Characterization of the canopy structure of Dutch grasslands

In order to characterize the canopy structure of different grassland types 50 stands, representing 15 syntaxonomically distinct types, were examined by harvesting the standing crop during the main flowering period. The types differ in water and nutrient conditions and vary largely in aboveground phytomass (0.5–23 t·ha^?1) and Leaf Area Index (0.4–21: bifacial). Living aboveground phytomass and canopy structure are almost entirely determined by phanerogams. The graminoids generally dominate over the forbs. Variation in aboveground phytomass is related to foliage characteristics such as Aboveground Leaf Area Ratio, Specific Foliage Weight and Leaf Area Development. A PCA with these canopy variables and the variables canopy height, phytomass density and the phytomass ratios stem-leaf-inflorescence shows a clear arrangement related to aboveground phytomass and LAI. The grassland stands can be divided into four groups of different productivity levels for which various combinations of canopy variables are characteristic. Leaf size and leaf inclination are also used for characterization of the different grasslands. Ordinations with these variables by means of PCA and CCA resemble partly to those computed with the canopy variables of aboveground phytomass and LAI. Small leaf sizes are characteristic for low productive grasslands, while the largest leaves occur in high productive grasslands, although they mostly do not belong to the species contributing most strongly to the phytomass of the stand. The leaf inclinations erect and erecto-patent are most common in each grassland. Horizontal leaf areas occur less frequent, but they are relatively well presented in some high productive grasslands. Ranges in leaf size vary more than ranges in leaf inclination do for this series of grasslands. However, leaf size and leaf inclination are useful variables for characterization of grassland canopies in a hierarchical way, where phytomass and leaf area are the first criteria for such a characterization.     

Impact of grazing and atmospheric nitrogen deposition on the vegetation of dry coastal dune grasslands

Abstract. A five-year experimental study was carried out to examine the combined effects of grazing and atmospheric nitrogen deposition on the vegetation of three dry dune grasslands: one short species-rich, one short species-poor, and one predominated by tall graminoids. Additional fertilization with nitrogen had no significant effect, neither in grazed nor in non-grazed plots. Exclusion of grazing by rabbits resulted in an increase in the frequency of perennial graminoids and a decrease in the frequency of annual graminoids and herbs. Nevertheless, species diversity remained the same in the species-rich grassland. During the experiment, the above-ground biomass increased in all nongrazed plots and the amount of bare soil and mosses decreased. The vegetation changes occurred mainly within one year after the exclusion of grazing. An exception is the grass-dominated site where the amount of Calamagrostis epigejos increased gradually from ca. 20 % in the first two years to about 50 % in the fourth and fifth year. Grazing by rabbits seems essential to prevent graminoids to become predominant in the dry dunes. If graminoids are dominant, grazing by horses can be an appropriate method to restore the original grassland vegetation. After six months of grazing by horses the grass-dominated site showed a decrease of the frequency of perennial graminoids, from 95 % to 80 %, and an increase of the frequency of perennial herbs, from 2.5 % to between 13 and 20 %.     

Restoration management of abandoned chalk grassland in the Netherlands

After abandonment of chalk grassland in the Netherlands, Brachypodium pinnatum had become very dominant and this resulted in a dramatic decrease of plant diversity. Restoration of these abandoned sites is important, because of its former high diversity with many nationally endangered species. To restore the chalk grassland vegetation, the impact of the introduction of different experimental cutting regimes was investigated during three years. The thick litter layer and the dominance of Brachypodium was strongly reduced after introduction of all cutting regimes. In all cutting treatments forb phytomass increased considerably, especially in the twice-a-year cutting vegetation, where the forb phytomass became as high as that of Brachypodium. Phanerogamic species diversity was stimulated in all cutting regimes. Especially the number of short-lived forbs. with a persistent seed bank, increased markedly in the twice-a-year cutting treatment. It is concluded that cutting the vegetation twice is adequate to reduce the negative effects of the thick litter layer and the abundant growth of Brachypodium. It creates an appropriate starting point for more detinite management, whether mowing or grazing. However, the restoration of a more complete species assortment strongly depends on the re-invasion possibilities of these species.     

Fire regimes and phytomass growth dynamics in a Quercus coccifera garrigue

An experiment was set up in a Quercus coccifera garrigue in southern France to analyze the effect of burning frequency and season on phytomass production. Fire regimes consisted of late spring or early autumn burns, every 6 yr, every 3 yr, or every 2 yr. The experiment started in 1969 and lasted for 19 yr. In May 1981 and May 1987, 10 samples, each 1 m², were harvested per treatment. Fire frequency had an effect on the quantity of phytomass which was produced: above-ground phytomass decreased with increasing fire frequency. This was mainly due to the lower biomass of woody plants. In all burning treatments the phytomass of herbs was higher than in the unburned vegetation. Within each burning frequency, the total phytomass of the spring-burned vegetation was always higher than that of the autumn-burned community. Generally, the herb phytomass produced was higher in the autumn-burned plots. There were two fairly distinct phases in the period following fire, each with a different level of annual phytomass production. For the first six years it was about 300 g m^?2 yr^?1, falling thereafter to about 50 g m^?2 yr^?1. This and other studies on Q. coccifera garrigue indicate that this community is very resilient with respect to fire, but possesses a low productive capacity and does not show any sign of degeneration up to 30 yr old.     

UV-B and PAR in single and mixed canopies grown under different UV-B exclusions in the field

The purpose of this study was to investigate whether differences in canopy architecture due to the investigated species (planophile versus erectophile, single versus mixed canopies) or to UV-B effects on plant morphology, lead to differences in UV-B and UV-B/PAR doses within canopies.The development of a very small (10 mm diameter) UV-B and PAR sensor on a long 5 mm wide stick allowed us to measure the penetration of UV-B and PAR in single and mixed canopies of the grass Dactylis glomerata and white clover, Trifolium repens. The plants were grown in greenhouses covered with different thicknesses (3 and 5 mm) of UV-transmittant plexi (12 and 18% UV-B exclusion).For clover, a planophile vegetation, radiation penetration was very low for both UV-B and PAR. UV-B penetration was much less than for PAR, resulting in low UV-B/PAR ratio's within the canopy. This is explained by the low UV-B transmittance of the leaves (<0.1 %) in combination with the planophile leaves.In the grass species, both UV-B and PAR penetrated much deeper into the canopy due to the erectophile structure. The difference between UV-B and PAR penetration was generally quite small except in very tall canopies.The mixed species canopies showed results comparable to the clover canopies. Due to the strongly increased grass growth in these plots, light penetration was generally much lower than in the single species cultures. The increased growth of grass in these mixed plots could be linked to the lower UV-B/PAR dose they received.In plots grown under the higher UV-B level there was a relative decrease in UV-B/PAR ratio within the canopy for both species, compared to canopies from the lower UV-B greenhouses. This could not be explained by changes in leaf angle or biomass, but might be linked to the increase in leaf transmittance of PAR.     

A Bryophyte flora of Kent. II. Hepaticae

Abstract

The military training area on Salisbury Plain contains the largest area of chalk grassland in western Europe. The grassland swards, though relatively tall, are often rich in flowering plant species. The bryophyte flora of areas disturbed in five different ways (prehistoric earthworks, twentieth century rifle ranges, ant-hills, vehicle tracks and shell-holes) was compared to that of adjacent, relatively undisturbed grassland. There was no significant difference in bryophyte cover between quadrats on disturbed ground and the controls, but the disturbed sites supported more bryophyte species per quadrat. Of the 55 taxa recorded, 38 species were at least three times more frequent in the disturbed than the undisturbed sites, compared to four which were at least three times more frequent in the control sites. The species favouring disturbed conditions included several bryophytes characteristically associated with chalk soils in southern England, including some that fruit freely (e.g. Microbryum curvicollum, Tortula lanceola) and others that fruit very rarely (e.g. Abietinella abietina, Entodon concinnus). These results are discussed in relation to the conservation of bryophytes and other disturbance-tolerant and disturbance-dependent species on Salisbury Plain and in the wider context of the protection of the bryophytes of chalkland habitats.     

Fine-scale spatial distribution of leaves and shoots of two chalk grassland perennials

The fine-scale spatial distribution of leaves and shoots of Brachypodium pinnatum and Carex flacca, two rhizomatous graminoids, was investigated in two chalk grasslands in South Limburg (The Netherlands). The objective was to examine whether leaves and shoots of Brachypodium, a dominant species, had a regular distribution on a small scale, as has been suggested for other clonal species that form high-density stands. Patterns were compared to Carex, which is never found to be as abundant as Brachypodium.The number of shoots and leaf contacts were counted in small quadrats, grouped in a grid. Using Moran's I analysis for autocorrelation, it appeared that leaves and shoots of both species were arranged in clumps, and that these clumps were randomly distributed across the soil surface. Shoot clumps in Carex were smaller in diameter and not as pronounced as those in Brachypodium.In most cases, patterns of leaves and shoots were positively correlated, indicating that leaves were predominantly positioned above and around the groups of quadrats where the shoots were attached. However, in dense stands of Brachypodium the positions of leaf clumps were not correlated to those of shoot clumps. This is a result of the tall growth form of this species and its high shoot densities, and it is suggested that this will be a characteristic of any species that dominates a dense stand.     

Vertical structure of wet grasslands under grazed and non‐grazed conditions in Tierra del Fuego

Abstract. We studied the vertical structure of wet grazed grasslands in Tierra del Fuego (southern Argentina). A point quadrat method was developed using a fine needle graduated in cm. The vertical and horizontal frequency of species and organs was quantified in samples collected from non‐grazed and grazed plots in the field. There was vertical stratification in both types of samples, but only in the first eight cm above the ground in grazed samples, with a dominance of Caltha sagittata. In non‐grazed samples graminoids grew taller than forbs and their inflorescences were an important element of the canopy structure. In both treatments, vertical species diversity was maximum in the lower part of the canopy, although diversity was significantly higher in grazed pots. In grazed samples, Caltha sagittata was the dominant species in 46% of samples and its leaves occupied 35% of the upper canopy. In the non‐grazed samples, Hordeum pubiflorum and Festuca magellanica were dominant in 63% of the samples, with H. pubiflorum leaves occupying 55% of the upper canopy. Comparing species by pairs, significant differences in vertical position were maintained in non‐grazed versus grazed pots. It is concluded that vertical stratification occurs even in the shortest communities. In this community, forb species grew close to the ground in the grazed areas, while forbs grew in the gaps and grasses above them in the non‐grazed areas. The main differences were in the relative dominance of forb and grass species and the presence of inflorescences.     

深圳市居住区绿地植物冠层格局对微气候的影响

选取深圳市3个典型居住小区为研究对象,利用实地观测及景观格局分析方法,探讨居住区绿地植物冠层格局特征对微气候的影响.各小区内布设22～26个观测点,以测点周边20 m×20 m范围为一个样区,将样区划分为有植物冠层遮盖和无植物冠层遮盖两种斑块类型,计算了斑块密度和景观聚集度两种格局指数,分析了样区内绿化植物冠层格局指数与温湿度之间的关系.结果表明:深圳市绿地冠层格局显著影响其温、湿度调控效应,温度调控能力主要来自绿地植物的遮荫作用,还与绿地的植物绿量有关;裸露地表聚集度的增加对绿地温湿度调控功能有显著的负面影响;居住小区绿地的温湿度调控能力主要来自乔木树种.     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Limited effect of hare grazing and short-term climatic variations on the most common alpine vegetation community in the Snowy Mountains,Australia

Background: In the alpine zone of the Snowy Mountains, grazing by mammals is limited. However, introduced European hare numbers have increased since the 1970s.

Aims: To estimate the density of hares and hence grazing pressure among years. To assess the response of biomass, vegetation height and composition to a cessation of hare grazing.

Methods: We used indices of hare abundance based on spotlighting and counts of hare pellets on a transect. The effect of hare grazing on tall alpine herbfield was assessed by using 15 paired exclosure and control quadrats for six years.

Results: The indices of hare abundance suggested densities similar to those in upland areas of Britain. Grazing did not affect the composition, cover of herbs or graminoids or, for 2010, vegetation height or biomass. Variation in vegetation and hare numbers among years was not correlated with climatic variables. Observations of selective grazing suggested that impacts on vegetation may be localised and restricted to certain species. Prior analyses of hare pellets indicated that hares might spread seed of native and exotic species.

Conclusions: Hares are having no general effect on tall alpine herbfield but may affect certain plant species via selective grazing or by spread of viable seed.     

Edaphic patchiness influences grassland regeneration from the soil seed‐bank in mountain grasslands of central Argentina

The soil seed‐banks in the main natural vegetation patches that make up mountain grasslands on granite substrates in central Argentina were studied. The main natural vegetation types are moist swards, tall‐tussock grasslands and stony grasslands. Ten compound soil samples from each community at two soil depths (0–5 and 5–10 cm) were taken. The density of soil seed‐banks was highest in moist swards, intermediate in stony grasslands and lowest in tall‐tussock grasslands. Low levels of similarity were found between the established vegetation and total soil seed‐bank in tall‐tussock grasslands and stony grasslands, but the similarity was higher in swards. In all three communities the seed‐bank was most frequently transient in nature. Short‐term persistent and long‐term persistent seed‐banks were less frequent. Regeneration from the seed‐banks after disturbance is expected to differ among communities on different edaphic patches. On the basis of the density and longevity of the soil seed‐banks and the similarity to the established vegetation, potential for in situ regeneration should be lower in tall‐tussock grasslands, intermediate in stony grasslands and higher in moist swards.     

Yield and species composition of a mesic grassland savanna in South Africa are influenced by long‐term nutrient addition

Species composition and productivity of natural grasslands are influenced by soil nutrient status. With high resource availability, productivity is expected to increase, and competition is assumed to gain prominence with predicted exclusion of species of lower competitive ability. During 2010 and 2011 we used the dry weight rank method to measure above‐ground phytomass production of herbage in 96 plots (9 m × 2.7 m) fertilized for 60 years with two forms of nitrogen (N as limestone ammonium nitrate or ammonium sulphate at four levels: 0, 7.1, 14.1, 21.2 g m^?2), phosphorus (P as superphosphate at two levels: 0, 33.6 g m^?2), and lime (two levels: 0, 225 g m^?2). Light attenuation was measured as the proportion of photosynthetically active radiation reaching the lower leaf layers of the grasses and the ground surface. Light conditions beneath the grass layer were reduced by nutrient addition to 30% of full sunlight but remained above 60% in non‐fertilized plots. Grass total above‐ground phytomass production increased with nutrient addition. The strongest yield responses were attained with N plus P addition. Species responses showed that Themeda triandra and Hyparrhenia hirta decreased in above‐ground phytomass production with nutrient addition while Panicum maximum, Eragrostis curvula and E. plana increased. These findings are discussed in terms of competitive interactions among species, their position in the grass canopy and their physiological tolerances to high nitrogen environments.     

Effect of Low Vegetation on the Recruitment of Plants in Successional Habitat Types1

I assessed the role of low vegetation (plants ca 1 yr old and ≤50 cm tall) as a biotic facilitator or barrier in the recruitment of different growth forms and species in primary forests, secondary forests, and old‐fields (abandoned pastures) in southeastern Mexico. I removed by hand all plants (≤100 cm tall, including roots) and litter from 20, 0.25 m² plots in each habitat. For 1 yr, I counted the number of plant species (5–50 cm tall) recruited, grouped them into different growth forms, and compared them to undisturbed control plots. Prior to manipulation, the standing density of trees and lianas was highest in primary and secondary forests. Shrubs were more abundant in secondary forests, whereas herbs, epiphytes, and hemi‐epiphytes were more abundant in old‐fields. Herbaceous plants appeared as important components of the community in all habitats. The removal of low vegetation increased total plant recruitment in all habitats. Considering each growth form, the absence of vegetation increased recruitment in primary forests for herbs, in secondary forests for epiphytes and hemi‐epiphytes, in old‐fields for trees, and for lianas in primary forests and old‐fields. In vegetation removal plots, recruitment of species was greater in pastures, lower in secondary forest, and similar in primary forest with respect to control plots. Depending on habitat type, species, and growth form, the presence of low vegetation may act as a recruitment barrier or facilitator for different species, affecting plant community structure, diversity, and composition in different habitats.     

Leaf dispersion and light partitioning in three-dimensionally digitized tall fescue–white clover mixtures

A three‐dimensional digitizing method was used to assess the canopy structure of six Festuca arundinacea (FA)–Trifolium repens (TR) mixtures during the installation stage. Virtual canopy images were synthesized and used to derive light interception and partitioning between species. Computations from images were compared with a simple light model based on Beer’s law, in order to analyse within‐ and between‐species foliage dispersion. The total leaf area index of the mixtures ranged from 0·6 to 4·5. The fraction of FA foliage overtopping TR was 9–30%. The mean inclination of FA and TR was 66 and 57°, respectively. Within‐species dispersion parameters of FA and TR were about 0·8 and 1·0, namely clumped and random foliage dispersion, respectively. Although FA was sown in rows, between‐species dispersion was random. Lower leaf inclination and lesser clumping in TR compensated foliage overtopping by FA, so that light partitioning between FA and TR (about 80 and 20%, respectively) was similar to the species contribution to total canopy foliage. Since between‐species dispersion was random, a simple two‐layer light model based on Beer’s law provided correct estimations of light partitioning (RMSE = 0·05), although light interception by FA was slightly overestimated because of its clumped dispersion.     

Effects of selective clipping and mowing time on species diversity in chalk grassland

Phytomass distribution and species diversity were followed in a chalk grassland with different selective clipping treatments during 4 years. Removal of the dominantBrachypodium resulted in a sharp increase in forb phytomass and in species number. The dominance-diversity curve changed in shape from geometric to log-normal. Obviously,Brachypodium restricted the growth of most other species. Removal of all species except the dominant showed, in contrast to the niche pre-emption theory, a marked increase in phytomass and reproductive effort ofBrachypodium. It seems that other mechanisms can also lead to a geometric distribution. Removal in a type of vegetation without obvious dominants and with a log-normal species distribution revealed that interspecific competition is not an important factor in this vegetation. This suggested a low intensity of species interaction and a large influence of restricting density-independent factors. Finally, to reduce the dominance ofBrachypodium we implemented a 4-year mowing experiment. Mowing earlier in the season resulted in a reduced exclusion of species byBrachypodium. This alternative mowing time may be sufficient to restore species diversity in chalk grassland. Experiments are under way to further substantiate this conclusion.     

Alpine and subalpine snow patch vegetation on the Bogong High Plains,SE Australia

Abstract. Snow patch vegetation in Australia is rare, being restricted to the relatively small area of alpine and subalpine country in the highlands of southeastern Australia. Snow patch vegetation occurs on steeper, sheltered southeastern slopes, where snow persists until well into the growing season (December/January). We surveyed the vegetation of 33 snow patch sites in the alpine and subalpine tracts of the Bogong High Plains, within the Alpine National Park, in Victoria. The vegetation was dominated by herbs and graminoids, with few shrubs and mosses. Major structural assemblages identified included closed herb‐fields dominated by Celmisia spp, and grasslands dominated by Poa fawcettiae or Poa costiniana. These assemblages occurred on mineral soils. Open herb‐fields dominated by Caltha introloba and several sedge species occurred on rocky and stony substrata. Vegetation‐environment relationships were explored by ordination and vector fitting. There was significant variation in the floristic composition of snow patch vegetation as a function of duration of snow cover, altitude, slope and site rockiness. Alpine sites were floristically distinct from subalpine sites, with a greater cover of Celmisia spp. and a lesser cover of low shrubs in the former. There was floristic variation within some snow patches as a function of slope position (upper, middle or lower slope) but this was not consistent across sites. The current condition of snow patch vegetation on the Bogong High Plains is degraded, with bare ground exceeding 20% cover at most sites. Snow patch vegetation is utilized preferentially by domestic cattle, which graze parts of the Bogong High Plains in summer. Such grazing is a potential threat to this rare vegetation type.     

Structure and dynamics of Japanese beech (Fagus japonica Maxim.) stools and sprouts in the regeneration of the natural forests

Structure and spatial distribution of stools and root-collar sprouts of Japanese beech (Fagus japonica) were studied to clarify the regeneration processes of the stool and the population, and the ecological importance of this stool formation in five quadrats of the natural forests with different forest floor vegetation on the Pacific side of Japan. F. japonica dominates in the canopy of each quadrat. Most of sprouts formed a circle around the root-collar and lowest parts of the parent stems of the stool with the youngest sprouts at the periphery. The regeneration by seedlings was slight especially on the forest floor vegetation of the dwarf bamboo Sasa. The variety of size structure of stems and the existence of dead traces and/or dead center in each stool, the apparent difference in stool size, and the aggregations of stools in the forests suggest that stool expansion and long persistence of the stool at a given location may contribute to compensate for the scarcity of regeneration by seedlings inhibited by dwarf bamboo, and by the other shrubs and herbs.     

Productivity and mineral cycling in an oak coppice forest 1. Structure and phytomass of the forest

Ecological studies were made on the structure and phytomass of the secondary coppice forest near Tokyo, which was dominated by a deciduous oak,Quercus serrata Thunb. Average height of dominant trees was about 10 m. The shoot density at the beginning of the study was 4,600 ha⁻¹ of which 89.5% belonged toQ. serrata. During the subsequent two growing seasons 8.3% of shoots, mainly small ones, died. All the tree shoots in a 10 m×10 m quadrat were cut and most of the underground parts were dug out. The phytomass calculated by the allometric relations of the dry weight of each plant organ to the square of DBH (D ²) agreed fairly well with the values directly weighed. The average phytomass of the overstory trees was 906 kg a⁻¹. The leaf area index (LAI) of the canopy was 3.85. Phytomass of the undergrowth, mainly a dwarf bamboo,Pleioblastus chino Makino, was 91.8 kg a⁻¹ with LAI of 3.46. The dead phytomass in the stand was 177 kg a⁻¹, so the phytomass alive and dead amounted to 1,170 kg a⁻¹. Heavy self-thinning of the coppice forest is discussed in relation to the rapid development of the log-normal distribution of tree sizes and to the large allometric constants for leaf and branch weight toD ².     

Population and community-level compositional patterns shape the realized niche of the rare arctic-alpine species Carex lachenalii Schkuhr

The ability for vegetative growth and development of generative organs often reflects an adaptation to the environment and may be a suitable proxy for understanding population dynamics of rare relict species. An example of such a plant is Carex lachenalii Schkuhr, an arctic-alpine species, in the temperate zone of Europe only occurring in isolated localities of high-elevation mountain ranges. We aimed to assess whether there were relationships between flower production and clonal growth of C. lachenalii, both at the tuft and plot level, and how co-occurring vegetation could modify this relationship. In the study we focused on population-level traits of C. lachenalii, vegetation traits and components of functional diversity. At the tuft level we found that the proportion of flowering ramets of C. lachenalii decreased with increasing diameter of the tuft. At the plot level, in snowbed vegetation C. lachenalii produced more flowering ramets. We suggest this is due to higher environmental stress, expressed by high importance of habitat filtering (low functional dispersion) in shaping species composition of co-occurring vegetation. In granite grasslands and milder environment (expressed by higher functional dispersion), C. lachenalii produced more vegetative ramets, which we suggest is a result of a more competitive environment. While in snowbeds investment in flowering ramets could promote successful persistence of C. lachenalii, survival of subpopulations occurring in the highly competitive conditions of granite grasslands may be uncertain due to potentially weak adaptation to competition with graminoids and dwarf shrubs.