Mapping the Microenvironment for Seed Germination in the Field

Gaps in the canopy have a major influence upon the microenvironmentfor seed germination in natural vegetation. Two methods formeasuring gaps in grasslands are described. The first uses afield-portable instrument for mapping R/IR ratios under thecanopy, and the second employs infrared photography and imageanalysis to measure gaps from above. The methods are illustratedwith results from an experiment in which grazing sheep are theprinciple influence upon the structure of the grass canopy.Results of both methods show that the effects of current andpast grazing regimes may be detected in the canopy structureof the grassland. Seeds of Geranium dissection sown into gapsof different sizes showed greater establishment in small gaps(1–2 cm²) than under a closed canopy Canopy gap, germination, grassland, grazing, Geranium dissectum     

Swards and structure: the interactions between farming practices and bird food resources in lowland grasslands

The ideal grassland to encourage plants and invertebrates suitable as food for farmland birds contains a mixture of grasses and broad-leaved plants with a range of vegetation heights and structures. However, as a result of agricultural intensification the majority of lowland grasslands in the UK now lack botanical and structural complexity. Plants within intensively managed swards are allowed little capacity to set seed, and the majority of invertebrates within these grasslands are either too small to be utilized by birds or inaccessible because of dense vegetation or impenetrable soils. Further research is required into the impact of different grassland management practices on the abundance and accessibility of bird food resources at both the field and the landscape level. The major challenge will be to identify changes to grassland management practices that provide significant biodiversity benefits and yet allow productive livestock farming systems to continue.     

Gap disturbance triggers the recolonization of the clonal plant Ambrosia tenuifolia in a flooding grassland of Argentina

Abstract Flooding provokes the death of many dicotyledonous species in grazed grasslands of the Flooding Pampa in Argentina, including the clonal plant Ambrosia tenuifolia, which produce the opening of numerous gaps. The objective of this study was to investigate the recolonization of grassland by A. tenuifolia after this species disappeared due to the occurrence of prolonged flooding events. To this end, responses of seed germination to environmental factors associated with gaps, such as light quality and temperature regime, conditions related to seedling survival, and clonal growth of ramets outside the gaps were studied in two different experiments in the field. Environmental factors related to gaps promoted the recruitment of new genets. The combined effect of alternating temperatures and the high red : far‐red ratio set off germination from the soil seed bank; germination also was enhanced when signals were generated artificially under the intact canopy in the field. Higher resource availabilities and maximum seedling survival were recorded in canopy gaps, which were the focus of invasion. Grassland recolonization outside the gaps continued rapidly by clonal growth, from small gaps and large ones, even within the dense surrounding canopy. This provoked an intense competition with the other species. Gap opening by disturbances, seed germination in gaps and clonal growth were decisive for the recolonization of A. tenuifolia populations. This sequence of events triggered the recolonization of the plant community by this species, in sites where it had been eliminated by prolonged flooding. This process represents one of the most significant fluctuations in the vegetation dynamics of the Flooding Pampa Grasslands.     

Do seedlings in gaps interact? A field test of assumptions in ESS seed size models

ESS models for the evolution of seed size variation assume that seedlings compete with each other for the occupancy of 'safe sites' or vegetation gaps. If mortality rates are high and/or frequency-independent, ESS models reduce to the classical model of Smith and Fretwell which predicts that a single, optimum seed size should occur. We tested whether seedlings compete with one another by following the survival of seedlings colonizing experimental gaps in a grazed grassland community. In small gaps (3 cm diameter) the proximity of established plants slightly, though significantly reduced seedling survival, but density-dependent mortality also occurred among seedlings in these gaps. In larger gaps (6 cm, 9 cm diameter) survival was significantly positively frequency-dependent. These results strongly support the validity of ESS models.     

Does Germination Success Differ with Respect to Seed Mass and Germination Season? Experimental Testing of Plant Functional Trait Responses to Grassland Management

BACKGROUND AND AIMS: Plant functional trait responses to processes such as grassland management have been analysed frequently; however, the scaling-up from individual traits to the outcomes of vegetation dynamics has seldom been tested. In this experiment, germination success was studied with respect to the relationships between grassland management (mowing and grazing), as well as abandonment, and two traits that are relevant for seedling recruitment: seed mass and germination season. On the basis of discussions in the literature and indirect trait analyses in our previous studies, the following hypotheses are proposed: (1) with respect to seed mass, mowing and grazing favour the germination of small seeds, whereas after abandonment the germination success of larger seeds is higher; and (2) with respect to germination season, mowing and grazing favour autumn-germinating seeds, whereas succession promotes spring-germinating seeds. METHODS: The germination experiment took place in a semi-natural, dry grassland in north-east Germany. Seeds of eight herbaceous species that differ with respect to seed mass and germination season were sown in mown, grazed and abandoned plots. Germination success was documented during the following year. KEY RESULTS AND CONCLUSIONS: Contrary to the hypothesis, germination of small seeds was not promoted by mowing or grazing and they germinated relatively more often than expected in the abandoned plots. A relationship between abandonment and gaps of bare soil below the vegetation cover that favour germination of small seeds was likely, but could not be proved statistically. It is possible that the small seeds suffered less from predation. Mowing favoured autumn germination, which could be explained by the removal of biomass in late summer. Contrary to our expectation, there was relatively more spring germination after grazing than after mowing, yet vegetation height was smallest in spring. Generally, germination season was found to be related to the temporal occurrence of favourable light conditions.     

Untangling the roles of fire,grazing and rainfall on small mammal communities in grassland ecosystems

In grassland systems across the globe, ecologists have been attempting to understand the complex role of fire, grazing and rainfall in creating habitat heterogeneity and the consequences of anthropogenic control of these factors on ecosystem integrity and functioning. Using a South African grassland ecosystem as a model, we investigated the impact of fire and grazing pressure on small mammal communities during three differing periods of a rainfall cycle. Over 2 years, 15,203 trap nights revealed 1598 captures of 11 species (nine rodents, one macroscelid and one insectivore). Results highlighted the importance of the interplay between factors and showed that the role of fire, grazing and rainfall in determining small mammal abundance was species-dependant. While no two species were affected by the same environmental variables, grass cover or height was important to 56% of species. Considered independently, high rainfall had a positive influence on small mammal abundance and diversity, although the lag period in population response was species-specific. High grazing negatively affected overall abundance, but specifically in Mastomys coucha; fire alone had little immediate impact on small mammal diversity. Six months after the fire, vegetation cover had recovered to similar levels as unburned areas, although small mammal diversity and richness were higher in burned areas than unburned areas. Grazing levels influenced the rate of vegetation recovery. In conclusion, low-level grazing and burning can help to maintain small mammal biodiversity, if conducted under appropriate rainfall levels. A too high grazing pressure, combined with fire, and/or fire conducted under drought conditions can have a negative impact on small mammal biodiversity. To maintain small mammal diversity in grassland ecosystems, the combined effects of the previous year’s rainfall and existing population level as well as the inhibition of vegetation recovery via grazing pressure need to be taken into consideration before fire management is applied. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Vegetation change after artificial disturbance in an alpine

We describe the colonisation of artificially created gaps in an alpine grassland dominated by Chionochloa pallens. Twelve years after their creation, the 50 cm _ 50 cm gaps supported a distinctive vegetation composed of a mixture of perennial forbs, grasses and mosses. Three species (Bryum sp., Epilobium alsinoides and Plantago novae-zelandiae) were recorded only in the gaps. Plantago novae-zelandiae and Polytrichum juniperinum had their highest frequency in the centres of the gaps, while Celmisia gracilenta and Geranium microphyllum were most frequent at the edges. The distinctive species composition of the gap plots indicates that they are still in an early stage of succession, assuming an eventual reversion to the surrounding climax grassland.     

Effects of slug herbivory on the seedling establishment of two montane Asteraceae species

Seedling establishment as the life stage transition most sensitive to herbivory might be impossible even if the herbivory losses suffered by adult plants of the same species are tolerable. We tested the hypothesis that herbivory impedes seedling establishment of two montane Asteraceae species on their lower altitudinal distribution border.In a submontane meadow in the Harz Mountains, Germany, the montane grassland perennials Arnica montana and Centaurea pseudophrygia (Asteraceae) were sown into artificially created gaps of two different sizes, half of which were protected from slug grazing by molluscicide application in the initial phase of the experiments. The Arnica experiments started in spring and late summer of 1999 and in spring of 2000; whereas Centaurea was only sown in the spring of 2000. Seedling survival was monitored until the autumn of 2000.Ten weeks after sowing, mollusc exclusion resulted in significantly higher Arnica or Centaurea seedling numbers in all experiments; whereas grasses and dicotyledons that emerged from the seedbank were promoted by slug exclosure in only one experiment. Species diversity was not affected by molluscicide application. By the end of the second growing season, the survivorship of Arnica sown in the spring of 1999 was reduced to less than 1% of the number of sown seeds in the gaps with natural slug grazing, compared to 7–15% in the gaps with molluscicide treatment. Plant density of most species was higher in the small than in the large gaps, indicating a microclimatic shelter effect of the surrounding vegetation.The results show that slug herbivory might exert large effects on species composition in grassland; it does strongly influence the seedling establishment of the highly palatable, slow-growing A. montana, while the seedlings of C. pseudophrygia, although of similar palatability, are more able to tolerate slug feeding because of their higher growth rate.     

A test of concordance in community structure between leafhoppers and grasslands in the central Tien Shan Mountains

Leafhoppers and related Auchenorrhynchous Hemiptera (AH) are among the most diverse grassland herbivores, and many have been linked inexorably to grassland vegetation through diet and shelter for millions of years. Are AH–plant communities in natural grasslands tightly integrated, how does the interaction differ across major ecological gradients, and do habitat or environmental factors explain the most variance in AH community structure? These questions have implications for the conservation of biodiversity and in evaluating effects of a warming climate. Using grasslands of the central Tien Shan Mountains as a natural laboratory, we examine whether AH species assemblages are concordant with vegetation in terms of community structure using closely associated species-level samples. Data were recorded from a nearly 3000-m elevation gradient crossing four arid and three montane grassland vegetation classes. We found elements of AH–plant community classification and structure to be closely correlated except for at the arid–montane habitat transition where a small group of widespread AH species were significant indicators for vegetation classes in both major grassland types. AH species richness and abundance are positively correlated with plant species density and percent cover and, correspondingly, peak at mid-elevations in association with montane grasslands. While overall elevation (and covariate mean annual temperature) explains the most variance in AH species assemblages, the sum total of habitat factors explain more variance than environmental factors when arid and montane grasslands were examined separately, but environmental factors are co-equal with habitat factors when the grassland types are combined. Unexplained variance in the AH community assemblages, attributable to individualistic species responses to environmental and habitat factors, slightly exceeds the total accounted for by the model.     

Experimental demography of the rare Gentianella germanica: seed bank formation and microsite effects on seedling establishment

The persistence of populations of short-lived species requires regular reproduction and seedling establishment. A persistent seed bank can buffer populations against extinction in unfavourable years. We experimentally investigated seed fate in Gentianella germanica, an endangered biennial species characteristic for species-rich nutrient-poor calcareous grasslands in central Europe. We studied the effect of experimental gaps on seedling establishment from sown seeds and the fate of seeds buried in bags over two years. In December 1993 experiments were established at seven calcareous grassland sites in the Swiss Jura mountains. In spring 1994 seedlings emerged in all plots where seeds had been sown, including previously unoccupied patches. This suggests that limited dispersal within sites contributes to small population sizes. Significantly more seedlings emerged at sites with current populations of G. germanica than at unoccupied sites (5.95% vs 3.40%). Because this difference was not explained by germinations from the natural seed bank it indicates differences in habitat quality. Clipping of the vegetation and disturbance of the soil reduced vegetation cover in the following spring and enhanced seedling emergence. In undisturbed plots 4.5% of seeds sown produced a seedling in spring 1994, whereas in plots with clipped vegetation 9.9% and in disturbed plots 12.7% produced seedlings (p>0.01). This suggests that management measures which create gaps in the vegetation (e.g. grazing) could positively influence population size and persistence of G. geymanica. On average, we recovered 7.55% viable seeds after one year of burial in bags, and 4.05% after two years, indicating that G. geymanica has a persistent seed bank. The demographic data indicate that the number of viable seeds in the seed bank exceeds the number of established plants in a population at least by a factor of 20. Restoration of extinct populations of the species from the seed bank may thus be possible if appropriate management measures are taken within a few years.     

Immediate responses in soil chemistry,vegetation and ground beetles to soil perturbation when implemented as a restoration measure in decalcified sandy grassland

The species-rich calcareous grassland communities in Europe are gradually disappearing due to lack of management such as grazing or cultivation, resulting in decalcification and reduction of gaps in the vegetation. In this study, experimental soil perturbation (deep and shallow) was performed in degenerated sandy grassland in plots with a size of 8 × 8 m, using a randomised block design. The hypothesis that soil perturbation that inverts the soil layers decreases nutrient availability, creates vegetation gaps and thereby selects for desirable species was tested through comparisons with untreated controls as well as with nearby target habitats. The deep perturbation was designed to bring CaCO₃ up to the surface, whereas the shallow perturbation tested the effect of disturbance alone. The effects of soil perturbation on soil chemistry, vegetation and beetle communities were analysed for the 2 years following the treatments. Increased pH and calcium concentration, and decreased nitrogen and phosphorus availability, showed that deep perturbation was successful in restoring the soil chemistry to levels similar to those of the target habitat. Perturbated plots were rapidly colonised by the acid tolerant grass Corynephorus canescens, but the slow colonisation of the threatened calcicole species Koeleria glauca was an indication that the vegetation could be evolving towards the target vegetation. Six red-listed beetle species associated with open, dry grasslands were found, out of which four were found only in perturbated plots, although this could not be statistically tested. In conclusion, it may take many years or even decades for the establishment of desirable flora, and seeding could therefore be a suitable method of increasing the rate of succession.     

Cultural monuments and nature conservation: a review of the role of kurgans in the conservation and restoration of steppe vegetation

Steppe is among the most endangered biomes of the world, especially in Eastern Europe, where more than 90 % of original steppes have been destroyed due to conversion into croplands, afforestation and other human activities. Currently, steppe vegetation is often restricted to places unsuitable for ploughing, such as ancient burial mounds called kurgans. The aim of our study was to collect and synthesise existing knowledge on kurgans by a review of research papers and grey literature. The proportion of kurgans covered by steppe vegetation increases from west to east and from lowlands to uplands. Despite their small size, kurgans act as biodiversity hotspots and harbour many red-listed species. High overall species richness and a high proportion of grassland specialists are maintained by a pronounced fine-scale environmental heterogeneity. The main factors threatening the biodiversity of kurgans are intensified agriculture and construction works. We conclude that kurgans can play a crucial role in preserving steppe vegetation, especially in intensively used agricultural landscapes in the western part of the steppe zone. Despite the vital role of kurgans in sustaining steppe vegetation, we identified serious knowledge gaps on their distribution, vegetation, flora and fauna and their potential role in steppe restoration.     

Canopy manipulations of exotic Bitter Willow (Salix elaeagnos) forest for indigenous seedling recruitment: A pilot study

The invasive exotic tree species Bitter Willow (Salix elaeagnos; Salicaceae) has colonised areas of rank exotic grassland and has been found to contain indigenous seed, dispersed by frugivorous birds into the monospecific stands. This small pilot study examined whether indigenous seedlings that have germinated in the understorey of exotic Bitter Willow stands could be stimulated to establish through the creation of small‐scale canopy gaps. In Bitter Willow forest, four single Bitter Willow trees were poisoned to create canopy gaps. Light transmission and seedling regeneration of tree and shrub species were assessed beneath both the four manipulated and three comparable intact Bitter Willow canopies. Over 3 years, seedling height and density increased more beneath opened compared to intact Bitter Willow canopies. These results suggest that Bitter Willow can fill the roles of both a facilitative nurse and a perch tree. Larger‐scale canopy manipulation experiments of both Bitter Willow and other Salix species are needed to determine the full potential of canopy manipulations for forest restoration.     

Small seed bank in grasslands and tree plantations in former grassland sites in the South Brazilian highlands

The soil seed bank can be an important source for vegetation regeneration, and data on the similarity between aboveground vegetation and the seed bank can provide information about successional pathways after disturbances or land-use change. We conducted this study in natural grasslands in the subtropical highland region in southern Brazil. We evaluated the effect of silviculture on richness, density, and composition of the seed bank at former grassland sites converted to pine plantations 25 years ago. We worked at six grassland sites and three pine plantation sites and used the seedling emergence method. Seed bank density and richness in grasslands were lower than those reported in similar environments in other regions. Species richness and density varied considerably within each vegetation type; therefore, richness and density were not statistically significant, while composition varied among vegetation types. In terms of species, the pine plantation seed bank was a small subset of the grassland seed bank. Seeds of typical grassland species were missing in the pine plantation, but also had only low abundances in the grassland, and similarity of seed bank and vegetation were low (less than 20%). The low seed density found in this study, including in grasslands areas, indicates that regeneration of species from the soil seed bank likely is of a limited role for the maintenance of plant populations after disturbances in this system. Our data further suggest that natural regeneration after tree planting in grasslands is reduced due to seed limitation.     

Static and dynamic approaches to landscape heterogeneity in the Hungarian forest‐steppe zone

Abstract. This paper describes the successional status of the vegetation in a clear‐felled dry oak woodland at the edge of the Hungarian forest‐steppe zone on the basis of a vegetation map. Due to a varied geomorphology of the colline landscape several so‐called landscape units can be distinguished. The patchwork on the vegetation map is evaluated using several, morphology‐based attributes (static morphological indices) traditionally applied in landscape ecology. In the ca. 100 years that elapsed since forest clear‐cut, xeric grassland species and steppe elements became more abundant and the former xeromesophilous vegetation – containing even some woodland components – is slowly turning into xeric grassland communities. The vegetation units mapped can be arranged into a hypothetical succession scheme in which successional distances (the number of steps between two stages) are determined. Based on the distances thus obtained, a new dynamic morphological index is introduced. This is applied to each landscape unit for the dynamic evaluation of successional vegetation, its results being compared with those obtained by static morphological indices.     

Phosphorus Translocation by Red Deer on a Subalpine Grassland in the Central European Alps

We examined the role of red deer (Cervus elaphus L.) in translocating phosphorus (P) from their preferred grazing sites (short-grass vegetation on subalpine grasslands) to their wider home range in a subalpine grassland ecosystem in the Central European Alps. Phosphorus was used because it is the limiting nutrient in these grasslands. When we compared P removal of aboveground biomass due to grazing with P input due to the deposit of feces on a grid of 268 cells (20 m × 20 m) covering the entire grassland, we detected distinct spatial patterns: the proportion of heavily grazed short-grass vegetation increased with increasing soil-P pool, suggesting that red deer preferably grazed on grid cells with a higher soil-P pool. Biomass consumption related to increased proportion of short-grass vegetation, and therefore P removal, increased with increasing soil-P pool. However, within the two vegetation types (short-grass and tall-grass), consumption was independent from soil-P pool. In addition, P input rates from defecation increased with increasing soil-P pool, resulting in a constant mean net P loss of 0.083 kg ha⁻¹ y⁻¹ (0.03%–0.07% of soil-P pool) independent of both soil-P pool and vegetation type. Thus, there was no P translocation between grid cells with different soil-P pools or between short-grass and tall-grass vegetation. Based on these results, it is likely that the net rate of P loss is too small to explain the observed changes in vegetation composition from tall-herb/meadow communities to short-grass and from tall-grass to short-grass on the grassland since 1917. Instead, we suggest that the grazing patterns of red deer directly induced succession from tall-herb/meadow communities to short-grass vegetation. Yet, it is also possible that long-term net soil-P losses indirectly drive plant succession from short-grass to tall-grass vegetation, because nutrient depletion could reduce grazing pressure in short-grass vegetation and enable the characteristic tall-grass species Carex sempervirens Vill. to establish.     

How does size and isolation affect patches of steppe-like vegetation on slumping hills in Transylvania,Romania?

The region of Southern Transylvania in Romania contains large expanses of species-rich grassland and mixed farmland. Within these landscapes, clusters of small, steep-sided slumping hills support mosaics of herbaceous vegetation caused by the heterogeneity in temperature and water supply and maintained by mowing or grazing. These are of conservation interest not only for the high species density, but also the relict steppe-like vegetation types especially found on their south-facing slopes. Usually surrounded by more intensively used mesic grassland, these hills can be considered patches of species-rich grassland vegetation. We therefore surveyed the vascular plants in 10 m² plots on the south- and north-facing slopes of 50 hills in 12 clusters in order to investigate the influence of their size and isolation on the species richness and composition of the vegetation. We found that larger hills had higher plot-scale species richness, a greater proportion of competitive species and a lower proportion of species of steppe-like vegetation, but only on their south-facing slopes where the conditions are more extreme. Both large and small hills should be considered as important habitats and potential sources of propagules for plant communities in the surrounding areas.     

草原露天煤矿区植被对地下水位埋深变化的响应

草原采矿活动疏排地下水导致区域地下水位下降,植被退化严重。明确草原采矿活动影响区域地下水位埋深的强度和范围,确定影响植被生长的地下水位埋深阈值,对草原矿区生态环境保护有着重要意义。以呼伦贝尔草原伊敏露天煤矿为例,利用遥感方法,建立采矿前后区域地下水埋深变化与NDVI的定量响应关系;通过样方调查,确定研究区植被群落随地下水位埋深变化的演替模式,分析草原矿区地下水位变化对草原植被类型、物种丰富度、植被覆盖度、地上生物量和综合优势比的影响;结合两种方法,确定维持研究区植被正常生长的地下水位埋深阈值。其结果如下：从柴达敏诺尔湖至采坑边缘,地下水位埋深从0 m逐渐下降至60 m,植被群落演替为盐化草地→典型草原→退化典型草原→退化草甸草原→盐化草甸草原;研究区最适宜植被生长的地下水位埋深约为1 m;1-30 m为维持研究区植被正常生长的阈值地下水位埋深。最后根据以上结论,将研究区划分为地下水开发的一级敏感区、二级敏感区和三级敏感区,针对不同敏感区提出了不同的地下水开发政策,以防止采矿活动疏排地下水引起草原退化。     

Influences of neighbouring plants on seedling establishment in limestone grassland

Abstract. Influences of neighbouring plants on seedling establishment of six dicotyledonous species was investigated in a nutrient-poor limestone grassland (Mesobrometum) in northern Switzerland. Microsites with different vegetational structure were created and seeds sown in them. The fate of emerging seedlings was followed for one or two years. A plant surviving its first growing season was regarded as established. Shelter by neighbouring plants appeared to be necessary for the establishment of Arabis hirsuta and Primula veris. Because of drought and frost heave, hardly any seedlings of these species established in gaps (4 - 21 %). In microsites with vegetation, their survival was significantly higher (40–57 %). Neighbouring plants slightly reduced the survival of Plantago lanceolata and Sanguisorba minor, but these species established well both in gaps (74 - 81 %) and in vegetated microsites (54 - 67 %). Medicago lupulina established well in all microsites in one year (71 - 79 %), but poorly in the next year (18–32 %). Linum catharticum emerged poorly in one year and was completely extinguished by a fungal pathogen the following year. The overall conclusion is, that gaps are of minor importance for recruitment of these species in this grassland. Physical hazards and pathogens control seedling establishment to a greater extent than competition by neighbouring plants. Some species are hardly able to establish without shelter of vegetation. Seed size is an important factor for success of establishment, especially in gaps. It is suggested, that the relatively low productivity and the absence of litter accumulation (due to mowing and biomass removal) are important conditions for the observed behaviour of the seedlings and juvenile plants in this community.     

高寒草甸地区小哺乳动物群落多样性的初步研究

高寒苹甸地区的人工草场、半人工草场、次生杂类草草场和撩荒地中共有4个小哺乳类群落;根田鼠+高原鼢鼠群落;根田鼠+甘肃鼠兔群落;高原鼠兔+高原鼢鼠群落;长尾仓鼠+高原鼠兔群落。小哺乳类群落种的多样性与植物群落种的多样性无显著相关关系,而与植被的高度、盖度呈显著负相关关系。高原鼢鼠是各生境中的广布种。根田鼠与甘肃鼠兔在植被郁闭度高的环境中为群落的主要组成者;植被郁闭度低的环境中,高原鼠兔和长尾仓鼠是群落的主要成员。它们的空间格局主要反映空间环境条件的差异。