Influence of herbivory and abiotic factors on the distribution of tall forbs along a productivity gradient: a transplantation experiment

In arctic-alpine areas tall herb vegetation is restricted to sites with high productivity. At higher elevation, low prostate forbs and narrow-leaved graminoids dominate the vegetation in sites with a protecting snow cover during winter. In this study, I test whether herbivory or abiotic factors prevent tall forbs from growing at higher altitudes. Vegetation blocks from a tall herb meadow were transplanted to herbivore exclosures and open plots in a low-productive snowbed and a productive tall herb meadow. The tall forbs performed equally well in the exclosures on the low-productive snowbed as in the tall herb meadow, but decreased in the open plots on the low-productive snowbed. Thus, even if abiotic factors are ultimately causing many of the vegetation patterns observed in arctic-alpine plant communities, herbivory appears to be the main proximate factor responsible for the decreasing abundance of tall forbs along gradients of decreasing productivity in arctic-alpine areas.     

Long-term effects of grazing on subalpine and alpine grasslands in the Central Alps,Austria

The effects of grazing exclusion on species diversity and functional diversity were analyzed along an elevation gradient from the subalpine (1960 m a.s.l.) to the lower and upper alpine zone (2275 m–2650 m a.s.l.) in the Austrian Central Alps for 15 years. Nine sites were chosen, including grasslands at different elevations, a bog and a glacier foreland site at the lower alpine zone and a snowbed at the upper alpine zone. Data were acquired by frequency counts in 1 m² permanent plots inside each fenced area (three plots per site) and outside in grazed areas (three plots). Diversity indices and functional diversity were analyzed by means of generalized linear mixed models (GLMMs). Exclosure, duration of exclusion and exclosure*years (interaction effect) were defined as predictor variables. Multivariate ordination techniques were used to (i) determine species responses to grazing exclusion (pRDA, partial redundancy analysis) and (ii) to create a distance matrix representing the changes between exclosures and control plots per year (NMDS, non-metric multidimensional scaling). At the subalpine grassland, first differences between exclosure and control plots occurred already only after three years, at the upper alpine zone after four and five years. Contrary to our expectation, dwarf shrubs did not increase within the exclosures of the subalpine grassland. Instead, mainly the tall forb Geranium sylvaticum increased. Species richness significantly decreased at the exclosures of the subalpine zone, the snowbed and at one upper alpine grassland sites. The communities of the glacier foreland and the bog were hardly affected by grazing exclusion.We conclude that plant species and communities react individually depending on elevation and grazing animals. Grazing exclusion studies at high elevations should definitely be carried out in the long-term.     

Influence of habitat characteristics on small mammals in a Mexican high-altitude grassland

Small mammals in a high-altitude grassland area close to Mexico City were studied. Populations of 10 species were censused using live traps in 48 sample quadrats. Within each quadrat, vegetation characterization, including complete floristic listings, cover values for species and layers and values of habitat modification, were assessed. Habitats were described according to plant communities identified using ordination and classification methods. Nine different plant communities were obtained. Densities and abundance of all small mammal species were calculated for each of the habitats classified. Peromyscus alsloni was the most abundant species in all habitats, reaching maximum densities of 55 ha⁻¹ in pine forest with dense ground and herb layer. Peromyscus melanotis also occurred in all habitats but at lower densities (maximum 29 ha⁻¹). Reithrodontomys megalotis was found in all habitats except in tall dense grassland. Densities for this species were generally low (1-9 ha⁻¹) but reached 19 ha⁻¹ in short dense grassland. All other species were largely absent from 4–8 habitats and showed very low densities (0.75–4 ha⁻¹). The densities of the more abundant species were largely correlated with more open habitats and higher indices of habitat modification. Lower altitude grassland habitats have a greater abundance of small mammals and a higher species richness than the medium and higher altitude, physiognomically more complex habitats. Species richness was highest in tall pine-alder forest with a species-rich, dense herb layer and lowest in pine forest with dense ground and herb layers. Species richness was positively correlated with overall small mammal density.     

Summer grazing by voles and lemmings upon subarctic snow-bed and tall herb meadow vegetation – an enclosure experiment

Ther hypothesis of the exploitation of ecosystems predicts that plant communities in productive habitats should experience a low grazing pressure, while communities in less productive habitats should be structured by intense grazing. Thus, if a high population density of rodents was to be established in these communities and they could graze without predation, the plant community in the productive habitat should be more affected than the community in the less productive habitat. This was tested by building enclosures for grey-sided voles Clethrionomys rufocanus and lemmings Lemmus lemmus in a productive tall herb meadow and a less productive snow-bed on Finnmarksvidda, northern Norway. The simulated rodent population densities varied from 150 to 750 ind. ha^-1, and the rodents were allowed to graze for 18 to 55 d. The shoot mortality of relatively abundant plant species were monitored together with plant community structure, and the above-ground biomass were harvested at the end of the experiment. The shoot mortality were generally low, the significantly grazed species were either known preferred species or species with a grazing-sensitive morphology. The community structure did not show any major changes. The aboveground biomass of woody plants was significantly lower in the enclosures on the snow-bed, but not in the tall herb meadow. The total biomass did not differ significantly. These results are somewhat ambiguous with reference to the predictions from the food chain hypothesis, and the conclusion were drawn that intense grazing by voles and lemmings during 18 to 55 d during the growing season does not seem to be sufficient to greatly influence shoot mortality and structure of the studied plant communities.     

Local and regional processes in low-productive mountain plant communities: the roles of seed and microsite limitation in relation to grazing

It is becoming widely accepted that plant community structure is determined not only by local scale factors, but that regional factors may play considerable role. The research studying the associated processes in different environments with different species assemblages is still limited. We conducted a two-year seed sowing experiment to test whether a plant community in a low-productive mountain snowbed is limited by seed or microsite availability and how these variables depend on natural grazing. In a factorial design, half of the plots received a mixture of seeds of fourteen species naturally occurring at the study site and above ground biomass was removed from half of the plots. These treatments were applied to plots with long term grazer exclosures and to plots accessible to grazers. Both sowing and biomass removal increased the number of seedlings, the species richness of seedlings and total species richness. The number of seedlings was higher in open plots than in exclosures in the second year. Both seedling richness and total species richness were higher in open plots. Seedling recruitment was negatively related to the amount of above ground biomass and positively to the initial species richness. These results suggest that even fairly low-productive environments can be both seed and microsite limited and that these depend on grazing pressure. Natural grazing by mammal herbivores (e.g. lemmings and reindeer) favours species colonization and seedling emergence. Low-productive mountain snowbeds are prone to colonization from the local species pool and even high species richness may not constrain ingression of new species.     

The interacting effects of herbivore exclosures and seed addition in a wet meadow

Seed limitation may prevent successful restoration of native plant communities. Seed addition is a common restoration practice but the role of small mammals in affecting seedling recruitment is not well understood. The purpose of this investigation was to test the relative effect of seed introduction in combination with small mammal and bird exclosures in an Ohio wet meadow. We ask whether the ambient population of Microtus pennsylvanicus (1) alters species composition (e.g. forb/grass/sedge, invasive, non-native); (2) influences plant diversity; and (3) reduces the effect of increasing local plant richness through seed introductions. We established a 2×2 factorial design including a seed addition treatment (0 and 20 seed species added) and an exclosure treatment (open and fenced to exclude all mammalian and bird herbivores and granivores). Seeds from twenty native species were selected to represent a broad range of plant life forms typically found in temperate eastern North American wet meadow communities. All species were obligate or facultative wetland species with forbs, grasses and sedges represented. We found that forb species increased inside exclosures, especially in the seed addition treatment. We also found that relative biomass of invasive species was reduced in exclosures and with seed addition. Species richness increased with seed addition; however, exclosures significantly increased species richness and diversity, particularly of those species that were experimentally introduced by seed. Our results support the seed limitation hypotheses. It is also evident that seed and seedling predation are important factors that can control wet meadow community composition and diversity.     

Progressively excluding mammals of different body size affects community and trait structure of ground beetles

Mammalian grazing induces changes in vegetation properties in grasslands, which can affect a wide variety of other animals including many arthropods. However, the impacts may depend on the type and body size of these mammals. Furthermore, how mammals influence functional trait syndromes of arthropod communities is not well known. We progressively excluded large (e.g. red deer, chamois), medium (e.g. alpine marmot, mountain hare), and small (e.g. mice) mammals using size‐selective fences in two vegetation types (short‐ and tall‐grass vegetation) of subalpine grasslands. We then assessed how these exclusions affected the community composition and functional traits of ground beetles (Coleoptera, Carabidae), and which vegetation characteristic mediated the observed effects. Total carabid biomass, the activity densities of carabids with specific traits (i.e. small eyes, short wings), the richness of small‐eyed species and the richness of herbivorous species were significantly higher when certain mammals were excluded compared to when all mammals had access, regardless of vegetation type. Excluding large and medium mammals increased the activity density of herbivorous carabid species, but only in short‐grass vegetation. Similarly, excluding large mammals (ungulates) altered carabid species composition in the short‐, but not in the tall‐grass vegetation. All these responses were related to aboveground plant biomass, but not to plant Shannon diversity or vegetation structural heterogeneity. Our results indicate that changes in aboveground plant biomass are key drivers of mammalian grazers’ influence on carabids, suggesting that bottom–up forces are important in subalpine grassland systems. The exclusion of ungulates provoked the strongest carabid response. Our results, however, also highlight the ecological significance of smaller herbivorous mammals. Our study furthermore shows that mammalian grazing not only altered carabid community composition, but also caused community‐wide functional trait shifts, which could potentially have a wider impact on species interactions and ecosystem functioning.     

Beyond the browse line: complex cascade effects mediated by white‐tailed deer

In many ecosystems, browsing of large mammals can affect plant species compositions. However, much less is known about potential above‐ and below‐ground trophic interactions of large browsing mammals. This study focused on the direct and indirect effects of browsing on trophic and abiotic interactions within forest ecosystems. To quantify above‐ and below‐ground cascade effects, white‐tailed deer have been excluded for over 18 years from three 4‐ha plots, which were paired with same sized deer access plots. Our results demonstrate complex direct and indirect cascade effects on forest food webs. Deer exclusion directly altered woody species composition and significantly increased shrub and sapling density. Above‐ground cascade effects include greater leaf litter accumulation and higher arthropod density and biomass within the exclosures. Below‐ground indirect effects include significant decrease in soil nutrients, and higher arbuscular mycorrhizal fungal inoculum potential in the exclosures. Because ecosystems have finite resource availability, high deer density may imbalance the system by redirecting resources toward maintaining deer biomass at the expense of multiple trophic levels throughout the forest community. Both complex bottom up and top down trophic cascade effects demonstrated largely unidirectional negative responses suggesting that high deer density has reduced the biodiversity of the forest community.     

Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

The vegetation of the Øvre Heimdalen valley

The catchment area of Øvre Heimdalsvatn reaches from the upper half of the subalpine birch forest zone to the lowermost part of the high-alpine subzone within the alpine zone. The units dominating the area include mountain birch forest, tall herb meadow, mire vegetation, bilberry heath, snowbed communities, chionophobous heath vegetation, and areas without a continuous vegetation layer. It is shown that the potential natural vegetation corresponds closely to the actual vegetation distribution.     

The herb community of a tropical forest in central Panamá: dynamics and impact of mammalian herbivores

Mammals are hypothesized to either promote plant diversity by preventing competitive exclusion or limit diversity by reducing the abundance of sensitive plant species through their activities as browsers or disturbance agents. Previous studies of herbivore impacts in plant communities have focused on tree species and ignored the herbaceous community. In an experiment in mature-phase, tropical moist forest sites in central Panamá, we studied the impact of excluding ground-dwelling mammals on the richness and abundance of herbs in 16, 30×45-m plots. Within each plot, we censused the herbaceous community in 28, 2×2-m subplots (1,792 m² total area sampled). We identified over 54 species of herbs averaging 1.21 ramets m⁻² and covering approximately 4.25% of the forest floor. Excluding mammals for 5 years had no impact on overall species richness. Within exclosures, however, there was a significant two-fold increase in the density of rare species. Overall herbaceous density and percent cover did not differ between exclosures and adjacent control plots, although cover did increase over time. Mammalian exclusion significantly increased the total cover of three-dominant herb species, Pharus latifolius, Calathea inocephala, and Adiantum lucidum, but did not affect their density. This study represents one of the most extensive herbaceous community censuses conducted in tropical forests and is among a few that quantify herbaceous distribution and abundance in terms of both density and cover. Additionally, this work represents the first community level test of mammalian impacts on the herbaceous community in a tropical forest to date. Our results suggest that ground dwelling mammals do not play a key role in altering the relative abundance patterns of tropical herbs in the short term. Furthermore, our results contrast sharply with prior studies on similar temporal and spatial scales that demonstrate mammals strongly alter tree seedling composition and reduce seedling density. Thus, we question the pervasiveness of top–down control on tropical plant communities and the paradigm that defaunation will inexorably lead to widespread, catastrophic shifts in plant communities.     

Spatial patterns and dynamic responses of arctic food webs corroborate the exploitation ecosystems hypothesis (EEH)

According to the exploitation ecosystems hypothesis (EEH), productive terrestrial ecosystems are characterized by community-level trophic cascades, whereas unproductive ecosystems harbor food-limited grazers, which regulate community-level plant biomass. We tested this hypothesis along arctic-alpine productivity gradients at the Joatka field base, Finnmark, Norway. In unproductive habitats, mammalian predators were absent and plant biomass was constant, whereas herbivore biomass varied, reflecting the productivity of the habitat. In productive habitats, predatory mammals were persistently present and plant biomass varied in space, but herbivore biomass did not. Plant biomass of productive tundra scrublands declined by 40% when vegetation blocks were transferred to predation-free islands. Corresponding transfer to herbivore-free islands triggered an increase in plant biomass. Fertilization of an unproductive tundra heath resulted in a fourfold increase in rodent density and a corresponding increase in winter grazing activity, whereas the total aboveground plant biomass remained unchanged. These results corroborate the predictions of the EEH, implying that the endotherm community and the vegetation of the North European tundra behaves dynamically as if each trophic level consisted of a single population, in spite of local co-occurrence of >20 plant species representing different major taxonomic groups, growth forms, and defensive strategies.     

Mesoherbivores affect grasshopper communities in a megaherbivore-dominated South African savannah

African savannahs are among the few places on earth where diverse communities of mega- and meso-sized ungulate grazers dominate ecosystem functioning. Less conspicuous, but even more diverse, are the communities of herbivorous insects such as grasshoppers, which share the same food. Various studies investigated the community assembly of these groups separately, but it is poorly known how ungulate communities shape grasshopper communities. Here, we investigated how ungulate species of different body size alter grasshopper communities in a South African savannah. White rhino is the most abundant vertebrate herbivore in our study site. Other common mesoherbivores include buffalo, zebra and impala. We hypothesized that white rhinos would have greater impact than mesoherbivores on grasshopper communities. Using 10-year-old exclosures, at eight sites we compared the effects of ungulates on grasshopper communities in three nested treatments: (i) unfenced plots (‘control plots’) with all vertebrate herbivores present, (ii) plots with a low cable fence, excluding white rhino (‘megaherbivore exclosures’), and (iii) plots with tall fences, excluding all herbivores larger than rodents (‘complete ungulate exclosures’). In each plot, we collected data of vegetation structure, grass and grasshopper community composition. Complete ungulate exclosures contained 30 % taller vegetation than megaherbivore exclosures and they were dominated by different grass and grasshopper species. Grasshoppers in complete ungulate exclosures were on average 3.5 mm longer than grasshoppers in megaherbivore exclosures, possibly due to changes in plant communities or vegetation structure. We conclude that surprisingly, in this megaherbivore hotspot, mesoherbivores, instead of megaherbivores, most strongly affect grasshopper communities.     

内蒙古典型草原区3种不同草地利用模式下植物功能群及其多样性的变化

以呼伦贝尔草原辉河国家级自然保护区草原群落为研究对象, 选取3块生境条件基本相同的样地(250 m × 250 m) ——放牧场、割草场及公共草场, 每块样地设置3条样带, 每条样带上设置8个样方(1 m × 1 m), 研究了不同草地利用模式下草原群落的物种组成、生物多样性及群落的功能群特征。结果表明: 在放牧场、割草场和公共草场3种不同利用模式下, 植物功能群结构及其多样性发生了变化。灌木、半灌木的累积优势度在各利用模式间差异显著( p< 0.05), 在割草场中最高; 多年生禾草的累积优势度在各利用模式间差异显著(p< 0.05); 一、二年生草本的累积优势度在各利用模式间差异显著(p< 0.05)。旱生植物功能群在公共草场中所占比例最高, 为50%; 中旱生植物在割草场中的百分比最低, 为19%, 且在各利用模式间差异显著(p< 0.05); 各利用模式下, 灌木、半灌木功能群生物量均较低; 随着割草场、公共草场及放牧场3种草地利用模式下利用强度的增加, 多年生禾草和多年生杂类草两大功能群的生物量递减, 而一、二年生草本功能群生物量递增。草地利用强度的增加, 使得草地植物群落结构简单、脆弱。丰富度指数和多样性指数在放牧场最低, 在割草场最高; 均匀度变化并不明显。割草场利用模式有助于改善草原的健康状况, 而放牧场及公共草场利用模式均存在不同程度的过度放牧现象, 尤其是公共草场利用模式由于所有权不明确而缺乏有效的管理, 导致草原退化明显。     

Quantifying ecological filters: the relative impact of herbivory, neighbours, and sediment on an oligohaline marsh

The objective of this experiment was to quantify the relative importance of herbivory, interactions with neighbours and sediment addition as filters in controlling the species present in an oligohaline marsh in southeastern Louisiana, USA. We planted 16 species of wetland plants in 3×3 m plots either inside or outside herbivore exclosures, with or without presence of established vegetation, and with or without added sediment. These species, representing 12 families and diverse morphologies, included herbaceous and woody plants. At the end of the growing season above‐ and below ground biomass were measured. In cleared plots (no neighbours) inside herbivore exclosures, 12 of the 16 species grew well, indicating that they could tolerate the physical conditions in the habitat. Competition significantly reduced the biomass of four of these species (A. calamus, C. occidentalis, P. hemitomon, P. cordata), suggesting that it is the strongest species‐specific filter operating in the marsh. These results confirm the general consensus that competition from existing plants is the strongest filter operating in high biomass habitats, and further imply that competition has the potential to remove 33% of the species from the community. Facilitation occurred for one species, R. corniculata. Herbivory had the largest general effect of reducing biomass, as determined in the full ANOVA model, but on a species‐specific basis, significantly reduced the biomass of only two species (T. distichum, T. domingensis). These results support the emerging view that herbivory is an important filter in coastal wetlands. Although several other studies have argued for the importance of sedimentation in controlling plant species composition in wetlands, we found no effect of sediment addition. Competition and herbivory appear to be two critical biological filters that control plant composition of coastal wetlands, and they must be considered in future studies and restoration efforts along the Gulf of Mexico.     

Effects of grazing exclusion on species composition in high-altitude grasslands of the Central Alps

Our study examines the effects of grazing exclusion on low-productive subalpine and alpine grasslands of the Central Alps (UNESCO Biosphere Park Gurgler Kamm, Obergurgl, Austria). A long-term exclusion experiment was established in 2000 in the subalpine, the lower, and the upper alpine zone. With exception of the subalpine zone, domestic herbivores have been grazing during the whole growing season. In grazed and exclosure plots species frequencies were recorded for 7 years. We analysed exclosure effects on species number, community composition, life forms, and functional groups.Species richness did not decrease significantly within the exclosures, but changes in species composition occurred in each zone, although some were transitory in nature. The dynamic trends of the plots were significantly explained by the ‘treatment×year’ effect along the whole altitudinal gradient, but the effects decreased considerably with altitude. In the subalpine and upper alpine exclosures, stress-tolerators, species of low or no nutritive value, and mosses showed a decreasing trend, whereas tall grasses (subalpine exclosures), competitors, and species with high or medium nutritive values (lower alpine exclosures) tended to increase.Overall, our 7-year study revealed that several functional groups reacted to grazing, according to our main expectations. We suggest that these effects will intensify in the long term.     

放牧干扰下高寒草甸物种、生活型和功能群多样性与生物量的关系

为明晰放牧强度对高寒草甸物种、生活型以及功能群多样性和生物量的时间累积作用以及对多样性与生物量之间相互关系的影响。选择青藏高原东北缘高寒草甸,在6个放牧强度样地连续4 a(2012—2015)进行物种、生活型、功能群多样性和生物量调查。采用重复测量方差分析和线性回归法分析放牧强度和放牧年份对物种、生活型、功能群多样性以及多样性与生物量之间的相关关系的影响。结果表明:(1)放牧强度和放牧年份均对生物量产生显著影响。随放牧强度增加,生物量显著降低。但随放牧年份延长,不同放牧强度区生物量无一致的变化规律。(2)放牧年份对物种、生活型和功能群丰富度、均匀度和优势度的影响均达到显著水平。但放牧强度和放牧年份只对物种丰富度产生交互作用,放牧强度对物种丰富度的影响具有时间的累积效应。(3)放牧干扰下不同层次多样性,仅物种丰富度与所有多样性指数显著相关,物种丰富度可以作为物种多样性测度的代表性指标。(4)多样性与生物量关系的研究,对丰富度而言,仅低放牧强度样地(Plot2)中物种丰富度随生物量增加而显著降低,其余放牧强度样地的物种、生活型和功能群丰富度均与生物量无关。对均匀度而言,高放牧强度样地(Plot4、Plot5、Plot6)生活型均匀度随生物量的增加而显著降低。对优势度而言,高放牧强度样地生活型优势度随生物量的增加而显著增加。生活型多样性可作为放牧干扰下生物量变化快速预测的有效指标。     

How is Regeneration of Plants after Mowing Affected by Shoot Size in Two Species-Rich Meadows with Different Water Supply?

Mowing a meadow is an example of an equalizing process that reduces differences among species by removing aboveground biomass approximately 5 cm above ground. This regular disturbance that affects all plants prevents competitive exclusion of small species and thus allows coexistence of numerous species differing in shoot size. In this paper we search for the mechanism behind this by comparing the shoot biomass of 41 common species in dry and wet species-rich meadows in mown and recently abandoned plots in June (before mowing) and in October. We asked the following questions: i) Do the plants differ in proportion of biomass lost by mowing? ii) Are the mown plants able to compensate for biomass lost by mowing? iii) Is the compensatory ability of mown plants related to their size? iv) Is the compensatory ability of plants related to severity of disturbance (removed biomass)? v) Does water availability in meadows affect these features? Our results revealed that the earlier explanation of equalization of meadow plants after mowing due to the proportionally larger biomass loss in larger plants than small plants does not represent the entire mechanism. Even when larger plants in the wet meadow lost more biomass, the proportion of lost biomass was not dependent on plant size, and compensation ability (growth of mown in comparison with unmown plants) was not related to the lost biomass in this meadow type. On the contrary, the observed pattern could be explained by different compensation abilities of small versus tall plants. In addition, according to our expectations, the compensation for lost biomass in the wet meadow was higher than in the dry one.     

Bird predation on herbivorous insects: indirect effects on sugar maple saplings

Insectivorous birds have been shown to have direct effects on abundances of herbivorous arthropods, but few studies have tested the indirect effects of birds on plant performance through consumption of herbivorous insects. In a 3-year study at the Hubbard Brook Experimental Forest, New Hampshire, we tested whether bird predation indirectly affects leaf herbivory levels and leaf and shoot biomass production of understory sugar maple (Acer saccharum) saplings. Trees were randomly assigned to one of four treatments: an insecticide application to reduce herbivory levels, exclosures that prevented bird access, addition of Lepidoptera larvae, and controls. Trees sprayed with an insecticide supported significantly fewer Lepidoptera larvae than other treatments throughout the study. Also, trees in exclosures supported more Lepidoptera larvae than controls during one count each year, and pooled across all counts during the second year. As predicted, the mean proportion of leaf area consumed varied significantly among treatments and was least in the insecticide treatment, followed by controls, exclosures, and Lepidoptera additions. Significant differences among treatments in herbivory levels, however, did not lead to differences in leaf or shoot biomass production. Thus, bird predation decreased Lepidoptera abundances and decreased herbivory levels, but did not increase biomass production during the following year. Over 85% of the herbivores in our study were Homoptera nymphs that were not folivorous and are not important bird prey items, potentially dampening the indirect effects of bird predation on biomass production. A comparison of these results with previous studies suggests that the indirect effects of bird predation on plant biomass production may depend on the plant species, abundance and composition of the herbivore community, and primary productivity of the ecosystem.     

放牧干扰下高寒草甸物种和生活型丰富度与地上及地下生物量的关系

明晰放牧干扰下高寒草甸植物丰富度与生物量的相关关系,为草地植物不同生长时期生物量的预测提供依据。设置6个放牧强度样地,连续3a放牧,2014年进行3个季节(6月、8月、10月)的植物丰富度和地上、地下生物量调查,对比分析放牧干扰下物种和生活型丰富度(生活型的种类)分别与地上、地下生物量的相关关系。结果表明:(1)物种和生活型丰富度与地上生物量均受放牧强度的显著影响,物种丰富度仅在8月与放牧强度显著负相关,生活型丰富度在10月随放牧强度单峰变化,地上生物量在不同季节均与放牧强度显著负相关,而地下生物量与放牧强度无关。(2)物种丰富度与地上和地下生物量均受季节的显著影响,物种丰富度和地上生物量仅在低强度放牧区随季节呈单峰变化,地下生物量在中等强度放牧区随季节呈单峰变化;生活型丰富度与季节无关。(3)放牧干扰前物种和生活型丰富度与地上和地下生物量均显著正相关。3a放牧后仅在8月,物种丰富度只与地上生物量显著正相关,生活型丰富度与地上和地下生物量均显著正相关。(4)对于不同放牧强度,物种丰富度仅在低强度放牧区与地上生物量显著正相关,而生活型丰富度在所有放牧强度区均与地上生物量显著正相关。综上所述,放牧干扰扰乱了高寒草甸丰富度与生物量之间的关系,尤其影响了物种丰富度与地下生物量之间的相关关系。生活型丰富度与地上生物量之间的显著关系不受放牧强度干扰,使生活型丰富度在预测生物量方面表现出优势。