Effects of harvester ants on seed availability and dispersal of Lavandula stoechas subsp. pedunculata in a Mediterranean grassland-scrubland mosaic

The role of harvester ants in Mediterranean grassland and scrubland has mostly focused on seed consumption. However, recent studies have reported their role as accidental dispersal agents of some of the collected seeds via refuse piles. The objective of this study is to examine the effect of the ant Messor barbarus on seed availability and dispersal of one of its major diet components, Lavandula stoechas subsp. pedunculata, in scrubland, grassland and the ecotones between them. After confirming and quantifying the Lavandula contribution to M. barbarus diet, we described the spatial and temporal patterns of pre- and post-dispersal seed predation, seed content and seedling occurrence in the refuse piles. Our results show that: (1) Lavandula propagules constitute a high proportion of the prey items collected by M. barbarus, with particularly intense collection activity in mid-summer, spring and autumn, in decreasing order. (2) Pre-dispersal predation rate was significantly higher in the ecotone than in the scrubland (76% and 13.5% of total seed production lost respectively). (3) Season and propagule type (seed vs. fruit) were the most significant variables explaining the post-dispersal predation probability, which approached 100% of seeds after 48 h in mid-summer. (4) Viable Lavandula seeds were found in refuse piles at densities of 0.06–0.2 per g of refuse pile material, or 58.8–207.2 per refuse pile. On the one hand, these results indicate that the ecotones are most affected by M. barbarus pre-dispersal consumption, which may locally limit Lavandula colonisation. On the other hand, the small proportion of consumed seeds that is dispersed to refuse piles may be relevant at the population level, as this dispersal implies arrival at potentially favourable sites for establishment.     

Post-fire seedling establishment in Florida sand pine scrub

Abstract. This study deals with a quantification of pre- and post-fire seedling establishment and microsite characteristics in two Florida sand pine scrub sites burned in May 1993. In addition, life history characteristics related to seedling establishment are described for five perennial species –Calamintha ashei, Chapmannia floridana, Eriogonum floridanum, Garberia heterophylla and Palafoxia feayi. Post-fire seedling establishment in sand pine scrub was sparse (median = 1, 12 seedling/m²), with 17 of 35 species establishing seedlings. Chapmannia, Eriogonum, Garberia and Palafoxia resprouted and flowered after fire; Eriogonum and Garberia had strong post-fire seedling establishment responses within 19 months post-fire. Calamintha individuals were killed by fire, but this species had a strong post-fire seedling establishment response, presumably from seeds in a soil seed bank. Eriogonum and Calamintha seedlings established preferentially in plots centered on conspecific adults. For these species with poor seed dispersal, spatial patterns of seedling establishment may be influenced more by pre-fire adult plant location than by post-fire microsite conditions. Post-fire seedling density in sand pine scrub was much lower than in California chaparral and South African sand plain lowland fynbos.     

Effects of intraspecific and interspecific density on the demography of a perennial herb, Sanicula europaea

We examined the effects of intraspecific and interspecific competition on demographic processes in the perennial herb Sanicula europaea by manipulating the density of neighbouring plants. We followed the response in terms of survival, growth and reproduction and in terms of seedling recruitment. The demographic data from all phases of the life cycle enabled us to assess also the overall effects of treatments on population growth rate (λ) by transition matrix models. We also decomposed the differences in λ between control and treatments, using life table response experiments (LTRE). To study the effects of competition on recruitment in more detail and to evaluate the role of seed availability, we sowed seeds at different densities with or without vegetation removal.
Vegetative growth and flowering frequency of established individuals was not significantly affected by removal treatments, which suggest no, or a delayed response to released competition. Neighbour removal had no effect on seedling emergence but enhanced recruitment through a higher seedling survival. Conspecific and simultaneous conspecific and heterospecific removal of plants led to an increase in population growth rate (λ), whereas heterospecific removal alone led to a decrease. Emergence of seedlings and fate of vegetative established individuals contributed most to differences in λ between the control and the different treatments. Seed addition enhanced seedling emergence but, as seedling and juvenile survival were density dependent, densities of established individuals appear not to be seed limited.
In S. europaea removal treatments had different effects on established individuals and recruitment. This suggests that studies quantifying the effects of competition over the entire life cycle and performed in a natural environment are necessary to assess the importance of competition in perennial plant populations.     

Effects of grassland extensification on the population dynamics of the invasive species Chaerophyllum aureum

Abstract. The consequences of extensification on grassland invasion by weeds was studied in French Pyrenean hay-meadows through a survey of the population dynamics of one very successful colonizer: Chaerophyllum aureum. Experiments established in natural permanent grasslands allow us to test the effect of different intensities of cutting on the demography of adult and seedling populations and on the production of seeds and their survival in soil. Although an early cut can control population density by drastically reducing invasion by new genotypes, the results reveal a positive effect of early cutting on adult and seedling survival probably by decreasing intra-specific competition. Allocation of resources to sexual reproduction in response to cutting is limited, affecting especially the seed viability in soil. Nevertheless, the invasive ability of this species is due largely to a strategy whereby those adults showing a high survival rate tend to be perennial.     

Effect of variation in self-incompatibility on pollen limitation and inbreeding depression in Flourensia cernua (Asteraceae) scrubs of contrasting density

Background and Aims

Selection may favour a partial or complete loss of self-incompatibility (SI) if it increases the reproductive output of individuals in the presence of low mate availability. The reproductive output of individuals varying in their strength of SI may also be affected by population density via its affect on the spatial structuring and number of S-alleles in populations. Modifiers increasing levels of self-compatibility can be selected when self-compatible individuals receive reproductive compensation by, for example, increasing seed set and/or when they become associated with high fitness genotypes.

Methods

The effect of variation in the strength of SI and scrub density (low versus high) on seed set, seed germination and inbreeding depression in seed germination (δ_germ) was investigated in the partially self-incompatible species Flourensia cernua by analysing data from self-, cross- and open-pollinated florets.

Key Results

Examination of 100 plants in both high and low scrub densities revealed that 51% of plants were strongly self-incompatible and 49 % varied from being self-incompatible to self-compatible. Seed set after hand cross-pollination was higher than after open-pollination for self-incompatible, partially self-incompatible and self-compatible plants but was uniformly low for strongly self-incompatible plants. Strongly self-incompatible and self-incompatible plants exhibited lower seed set, seed germination and multiplicative female fitness (floral display × seed set × seed germination) in open-pollinated florets compared with partially self-incompatible and self-compatible plants. Scrub density also had an effect on seed set and inbreeding depression: in low-density scrubs seed set was higher after open-pollination and δ_germ was lower.

Conclusions

These data suggest that (a) plants suffered outcross pollen limitation, (b) female fitness in partially self-incompatible and self-compatible plants is enhanced by increased mate-compatibility and (c) plants in low-density scrubs received higher quality pollen via open-pollination than plants in high-density scrubs.Key words: Flourensia cernua, population density, seed set, seed germination, female fitness, partial self-incompatibility, Mapimí Biosphere Reserve     

Old-field seedling responses to insecticide,seed addition,and competition

The controls of seedling emergence and survival determine the potential distribution of adult plants and, thereby, plant community structure. Seed availability, competition from established neighbors, and seedling predation may all limit seedling recruitment. In this field experiment, we followed the emergence and survival of seedlings of three perennial forbs, Achillea millefolium, Hypericum perforatum, and Monarda fistulosa, in old-fields in southeastern Michigan, USA. As adults, all three have aromatic foliage that may deter herbivory, but seedlings may be more susceptible than adults. To establish the relative importance of potential controls on seedling numbers, we manipulated seed availability through seed additions, the influence of competitors by neighbor-removals, and the influence of insect herbivores with insecticide in a fully factorial field experiment. Seed addition and insecticide never affected seedling emergence for any species. Competition from established neighbors controlled seedling emergence for all three species and decreased Achillea survival. Insecticide significantly increased Monarda seedling survival in competition plots, significantly increased Hypericum survival in open plots, and had no effect on Achillea. Notably, insecticide increased survival of the native Monarda fistulosa more than the two introduced species. While neighbors strongly reduced emergence and survival of all three species, herbivores acted on a species-specific basis. These results suggest the differential effects of insects may contribute more to the seedling species composition and abundance patterns than the less-selective influence of competition.     

Spatial patterns of seed predation by harvester ants (<Emphasis Type="Italic">Messor</Emphasis> Forel) in Mediterranean grassland and scrubland

Summary. Seed predation by harvester ants is one of the main processes involved in the seed bank dynamics of Mediterranean grassland and scrubland. We investigated the spatial patterns of seed predation by Messor spp. in these systems from a multi-level approach (nest-site selection, foraging patch selection and seed predation risk), using ten experimental plots (20 m × 40 m) set in central Spain. All habitats in grasslands are dominated by M. barbarus, while in scrubland this species shares seed exploitation with M. bouvieri. In this type of system, the nest-hole and foraging territory distribution of the two species show a certain degree of segregation. Messor bouvieri focuses on the exploitation of dry scrub, while M. barbarus also forages on subhumid and humid patches. We interpret this as a consequence of the different foraging behaviour and nest site requirements of the two species. In general, the spatial distribution of seed predation risk reflects the patterns detected for the nest-holes and foraging areas, although scrubland has a more heterogeneous predation pattern than grassland.     

Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophaga ruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males, appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.     

Ecological resistance, seed density and their interactions determine patterns of invasion in a California coastal grassland

Relatively little experimental evidence is available regarding how ecological resistance and propagule density interact in their effects on the establishment of invasive exotic species. We examined the independent and interactive effects of neighbour cover (biotic resistance), winter vs. spring water addition (abiotic resistance) and seed density on the invasion of the European perennial grass Holcus lanatus into a California coastal grassland dominated by exotic annual grasses. We found that decreased competition from resident exotic grasses had no effect. In contrast, increased late-season water availability eroded the abiotic resistance offered by naturally dry conditions, facilitating invasion. Finally, watering treatment and seed density interacted strongly in determining seedling survival: while seedling mortality was close to 100% in ambient and winter water addition plots, survivor numbers increased with seed density in spring-watered plots. Thus, decreased abiotic resistance can amplify the effect of increased propagule density on seedling establishment, thereby increasing the likelihood of invasion.     

Nitrogen addition decreases seed germination in a temperate steppe

Seed germination and seedling establishment play an important role in driving the responses of plant community structure and function to global change. Nitrogen (N) deposition is one of the driving factors of global change, which often leads to a loss in species richness in grassland ecosystems. However, how seed germination responds to N addition remains unclear. A pot incubation test was conducted in a semi‐arid grassland in the Mongolian Plateau, Northern China, to investigate the effect of N addition (0, 5, 10, 20, 40, and 80 g N/m²) on seed germination from May to October 2016. Twenty species germinated under all treatments; however, the responses of the 20 species to N addition were different. The densities of Stipa krylovii, Leymus chinensis, and Artemisia frigida, which are the dominant species in this temperate steppe, decreased significantly as the amount of N addition. Moreover, N addition significantly suppressed seedling densities of the community, perennial forbs, perennial grasses, and annuals and biennials. Furthermore, species richness of the community, perennial forbs, and annuals and biennials decreased sharply with increasing N addition level, but perennial grass species richness did not change. The Shannon–Wiener diversity index also decreased as the amount of N addition increased. Our results suggest that N enrichment plays an important role in the seed germination stage and decreases supplements of seedlings to adult plants. These findings may help explain the causes of species loss by atmospheric N deposition in grassland ecosystems.     

Recruitment limitations in Primula vulgaris in a fragmented landscape

Plant recruitment is limited by dispersal, if seeds cannot arrive at potential recruitment sites, and by establishment, due to a low availability of safe sites for recruitment. Seed-sowing experiments, scarcely applied along gradients of landscape alteration, are very useful to assess these limitations. Habitat loss and fragmentation may foster recruitment limitations by affecting all the processes from seed dispersal to seedling establishment. In this study, we perform a seed-sowing experiment to disentangle the importance of dispersal and establishment limitations in different stages of recruitment of the perennial herb Primula vulgaris in fragmented forests of the Cantabrian Range (Northwestern Spain). We evaluated the influence of ecological gradients resulting from habitat loss and fragmentation (modifications of habitat amount at the landscape and microhabitat scales, changes in the species’ population size, changes in seed predation and seedling herbivory) on seedling emergence, survival and early growth. We found strong evidence of dispersal limitation, as seedling emergence was very low in experimental replicates where no seeds were added. This limitation was independent of landscape alterations, as we found no relation with any of the ecological gradients studied. Establishment limitations at the germination phase were also unrelated to ecological gradients, probably because these limitations are more related to fine-scale environmental gradients. However, further monitoring revealed that seedling survival after summer and winter periods and seedling growth were conditioned by landscape alteration, as we found effects of habitat amount at the landscape and microhabitat scales, of presence of populations of P. vulgaris and of seedling herbivory. These effects were complex and sometimes opposite to what can be expected for adult plants, revealing the presence of different requirements between life stages.     

Does An Asexual Endophyte Symbiont Alter Life Stage and Long-Term Survival in a Perennial Host Grass?

Asexual, seedborne endophytic fungi in perennial grasses are often viewed as strong mutualists because fitness of the symbiont and host grass are closely coupled. However, at least for some native grasses, the asexual endophyte, Neotyphodium, acts parasitically, yet remains at high frequencies in natural populations. Most previous studies of Neotyphodium effects on host survival have been short term relative to the long life span of the perennial grass host. We therefore tested the hypothesis that Neotyphodium alters the survival in various life stages and long-term survival of adult native Arizona fescue (Festuca arizonica). To test the former, we planted 40 infected (E+) and 40 uninfected (E−, endophyte removed) seeds from four different maternal plants in the field under ambient conditions. We followed survival of seeds, seedlings, and adult plants over a 5-year period. To test the latter, we determined the infection of 1633 adult plants and followed their survival over the next 5–7 years. E+ seeds did not differ from uninfected seeds in terms of overall survival from seed germination to seedling to adult. However, the shape of the survival curve differed, with E+ plants showing higher mortality in early life stages. E+ adult plants did not differ from E− plants in long-term survival. Survival was generally very high during the study, which included a severe and prolonged drought. Infection by asexual Neotyphodium does not increase survival in early life stages or that of adult plants. Because asexual, vertically transmitted symbionts are predicted by evolutionary theory to be strong mutualists, the persistence of high infection frequencies in natural populations without long-term benefits to the host remains enigmatic. One possible explanation is that the long life span of the perennial host and low seedling recruitment may obscure either the costs or benefits of endophyte infection.     

Competition as a determinant of the upper limit of subtidal kelp Ecklonia stolonifera Okamura in the southern coast of Korea

Experiments in the subtidal zone of the southeastern coast of Korea examined the role of competition in determining the upper limit of perennial kelp, Ecklonia stolonifera, which is a dominant species in the subtidal community, but rarely found shallower than 5 m depth. Replicate clearings of dense algal turfs simulating natural disturbance at 1 and 3 m depth zones showed that E. stolonifera is able to settle and develop adult sporophytes in the upper zone. However, these adult sporophytes were eventually excluded from the clearings within one year after clearing. Two-year succession patterns at the clearings fell into three sequential categories: E. stolonifera and polychaete worms (Serpula vermicularis Linnaeus), mussels (Mytilus edulis Linnaeus), and turf-forming algae. A competitor removal treatment revealed that competition with M. edulis for primary space is the direct cause resulting in exclusion of E. stolonifera from the clearings. Recolonization of E. stolonifera after exclusion did not occur at clearings due to preemptive competition with other sessile organisms, which occupied 75-99% of bottom cover during the reproduction period of E. stolonifera (Autumn). Algal turfs, the final colonizer of the clearings, recovered more than 80% of bottom cover within 10 months after exclusion of M. edulis, and their species composition and abundance were not significantly different to those of adjacent controls. Experiments using artificial substrata showed that recruitment of E. stolonifera from spores is essentially impossible on the plates occupied by other sessile organisms (>86% cover). These results indicate that preemptive competition with dense algal turfs for the settlement and competition with M. edulis for survival in the course of succession are two most important factors determining the vertical distribution limits of E. stolonifera.     

Experimental evaluation of seed limitation in alpine snowbed plants

Background

The distribution and abundance of plants is controlled by the availability of seeds and of sites suitable for establishment. The relative importance of these two constraints is still contentious and possibly varies among species and ecosystems. In alpine landscapes, the role of seed limitation has traditionally been neglected, and the role of abiotic gradients emphasized.

Methodology/Principal Findings

We evaluated the importance of seed limitation for the incidence of four alpine snowbed species (Achillea atrata L., Achillea clusiana Tausch, Arabis caerulea L., Gnaphalium hoppeanum W. D. J. Koch) in local plant communities by comparing seedling emergence, seedling, juvenile and adult survival, juvenile and adult growth, flowering frequency as well as population growth rates λ of experimental plants transplanted into snowbed patches which were either occupied or unoccupied by the focal species. In addition, we accounted for possible effects of competition or facilitation on these rates by including a measure of neighbourhood biomass into the analysis. We found that only A. caerulea had significantly lower seedling and adult survival as well as a lower population growth rate in unoccupied sites whereas the vital rates of the other three species did not differ among occupied and unoccupied sites. By contrast, all species were sensitive to competitive effects of the surrounding vegetation in terms of at least one of the studied rates.

Conclusions/Significance

We conclude that seed and site limitation jointly determine the species composition of these snowbed plant communities and that constraining site factors include both abiotic conditions and biotic interactions. The traditional focus on abiotic gradients for explaining alpine plant distribution hence appears lopsided. The influence of seed limitation on the current distribution of these plants casts doubt on their ability to readily track shifting habitats under climate change unless seed production is considerably enhanced under a warmer climate.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Comparative water relations of adjacent california shrub and grassland communities

Summary Much of the coastal mountains and foothills of central and southern California are covered by a mosaic of grassland, coastal sage scrub, and evergreen sclerophyllous shrubs (chaparral). In many cases, the borders between adjacent plant communities are stable. The cause of this stability is unknown. The purpose of our study was to examine the water use patterns of representative grasses, herbs, and shrubs across a grassland/chaparrel ecotone and determine the extent to which patterns of water use contribute to ecotone stability. In addition, we examined the effects of seed dispersal and animal herbivory. We found during spring months, when water was not limited, grassland species had a much higher leaf conductance to water vapor diffusion than chaparral plants. As the summer drought progressed, grassland species depleted available soil moisture first, bare zone plants second, and chaparral third, with one chaparral species (Quercus durata) showing no evidence of water stress. Soil moisture depletion patterns with depth and time corresponded to plant water status and root depth. Rabbit herbivory was highest in the chaparral and bare zone as indicated by high densities of rabbit pellets. Dispersal of grassland seeds into the chaparral and bare zone was low. Our results support the hypothesis that grassland species deplete soil moisture in the upper soil horizon early in the drought, preventing the establishment of chaparral seedlings or bare zone herbs. Also, grassland plants are prevented from invading the chaparral because of low seed dispersability and high animal herbivory in these regions.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.     

Negative effects of habitat degradation and fragmentation on the declining grassland plant Trifolium montanum

Changes in land use have resulted in a strong decline in the plant diversity of nutrient-poor grasslands, but little is known about the combined effects of habitat degradation and fragmentation on populations of individual species. We studied these effects on stage structure, recruitment, reproduction and offspring fitness in populations of the declining perennial grassland plant Trifolium montanum in central Germany. Density and survival probability of juvenile plants decreased with light competition, measured as leaf area index (LAI) above T. montanum plants, resulting in aged populations with few juvenile plants at unmanaged sites with higher LAI. Reproduction of T. montanum was not related to LAI, but increased strongly with local density, suggesting pollinator limitation in fragmented populations with a low density of flowering plants. In the common garden, the survival of sown offspring increased with mean seed size, whereas seed production of offspring decreased with isolation, and in strong contrast to previous studies, also decreased with size and density of the population of origin. This could be due to increased inbreeding because of pollination between closely related neighbouring plants in dense and large populations. Our results indicate that both habitat degradation and fragmentation have negative effects on populations of T. montanum, but affect different phases of the life cycle. In the short term, the effects of habitat degradation are more important than those of fragmentation, and populations of T. montanum are primarily threatened by an increase in light competition in unmanaged sites, which rapidly affects the dynamics of the populations. The observed opposite effects of habitat fragmentation on reproduction and offspring fitness indicate that the effects of population size, density and isolation on plant fitness and population viability may be complex.