Herbivory limits recruitment in an old-field seed addition experiment

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.     

Season- and herbivore-dependent competition and facilitation in a semiarid savanna

Empirical and theoretical evidence suggests that facilitation between plants, when it occurs, is more likely during periods of abiotic stress, while competition predominates under more moderate conditions. Therefore, one might expect the relative importance of competition vs. facilitation to vary seasonally in ecosystems characterized by pronounced dry (abiotically stressful) and wet (benign) seasons. Herbivory also varies seasonally and can affect the net outcome of plant-plant interactions, but the interactive effects of seasonality and herbivory on the competition-facilitation balance are not known. I experimentally manipulated neighboring plants and herbivory during wet and dry periods for two species of grass: Cynodon plectostachyus and Pennisetum stramineum, in the semiarid Laikipia District of Kenya. These experiments indicate that Pennisetum was competitively dominant during the wet season and that it responded negatively to grazing, especially during the dry season. Cynodon showed more complex season- and herbivore-dependent responses. Cynodon experienced facilitation that was simultaneously dependent on presence of herbivores and on dry season. During the wet season Cynodon experienced net competition. These results illustrate how herbivory and seasonality can interact in complex ways to shift species-species competition-facilitation balance. Additionally, because Cynodon and Pennisetum are key players in a local successional process, these results indicate that herbivory can affect the direction and pace of succession.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Differing Effects of Cattle Grazing on Native and Alien Plants

Abstract:   Habitat managers use cattle grazing to reduce alien plant cover and promote native species in California grasslands and elsewhere in the western United States. We tested the effectiveness of grazing as a restoration method by examining the effects of herbivory on native and alien plants. At Carrizo Plain National Monument, California, we surveyed native and alien species cover in adjacent grazed and ungrazed areas. We also established experimental plots in which plants were clipped or mulch (dead biomass) was removed. In addition, we clipped plants grown in pots and plants in the field that grew with and without competitors. Native species were negatively affected by clipping in 1999, 2000, and 2001, whereas alien species were unaffected. In the experimental field plots, the European annual forb Erodium cicutarium compensated in growth and reproduction following simulated herbivory. In contrast, growth and reproduction of the native perennial bunchgrass Poa secunda were reduced 1 year after clipping. In pots, E. cicutarium overcompensated and grasses undercompensated. In the field, European grasses were unaffected by the removal of competitors. It is unclear by what mechanism E. cicutarium was able to compensate, but the ability may be related to its basal rosette growth form and indeterminately growing inflorescences. The native California grassland community assembled in the absence of grazing herds, whereas invasive European species have been exposed to grazing for centuries. It may be that these invaders have adaptations that better enable them to recover from grazing. In the grassland we studied, the strategy of livestock grazing for restoration is counterproductive. It harms native species and promotes alien plant growth.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Effects of prey quantity on predatory wasps (<Emphasis Type="Italic">Polistes dominulus</Emphasis>) when patch quality differs

To determine the effects of prey quantity on central-place foraging of predatory wasps ( Polistes dominulus), prey of varying quality were distributed in patches. A field experiment was conducted, which controlled the amount and quality of prey available. 'Low-fed' colonies were provided with one-third the quantity of prey as that of 'high-fed' colonies. Both were provided with a 1:1 ratio of palatable:unpalatable prey. Experienced wasps of the high-fed colonies never selected unpalatable prey as their first choice of the day, whereas those of the low-fed colonies selected unpalatable prey as their first choice about 1:4 times. In general, wasps from the high-fed colonies reduced palatable patches to zero prey before exploiting unpalatable patches. Foundresses of high-fed colonies captured disproportionately more palatable prey than those of low-fed nests, but there was no correlation between ratio of palatable to unpalatable prey taken and the number of offspring produced. Wasps from low-fed colonies attacked unpalatable prey sooner, but not without considerable effort to avoid use of those patches. Foundresses of low-fed colonies also spent a greater proportion of time overall in unsuccessful search, which may explain why only wasps from low-fed colonies foraged on cool days. High-fed colonies produced more cells and more and heavier offspring than low-fed colonies. But the productivity of the low-fed colonies was greater than that of the 'natural' colonies, which had to find their own prey in a field-woodland area. These results indicate that prey scarcity changes foraging behavior and affects prey choice. These changes may not totally alleviate negative effects of unpalatable prey on colony development and offspring production. The results of this study increase understanding of the central-place foraging behavior of paper wasps, which are important biocontrol agents in natural and agricultural settings.     

Herbivory reduces plant interactions with above- and belowground antagonists and mutualists

Herbivores affect plants through direct effects, such as tissue damage, and through indirect effects that alter species interactions. Interactions may be positive or negative, so indirect effects have the potential to enhance or lessen the net impacts of herbivores. Despite the ubiquity of these interactions, the indirect pathways are considerably less understood than the direct effects of herbivores, and multiple indirect pathways are rarely studied simultaneously. We placed herbivore effects in a comprehensive community context by studying how herbivory influences plant interactions with antagonists and mutualists both aboveground and belowground. We manipulated early-season aboveground herbivore damage to Cucumis sativus (cucumber, Cucurbitaceae) and measured interactions with subsequent aboveground herbivores, root-feeding herbivores, pollinators, and arbuscular mycorrhizal fungi (AMF). We quantified plant growth and reproduction and used an enhanced pollination treatment to determine if plants were pollen limited. Increased herbivory reduced interactions with both antagonists and mutualists. Plants with high levels of early herbivory were significantly less likely to suffer leaf damage later in the summer and tended to be less attacked by root herbivores. Herbivory also reduced pollinator visitation, likely due to fewer and smaller flowers, and reduced AMF colonization. The net effect of herbivory on plant growth and reproduction was strongly negative, but lower fruit and seed production were not due to reduced pollinator visits, because reproduction was not pollen limited. Although herbivores influenced interactions between plants and other organisms, these effects appear to be weaker than the direct negative effects of early-season tissue loss.     

Pulses of dead periodical cicadas increase herbivory of American bellflowers

Resource pulses can have both direct bottom-up and indirect top-down effects on their consumers, but comparatively few studies have investigated the top-down effects of naturally occurring resource pulses on plants. This study describes two years of field experiments conducted to determine the indirect effects of 17-year periodical cicadas (Magicicada spp.) on herbivory in American bellflowers (Campanulastrum americanum). In 2004, the area of damaged leaves on cicada-supplemented plants was 78% greater than the area of damaged leaves on control plants. In 2005, cicada-supplemented plants were more likely to experience herbivory by mammalian herbivores than control plants. When large herbivores were excluded, similar patterns of leaf herbivory were observed, but these differences were not statistically significant. These results suggest that the pulsed input of dead periodical cicada bodies increased rates of herbivory on bellflowers, and that this effect was largely mediated by the selective foraging of large mammalian herbivores. More broadly, this study suggests that pulses of limiting resources can have both positive direct effects on plants and negative indirect effects due to selective herbivory, and that the net effects of pulsed resources on plants may depend on the composition and behavior of the surrounding herbivore community.     

Does colour matter? The importance of colour in avoidance learning, memorability and generalisation

Aposematic species exploit the ability of predators to associate, for example, conspicuous colouration with the unprofitability of prey. We tested the importance of colour for avoidance learning, memory and generalisation in wild-caught great tits (Parus major). First, we determined the birds’ initial colour preferences for red, yellow, orange and grey artificial prey items. The birds showed some preferences, as they were more willing to eat grey prey as their first choice, but these were not strong preferences. We then trained birds to discriminate red, yellow or variable (red and yellow) signals from grey where colours signalled palatable and unpalatable food. In general, the birds learned the discrimination task equally well, irrespective of which colours signalled unpalatability, and subsequently remembered the distinction between previously palatable and previously unpalatable colours in the memorability test. We did not find strong evidence that variability in the signal affected learning or memory. Our results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent. To study whether training also affects responses to unconditioned stimuli, we included orange prey items in the memorability test. Although orange had been palatable in the initial preference test, the birds ate fewer orange prey items after they had been trained to avoid red, yellow or both colours (variable signal) as unpalatable prey, but did not change their preference when trained that these colourful signals were palatable. This indicates that generalisation occurred more readily after a negative experience than a positive experience, a situation that would potentially allow imperfect mimicry to occur.     

Acute effects of removing large fish from a near-pristine coral reef

Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems.     

Trait similarity patterns within grass and grasshopper communities: multitrophic community assembly at work

Trait-based community assembly theory suggests that trait variation among co-occurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and (3) top-down control of grass trait variation. To test this, we collected data on the community composition and trait variation of grasses (41 species) and grasshoppers (53 species) in 50 plots in a South African savanna. Structural equation models were used to investigate how the range and spacing of within-community functional trait values of both grasses and their insect herbivores (grasshoppers; Acrididae) respond to (1) rainfall and fire frequency gradients and (2) the trait similarity patterns of the other trophic level. The analyses revealed that traits of co-occurring grasses became more similar toward lower rainfall and higher fire frequency (environmental control), while showing little evidence for top-down control. Grasshopper trait range patterns, on the other hand, were mostly directly driven by vegetation structure and grass trait range patterns (bottom-up control), while environmental factors had mostly indirect effects via plant traits. Our study shows the potential to expand trait-based community assembly theory to include trophic interactions.     

Facilitation drives local abundance and regional distribution of a rare plant in a harsh environment

The importance of facilitation to local community dynamics is becoming increasingly recognized. However, the predictability of positive interactions in stressful environments, the balance of competition and facilitation along environmental gradients, and the scaling of local positive interactions to regional distributions are aspects of facilitation that remain unresolved. I explored these questions in a habitat specialist, Delphinium uliginosum, and a moss, Didymodon tophaceus, both found in small serpentine wetlands. I tested three hypotheses: (1) moss facilitates germination, growth, and/or fecundity of D. uliginosum; (2) facilitation is stronger at the harsher ends of gradients in soil moisture, toxicity, and/or biomass; and (3) facilitation is reflected in positive associations at the levels of local abundance and regional occurrence. Although considerable competitive interactions occurred in later life stages, moss strongly facilitated D. uliginosum seedling emergence. There was no evidence that this facilitative effect weakened, or switched to competition, in benign environments. D. uliginosum was more locally abundant and more frequently present, across a large portion of its range, with than without moss, indicating a net facilitative effect in the face of competitive influences. Facilitated recruitment, possibly by seed retention, was found to be an important control on abundance and distribution in this rare species.     

Cascading effects of long-term land-use changes on plant traits and ecosystem functioning

There is much concern that the functioning of ecosystems will be affected by human-induced changes in biodiversity, of which land-use change is the most important driver. However, changes in biodiversity may be only one of many pathways through which land use alters ecosystem functioning, and its importance relative to other pathways remains unclear. In particular, although biodiversity-ecosystem function research has focused primarily on grasslands, the increases in agricultural inputs (e.g., fertilization, irrigation) and grazing pressure that drive change in grasslands worldwide have been largely ignored. Here we show that long-term (27-year) manipulations of soil resource availability and sheep grazing intensity caused marked, consistent shifts in grassland plant functional composition and diversity, with cascading (i.e., causal chains of) direct, indirect, and interactive effects on multiple ecosystem functions. Resource availability exerted dominant control over above-ground net primary production (ANPP), both directly and indirectly via shifts in plant functional composition. Importantly, the effects of plant functional diversity and grazing intensity on ANPP shifted from negative to positive as agricultural inputs increased, providing strong evidence that soil resource availability modulates the impacts of plant diversity and herbivory on primary production. These changes in turn altered litter decomposition and, ultimately, soil carbon sequestration, highlighting the relevance of ANPP as a key integrator of ecosystem functioning. Our study reveals how human alterations of bottom-up (resources) and top-down (herbivory) forces together interact to control the functioning of grazing systems, the most extensive land use on Earth.     

Herbivores and edaphic factors constrain the realized niche of a native plant

Biotic interactions, such as competition and herbivory, can limit plant species ranges to a subset of edaphically suitable habitats, termed the realized niche. Here we explored the role that herbivores play in restricting the niche of serpentine ecotypes of the native California annual Collinsia sparsiflora. We planted seeds from four populations into a range of natural field environments that varied in the presence/absence of naturally occurring C. sparsiflora and in predicted suitability for growth and survival of the serpentine ecotype of C. sparsiflora. Path analysis was then used to model the direct and herbivore-mediated indirect effects of environmental variables on the survival of C. sparsiflora serpentine ecotypes. We found that C. sparsiflora received more herbivory when planted into areas where serpentine ecotypes of C. sparsiflora were not predicted to persist, and that increased herbivory was associated with decreased survival, suggesting that herbivores may limit the distribution of C. sparsiflora serpentine ecotypes. Additionally, we demonstrated that edaphic environmental variables impacted the survival of C. sparsiflora serpentine ecotypes both directly and indirectly, by altering interactions with herbivores. These indirect effects were probably trait-mediated and probably occurred because edaphic factors may influence plant traits that, in turn, alter attractiveness to herbivores. Although the magnitude of direct effects exceeded the magnitude of indirect effects, many strong herbivore-mediated indirect effects were detected. Thus, interactions between the abiotic environment and insect herbivory contributed to restricting the niche of C. sparsiflora serpentine ecotypes to a subset of available habitat.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Do plant density, nutrient availability, and herbivore grazing interact to affect phlorotannin plasticity in the brown seaweed Ascophyllum nodosum

Plants have different strategies to cope with herbivory, including induction of chemical defences and compensatory growth. The most favourable strategy for an individual plant may depend on the density at which the plants are growing and on the availability of nutrients, but this has not been tested previously for marine plant–herbivore interactions. We investigated the separate and interactive effects of plant density, nutrient availability, and herbivore grazing on the phlorotannin (polyphenolic) production in the brown seaweed Ascophyllum nodosum. Seaweed plants grown at low or high densities were exposed either to nutrient enrichment, herbivorous littorinid gastropods (Littorina obtusata), or a combination of nutrients and herbivores in an outdoor mesocosm experiment for 2 weeks. Seaweeds grown at a low density tended to have higher tissue nitrogen content compared to plants grown at a high density when exposed to elevated nutrient levels, indicating that there was a density dependent competition for nitrogen. Herbivore grazing induced a higher phlorotannin content in plants grown under ambient, but not enriched, nutrient levels, indicting either that phlorotannin plasticity is more costly when nutrients are abundant or that plants responded to herbivory by compensatory growth. However, there were no significant interactive or main effects of plant density on the seaweed phlorotannin content. The results indicate that plants in both high and low densities induce chemical defence, and that eutrophication may have indirect effects on marine plant–herbivore interactions through alterations of plant chemical defence allocation.     

Taste-rejection by predators and the evolution of unpalatability in prey

Aposematic species advertise their unpalatability to potential predators using conspicuous warning colouration. The initial evolution of aposematism is thought to occur by warningly coloured mutants emerging in an already unpalatable cryptic species. However, possessing defence chemicals is often costly, and it is difficult to understand what the selective benefits might be for a mutation causing its bearer to be defended in a population of otherwise palatable cryptic prey. One solution to this problem is that chemically defended individuals are tasted and rejected by predators, and are, therefore, more likely to survive predatory attacks than undefended individuals. Using naïve domestic chicks Gallus gallus domesticus as predators and cryptic green chick crumbs as prey, we asked whether the accuracy with which birds discriminated between palatable and unpalatable prey was affected by the palatability of the unpalatable prey (moderately or highly defended), or their frequency in the population (10 or 25%). Birds could discriminate between green prey on the basis of their defences, and showed better discrimination between palatable and unpalatable prey when defended crumbs were highly unpalatable, compared to when they were moderately unpalatable. Although there was no detectable effect of the frequency of unpalatable prey in the population on predator taste-rejection behaviour in our main analysis, frequency did appear to affect the strategies that birds used in their foraging decisions when prey were only moderately unpalatable. How birds used taste to reject prey also suggests that birds may be able to monitor and regulate their chemical intake according to the frequency and defence levels of the unpalatable prey. Taken together, these results show that avian predators can generate selection for unpalatability in cryptic prey by sampling and taste-rejecting prey, but that a relatively large chemical difference between palatable and unpalatable prey may be necessary before unpalatable prey can enjoy a selective advantage. The exact nature of this evolutionary dynamic will depend on other environmental factors, such as defence costs and prey availability, but it provides a mechanism by which defences can evolve in a cryptic population.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

The community effects of phenotypic and genetic variation within a predator population

Natural populations are heterogeneous mixtures of individuals differing in physiology, morphology, and behavior. Despite the ubiquity of phenotypic variation within natural populations, its effects on the dynamics of ecological communities are not well understood. Here, we use a quantitative genetics framework to examine how phenotypic variation in a predator affects the outcome of apparent competition between its two prey species. Classical apparent competition theory predicts that prey have reciprocally negative effects on each other. The addition of phenotypic trait variation in predation can marginalize these negative effects, mediate coexistence, or generate positive indirect effects between the prey species. Long-term coexistence or facilitation, however, can be preceded by long transients of extinction risk whenever the heritability of phenotypic variation is low. Greater heritability can circumvent these ecological transients but also can generate oscillatory and chaotic dynamics. These dramatic changes in ecological outcomes, in the sign of indirect effects, and in stability suggest that studies which ignore intraspecific trait variation may reach fundamentally incorrect conclusions regarding ecological dynamics.     

喀斯特地区长期草地利用制度对群落稳定性的影响

草地利用方式和混播组合对群落特性的长期影响不同。利用方式对白三叶（Trifolium repens）盖度没有显著影响,而混播组合与利用方式的交互作用影响显著。白三叶在宿营处理和刈割为主利用下的与鸭茅（Dactylis glomerata）混播中的比例最低,而在刈割利用和宿营处理的多年生黑麦草（Lolium perenne）组合中的盖度最高。混播组合对禾本科组分盖度影响不显著,但利用方式和交互作用有显著的作用,长期放牧绵羊的混播草地多年生黑麦草很少。混播组合对杂草密度、盖度和多样性指数的影响不明显,而利用方式和交互作用显著地影响杂草盖度和多样性指数。放牧草地中的杂草密度高于刈割草地的,其中放牧利用的白三叶/多年生黑麦草草地最高。放牧牛的草地杂草密度较少,而放牧绵羊的草地杂草较多。白三叶、禾本科牧草以及杂草的密度都显著地受混播组合、利用方式及其交互作用的影响,说明密度是一个敏感的指标。