Grass competition is more important than seed ingestion by livestock for Acacia recruitment in South Africa

Although grazing livestock may have direct negative effects on woody species through herbivory and trampling, (heavy-)grazing is often associated with woody plant encroachment. Two main mechanisms can explain the positive effects of grazers on woody cover: (1) Grazers reduce the interspecific competition with trees and may reduce fuel load, and (2) gut passage through livestock increases seedling establishment by scarification (increased germination) and fertilization by dung (higher survival and growth of seedlings). We tested the effects of fire, grass, cattle ingestion (transit) and dung on germination, seedling height and survival as well as on recruitment of Acacia sieberiana in a sub-humid grassland of South Africa. About 8,000 seeds were planted in the field in a randomized block design. The removal of grass by grazing and/or fire had the most important effect on Acacia recruitment in savanna. Our findings highlight the hierarchy of the main factors affecting Acacia recruitment, which provides new insights to the understanding of woody plant encroachment.     

Herbivory effects on saplings are influenced by nutrients and grass competition in a humid South African savanna

Woody plant encroachment is a common consequence of disturbance in savannas. Grazers and browsers interfere with sapling establishment dynamics by direct consumption of plant tissue, changing soil nutrient status (through fertilization and trampling) and grass competition. Studies evaluating the effects of herbivory on sapling establishment have mostly been extrapolated from single species. In a controlled field experiment, we studied the effects of clipping (simulating grazing and browsing), nutrients, grass competition, and their interactive effects on sapling survival and growth of four dominant humid and four dominant mesic savanna species. We conducted this experiment in a humid South African savanna. We found no effects on sapling survival by the treatments provided. However, clipped saplings of all species increased their investment in relative growth rate of stem length (RGR_L). Clipping had a greater negative impact on relative growth rate of more humid than mesic species in terms of stem diameter (RGR_D), total dry biomass and proportion of leaf biomass. Nutrients had a positive effect on the RGR_L and sapling biomass of three mesic species. Positive effects of nutrients on RGR_L of one humid and two mesic species were observed in their clipped saplings only. Grass competition had a strong negative impact on all growth parameters measured. Clipped saplings of one humid and two mesic species had lower RGR_L with grass competition whereas intact saplings showed no significant response. After clipping, humid savanna species were more vulnerable to grass competition than mesic species, with reduced ability to use nutrients. In conclusion, herbivory increases sapling vulnerability to grass competition, with humid species being more susceptible than mesic species, indicating that woody-plant control strategies are more likely to be effective in humid savannas.     

Effects of species interactions on community organization along a trampling gradient

Abstract. Effects of interspecific interactions on the organization of two trampled communities, Eragrostio ferrugineae - Plantaginetum asiaticae (EP) and Eleusino indicae - Digitarietum violascentis (ED), were examined by 4-yr field experiments. We compared changes in the relative abundance of the main component species of the communities in monoculture and mixed culture along a trampling gradient. At no trampling in mixed culture, Ambrosia artemisiifolia var. elatior and Erigeron annuus (roadside herbs, RH) predominated and excluded the trampled community species. Severe trampling markedly reduced the predominance of these roadside herbs, promoted Eragrostis ferruginea (a perennial grass of EP), but suppressed Eleusine indica (an annual grass of ED). These results suggest that the differentiation of trampled and roadside herb communities under heavy trampling are caused by asymmetric competition between their main species. Several species pairs showed a constant rank order of relative cover at high trampling levels. Pennisetum alopecuroides (a perennial grass of both RH and EP) and Eragrostis coexisted, indicating symmetric competition. Eragrostis and Plantago asiatica (a perennial herb of EP) or Poa annua (a winter-annual grass of both EP and ED) coexisted through separation in phenology. Eragrostis promoted the survival of Plantago by moderating unpredictable drought (commensalistic positive interaction). This suggests that community composition is maintained by several interspecific interactions.     

Loss of a large grazer impacts savanna grassland plant communities similarly in North America and South Africa

Large herbivore grazing is a widespread disturbance in mesic savanna grasslands which increases herbaceous plant community richness and diversity. However, humans are modifying the impacts of grazing on these ecosystems by removing grazers. A more general understanding of how grazer loss will impact these ecosystems is hampered by differences in the diversity of large herbivore assemblages among savanna grasslands, which can affect the way that grazing influences plant communities. To avoid this we used two unique enclosures each containing a single, functionally similar large herbivore species. Specifically, we studied a bison (Bos bison) enclosure at Konza Prairie Biological Station, USA and an African buffalo (Syncerus caffer) enclosure in Kruger National Park, South Africa. Within these enclosures we erected exclosures in annually burned and unburned sites to determine how grazer loss would impact herbaceous plant communities, while controlling for potential fire-grazing interactions. At both sites, removal of the only grazer decreased grass and forb richness, evenness and diversity, over time. However, in Kruger these changes only occurred with burning. At both sites, changes in plant communities were driven by increased dominance with herbivore exclusion. At Konza, this was caused by increased abundance of one grass species, Andropogon gerardii, while at Kruger, three grasses, Themeda triandra, Panicum coloratum, and Digitaria eriantha increased in abundance.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Grazing lawns contribute to the subsistence of mesoherbivores on dystrophic savannas

Resource manipulation, such as the creation and maintenance of grazing lawns, may shape the structure of herbivore communities. We tested the hypothesis that grazing lawns contribute towards the subsistence of the Kobus kob kob in a dystrophic West African savanna, where kob and Hippopotamus amphibius both occur. Comparison of the foliage of grazing lawns and ungrazed swards shows that hippo lawns are more nutritious with regard to both structure and nutrients; kob lawns are higher in nutrients only. Up to the early dry season, hippo lawns meet kob energy and protein demand, thereafter, the shortness of the sward limits intake. Kob lawns always provide sub-maintenance values. Grazing on ungrazed swards is least profitable. We suggest that grazing lawns are essential for the daily subsistence of mesoherbivores, particularly on nutrient-poor soil, and that megaherbivores facilitate their food supply, for at least part of the year.     

Disturbance and the (surprising?) role of ecosystem engineering in explaining spatial patterns of non‐native plant establishment

Different conceptions of disturbance differ in the degree to which they appeal to mechanisms that are general and equivalent, or species‐, functional group‐, or interaction‐specific. Some concepts of disturbance, for example, predict that soil disturbances and herbivory have identical impacts on species richness via identical mechanisms (reduction in biomass and in competition). An alternative hypothesis is that the specific traits of disturbance agents (small mammals) and plants differentially affect the richness or abundance of different plant groups. We tested these hypotheses on a degu (Octodon degus) colony in central Chile. We ask whether native and non‐native forbs respond differently to degu bioturbation on runways versus herbivory on grazing lawns. We ask whether this can explain the increase in non‐native plants on degu colonies. We found that biopedturbation did not explain the locations of non‐native plants. We did not find direct evidence of grazing increasing non‐native herbs either, but a grazing effect appears to be mediated by grass, which is the dominant cover. Further, we provide supplementary evidence to support our interpretation that a key mechanism of non‐native spread is the formation of dry soil conditions on grazing lawns. Thus, ecosystem engineering (alteration of soil qualities) may be an outcome of disturbances, in which each interacts with specific plant traits, to create the observed pattern of non‐native spread in the colony. Based on these results, we propose to extend Jentsch and White (Ecology, 100, 2019, e02734) concept of combined pulse/ disturbance events to the long‐term process duality of ecosystem engineering/ disturbance.     

Herbaceous Forage and Selection Patterns by Ungulates across Varying Herbivore Assemblages in a South African Savanna

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.     

Biomass allocation in herbaceous plants under grazing impact in the high semi-arid Andes

Among the plant traits that affect performance, vitality and herbivore resistance in rangeland vegetation, biomass partitioning ranks top, commonly outweighing processes at single leaves (e.g., photosynthesis, respiration). We explored the allometry of a broad sample of herbaceous species from two high elevation sites in the Andes in order to explain biomass partitioning under harsh climatic conditions and risk of biomass losses under strong camelid grazing pressure. We combined data from NW-Argentina and W-Bolivia from elevations between 4200 and 4250 m in a landscape dominated by tall Festuca orthophylla tussocks and a drought driven seasonality (rainfall only between November and March). Across 10–20 taxa per region we found less investment in leaves with a mean leaf mass fraction of only 11% and instead a massive storage in below-ground compartments (rhizomes, tap roots), particularly at the colder Argentinean site. Though grazing pressure was much greater in Bolivia, the foliage mass fraction was there larger than in Argentina. The inter-tussock space in these open, dry plains (‘pajonal’) was dominated by rosette forming species with a below-ground shoot apex and massive tap roots (70% of all species), rendering these species less sensitive to grazing and trampling. The storage organs of these species represented more than 50% of total biomass. Llamas, which represented the main vertebrate herbivore in these open plains at the Bolivian site, preferred non tap-root herbs and species with low leaf nitrogen concentration. Palatable forbs for llamas (22% of all species at the Bolivian site) only survived when nested (facilitated) in the rigid, tall Festuca orthophylla tussocks or thorny shrubs. In conclusion, these extremely high elevation rangeland herbs invest preferentially in structures for persistence (K-strategy) rather than maximizing carbon gain.     

Habitat preference of the striped legless lizard: Implications of grazing by native herbivores and livestock for conservation of grassland biota

Across the globe, many species of reptile are threatened with extinction, with changes in grazing pressure as a significant factor in their decline. Few studies have investigated the role of native herbivores, yet studying natural grazers may provide insight into natural grazing regimes, not apparent in studies of domestic livestock. In this study, we investigate the habitat requirements of a threatened Australian grassland reptile, the striped legless lizard, Delma impar, in grasslands grazed by a native herbivore, the eastern grey kangaroo Macropus giganteus. Delma impar appears sensitive to habitat change resulting from altered grazing intensity, but a lack of information hinders implementation of appropriate grazing regimes. To address this gap, we investigated habitat preferences of D. impar at multiple spatial scales across a grazing gradient. We found that the occurrence of D. impar was not affected by the size of grassland remnants, but was negatively related to the density of native grazers. This result was likely a consequence of the negative effect of high grazing intensity on grass structural complexity, as the probability of encountering a D. impar was positively related to grass structural complexity at the fine scale (1 m²). We recommend that conservation efforts should avoid high intensity grazing (equivalent to > 1.3 kangaroos/ha), yet ensure enough grazing disturbance is maintained to promote the formation of complex grass structures. We also recommend that small floristically degraded and fragmented grassland habitat should be included in conservation efforts. These recommendations will likely benefit a number of faunal species in grasslands grazed by domestic and native grazers. Importantly, our data highlight the need for managing grazing regimes, even in environments dominated by native herbivores.     

Herbivore and nutrient control of lawn and bunch grass distributions in a southern African savanna

In African savannas, bottom-up soil nutritional factors and top-down herbivory have both been suggested to control the distribution patterns of bunch and lawn grasses. We tested the separate and combined roles of these factors on grassland distribution in a South African savanna by focusing on three main objectives, namely (1) are grazing patches and lawns restricted to specific soils or sites, (2) does herbivory alter rates of nutrient cycling which facilitates lawns and (3) are there any differences in foliage quality between lawn and bunch grass-dominated sites that might influence animals in choosing to feed on lawns? We set up ten sites along a grazing gradient in the Hluhluwe-iMfolozi Park in KwaZulu-Natal, South Africa. At these sites, we measured total soil nutrients, soil nutrient turnover rates and grass foliage nutrient concentrations. We found that in the Hluhluwe-iMfolozi Park the spatial distribution of lawn and bunch grassland types appears to be an animal-driven phenomenon and not dependent on specific soil properties. The short-structured and distinct species found in lawns do not appear to be restricted to nutrient-enriched patches. However, the grasses of these lawns had significantly higher nutrient concentrations, in their foliage, which might explain the high attraction to these patches by herbivores. We also did not find any animal-induced stimulation of nutrient cycling rates that are often associated with lawn grass species.     

Ecology of grazing lawns in Africa

Grazing lawns are a distinct grassland community type, characterised by short‐stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co‐evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low‐biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall‐grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short‐grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall‐grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.     

Tracking nutrients in space and time: Interactions between grazing lawns and drought drive abundances of tallgrass prairie grasshoppers

1. We contrast the response of arthropod abundance and composition to bison grazing lawns during a drought and non‐drought year, with an emphasis on acridid grasshoppers, an important grassland herbivore.
2. Grazing lawns are grassland areas where regular grazing by mammalian herbivores creates patches of short‐statured, high nutrient vegetation. Grazing lawns are predictable microsites that modify microclimate, plant structure, community composition, and nutrient availability, with likely repercussions for arthropod communities.
3. One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
4. We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
5. Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.

     

Morphology and growth of stolons and rhizomes in three clonal grasses,as affected by different light supply

In this paper, the hypothesis is tested that, in clonal grasses producing stolons and/or rhizomes, stolons always show a higher morphological plasticity than rhizomes in response to variation in light availability.Agrostis stolonifera (a stoloniferous grass),Holcus mollis (a rhizomatous grass) andCynodon dactylon (a grass forming both stolons and rhizomes), were grown in pots and subjected to three levels of light intensities. Both stolons and rhizomes branched more intensively under higher light levels. Irrespective of species, stolons consisted of longer internodes under lower light levels, while rhizome morphology did not respond significantly. Biomass partitioning to rhizomes was lower under lower light intensities while partitioning to stolons was not affected. Rhizomes usually had more dormant buds than did stolons. Our results suggest that stolons serve primarily as foraging organs for light, whereas the main function of rhizomes is storage of meristem and carbohydrates, irrespective of whether the grass species involved produces both rhizomes and stolons or only one type of spacer.     

Differential responses of three grasses to defoliation,water and light availability

Grazing not only modifies the structure and functioning of grasslands, it also changes micro-environmental conditions that alter the availability of resources. The aim of this study was to analyze the response of grasses with different photosynthetic pathways (C₃/C₄), growth forms (prostrate/erect), and grazing responses (increaser/decreaser) to defoliation and resource availability. In a greenhouse, we performed a factorial experiment with three factors: defoliation, light, and water and three species: Axonopus affinis (C₄ prostrate, increaser), Coelorachis selloana (C₄ erect, decreaser), and Bromus auleticus (C₃ erect, decreaser). We measured the relative growth rate (RGR), biomass assignment, and specific leaf area. The RGR of both C₄ species was affected by light availability, while the decreaser C₃ did not respond to any factor. Biomass allocation to leaves and stolons changed with the interaction between light and water in the C₄ prostrate species (increaser). In the C₄ erect grass (decreaser), biomass allocation was more affected by defoliation under low levels of light and water. Low light availability and defoliation reduced the assignment to leaves, while the allocation to rhizomes increased. Species-specific responses to resources availability that are modified by grazing were related to photosynthetic pathway, growth form, and grazing responses. Biomass allocation was related to strategies to avoid and/or tolerate grazing. The investment to leaves was limited by light and water availability in prostrate species, while in erect grasses it was controlled by defoliation and water availability. Our results highlight the importance of species responses to changes in resource availability associated to grazing regimes.     

The Effects of Seed Ingestion by Livestock,Dung Fertilization,Trampling, Grass Competition and Fire on Seedling Establishment of Two Woody Plant Species

The increasing rate of woody plant encroachment in grasslands or savannas remains a challenge to livestock farmers. The causes and control measures of woody plant encroachment are of common interest, especially where it negatively affects the objectives of an agricultural enterprise. The objectives of this study were to determine the effects of gut passage (goats, cattle), dung (nutrients), fire, grass competition and trampling on establishment of A. nilotica and D. cinerea seedlings. Germination trials were subjected to the following treatments: 1) seed passage through the gut of cattle and goats and unpassed/ untreated seeds (i.e. not ingested), 2) dung and control (no dung), 3) grass and control (mowed grass), 4) fire and control (no fire), 5) trampling and control (no trampling). The interaction of animal species, grass and fire had an effect on seedling recruitment (P < 0.0052). Seeds retrieved from goats and planted with no grass and with fire (6.81% ± 0.33) had a significant effect on seedling recruitment than seeds retrieved from goats and planted with grass and no fire (2.98% ± 0.33). Significantly more D. cinerea and A. nilotica seeds germinated following seed ingestion by goats (3.59% ± 0.16) than cattle (1.93% ± 0.09) and control or untreated seeds (1.69% ± 0.11). Less dense grass cover, which resulted in reduced grass competition with tree seedlings for light, space and water, and improved seed scarification due to gut passage were vital for emergence and recruitment of Acacia seedlings. These results will contribute considerably to the understanding of the recruitment phase of woody plant encroachment.     

Dynamics of grazing lawn formation: an experimental test of the role of scale‐dependent processes

Grazing lawns are characteristic for African savanna grasslands, standing out as intensely grazed patches of stoloniferous grazing‐tolerant grass species. Grazing lawn development has been associated with grazing and increased nutrient input by large migratory herds. However, we argue that in systems without mass migrations disturbances, other than direct grazing, drive lawn development. Such disturbances, e.g. termite activity or megaherbivore middens, also increase nutrient input and keep the bunch vegetation down for a prolonged time period. However, field observations show that not all such disturbances lead to grazing lawns. We hypothesize that the initial disturbance has to be of a minimal threshold spatial scale, for grazing intensity to be high enough to induce lawn formation. We experimentally tested this idea in natural tall savanna grassland. We mowed different‐sized plots to simulate initial disturbances of different scales (six times during one year) and applied fertilizer to half of the plots during two years to simulate increased nutrient input by herbivores or termite activity. Allowing grazing by naturally occurring herbivores, we followed the vegetation development over more than three years. Grazing kept bunch grass short in coarser, fertilized plots, while grasses grew out toward their initial height in fine‐scale and unfertilized plots. Moreover, lawn grasses strongly increased in cover in plots with an increased nutrient input but only after coarser scale disturbance. These results support our hypothesis that an increased nutrient input in combination with grazing indeed induces grazing lawn formation, but only above a threshold scale of the initial disturbance. Our results provide an alternative mechanism for the development of grazing lawns in systems that lack mass migrating herds. Moreover, it gives a new spatial dimension to the processes behind grazing lawn development, and hence help to understand how herbivores might create and maintain spatial heterogeneity in grassland systems.     

Phenotypic and Genotypic Analysis of Newly Obtained Interspecific Hybrids in the Campanula Genus

Interspecific hybridisation creates new phenotypes within several ornamental plant species including the Campanula genus. We have employed phenotypic and genotypic methods to analyse and evaluate interspecific hybridisation among cultivars of four Campanula species, i.e. C. cochleariifolia, C. isophylla, C. medium and C. formanekiana. Hybrids were analysed using amplified fragment length polymorphism (AFLP), flow cytometry and biometrical measurements. Results of correlation matrices demonstrated heterogeneous phenotypes for the parental species, which confirmed our basic premise for new phenotypes of interspecific hybrids. AFLP assays confirmed the hybridity and identified self-pollinated plants. Limitation of flow cytometry analysis detection was observed while detecting the hybridity status of two closely related parents, e.g. C. cochleariiafolia × C. isophylla. Phenotypic characteristics such as shoot habitus and flower colour were strongly influenced by one of the parental species in most crosses. Rooting analysis revealed that inferior rooting quality occurred more often in interspecific hybrids than in the parental species. Only interspecific hybrid lines of C. formanekiana ‘White’ × C. medium ‘Pink’ showed a high rooting level. Phenotype analyses demonstrated a separation from the interspecific hybrid lines of C. formanekiana ‘White’ × C. medium ‘Pink’ to the other clustered hybrids of C. formanekiana and C. medium. In our study we demonstrated that the use of correlation matrices is a suitable tool for identifying suitable cross material. This study presents a comprehensive overview for analysing newly obtained interspecific hybrids. The chosen methods can be used as guidance for analyses for further interspecific hybrids in Campanula, as well as in other ornamental species.     

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Differential herbivory tolerance of dominant and subordinate plant species along gradients of nutrient availability and competition

We tested whether differences in the herbivory tolerance of plant species is related to their abundance in grassland communities and how herbivory and nutrient availability affect competitive balances among plant species through changes in their tolerance. The experimental approach involved a simulated grazing treatment (clipping) of two competitive grass species (Arrhenatherum elatius and Holcus lanatus) and two subordinate forb species (Prunella vulgaris and Lotus corniculatus) along a gradient of nutrient availability and under conditions of competition. Total standing, aboveground, root, and regrowth biomass were evaluated at the end of the experiment as an estimate of the capacity to compensate for twice removing aboveground biomass at different nutrient levels (NPK). Although clipping had a more pronounced negative effect on dominant plant species (Arrhenatherum and Holcus) than on subordinate species, the negative effects on dominant species were offset by the application of fertilizer. The combined effect of fertilizer and competition had more negative effects on the performance of Lotus and Prunella than on the dominant species. In terms of competition, the regrowth ability of Arrhenatherum and Holcus increased with the application of fertilizer, while the opposite pattern was observed for Lotus and Prunella. The addition of fertilizer has a positive effect on both grass species in terms of growth in clipped pots and competition, while subordinate species did not respond to the addition of fertilizer to the clipped pots and were negatively affected by competition with both grass species. The results suggest (1) that species replacement towards subordinate species as a function of herbivory is partially dependent on the herbivory tolerance of that species, (2) competitive relations between competitive grass species and subordinate forb species change under different environmental conditions, and (3) although grazing disturbance significantly influences competitive relations in favor of less competitive species, increasing nutrient levels counteract the negative effect of grazing on dominant competitive plant species.