Differential herbivory tolerance of dominant and subordinate plant species along gradients of nutrient availability and competition

We tested whether differences in the herbivory tolerance of plant species is related to their abundance in grassland communities and how herbivory and nutrient availability affect competitive balances among plant species through changes in their tolerance. The experimental approach involved a simulated grazing treatment (clipping) of two competitive grass species (Arrhenatherum elatius and Holcus lanatus) and two subordinate forb species (Prunella vulgaris and Lotus corniculatus) along a gradient of nutrient availability and under conditions of competition. Total standing, aboveground, root, and regrowth biomass were evaluated at the end of the experiment as an estimate of the capacity to compensate for twice removing aboveground biomass at different nutrient levels (NPK). Although clipping had a more pronounced negative effect on dominant plant species (Arrhenatherum and Holcus) than on subordinate species, the negative effects on dominant species were offset by the application of fertilizer. The combined effect of fertilizer and competition had more negative effects on the performance of Lotus and Prunella than on the dominant species. In terms of competition, the regrowth ability of Arrhenatherum and Holcus increased with the application of fertilizer, while the opposite pattern was observed for Lotus and Prunella. The addition of fertilizer has a positive effect on both grass species in terms of growth in clipped pots and competition, while subordinate species did not respond to the addition of fertilizer to the clipped pots and were negatively affected by competition with both grass species. The results suggest (1) that species replacement towards subordinate species as a function of herbivory is partially dependent on the herbivory tolerance of that species, (2) competitive relations between competitive grass species and subordinate forb species change under different environmental conditions, and (3) although grazing disturbance significantly influences competitive relations in favor of less competitive species, increasing nutrient levels counteract the negative effect of grazing on dominant competitive plant species.     

Productivity and Subordinate Species Response to Dominant Grass Species and Seed Source during Restoration

Grasses can be important regulators of species diversity and ecosystem processes in prairie systems. Although C₄ grasses are usually assumed to be ecologically similar because they are in the same functional group, there may be important differences among species or between seed sources that could impact restorations. I tested whether C₄ grass species identity, seed source, or grass species richness scales to influence aboveground net primary productivity (ANPP), resistance to weed invasion, or establishment of subordinate prairie species during restoration. Plots in western Iowa, United States, were planted with equal‐sized transplants of one of five common grass species (Panicum virgatum L., Sorghastrum nutans (L.) Nash, Andropogon gerardii Vitman, Schizachyrium scoparium (Michx.) Nash, and Bouteloua curtipendula (Michx.) Torrey) either from local seed or from cultivar seed sources. These plots were compared to plots containing all five species in mixture and to nonplanted plots. Differences in ANPP were found among species but not between cultivars and noncultivars or between monocultures and mixtures. Panicum virgatum, S. nutans, and S. scoparium were more productive than A. gerardii and B. curtipendula. Weed invasion was much higher when plots were not planted with grasses. Schizachyrium scoparium allowed greater establishment of subordinant prairie species than all other focal grass species. There were two separate mechanisms by which grasses suppressed prairie species establishment either (1) by growing tall and capturing light or (2) by quickly filling in bare space by spreading horizontally through rhizome growth in short species. These results suggest that high ANPP can be found with noncultivar plantings during the first 2 years after planting and that subordinate species establishment is most likely when shorter bunchgrasses such as S. scoparium are dominant.     

Importance and intensity of competition along a fertility gradient and across species

Questions: 1. Can the importance and the intensity of competition vary independently along a nutrient gradient? 2. Are these variations species dependent? Location: Sub‐alpine pastures of the northern French Alps. Methods: Competition intensity measures how much competition decreases the performances of an organism. Competition importance measures how much competition contributes to affect performance, among other processes (such as environmental stress or disturbance). Competition intensity and importance were measured on three co‐occurring species: Festuca rubra, a perennial grass, and two forbs of contrasting basal area, Chaerophyllum hirsutum and Alchemilla xantho‐chlora. A neighbour removal experiment was performed on Festuca rubra in three sub‐alpine grassland communities differing in fertility and on Chaerophyllum hirsutum and Alchemilla xanthochlora in the two more fertile of these communities. The importance of competition was quantified using an index proposed by Brooker et al. (2005). Results: Competition intensity and importance showed different patterns of variation along the fertility gradient for Festuca rubra: competition importance decreased with decreasing fertility whereas competition intensity did not change. The largest forb was the least affected by competition. Our results suggest that the importance of competition for all three species depended on their individual tolerance to low nutrient availability. Conclusions: 1. The distinction between the importance and the intensity of competition is helpful to explain conflicting results obtained on the variations of competition indices along productivity gradients. 2. The choice of a phytometer can affect the conclusions drawn from empirical studies.     

Vertically transmitted fungal endophytes: different responses of host-parasite systems to environmental conditions

The relationship between vertically transmitted asexual fungal grass endophytes and their hosts is considered to be mutualistic. Results from agronomic field support this line of reasoning but recent studies have shown more variable results in natural systems. We investigated how high and low nutrient and water treatments affected biomass allocation patterns of endophyte‐infected and uninfected Festuca pratensis and F. rubra in greenhouse experiments over two growing seasons. Irrespective of infection status, both grass species showed improved performance on highly fertilized and watered soils. However, infected F. pratensis plants produced larger tillers than endophyte‐free plants on soil low in nutrients and water in the first growing season, although they (E+) otherwise showed decreased performance on nutrient‐poor soil. In low nutrient and water conditions, endophyte‐infected plants produced less tillers and had lower total biomass compared to uninfected plants, and displayed a negative phenotypic correlation between seed production and vegetative growth. The latter indicates costs of reproduction when the plant shares common resources with the fungal endophyte. However, endophyte infection status (E+, E?) interacted significantly with the soil fertilisation in terms of plant growth, having a stronger positive effect on growth in infected F. pratensis plants. In F. rubra, endophyte‐infected plants showed higher vegetative growth in fertilized and watered soils compared to uninfected plants. However, infected plants tended to produce fewer inflorescences. This had no effect on seed production, perhaps because seed production was partly replaced by asexual pseudovivipary. Contrary to the general assumption in the literature that fungal endophytes are plant mutualists, these findings suggest that the costs of endophytes may outweigh their benefits in resource limited conditions. However, the costs of endophyte infections appear to differ among the grass species studied; costs of endophytes were mainly detected in F. pratensis under low nutrient conditions. We propose that differences in response to endophyte infection in these species may depend on the differences in life‐history strategies and environmental requirements of these two fescue and fungal species and may change during the life span of the plant.     

Root Dynamics of Cultivar and Non‐Cultivar Population Sources of Two Dominant Grasses during Initial Establishment of Tallgrass Prairie

Dominance of warm‐season grasses modulates tallgrass prairie ecosystem structure and function. Reintroduction of these grasses is a widespread practice to conserve soil and restore prairie ecosystems degraded from human land use changes. Seed sources for reintroduction of dominant prairie grass species include local (non‐cultivar) and selected (cultivar) populations. The primary objective of this study was to quantify whether intraspecific variation in developing root systems exists between population sources (non‐cultivar and cultivar) of two dominant grasses (Sorghastrum nutans and Schizachyrium scoparium) widely used in restoration. Non‐cultivar and cultivar grass seedlings of both species were isolated in an experimental prairie restoration at the Konza Prairie Biological Station. We measured above‐ and belowground net primary production (ANPP and BNPP, respectively), root architecture, and root tissue quality, as well as soil moisture and plant available inorganic nitrogen (N) in soil associated with each species and source at the end of the first growing season. Cultivars had greater root length, surface area, and volume than non‐cultivars. Available inorganic N and soil moisture were present in lower amounts in soil proximal to roots of cultivars than non‐cultivars. Additionally, soil NO₃–N was negatively correlated with root volume in S. nutans cultivars. While cultivars had greater BNPP than non‐cultivars, this was not reflected aboveground root structure, as ANPP was similar between cultivars and non‐cultivars. Intraspecific variation in belowground root structure and function exists between cultivar and non‐cultivar sources of the dominant prairie grasses during initial reestablishment of tallgrass prairie. Population source selection should be considered in setting restoration goals and objectives.     

Suppression of an invasive legume by a native grass — High impact of priority effects

Nitrogen-limited ecosystems are threatened by extensive spread of broom (Cytisus scoparius (L.) Link), a European leguminous shrub that is invasive in several countries. The establishment of invading species may, however, be suppressed by competition from native vegetation. The neighbor impact of the grass Festuca rubra subsp. commutata Gaudin on the performance of C. scoparius was studied in a greenhouse experiment with different arrival order, under low and high nitrogen supply, and with or without inoculation of nitrogen-fixing bacteria. Aboveground biomass of both species was measured after a six-months establishment period, and after a five-months regrowth period. In both periods, presence of F. rubra reduced the performance of C. scoparius as indicated by negative neighbor-effect intensity indices (NInt_A).During the establishment period the competitive impact of F. rubra was highest, when planted before C. scoparius, followed by synchronous and late planting. Inoculation with rhizobia and low fertilization decreased the competitive impact of F. rubra. After cutting and regrowth priority effects of F. rubra were still visible. Interaction between the two study species was not affected anymore by inoculation, but strongly by fertilization, with highest competitive impact of F. rubra on C. scoparius under high nitrogen fertilization. In both study periods biomass of C. scoparius was negatively correlated with biomass of F. rubra. Our study provides knowledge about competition processes, which help to improve conservation and restoration measures regarding the spread of C. scoparius. Early sowing of a native grass can help to suppress the invasive species at an early stage. Competitive impact of the grass might be strengthened by high nitrogen availability.     

Plant species and nutritional-mediated control over rhizodeposition and root decomposition

This study focuses on the influence of nitrogen (N) availability and species on rhizodeposition and on decomposition of rhizodeposits, roots and soil organic matter. Four perennial grass species were studied that are characteristic of grassland habitats that differ in nutrient availability. These perennial grass species, Holcus lanatus L., Festuca rubra L., Anthoxanthum odoratum L. and Festuca ovina L., were homogeneously labeled with ¹⁴CO₂. Plants were grown on soil without N addition and with N addition (14 g N m^–2). After 8 weeks, plants were harvested and root production and the remaining amount of rhizodeposits in the soil were measured. ¹⁴C-labelled roots were incubated in fresh soil. Decomposition was measured of 1) the labeled rhizodeposits in the soil in which the plants had been growing and 2) the labeled dead roots incubated in fresh soil, by trapping the evolved ¹⁴CO₂, over 69 days.In general, decomposability of both roots and rhizodeposits increased when nitrogen availability increased. Moreover, the species differed in their response to N. Higher N supply increased total rhizodeposition of H. lanatus and the decomposability of rhizodeposited carbon compounds of this high fertility species was greater than of the low fertility species F. ovina, but lower than of A. odoratum. The presented study gives no evidence for a relation between root decomposition rate and the nutrient availability of the habitat of the four species. Overall, we suggest on the basis of the results that species can affect nutrient cycling by differences in rates of rhizodeposition and litter production. This offers a mechanism whereby species can influence species replacement during succession.     

Testing the cultivar vigor hypothesis: comparisons of the competitive ability of wild and cultivated populations of Pascopyrum smithii along a restoration chronosequence

Restoration practitioners often rely on seeds of widely available cultivars representing native species but nonlocal germplasm. Cultivation improves the supply of plant materials and minimizes revegetation costs, but can also favor agronomic traits, and resulting vigor may affect the competitive ability and long‐term persistence of cultivated genotypes at restoration sites. We compared cultivated, restored, and wild populations of Pascopyrum smithii (western wheatgrass) in a greenhouse study to test the extent to which cultivars outcompete local plants in biomass production, and to determine if morphological differences (including height and number of leaves) among cultivated and wild populations persist at restoration sites over time. We found evidence of vigor and greater competitive ability of cultivars in seed mass, growth rate, plant height, and biomass and this advantage occurred when plants were grown alone or in competition with other seed sources. Cultivar vigor persisted at restoration sites over 30 years, but restored populations more closely resembled wild, local populations when cultivars were planted in closer proximity to nearby undisturbed sites. This study supports the cultivar vigor hypothesis and provides evidence for the long‐term persistence of cultivated traits in the environment.     

Variation in plant performance in a grassland: Species‐specific and neighbouring root mass effects

Question: What characteristics of local biotic neighbourhood is the best proxy of competitive effects experienced by plants in a herbaceous community: (1) total above‐ground biomass, (2) root mass or (3) relative above‐ground abundance of selected species? Location: Grassland at ca. 1100 m a.s.l. in the Krkono?e Mts., northern Czech Republic. Methods: We implanted two phytometer species, Antho‐xanthum alpinum and Festuca rubra, into a mountain grassland, and examined their response to local variation in (1) total above‐ground biomass, (2) root mass at three soil depths, and (3) relative abundance of individual species above‐ground. Results : Performance of both phytometer species was determined much more consistently by the mass of neighbouring roots and by species composition of neighbours than by the total above‐ground biomass. The two phytometer species showed different responses to these parameters. The most important relationships were (1) negative relationship between performance of Anthoxanthum and mass of neighbouring roots at 0–3 cm, (2) positive relationship between performance of Festuca and mass of neighbouring roots at 3–6 cm, and (3) negative relationship between performance of Festuca and relative abundance of Festuca in the neighbourhood. Conclusions: Neighbouring root mass and above‐ground species composition are better determinants of biotic interactions than total above‐ground biomass of neighbours in the studied mountain grassland. However, the relationships found are not necessarily due to variation in competitive intensity but can be due to other hidden factors as well, e.g. local availability of resources.     

Components of ‘competitive ability’ in the LHS model: Implication on coexistence for twelve co-occurring Mediterranean grasses

The leaf–height–seed (LHS) model has been proposed as a simple trait-based functional classification. We investigated whether the two components of competitive ability, i.e. competitive response (CR) and competitive effect (CE), are captured by the LHS model and whether these two components are independent for 12 coexisting Mediterranean grasses. Two greenhouse experiments were conducted to estimate competitive effect and response of 12 coexisting grass species from Mediterranean habitats in Jordan. We applied a phytometer design and calculated CR and CE using the relative interaction index (RII). Mature plant height, seed mass and leaf dry matter content (LDMC, used as the leaf trait) were measured for each species. Correlations and trade-offs between the three traits and the components of competitive ability, CR and CE, were analyzed with principal components analysis (PCA). The LHS model was a good predictor of competitive ability but CR and CE were independent and related to different traits. CR was positively correlated with seed mass and CE with plant height. LDMC was neither correlated to CR nor to CE. Based on these findings, we suggest that there are three primary strategies allowing coexistence in Mediterranean communities, which are related to competition: (1) large CE, i.e. large negative impact on other species associated with large stature, (2) large CR, i.e. resistance to competition associated with large seeds, and (3) competition avoidance associated with small seeds.     

Genetic Structure of Remnant Populations and Cultivars of Switchgrass (Panicum virgatum) in the Context of Prairie Conservation and Restoration

Switchgrass (Panicum virgatum L.) is a dominant, perennial C₄ grass of North American tallgrass prairies with cultivars that are widely used in grassland restoration, pastures, and landscaping. However, these cultivars may be genetically dissimilar to small, remnant populations, raising concerns about altered genetic composition of native populations through gene flow. To address this issue on a local scale in Ohio and Illinois, we used microsatellite markers to characterize genetic diversity and differentiation of 10 remnant prairie populations (5 in each state) and 8 common cultivars. The bulk of genetic variation was found to reside within rather than among wild populations, consistent with the outcrossing breeding system of switchgrass. Genetic diversity was similar among the remnant populations despite large differences in area (approximately 2–2,590 ha), highlighting the importance of small native populations as reservoirs of variation and potential seed sources for prairie restoration. Cultivars generally had similar levels of variation to the wild populations, but we found clear genetic dissimilarity between wild and cultivated gene pools (especially for Kanlow, but also Trailblazer, Blackwell, Dacotah, Summer, and Sunburst cultivars). This suggests that using cultivars in local prairie restoration efforts may alter the genetic composition of wild populations. Whether such changes are deemed as negative depends on the cultivar under consideration and specific conservation goals for preserving native switchgrass populations. Patterns of genetic variation in remnant prairie populations and potential cultivar sources can be used to develop guidelines for restoration as well as future planting of cultivars for biofuels.     

Population source affects competitive response and effect in a C4 grass (Panicum virgatum)

Grassland restoration success depends on the development of plant communities that accord with restoration goals. Intraspecific variation in competitiveness may affect community development. For some grassland species, germplasm can be obtained from sources ranging from wild collections to selectively bred cultivars. The extent to which population source affects competitive outcomes in restoration projects is unclear. We addressed this knowledge gap in a glasshouse experiment comparing competitive response and effect among three sources of switchgrass (Panicum virgatum) that are available for restoration: selectively bred cultivars, commercial ecotypes (commercially produced but not deliberately selected), and wild collections. Two strains per source type were grown with four associates chosen to encompass varied functional groups: conspecifics, Bromus inermis, Cirsium arvense, and Solanum ptycanthum. Switchgrass competitive response was evaluated for survival, height, biomass, and shoot:root biomass ratio; competitive effect was assessed as associate survival, height, biomass, and shoot:root ratio. Competitive responses of cultivars and commercial ecotypes were broadly similar, although cultivar biomass exceeded both that of ecotypes and wild collections, and ecotypes had the highest shoot:root ratio. Wild collections were most negatively affected by competition. The shoot:root ratios of all sources were highest when grown with S. ptycanthum, indicating that competitive responses were plastic; plasticity in fitness‐related traits can contribute to persistence in variable environments. Cultivars exerted negative effects on B. inermis. Secondary analyses indicated that all switchgrass sources were most inhibited by the annual S. ptycanthum. To summarize, population source affected multiple aspects of switchgrass competitive ability, when grown against functionally varied associates.     

Consequences of Seed Origin and Biological Invasion for Early Establishment in Restoration of a North American Grass Species

Local, wild-collected seeds of native plants are recommended for use in ecological restoration to maintain patterns of adaptive variation. However, some environments are so drastically altered by exotic, invasive weeds that original environmental conditions may no longer exist. Under these circumstances, cultivated varieties selected for improved germination and vigor may have a competitive advantage at highly disturbed sites. This study investigated differences in early establishment and seedling performance between wild and cultivated seed sources of the native grass, Poa secunda, both with and without competition from the invasive exotic grass, Bromus tectorum. We measured seedling survival and above-ground biomass at two experimental sites in western Montana, and found that the source of seeds selected for restoration can influence establishment at the restoration site. Cultivars had an overall advantage when compared with local genotypes, supporting evidence of greater vigor among cultivated varieties of native species. This advantage, however, declined rapidly in the presence of B. tectorum and most accessions were not significantly different for growth and survival in competition plots. Only one cultivar had a consistent advantage despite a strong decline in its performance when competing with invasive plants. As a result, cultivated varieties did not meet expectations for greater establishment and persistence relative to local genotypes in the presence of invasive, exotic species. We recommend the use of representative local or regional wild seed sources in restoration to minimize commercial selection, and a mix of individual accessions (wild, or cultivated when necessary) in highly invaded settings to capture vigorous genotypes and increase the odds native plants will establish at restoration sites.     

Effects of competition with Bromus tectorum on early establishment of Poa secunda accessions: can seed source impact restoration success?

When landscapes are heavily impacted by biological invasion, local populations of native plant species may no longer be adapted to altered environmental conditions. In these cases, it is useful to investigate alternative sources of germplasm, such as cultivated varieties, for planting at restoration sites. This study compared cultivated and wild (local) varieties of the native perennial bunchgrass, Poa secunda J. Presl, grown with and without the exotic, invasive Bromus tectorum L. in a greenhouse setting. While P. secunda cultivars emerged and grew more rapidly than wild seed sources, this advantage declined in the presence of B. tectorum and cultivated germplasm did not outperform wild accessions in the presence of an invasive species. Given the novel genetic background of cultivars and their potential to alter patterns of dominance in native plant communities, we recommend the use of local or regional wild seed sources when possible to conserve regional patterns of genetic diversity and adaptation. Use of multiple seed sources may increase the potential for capturing vigorous genotypes in the restoration seed mix. In cases where sites are heavily impacted by exotic, invasive species, other control measures will be necessary to improve establishment of native species in grassland restoration programs.     

Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.     

Infection with the fungal endophyte Epichloë festucae may alter the allelopathic potential of red fescue

Red fescue (Festuca rubra) is a perennial grass used as both forage and turfgrass. Asymptomatic plants of this species are systemically infected by the fungal endophyte Epichloë festucae, which has a beneficial effect on the infected plants. The aim of this study was to determine the effect of the endophyte Epichloë festucae on the allelopathic potential of F. rubra against four associated pasture species that are also considered as weeds in lawns, Trifolium pratense, Trifolium repens, Lotus corniculatus and Plantago lanceolata. Two experiments were designed to evaluate the allelopathic effect of extracts from the roots and leaves of endophyte‐infected (E+) and non‐infected (E?) plants on the germination and seedling growth of the four target species. Regardless of the endophyte status of the host plant, leaf extracts elicited a stronger reduction in germination and seedling growth than root extracts. Extracts from E+ plants reduced the speed of germination index of Trifolium spp. to a greater extent than those from E? plants. Radicle length of the target species was the parameter most affected by the presence of the endophyte in F. rubra. Root extracts from E+ plants had a greater inhibitory effect on the radicle growth of the target species than did root extracts from E? plants. A greater concentration in total phenolic compounds was found in the roots of E+ plants than of E?; however, this difference was not observed in the leaves. Thus, the allelopathic potential of F. rubra is altered in infected plants.     

Do plant parasitic nematodes have differential effects on the productivity of a fast- and a slow-growing grass species?

We have examined the interaction between plant parasitic nematodes and plant species from different stages of grassland succession. In these grasslands, fertiliser application was stopped in order to restore the former nutrient-poor ecosystems. This management resulted in a reversed succession of high- to low-productivity. Nematodes isolated from a high-productive early-successional field and a low-productive late-successional field were inoculated to sterilised soil planted with seedlings of either Lolium perenne (a fast-growing early-successional species) or Festuca rubra (a slow-growing late-successional species). The experiment was performed at low and high supply rates of nutrients. We hypothesised that at a low nutrient supply rate the growth of L. perenne will be more reduced by nematode herbivory than the growth of F. rubra. Furthermore, we hypothesised that higher numbers of plant parasitic nematodes will develop under L. perenne. We found no support for our first hypothesis, because nematodes did not affect plant growth. Our results suggest that changes in the nutrient availability rather than plant parasitic nematodes affect plant succession in impoverished grasslands. On the other hand, plant species and nutrient supply rate significantly affected the density and composition of the plant parasitic nematode community. In line with our second hypothesis, plant parasitic nematodes reproduced better on the fast-growing L. perenne than on the slow-growing F. rubra. Our results, therefore, suggest that the succession of the plant parasitic nematode community is probably more affected by changes in the plant community than the other way round.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

POLLINATION BIOLOGY,SEED PRODUCTION,AND POPULATION STRUCTURE IN QUEEN-OF-THE-PRAIRIE,FILIPENDULA RUBRA (ROSACEAE) AT BOTKIN FEN,MISSOURI

Queen-of-the-prairie, Filipendula rubra (Rosaceae), is a perennial, rhizomatous herb restricted to calcareous fens, spring seeps, and wet prairies, and is distributed in the northcentral United States from Iowa to New York. Because of the scarcity of adequate habitat, F. rubra is a rare and endangered species in several states. At Botkin Fen and other fens in Missouri, F. rubra possesses limited sexual reproduction. A study was conducted to determine causal mechanisms of its limited seed production that may be barriers to establishing this rare and endangered species in other fens in Missouri. A genetic-ecological hypothesis was supported. F. rubra was found to be self-incompatible. At Botkin Fen, most inflorescences produced very few seeds because most (85.4%) belonged to a single clone. Pollinators (Lassioglossum sp.) had a high probability of transporting pollen with the same self-incompatibility alleles. A few genetically different clones did produce significant quantities of seed. However, this seed, with its significant pool of genetic variability, was not recruited into the population. Apparently, competition from the dense sedges and grasses prevented the establishment of new seedlings.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.