Spring grazing by sheep: effects on seasonal changes during early old field succession

This paper describes early secondary succession on an old field on limestone released from cultivation four years previously. Seasonal changes in plant composition after spring grazing by sheep are compared with those in ungrazed controls. Grazed and ungrazed paddocks were laid out in Latin squares. Plants were sampled before and several times after grazing in April, at several spatial scales.Major changes in plant abundance and sward characters such as height and density persisted throughout the growing season. Annual herbs increased after grazing, but annual grasses declined, as did short-lived perennial herbs. Effects on perennial herbs were weak; perennial grasses usually increased but this depended on the species. This pattern confirms that sheep grazing affects the direction, as well as the rate of succession.Some effects, such as increases in biennial herbs and in species richness, were only evident at large scales of sampling, suggesting that they arose from changes in rare and widely dispersed species. Other species were affected at different spatial scales, and no one sampling method detected the full range of effects.These results indicate the potential power of manipulating grazing early in secondary succession for directing the course of community change, for conservation or other purposes.Nomenclature follows Clapham et al. (1962)This study is funded by a grant from the Nuffield Foundation to V.K.B. We thank the Wytham Management Committee, Oxford University, for permission to use the site, and the staff of the University Farm, especially Richard Bampton and Cheryl Howes, for making the grazing possible. Alan Gange, Nicola marples, Fiona Matthews, Bridget Peace, Anne Storr and others helped at various times with field recording.     

Top-down effects of insect herbivores during early succession: influence on biomass and plant dominance

We tested the hypothesis that phytophagous insects would have a strong top-down effect on early successional plant communities and would thus alter the course of succession. To test this hypothesis, we suppressed above-ground insects at regular intervals with a broad-spectrum insecticide through the first 3 years of old-field succession at three widely scattered locations in central New York State. Insect herbivory substantially reduced total plant biomass to a similar degree at all three sites by reducing the abundance of meadow goldenrod, Solidago altissima. As a result, Euthamia graminifolia dominated control plots whereas S. altissima dominated insecticide-treated plots by the third year of succession. S. altissima is the dominant old-field herbaceous species in this region but typically requires at least 5 years to become dominant. Past explanations for this delay have implicated colonization limitation whereas our data demonstrate that insect herbivory is a likely alternative explanation. A widespread, highly polyphagous insect, the xylem-tapping spittlebug, Philaenus spumarius, appeared to be the herbivore responsible for the reduction in standing crop biomass at all three sites. Insect herbivory typically caused little direct leaf tissue loss for the ten plant species we examined, including S. altissima. Consequently, the amount of leaf area removed was not a reliable indicator of the influence of insect herbivory on standing crop biomass or on early succession. Overall, we found a strong top-down effect of insect herbivores on biomass at several sites, so our results may be broadly applicable. These findings run counter to generalizations that top-down effects of herbivores, particularly insects, are weak in terrestrial systems. These generalizations may not apply to insects, such as spittlebugs, that can potentially mount an effective defense (i.e., spittle) against predators and subsequently reach relatively high abundance on common plant species. Our results suggest that insect herbivory may play an important but often overlooked role during early old-field succession. Received: 26 December 1998 / Accepted: 3 April 1999     

Insect herbivory and plant adaptation in an early successional community*

To address the role of insect herbivores in adaptation of plant populations and the persistence of selection through succession, we manipulated herbivory in a long‐term field experiment. We suppressed insects in half of 16 plots over nine years and examined the genotypic structure and chemical defense of common dandelion (Taraxacum officinale), a naturally colonizing perennial apomictic plant. Insect suppression doubled dandelion abundance in the first few years, but had negligible effects thereafter. Using microsatellite DNA markers, we genotyped >2500 plants and demonstrate that insect suppression altered the genotypic composition of plots in both sampling years. Phenotypic and genotypic estimates of defensive terpenes and phenolics from the field plots allowed us to infer phenotypic plasticity and the response of dandelion populations to insect‐mediated natural selection. The effects of insect suppression on plant chemistry were, indeed, driven both by plasticity and plant genotypic identity. In particular, di‐phenolic inositol esters were more abundant in plots exposed to herbivory (due to the genotypic composition of the plots) and were also induced in response to herbivory. This field experiment thus demonstrates evolutionary sorting of plant genotypes in response to insect herbivores that was in same direction as the plastic defensive response within genotypes.     

Palatability, decomposition and insect herbivory: patterns in a successional old-field plant community

We tested the hypothesis that selective feeding by insect herbivores in an old‐field plant community induces a shift of community structure towards less palatable plant species with lower leaf and litter tissue quality and may therefore affect nutrient cycling. Leaf palatability of 20 herbaceous plant species which are common during the early successional stages of an old‐field plant community was assayed using the generalist herbivores Deroceras reticulatum (Mollusca: Agriolomacidae) and Acheta domesticus (Ensifera: Gryllidae). Palatability was positively correlated with nitrogen content, specific leaf area and water content of leaves and negatively correlated with leaf carbon content and leaf C/N‐ratio. Specific decomposition rates were assessed in a litter bag experiment. Decomposition was positively correlated with nitrogen content of litter, specific leaf area and water content of living leaves and negatively correlated with leaf C/N‐ratio. When using phylogentically independent contrasts the correlations between palatability and decomposition versus leaf and litter traits remained significant (except for specific leaf area) and may therefore reflect functional relationships. As palatability and decomposition show similar correlations to leaf and litter traits, the correlation between leaf palatability and litter decomposition rate was also significant, and this held even in a phylogenetically controlled analysis. This correlation highlights the possible effects of invertebrate herbivory on resource dynamics. In a two‐year experiment we reduced the density of above‐ground and below‐ground insect herbivores in an early successional old‐field community in a two‐factorial design by insecticide application. The palatability ranking of plants showed no relationship with the specific change of cover abundance of plants due to the reduction of above‐ or below‐ground herbivory. Thus, changes in the dominance structure as well as potentially associated changes in the resource dynamics are not the result of differences in palatability between plant species. This highlights fundamental differences between the effects of insect herbivory on ecosystems and published results from vertebrate‐grazing systems.     

Leaf miner assemblies: effects of plant succession and grazing management

Abstract. 1. Changes in leaf-miner assemblies during 4 years of secondary succession, under different controlled sheep-grazing treatments, are described and compared to the miner fauna of older grazed grassland nearby.
2. Multivariate analyses were used in conjunction with examination of individual common species to assess the independent effects of time, grazing treatment, plant species composition and architecture on the leaf-miner assemblies.
3. Leaf-miner species composition was strongly related to plant species composition, but was modified by plant structure under different grazing treatments. There was a strong successional trend in miner assemblies, even when the effects of changes in plant composition had been taken into account. Conversely, local variation in miner species composition generally reflected foodplant distribution alone.
4. Grazed treatments had fewer mines than controls, but there were also species specializing in grazed areas, despite the abundance of their foodplants elsewhere. There was a weak indication that miner species in grazed treatments were more likely to fluctuate in abundance than those in controls.
5. The results are discussed in relation to the assembly of grassland insect communities during succession, and the use of 'indicator groups' in management for nature conservation.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of mammal and insect herbivory on population dynamics of a native Californian thistle, Cirsium occidentale

We explored consequences of spatial and temporal heterogeneity in herbivory on the survival, growth, and reproduction of the Californian native dune thistle, Cirsium occidentale, in coastal and inland sites, for 2 years. We assessed the relative impacts of insect and mammalian herbivores and compared the relative importance of herbivory in coastal and inland habitats and among locations with different microclimates across a coastal dune. Effects of insect and mammalian herbivores were tested with a combination of insecticidal spray and cage exclusion treatments in a factorial experiment at the coastal site. Mammalian herbivores strongly affected the population dynamics of C. occidentale in both years, and their effects were augmented by fungal infection (1991), herbivory by stem-borers (1990) and, to a lesser extent, by insect seed predators in both years. Mammals caused most plant deaths, but the mammal species responsible differed among sites. Rabbit herbivory altered the vegetative growth of coastal thistles and significantly modified other key aspects of Cirsium demography, including growth rate and timing of reproduction. Small, uncaged plants grazed by rabbits took at least 1 year longer to mature than did caged plants. Larvae of Pyrausta subsequalis were the only insects that killed established plants. In 1990 and 1991, the numbers of insects damaging seed heads before dispersal were low, but were sufficient to cause receptacle and seed damage. The number of mature, undamaged seeds (and percent successful seed production) was reduced significantly only for heads infested by fungi near the ocean in 1991: the fungus occurred in 37% of heads and caused a 77% reduction in mature seeds. Received: 21 October 1996 / Accepted: 27 March 1997     

高原鼠兔对退化草地植物群落结构及恢复演替的影响

2009 年4 月至2011 年9 月采用围栏封育和去除法,对去除高原鼠兔的处理组和鼠兔扰动(对照组)区域植物群落结构、物种组成和地上生物量进行了比较分析,旨在阐明高原鼠兔对草地植物群落结构和群落演替的影响。结果表明:去除高原鼠兔处理组和对照区域植物群落平均高度和种盖度逐年上升,均在2011 年达到最大值,平均高度分别为6. 5 cm 和4. 2 cm,二者差异极显著(F =127. 80,df = 1,10,P < 0.01),种盖度分别为126. 5%和117. 9% ,差异不显著;植物群落组成中,处理区域禾草和莎草地上生物量比对照区域分别增加了1 965.1% 和33. 2% ,二者差异极显著(F = 41. 29,df = 1,10,P < 0.001),豆科植物和杂类草分别下降了89. 9% 和30.7% ;处理区域植物群落生态优势度发生了显著变化,由以杂类草为主改变为以禾草为主,而对照组均以杂类草为主,变化不明显。相似性指数2009 - 2011 年在对照区域年间变化范围为0.7168 - 0. 7550,在去除高原鼠兔处理区域年间变化范围为0.6464 - 0. 6732;对照区域和处理区间变化范围为0. 5354 - 0. 8956。根据试验结果我们认为,高原鼠兔的扰动可以有效降低植物群落的平均高度和植物种盖度,但植物种类组成主要以杂类草为主,年间变化不明显,表明鼠兔的扰动延缓了植物群落的恢复演替。     

Interacting effects of herbivory and fertility on a synthesized plant community

1 Outdoor microcosms were used to investigate the effects of invertebrate herbivory on plant community composition, and thereby infer possible effects on the rate of secondary succession, at differing levels of soil fertility.
2 A mixture containing 24 grassland plant species of widely different functional types was established, with 12 microcosms at each of three fertility levels. Four generalist herbivores ( Helix aspersa , Cepaea hortensis , Arianta arbustorum and Sitobion avenae ) were added to half of the microcosms. Above-ground biomass of each species was harvested after 2 years. Reproductive variables were also measured for one species, Poa annua .
3 At both moderate and high soil fertility generalist invertebrate herbivores fed selectively on early successional, fast-growing species, thus increasing the relative abundance of later successional, slow-growing species. This supports the hypothesis that herbivory increases the rate of secondary succession. Flowering and viable seed production of early successional ephemerals was also reduced by the invertebrate herbivores across a wide range of soil fertility. This would seriously reduce the ability of a species to persist in the community, thereby further hastening the rate of succession.     

Phenolics,nutrition and insect herbivory in some garrigue and maquis plant species

Summary Garrigue plant species growing on a calcareous substrate in southern France had higher foliar N levels than the same species growing on a relatively lower nutrient siliceous substrate (maquis). However maquis species had significantly higher foliar levels of P, more water, higher phenolic concentrations and larger leaf areas. The cumulative amount of insect damage on garrigue and maquis plants was similar, presumably due to different nutritional advantages in each case. Soil fertilization signifincantly elevated N levels in Q. coccifera, increased total leaf areas, decreased condensed tannin levels, and these leaves showed significantly more insect damage. Some effects of burning on Q. coccifera are also described. In these shrublands, fertilization may render leaf material more nutritional for herbivores by increasing nitrogen content and decreasing condensed tannin concentration, although very heavy grazing pressure may increase levels of leaf phenolics.     

Relationships between successional situation and production of miscanthus sinensis communities at Kirigamine Heights,Central Japan

Summary In the Miscanthus sinensis type grasslands of the Kirigamine Heights situated in a high altitude area (1,600 m) of Central Japan, the biomass in various successional phases was measured and the degree of succession (DS) was calculated. The maximum biomass during the growing season was used to compare the productivity of different grassland communities.The range and maximum point of DS in the DS-biomass curve of Miscanthus grasslands at Kirigamine was small compared with that of other Miscanthus grasslands in Japan.Between successional phases and production in the Miscanthus type grassland we found, that the total biomass increases, as succession proceeds to the maximum point. The of DS, and decreases after the maximum point. The biomass of Miscanthus sinensis changes as a normal curve against the axis of DS.Accordingly, in this Miscanthus type grassland, the production of the community decreases temporarily when the dominants of seral stages change. These results confirm Numata's (1966) model.Nomenclature followsA Handbook for Plant Collectors by H. Okuyama (1974).     

Effects of small mammal and invertebrate herbivory on plant species richness and abundance in tallgrass prairie

Summary A factorial field experiment was designed to test the effects of small mammals and above- and below-ground invertebrates on plant species richness and composition in native tallgrass prairie at Konza Prairie Research Natural Area, northeast Kansas. Over a 4-year period, Microtus ochrogaster densities were maintained by live-trapping in fenced plots, and invertebrate levels were reduced using the pesticides carbaryl for above-ground invertebrates and an organophosphate (isofenphos) for belowground invertebrates. ANOVA according to a split-plot design of plant species biomass data harvested in 1984 and 1986 revealed few significant effects of either small mammal densities or pesticide application. Of 54 species harvested from both sample dates, only 10 were significantly affected by either treatment. Analysis of species richness according to 8 life-form classes provided a clearer pattern of response than did biomass either by species or life-form class. For example, numbers of C4 grasses were reduced by increasing small mammal densities, whereas numbers of C4 annual forbs were lowest when above-ground herbivory was reduced. While consumers have been shown to have strong effects on successional communities, the few significant results observed in this study suggests that the manipulated levels of small mammals and insects had few effects on a mature tallgrass prairie.Deceased May, 1986     

Effects of root herbivory by an insect on a foliar-feeding species,mediated through changes in the host plant

Summary The effects of root herbivory by larvae of the scarabaeid, Phyllopertha horticola, on the growth of Capsella bursa-pastoris were examined. Individuals of Aphis fabae were reared on the leaves to determine what effect, if any, root feeding has on the performance of this insect. The experiment was conducted under two watering regimes (low and high). Low watering and root feeding caused water stress in the plants and this was reflected in a reduction in vegetative biomass and an increase in the proportion of material allocated to reproduction. Supplying plants with ample water in the high treatment enabled the water stress caused by root herbivory to be offset, but not completely overcome. Low watering and root feeding caused an increase in aphid weight and growth rate, while root feeding also increased fecundity and adult longevity. These effects are attributed to an improvement in food quality, measured by total soluble nitrogen, and caused by amino acid mobilization due to the water stress. The implications of these results in agricultural and ecological situations are discussed.     

Effects of plant-plant interactions and herbivory on the plant community structure in an arid environment of Saudi Arabia

There is currently considerable evidence support that plant community structures are driven by plant-plant interactions (e.g., competition and facilitation). In contrast, there is also evidence demonstrating that plant community structure is affected by the impact of consumer pressure (e.g., grazing). In this study, 15 and 10 Acacia gerrardii nurse plants were selected inside and outside Sudyrah natural reserve (protected) area in western Saudi Arabia, respectively. The understory vegetation abundance (e.g. cover and density) was measured among quadrats around the nurse trees in both protected and unprotected areas to examine the impact of grazing and the positive interaction on the understory species. I found that understory vegetation associated with nurse trees (A. gerrardii) has been driven by both the positive impact of nurse plant and the grazing. Although the understory vegetation was positively affected by the impact of facilitation, the composition of such vegetation has been changed due to the impact of herbivory.     

Grassland dynamics under sheep grazing in an Australian Mediterranean type climate

Grassland dynamics in a degraded disclimax grassland dominated by Danthonia caespitosa Gaudich. are examined using both demographic and multivariate approaches in an experiment designed to determine the effect of grazing intensity and exclosure on pasture dynamics. The experiment ran for 20 years from 1949 to 1968, using permanent quadrats at 3 grazing intensities and within exclosures. Demographic studies of some perennial grass species demonstrated markedly different responses to grazing; Danthonia caespitosa was unaffected by grazing but responsive to seasonal rainfall differences. Enteropogon acicularis survived only on protected sites. Numerical classification of total species set (121 species) for six observation periods demonstrated that community types were sensitive to differences in winter rainfall, and time since the start of experiment. Principal component analysis of permanent quadrat observations for individual years demonstrates quadrat trajectories which confirm this and indicate progressive divergence of the successional trends of the grazed and ungrazed quadrats. Repeated analysis on grazed quadrats only, shows that three components of pasture dynamics can be recognized; these are trend (succession?) and seasonal differences, each of which account for about 20% of the variance, and differences due to soil heterogeneity in the experimental paddock (8% of variance accounted for). No effect of grazing intensity was detected. Multivariate techniques can provide a clear partitioning of types of dynamic behaviour present in grassland communities. It is concluded that partitioning of environmental heterogeneity prior to demographic studies would increase their sensitivity.

     

Diversity of an early successional plant community as influenced by ozone

An early successional plant community was exposed to various ozone concentrations for two growing seasons (1994–1995) in open-top chambers in Auburn, Alabama, USA. The ozone treatments were: AA, ambient air (open plots); CF, carbon-filtered air ( c . 0·5×ambient air), 1×, non-filtered air, and 2×, twice ambient air. Vegetative canopy cover exhibited a pattern of accumulation in the spring, with maximum canopy cover attained in summer, then senescence of foliage in the autumn 1994. This pattern was not observed in 1995 as a result of a drought during the spring and summer. Varying ozone exposures caused shifts in the competitive interactions between plants, thereby altering community structure. Higher canopy cover, vertical canopy density (layers of foliage), species richness, diversity, and evenness existed in the CF treatments than in the other treatments. In addition, winged sumac ( Rhus copallina L.) became a major component of the CF treatments only during 1995. Surprisingly, blackberry ( Rubus cuneifolius Pursh.), a species considered ozone-sensitive, based on visible injury, dominated canopy cover within the 2× treatments, 41 and 33% of total canopy cover in 1994 and 1995, respectively. From these results it is concluded that plant communities existing in areas where lower ozone concentrations are prevalent might be more complex and diverse than those existing in areas with higher ozone concentrations.     

Conflicting management objectives on the Colorado Plateau: Understanding the effects of bison and cattle grazing on plant community composition

The American Bison (Bison bison Linnaeus) in the Henry Mountains are one of the last free-roaming, genetically pure herds of bison remaining in North America. Anecdotal evidence indicates that this herd is utilising a cattle winter range during the summer and fall, creating a conflict between the state agency that manages the bison, and the Bureau of Land Management (BLM) and local ranchers. In theory, the addition of bison grazing pressure could reduce forage availability in the short term and lead to undesired changes in the plant community in the long term. Our objective was to determine whether bison have altered the plant species composition of the cattle winter range. We characterised plant species composition, percent cover, and grazing intensity on three adjacent, geomorphologically similar mesas. Grazing regimes were different on the three mesas, one with bison and cattle present, one with cattle only present, and the third with neither cattle nor bison present. Vegetation surveys were accompanied by a 28-year remote sensing time series to test for temporal shifts in an index of primary productivity. We found a higher grazing intensity on two dominant forage species on the bison plus cattle grazed mesa in fall, before the cattle were turned out to winter pasture. Despite this difference in grazing intensity, we found few differences in species composition, percent cover, or NDVI across the three grazing regimes. Our results suggest that high intensity summer bison grazing, while likely creating short-term reductions in forage availability, has not caused differences in plant community composition or productive potential. Shifts in community composition can take years to unfold and just as long to correct; therefore, continued monitoring of the combined effects of cattle and bison is needed.     

山地土壤肥力与植物群落次生演替关系研究

从土壤物理、化学和生物性质出发,通过土壤肥力综合指标体系,利用多元统计方法,对重庆缙云山亚热带常绿阔叶林不同演替阶段山地土壤肥力的变化特征进行综合评价。并分析了缙云山亚热带常绿阔叶林群落演替与山地土壤肥力变化的关系。     

The effects of sheep grazing on seedling establishment and survival in grassland

This paper examines the effects of sheep and insect grazing on the generation of natural gaps in turf and the establishment of seedlings therein. The site is on an old field on limestone. There were five sheep grazing treatments: ungrazed, and grazed briefly in spring, briefly in autumn, in autumn or in summer and autumn. The first three of these treatments also contained plots in which insecticide was applied regularly during the growing season. Pins of increasing size (1.6 mm–12.8 mm diameter) were lowered at grid intersections in permanent quadrats to record gap type and the occurrence of seedlings on seven dates over 15 months.Heavily-grazed plots had more bare ground in spring and more herb seedlings established in them. Seedlings of annual species of Bromus were able to establish in gaps (bare ground and litter) of smaller size than was required by herb seedlings (which had much smaller seeds). The use of an insecticide increased seedling establishment in October in ungrazed and in spring-grazed paddocks but decreased it in autumn-grazed paddocks. This may indicate the greater importance of insect herbivory in making gaps in the matted sward of autumn-grazed plots.Seedlings which established in October 1986 and March 1987 were more likely to die than those establishing earlier in 1986. This may be because of winter-kill and especially vigorous spring vegetation growth respectively. Seedling mortality was not affected by gap size or grazing treatment (unless such effects operated before we detected the seedlings).