Floral volatiles controlling ant behaviour

1 . Ants show complex interactions with plants, both facultative and mutualistic, ranging from grazers through seed predators and dispersers to herders of some herbivores and guards against others. But ants are rarely pollinators, and their visits to flowers may be detrimental to plant fitness.
2 . Plants therefore have various strategies to control ant distributions, and restrict them to foliage rather than flowers. These 'filters' may involve physical barriers on or around flowers, or 'decoys and bribes' sited on the foliage (usually extrafloral nectaries - EFNs). Alternatively, volatile organic compounds (VOCs) are used as signals to control ant behaviour, attracting ants to leaves and/or deterring them from functional flowers. Some of the past evidence that flowers repel ants by VOCs has been equivocal and we describe the shortcomings of some experimental approaches, which involve behavioural tests in artificial conditions.
3 . We review our previous study of myrmecophytic acacias, which used in situ experiments to show that volatiles derived from pollen can specifically and transiently deter ants during dehiscence, the effects being stronger in ant-guarded species and more effective on resident ants, both in African and Neotropical species. In these plants, repellence involves at least some volatiles that are known components of ant alarm pheromones, but are not repellent to beneficial bee visitors.
4 . We also present new evidence of ant repellence by VOCs in temperate flowers, which is usually pollen-based and active on common European ants. We use these data to indicate that across a wide range of plants there is an apparent trade-off in ant-controlling filter strategies between the use of defensive floral volatiles and the alternatives of decoying EFNs or physical barriers.     

The ecological function of extrafloral nectaries: herbivore deterrence by visiting ants and reproductive output in Caryocar brasiliense (Caryocaraceae)

1. Shrubs of Caryocar brasiliense are intensively visited by many ant species in the cerrado vegetation of south-east Brazil. Ants collect nectar from the extrafloral nectaries (EFNs) located on the buds at the apex of the plant.
2. The proportion of C. brasiliense shrubs occupied by ants greatly surpassed that of neighbouring plants without EFNs and this resulted in many more termites being attacked and removed on this plant than on plants lacking EFNs. Ant attacks on Caryocar were more frequent near the EFNs.
3. Ant-exclusion experiments with Caryocar shrubs revealed that ants significantly reduce the infestation levels by the butterfly Eunica bechina , the sucking bud Edessa rufomarginata , the bud-destroying fly Prodiplosis floricola and by a stem-galling wasp. Ants had no effect on the infestation by a leaf-galling wasp.
4. Control (ants present) and treatment (ants excluded) plants produced equal numbers of buds but flower and initial fruit production was significantly greater for ant-visited plants. Higher abortion rates of initial fruits by control plants resulted in similar final fruit and seed sets for plants in either experimental category.
5. The extremely low fertility of cerrado soils is suggested as a possible factor constraining the ability of Caryocar shrubs to provide the resources necessary for growth and development of their heavy fleshy fruits. Higher production of initial fruits by ant-visited shrubs would be neutralized by competition among developing fruits and subsequent abortion.
6. Possible ant-derived benefits to C. brasiliense include: protection of vegetative tissues during plant growth, larger floral display and increased attractiveness to bat pollinators, increased male contribution to fitness through pollen donation and selective abortion of genetically inferior progeny as the maternal investment is adjusted to match available resources.     

Plant-attracted ants affect arthropod community structure but not necessarily herbivory

Abstract.  1. The effectiveness of ants as plant defenders is equivocal for plants that attract ants via extrafloral nectaries (EFNs).
2. This study focused on the myrmecophilic savannah tree Pseudocedrela kotschyi that attracts ants to EFNs and on the arthropod fauna associated with P. kotschyi . Herbivory and arthropod community composition were compared between trees that were dominated by one of three congeneric ant species, Camponotus acvapimensis , C. rufoglaucus , and C. sericeus , and between trees where ants were experimentally excluded and untreated control trees.
3. Short-term ant-exclusion experiments failed to demonstrate a consistent effect of ants on herbivory.
4. Plants dominated by different ant species differed significantly in leaf damage caused by herbivorous insects. The relative ranking of herbivory levels of the trees dominated by different ant species was persistent in three consecutive years.
5. Ants significantly reduced the abundance of different arthropod groups (Araneae, Blattodea, Coleoptera, Homoptera, non-ant Hymenoptera). Other groups, including important herbivores, seemed not to be affected (Lepidoptera, Orthoptera, Thysanoptera, Heteroptera).
6. The study suggests that the presence of ants only benefits plants when specific ant species are attracted, and protection by these ants is not counterbalanced by their negative effect on other beneficial arthropods.     

Tree-shrub associations in grazed woodlands: first rodents,then cattle?

Spatial associations of tree saplings with spiny or toxic plants in grazed woodlands are generally explained by associational resistance, i.e., protection against grazing via a well-defended neighbor. In this study, we tested whether directed seed dispersal and post-dispersal seed removal by wood mice are additional explanations for the observed spatial association between thorny shrubs and trees, hence before associational resistance. We performed three studies in grazed woodlands in the Netherlands to test this idea. Our first seed dispersal experiment with tagged acorns showed indeed that wood mice disperse acorns directed towards shrubs. The majority of these dispersed acorns were, however, consumed. Our second experiment revealed that post-dispersal removal of cached acorns was higher under shrubs than in grassland and under trees, but also indicated the importance of within-shrub position: shrub centers were faster depleted than shrub edges. Also the number of freshly emerged seedlings and older saplings differed between micro-habitats, being higher under trees and at shrub inner-edges than at shrub outer-edges, shrub centers, and in grassland. The spatial associations with shrubs got stronger with the age of saplings, which probably reflect accumulated grazing effects over time, being higher in the open than in shrubs. We conclude that directed acorn dispersal and post-dispersal acorn removal by wood mice are two additional explaining mechanisms for the spatial associations between thorny shrubs and trees in grazed woodlands, before associational resistance. Our findings change the view that large herbivores are the sole main drivers behind tree recruitment patterns in grazed woodlands.     

Conspicuous extra-floral nectaries are inducible in Vicia faba

Mutualistic interactions are dynamic associations that vary depending on the costs and benefits to each of the interacting parties. Phenotypic plasticity in mutualistic interactions allows organisms to produce rewards to attract mutualists when the benefits of their presence outweigh the costs of producing the rewards. In ant–plant defensive mutualisms, defences are indirect as plants produce extra‐floral nectaries (EFN) to attract predatory ants to deter herbivores. Here we demonstrate that in broad bean, Vicia faba, the overall number of EFNs on a plant increases dramatically following leaf damage. In two damage treatments, removal of: (1) one‐third of one leaf in a single leaf pair or (2) one‐third of both halves of a single leaf pair, resulted in a 59 and 106% increase in the number of EFNs on the plants, respectively, over 1 week. We suggest that the increased production of visually conspicuous EFNs is an adaptive inducible response, to attract predatory arthropods when risk of herbivory increases.     

Effectiveness of protein-rich pea flour for the control of stored-product beetles

Abstract The association between visiting ants and the extrafloral nectaries (EFN)‐bearing shrub Hibiscus pernambucensis Arruda (Malvaceae) was investigated in two different coastal habitats – a permanently dry sandy forest and a regularly inundated mangrove forest. In both habitats the frequency of plants with ants and the mean number of ants per plant were much higher on H. pernambucensis than on non‐nectariferous neighbouring plants. In the sandy forest the proportion of live termite baits attacked by ants on H. pernambucensis was much higher than on plants lacking EFNs. In the mangrove, however, ants attacked equal numbers of termites on either plant class. Ant attendance to tuna/honey baits revealed that overall ant activity in the sandy forest is higher than in the mangrove area. The vertical distribution (ground vs. foliage) of ant activity also differed between habitats. While in the mangrove foraging ants were more frequent at baits placed on foliage, in the sandy forest ant attendance was higher at ground baits. Plants housing ant colonies were more common in the mangrove than in the sandy forest. Frequent flooding in the mangrove may have resulted in increased numbers of ant nests on vegetation and scattered ant activity across plant foliage, irrespective of possession of EFNs. Thus plants with EFNs in the mangrove may not experience increased ant aggression towards potential herbivores relative to plants lacking EFNs. The study suggests that the vertical distribution of ant activity, as related to different nest site distribution (ground vs. foliage) through a spatial scale, can mediate ant foraging patterns on plant foliage and probably affect the ants’ potential for herbivore deterrence on an EFN‐bearing plant species.     

Game theory provides no explanation for seed size variation in grasslands

Game theoretical models have been suggested to explain the maintenance of a remarkable variation in seed size across species in most types of vegetation. According to these models, which are based on the existence of a trade-off between seed size and seed number; smaller-seeded species can invade any species mixture due to their numerical advantage, and larger-seeded species can invade any species mixture due to their competitive superiority over smaller-seeded species during recruitment. However, till now, there is very limited evidence for seed size effects on recruitment interactions among different species. An experiment was designed using 16 species in Swedish grasslands, varying 384-fold in seed size. Species were sown pairwise and alone in disturbed versus undisturbed small plots in grazed versus ungrazed grassland, and the resulting recruitment was recorded. Seedling densities in the experimental plots were within the natural range. Both disturbance and grazing had a positive effect on recruitment. Seed size did not affect recruitment except from a tendency that increasing seed size made recruitment less dependent on disturbance. Recruitment of smaller-seeded species was not affected by the presence of larger-seeded species. Larger-seeded species did not generally win in direct contest. These results suggest that game theoretical models do not explain maintenance of seed size variation across species in these grasslands. Alternative explanations for seed size variation are that either small-scale heterogeneity provide conditions favouring a range of different seed sizes or other attributes than seed size effectively determine recruitment.     

Effects of ant behaviour and presence of extrafloral nectaries on seed dispersal of the Neotropical myrmecochore Turnera ulmifolia L. (Turneraceae)

This paper describes the myrmecochory system of Turnera ulmifolia in a coastal sand dune matorral in Mexico. Turnera ulmifolia has elaiosome‐bearing seeds and extrafloral nectaries (EFNs). In ten quadrants (4 × 15 m) ant–seed interaction was monitored, and an interaction intensity index calculated and correlated with the number of seedlings. Seed removal rates by ants were surveyed every 2 h for 24 h, the ants being observed both on and beneath the plants. The role of the elaiosome in seed removal was evaluated by offering seeds with and without elaiosomes, and elaiosomes only. Finally, the effect of ant manipulation in seed germination was evaluated. There were 25 ant species associated with seeds and/or EFNs, the most frequently recorded being Monomorium cyaneum and Forelius analis. There was a positive correlation between the intensity index and seedling number per quadrant. There was significantly higher mean seed removal during the day than during the night (19.3% and 12.3%, respectively), and from beneath than on the plant (21.9% and 9.5%, respectively). The preference for elaiosomes only was also greater during the diurnal period, and when gathered on, rather than beneath, the plant. Seed manipulation by F. analis enhanced germination by T. ulmifolia. Seed removal, dispersal distances, seed predation and germination were largely determined by ant behaviour. The presence of EFNs may be influencing seed removal on the plant by attracting a specific assemblage of omnivorous ants. Among such assemblages associated with T. ulmifolia we encountered a variety of behaviours, with ant species either good at defending plants but bad at dispersing seeds, or vice versa. We discuss the way in which these two rewards, and the processes involved (defence and dispersion), could have interacted with each other and evolved. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 67–77.     

Jumping spiders (Salticidae) enhance the seed production of a plant with extrafloral nectaries

Many plants secrete nectar from extrafloral nectaries (EFNs), specialized structures that usually attract ants which can act as plant defenders. We examined the nectar-mediated interactions between Chamaecrista nictitans (Caesalpineaceae) and jumping spiders (Araneae, Salticidae) for 2 years in old fields in New Jersey, USA. Previous research suggests that spiders are entirely carnivorous, yet jumping spiders (Eris sp. and Metaphidippus sp.) on C. nictitans collected nectar in addition to feeding on herbivores, ants, bees, and other spiders. In a controlled-environment experiment, when given a choice between C. nictitans with or without active EFNs, foraging spiders spent 86% of their time on plants with nectar. C. nictitans with resident jumping spiders did set significantly more seed than plants with no spiders, indicating a beneficial effect from these predators. However, the presence of jumping spiders did not decrease numbers of Sennius cruentatus (Bruchidae), a specialist seed predator of C. nictitans. Jumping spiders may provide additional, unexpected defense to plants possessing EFNs. Plants with EFNs may therefore have beneficial interactions with other arthropod predators in addition to nectar-collecting ants. Received: 27 May 1998 / Accepted: 23 December 1998     

Interaction between ants, extrafloral nectaries and insect herbivores in Neotropical coastal sand dunes: herbivore deterrence by visiting ants increases fruit set in Opuntia stricta (Cactaceae)

1. This study examines the anti-herbivore effect of ants visiting the extrafloral nectaries (EFNs) of Opuntia stricta (Cactaceae) and its possible influence on the plant's reproductive output in Mexican coastal sand dunes. Opuntia 's EFNs are located in the areoles of the developing tissue of emerging cladodes and flower buds.
2. Ants visited the EFNs of O . stricta on a round-the-clock basis. The associated ant assemblage was formed by nine species distributed in four subfamilies, and the species composition of the principal ant visitors changed markedly from day to night period.
3. Cladodes of control (ants present) and treatment (ants excluded) plants of Opuntia were equally infested by sucking bugs and mining dipterans. Damage to buds by a pyralid moth, however, was significantly higher on treatment than on control plants. Ant visitation to Opuntia 's EFNs translated into a 50% increase in the plant's reproductive output, as expressed by the number of fruits produced by experimental control and treatment branches. Moreover, fruit production by ant-visited branches was positively and significantly associated with the mean monthly rate of ant visitation to EFNs.
4. This is the first demonstration of ant protection leading to increased fruit set in the Cactaceae under natural conditions. Although the consequences of damage by sucking and mining insects remain unclear for Opuntia , the results show how the association of EFNs with vulnerable reproductive plant organs can result in a direct ant-derived benefit to plant fitness.     

Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Spatial variation in the demography and population dynamics of a perennial shrub (Atriplex vesicaria) under sheep grazing in semi‐arid Australian rangelands

Atriplex vesicaria Heward ex Benth. (Chenopodiaceae) is a widespread perennial shrub in southern Australia's chenopod rangelands but is sensitive to grazing. A detailed investigation of the demography and population dynamics of A. vesicaria under sheep grazing was conducted over 6.5 years at a range of sites across a typical paddock to assess the long‐term effects of grazing on the species and elucidate the mechanisms of population change under grazing. The effects of rainfall on recruitment and mortality were also examined. Six‐monthly censuses of all A. vesicaria individuals were conducted in permanent grazed and ungrazed plots at sites located across an 1100‐ha paddock. Grazing increased adult shrub mortality close to water and reduced recruitment over a broader area of the paddock, but seedling survival did not appear to be affected by grazing. As a result of these changes, the population declined on grazed plots up to 2200 m from water during the study, but the decline was greatest closer to water. The population was most dynamic at the sites furthest from the water point where it was unaffected by grazing because of the greater recruitment and mortality of young plants, but because these processes balanced out over time, population density was effectively unchanged by the end of the study. Although statistical models indicated that six‐monthly rainfall did not explain temporal variation in recruitment or mortality, rainfall nevertheless has a central role in both processes. In particular, longer periods of favourable rainfall and drought appear to have an important influence on recruitment and mortality, respectively, with heavy grazing during a drought period increasing mortality. Occasional shortages of seed or rains occurring during the warmer months when seed germination is limited possibly explain poor recruitment at sites unaffected by grazing following good rainfall.     

A fungal endosymbiont affects host plant recruitment through seed- and litter-mediated mechanisms

1.  Many grass species are associated with maternally transmitted fungal endophytes. Increasing evidence shows that endophytes enhance host plant success under varied conditions, yet studies have rarely considered alternative mechanisms whereby these mutualistic symbionts may affect regeneration from seed.
2.  We performed a microcosm experiment to evaluate whether infection with Neotyphodium occultans affects recruitment in the annual grass Lolium multiflorum either directly, by infecting the seeds, or indirectly, by altering the suitability of recruitment microsites through the litter shed by host plants. Endophyte effects on establishment were tested for different litter depths and watering regimes under natural herbivory by leaf-cutting ants.
3.  Seed infection increased seedling emergence through the litter as well as final recruitment, irrespective of microsite conditions. However, litter produced by infected plants delayed emergence and decreased density of both infected and non-infected grass populations.
4.  Individual plant biomass did not change with seed infection but was increased under deep litter from endophyte-infected plants. Although seed infection did not protect establishing plants from leaf-cutting ants, herbivory was reduced in the presence of deep litter shed by infected plants.
5.  We conclude that fungal endophytes may affect host plant recruitment across subsequent generations not only by infecting the seeds but also through the host's dead remains. While the former effect entailed an advantage to infected plants, litter-mediated effects did not discriminate by infection status, and generally promoted the establishment of fewer and larger plants. Thus hidden foliar symbionts may play an underappreciated role in maintaining host species dominance through the litter produced by prior patch occupants.     

Sheep grazing vs. cutting: regeneration and soil nutrient exploitation of the grassland weed Rumex obtusifolius

Broad-leaved dock (Rumex obtusifolius L., Polygonaceae) is an agronomically important perennial weed causing decreases in pasture yields and fodder quality. Non-chemical control measures for dock are often limited to frequent pulling and cutting, additionally it is usually avoided by grazing animals. Here, the regeneration of R. obtusifolius in a Rumex-infested grassland grazed by a sheep breed that explicitly feeds on dock (Ovis aries L. cv. East Prussian Skudden) was compared to cutting. Therefore, regeneration of 90 R. obtusifolius plants of three different size classes was monitored in three plots during three grazing and cutting cycles. Plant height and number of fruit-stands of regrown R. obtusifolius was significantly lower, number of leaves significantly higher after grazing than after cutting, while plant diameter was unaffected. Initially medium and large-sized plants (>40 cm diameter) were significantly more sensitive to grazing or cutting than initially smaller sized plants. Soil nitrate and ammonium concentrations in the vicinity of R. obtusifolius correlated with some regrowth parameters but were not affected by grazing or cutting. Sheep-grazed grassland communities comprised fewer legumes (p = 0.002), more grasses (p = 0.010) and fewer sward gaps (p = 0.025) than cut grassland. At the end of the experiment, abundance of R. obtusifolius in sheep grazed plots was lower than in cut plots (p = 0.089) suggesting that regrowth potential of this weed was depleted by continuous grazing and higher sward density. In conclusion, these data suggest that sheep could be considered in grassland management schemes to both directly and indirectly control Rumex infestations.     

Effects of grazing on seedling establishment: the role of seed and safe-site availability

Abstract. The first objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment. We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been experimentally defoliated. The species compared were Dan-thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodón taraxacoides, an invading exotic rosette species from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-term regeneration capacity after disturbance was lost.     

科尔沁沙质草地放牧和围封条件下的土壤种子库

该文研究了科尔沁沙质草地在放牧和围封条件下土壤种子库密度、组成及其与地上植被的关系。结果表明:1)放牧草地植物种数22种,围封草地植物种数30种,围封使土壤种子库植物种数增加了36%;2)放牧草地土壤种子库密度为16 149±1 900有效种子数·m^-2, 围封草地土壤种子库密度为20 657±3 342有效种子数·m^-2,比放牧草地增加了28%。放牧和围封草地种子库组成密度均以一年生植物为主(分别占99%和98%的比例),多年生植物所占的比例很小;3)放牧草地种子库的Shannon-Wiener指数和丰富度指数分别为0.836 3和4.954 9,明显小于围封草地的0.968 2和7.226 0,表明自由放牧导致物种多样性下降;4)放牧和围封草地土壤种子库密度与地上植被密度均存在显著的相关性(p<0.001)。表明了随着土壤种子库密度的增加, 地上植被密度随之增加,放牧草地地上植被密度78%的变异可归结为土壤种子库密度的变异, 而围封草地地上植被密度58%的变异可由土壤种子库密度的变异来解释。     

Response of Grassland Birds in Sand Shinnery Oak Communities Restored Using Tebuthiuron and Grazing in Eastern New Mexico

Sand shinnery oak (Quercus havardii) communities are a unique component of grassland bird habitat in eastern New Mexico and have been impacted by human activities for decades. These communities are frequently managed with livestock grazing and herbicide application for shrub control, strategies that potentially can be used to restore the historical shrub–grass composition of this plant community. During spring migration and the breeding seasons of 2004 and 2005, we compared density and community structure of grassland bird species among four combinations of tebuthiuron application and grazing treatments that were being evaluated for restoration of shinnery oak communities. We performed biweekly point transects on sixteen 65‐ha study plots in these communities. Density of all avian species combined did not differ between grazed and ungrazed plots. Tebuthiuron‐treated plots had a 40% higher average density for combined species than untreated plots. There was a 41% higher average density of all species during spring 2005 than 2004, but density was similar during the breeding season of both years. These trends were predominantly influenced by densities of migratory Cassin’s Sparrow (Aimophila cassinii), which were greater in tebuthiuron‐treated plots in both years. Densities of resident Meadowlarks (Sturnella spp.) exhibited little response to tebuthiuron or grazing treatments. Avian species richness, evenness, and diversity were only minimally affected by the tebuthiuron and grazing treatments. This study occurred over a period of highly variable precipitation, so future assessments, spanning longer wet–dry cycles and maturing plant communities, may be necessary to completely determine avian response to these restoration efforts.     

COMPONENTS OF SEED AND POLLEN YIELD OF LOBELIA CARDINALIS: VARIATION AND CORRELATIONS

Plants from three Lobelia cardinalis populations were grown under common garden conditions to assess intra- and interplant variation in seed and pollen production. Seed number per flower and mean seed weight varied systematically with floral position on the inflorescence (lowest values were from terminal flowers) but pollen grain number per flower did not vary systematically with floral position. Most of the remaining variance in seed and pollen grain number per flower and mean seed weight was distributed among plants; clones produced very similar amounts of pollen and seed. Seed yield was positively correlated with seed production per flower and with total flower production, but not with mean seed weight; pollen yield was also positively correlated with pollen grain production per flower and total flower production. Seed and pollen yield were simple linear functions of plant size but only pollen yield was a simple linear function of flower production; seed yield was a quadratic function in which the second order term was negative. This quadratic relationship resulted from a negative correlation between seed number per flower and total flower production. This correlation, in addition to the wide variation among plants in pollen number per flower, accounts for the weak correlation of seed and pollen yield. I conclude from these data that it is unlikely that plants in natural L. cardinalis populations transmit genes to the population's seed crop equally through pollen and ovules—emphasizing the importance of measuring both male and female components of reproductive success.     

Recreating Grasslands in Swedish Rural Landscapes – Effects of Seed Sowing and Management History

Recent loss of plant species richness in Swedish semi-natural grasslands has led to an increase in grassland recreation and restoration. To increase the establishment of declining species favoured by grazing and to re-establish original species richness, seed sowing has been discussed as a conservation tool. In this study, I examined to what extent seed sowing in former arable fields increases species richness and generates a species composition typical of semi-natural grasslands. Six grassland species favoured by grazing (target species) and six generalist species favoured by ceased grazing, were studied in a seed-addition experiment. Four different seed densities were used on four different grassland categories, two grazed former arable fields, one continuously grazed grassland and one abandoned grassland. Target and generalist species emerged in all grassland categories, but seedling emergence was higher in the grazed than in the abandoned grassland. Target species had higher emergence in the two grasslands with the longest grazing continuity. Seedling emergence and frequency of established plants of each target species were positively associated. The largest fraction of seeds germinated at an intermediate sowing density, 20–50 seeds/dm², suggesting that aggregation of seeds positively affects emergence up to a certain threshold. In conclusion, artificial seed sowing may induce the recreation of typical grassland communities on former arable fields, which may be an important contribution to increase the total grassland area and species richness in the landscape.     

Effects of grazing on soil seed bank dynamics: An approach with functional groups

Abstract. The relationship between intensity and timing of cattle grazing on changes in the size and composition of the soil seed bank were investigated in a 3‐yr study in a Mediterranean grassland in northeastern Israel. Treatments included manipulations of stocking rates and of grazing regimes, in a factorial design. The retrieved soil seed bank community was rich in species, with 133 species accounting for 80% of the 166 species recorded at the site. Within the seed bank, 89% of the species were annuals. Seed bank dynamics was analysed in terms of plant functional groups and germination strategies. In terms of total seed bank density and including all functional groups, 42% of the seeds present in the soil did not germinate under watering conditions. The dormancy level differed greatly among functional groups. The seed bank of annual legumes, crucifers, annual thistles and annual forbs had a large fraction of non‐germinated seeds and characterized areas grazed early in the growing season under high and very high grazing intensity. These functional groups were considered to have a higher potential for persistent seed banks production. In contrast, short and tall annual grasses and tall perennial grasses, that were dominant in ungrazed or moderately grazed paddocks, generally had seed banks with a very small fraction of non‐germinated seeds. Seed bank densities varied widely between grazing treatments and years. Under continuous grazing, heavy grazing pressure reduced seed bank densities of grasses and crucifers in comparison to moderate grazing. The greatest reduction on the seed bank densities resulted from heavy grazing concentrated during the seed‐set stages.