E. unicolor) that co-occur in the same grasslands and feed on the same food plant (the dominant grass Leymus chinensis). The two grasshopper species differ in phenology so that their peak abundances are separated into early- and late-growing seasons. We used an exclosure experiment to monitor grasshopper abundance and food quality in the field under grazed and ungrazed conditions, and performed feeding trials to examine grasshopper preference for grazed or ungrazed food plants in the laboratory. We found that the nitrogen content of L. chinensis leaves continuously declined in the ungrazed areas, but was significantly enhanced by cattle grazing over the growing season. Cattle grazing facilitated the early-season grasshopper E. cheui, whereas it suppressed the late-season grasshopper E. unicolor. Moreover, feeding trials showed that E. cheui preferred L. chinensis from grazed plots, while E. unicolor preferred the leaves from ungrazed plots. We conclude that livestock grazing has opposite effects on the two grasshopper species, and that these effects may be driven by grazing-induced changes in plant nutrient content and the unique nutritional niches of the grasshoppers. These results suggest that insects that belong to the same guild can have opposite nutrient requirements, related to their distinct phenologies, and that this can ultimately affect their response to cattle grazing. Our results show that phenology may link insect physiological needs to local resource availabilities, and should be given more attention in future work on interactions between large herbivores and insects.     

Comparative Feeding Ecology of Sympatric <Emphasis Type="Italic">Microcebus berthae and M. murinus</Emphasis>

Most Malagasy primate communities harbor a diverse assemblage of omnivorous species. The mechanisms allowing the coexistence of closely related species are poorly understood, partly because only preliminary data on the feeding ecology of most species are available. We provide an exemplary feeding ecology data set to illuminate coexistence mechanisms between sympatric gray and Madame Berthe’s mouse lemurs (Microcebus murinus, M. berthae). We studied their feeding ecology in Kirindy Forest/CFPF, a highly seasonal dry deciduous forest in western Madagascar. Between August 2002 and December 2007, we regularly (re-)captured, marked, and radiotracked females of both species. A combination of direct behavioral observations and fecal analyses revealed that both Microcebus species used fruit, arthropods, gum, insect secretions, and small vertebrates as food sources. However, Microcebus berthae and M. murinus differed in both composition and seasonal variation of their diets. Whereas the diet of Microcebus murinus varied seasonally and was generally more diverse, M. berthae relied mainly on insect secretions supplemented by animal matter. The differences were also reflected in a very narrow feeding niche of Microcebus berthae and a comparatively broad feeding niche of M. murinus. Resource use patterns of Madame Berthe’s and more so of opportunistic gray mouse lemurs broadly followed resource availability within the strongly seasonal dry forest. Feeding niche overlap between the 2 sympatric species was high, indicating that food resource usage patterns did not reflect niche partitioning, but can instead be explained by constraints due to food availability.     

Population differences in<Emphasis Type="Italic"> Trifolium repens</Emphasis> L. response to ultraviolet-B radiation: foliar chemistry and consequences for two lepidopteran herbivores

White clover growing in New Zealand is experiencing increasing levels of ultraviolet-B (UV-B) radiation as a result of ozone depletion. We evaluated the effects of UV-B radiation on the foliar chemistry of two populations of white clover (Trifolium repens L.), ’Huia’ and ’Tienshan,’ and the consequences for the performance of armyworms (Spodoptera litura) and cutworms (Graphania mutans). Plants were grown in controlled environment rooms with and without supplemental UV-B radiation at a dose of 13.3 kJ m^–2 day^–1, corresponding to a 25% mid-summer ozone depletion above Palmerston North, New Zealand. In both white clover populations, UV-B radiation elicited changes in foliar chemistry, including slight increases in nitrogen concentrations and decreases in carbohydrate concentrations. In addition, the ’Huia’ population showed decreases in fiber concentrations and marked increases in cyanogenic activity. No change in UV-absorbing compounds was detected in either population. Long- and short-term feeding trials were conducted to assess dietary effects on insect growth, consumption, and food utilization. Changes in the performance of both insect species were generally small. The most pronounced effect was a 36% reduction in weight of S. litura after 2 weeks of feeding on Huia grown at high UV, but larval development times were only slightly prolonged and pupal weights were unaffected. S. litura short-term performance was affected by differences in white clover population. The long-term performance of G. mutans was not affected and its short-term performance (stadium duration and consumption rate) was only marginally affected by the high-UV treatment. We conclude that the effects of elevated UV-B radiation on white clover plant chemistry can be specific to certain plant populations. The differences in sensitivity of the two generalist insect species suggest that effects may also be specific to certain plant-herbivore associations. These results indicate that future UV-B herbivory studies should examine genotypic effects in both plants and animals. Received: 18 May 1999 / Accepted: 25 July 1999     

The cost of mutualism in a fly-fungus interaction

The movement ability of individuals has become increasingly important to a variety of ecological questions. In this study, I investigate how plant structure and changes in body size through development affect the movement ability of a predaceous stinkbug, Podisus maculiventris, on three species of goldenrod (Solidago spp.) representing a wide range of surface complexities. I adapt existing techniques for quantifying movement in two dimensions to the study of movement on natural plant structures in three dimensions. These experiments indicate that plant structure and insect size are significant factors affecting the movement ability of P. maculiventris. Changes in movement ability due to factors of ontogeny and different habitat structures suggest that the scale of an individual’s ambit or ecological sphere of influence may vary within its lifespan. Considering the influence of ontogeny and habitat structure on movement ability may be useful to investigations of population dynamics, foraging behavior, and pest management. Received: 14 July 1999 / Accepted: 23 March 2000     

Dietary overlap of grasshoppers on sandhill rangeland in northeastern Colorado

Summary Overlap in the diets of 14 species of grasshoppers on mixed-grass prairie in northeastern Colorado was estimated by microscopic examination of crop contents. Food availability estimates, taken by a weight-estimate method, facilitated the determination of food preferences. Two cases of high overlap in diets and period of habitat occupancy occurred between species which consumed equal or almost equal numbers of different foods (foodniche breadth) and had almost identical foraging behavior. The first case occurred early in the season between Xanthippus corallipes and Arphia conspersa with foodniche breadths of 14 and 11, respectively. The second case occurred later in the season between Amphitornus coloradus and Trachyrhachys kiowa both of which have foodniche breadths of 6. The amount of movement between plants while feeding was equal or almost equal in both cases, thus we observed high dietary overlap between species with equal ecological efficiencies. Dietary overlap between the 14 grasshopper species decreased as foodniche dimensions increased and the grasshopper population as a whole utilized each food resource approximately proportional to its relative abundance. Factors influencing the interpretation and importance of dietary overlap in this study included 1. non-synchronous life cycles, 2. seasonal changes in diets and food preferences, 3. complexity of the food resources, 4. variations in foodniche dimensions among grasshopper species, and 5. variations in diets between males and females within a species. Our findings agree with MacArthur and Levins theory that the number of competing species which can coexist is proportional to the total range of the environment divided by the niche breadth of the species.     

Functional identity versus species richness: herbivory resistance in plant communities

The resistance of a plant community against herbivore attack may depend on plant species richness, with monocultures often much more severely affected than mixtures of plant species. Here, we used a plant–herbivore system to study the effects of selective herbivory on consumption resistance and recovery after herbivory in 81 experimental grassland plots. Communities were established from seed in 2002 and contained 1, 2, 4, 8, 16 or 60 plant species of 1, 2, 3 or 4 functional groups. In 2004, pairs of enclosure cages (1 m tall, 0.5 m diameter) were set up on all 81 plots. One randomly selected cage of each pair was stocked with 10 male and 10 female nymphs of the meadow grasshopper, Chorthippus parallelus. The grasshoppers fed for 2 months, and the vegetation was monitored over 1 year. Consumption resistance and recovery of vegetation were calculated as proportional changes in vegetation biomass. Overall, grasshopper herbivory averaged 6.8%. Herbivory resistance and recovery were influenced by plant functional group identity, but independent of plant species richness and number of functional groups. However, herbivory induced shifts in vegetation composition that depended on plant species richness. Grasshopper herbivory led to increases in herb cover at the expense of grasses. Herb cover increased more strongly in species-rich mixtures. We conclude that selective herbivory changes the functional composition of plant communities and that compositional changes due to selective herbivory depend on plant species richness.     

Spatial distributions of multiple plant species affect herbivore foraging selectivity

Spatial distribution of food resources is an important factor determining herbivore foraging. Previous studies have demonstrated that clumped distribution of preferred species increases its consumption by herbivores in single‐ or two‐species systems. However, the potential impact of distribution pattern of less preferred species on foraging was ignored. In natural grasslands with high species diversity and complexity, the spatial distribution of preferred species impacts on herbivore foraging may be strongly correlated with the distribution of less preferred species. Our aims were to determine the effect of distribution of both preferred and other plant species on herbivore foraging under conditions close to a native, multi‐species foraging environment, and conceptualize the relationships between spatial distribution of food resources and herbivore consumption. We hypothesized that random distribution of non‐preferred species reduces herbivore consumption of preferred species because the dispersion of less preferred species likely disturbs herbivore foraging. We conducted an experiment using three species with five combinations of clumped and random distribution patterns. Three species Lathyrus quinquenervius, Phragmites australis and Leymus chinensis, were of high, intermediate and low preferences by sheep, respectively. Results showed that distribution of low preferred species, but not that of high preferred one, affected the consumption of preferred species. Sheep obtained higher consumption of high preferred species when low preferred species followed a clumped distribution than a random distribution. Distance between aggregations of high and low preferred species did not affect sheep foraging. It was concluded that the effects of spatial distribution of preferred species on its consumption are dependent on herbivore foraging strategy, and sheep can consume more preferred species when there is a consistent spatial pattern between preferred species and the entire food resource, and that the random dispersion of low preferred species in grassland may reduce herbivore consumption of high preferred species, thus minimizing selective grazing.     

Flower colour variation across a hybrid zone in Antirrhinum as perceived by bumblebee pollinators

To assess if pollinators’ behaviour could explain the maintenance of hybrid zones between different flower colour morphs, we analyzed flower colour variation in an Antirrhinum hybrid zone using spectrometry and a model of bee perception. Some colours generated by hybridization were not observed in any Antirrhinum species and even appeared to be rare among angiosperms. Variation in flower colours within the hybrid zone was continuous; the most similar colours were predicted not to be discriminated from one another in natural foraging situations. However, when compared at a scale corresponding to bees’ foraging range, some flower colours could be discriminated from all colours displayed by neighbouring plants. This could affect pollinator behaviour and explain lower visitation rates within the centre of the hybrid zone. Behavioural studies involving bumblebees and plant mixtures of parental and hybrid flower colours carefully characterized with appropriate visual models will be necessary to test this hypothesis. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Lars Chittka     

Arthropod prey of imported fire ants (Hymenoptera: Formicidae) in Mississippi sweetpotato fields

The red imported fire ants, Solenopsis invicta (Buren), are generally considered pests. They have also been viewed as beneficial predators feeding on other insect pests of various agroecosystems. This study documents the foraging habits of fire ants in a sweetpotato field in Mississippi. Fire ant foraging trails connecting outside colonies to a sweetpotato field were exposed and foraging ants moving out of the field toward the direction of the colony were collected along with the solid food particles they were carrying. The food material was classified as arthropod or plant in origin. The arthropod particles were identified to orders. Fire ant foragers carried more arthropods than plant material. Coleoptera and Homoptera were the most abundant groups preyed upon. These insect orders contain various economically important pests of sweetpotato. Other major hexapod groups included the orders Hemiptera, Diptera and Collembola. The quantity of foraged material varied over the season. No damage to sweetpotato roots could be attributed to fire ant feeding. Imported fire ant foraging may reduce the number of insect pests in sweetpotato fields.     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Feeding of small rodents on seeds and fruits: a comparative analysis of three species of rodents of the<Emphasis Type="Italic">Araucaria</Emphasis> forest,southern Brazil

We investigated the feeding of three small-sized sigmodontine rodents (Oligoryzomys nigripes Olfers, 1818,Akodon montensis Thomas, 1913, andDelomys dorsalis Hensel, 1872) on fruiting plants in theAraucaria forest of southern Brazil. We offered to wild-captured animals fruits of 22 plant species in 14 families. For each individual, we recorded consumption patterns of pulp and seed. We also analyzed fecal samples to determine the relative abundance of invertebrates, fruit and seed parts, and fungi. The three rodents showed marked potential frugivory, feeding on most species offered. OnlyO. nigripes showed differences in relation to seed-size categories, consuming a high proportion of ‘small’ (diameter ≤2.5 mm) vs ‘large’ seeds. The rodents may be involved in seed dispersion of three plant species, by defecation of viable seeds of twoLeandra (Melastomataceae) species and also by removing the pulp ofMyrceugenia miersiana (Myrtaceae) fruits without swallowing or damaging their relatively large seeds, thus, potentially increasing germination rates ofM. miersiana. Diet analyses indicated no significant difference among proportions of seed/fruit in the fecal samples of the three species of rodents (33 to 41% of plant material). Our results suggest that the three focal species are the main small-mammal seed predators on the ground ofAraucaria forests.     

Increase of fast nutrient cycling in grassland microcosms through insect herbivory depends on plant functional composition and species diversity

Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear. In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers (Chorthippus parallelus) were added to the plant communities. Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO₄‐P, SO₄‐S) and pH in throughfall increased with herbivory. PO₄‐P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO₄‐P; frass may therefore be an important driver of throughfall pH and a main source of PO₄‐P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands.