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.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

Ecosystem Services from Keystone Species: Diversionary Seeding and Seed‐Caching Desert Rodents Can Enhance Indian Ricegrass Seedling Establishment

Seeds of Indian ricegrass (Achnatherum hymenoides), a native bunchgrass common to sandy soils on arid western rangelands, are naturally dispersed by seed‐caching rodent species, particularly Dipodomys spp. (kangaroo rats). These animals cache large quantities of seeds when mature seeds are available on or beneath plants and recover most of their caches for consumption during the remainder of the year. Unrecovered seeds in caches account for the vast majority of Indian ricegrass seedling recruitment. We applied three different densities of white millet (Panicum miliaceum) seeds as “diversionary foods” to plots at three Great Basin study sites in an attempt to reduce rodents' over‐winter cache recovery so that more Indian ricegrass seeds would remain in soil seedbanks and potentially establish new seedlings. One year after diversionary seed application, a moderate level of Indian ricegrass seedling recruitment occurred at two of our study sites in western Nevada, although there was no recruitment at the third site in eastern California. At both Nevada sites, the number of Indian ricegrass seedlings sampled along transects was significantly greater on all plots treated with diversionary seeds than on non‐seeded control plots. However, the density of diversionary seeds applied to plots had a marginally non‐significant effect on seedling recruitment, and it was not correlated with recruitment patterns among plots. Results suggest that application of a diversionary seed type that is preferred by seed‐caching rodents provides a promising passive restoration strategy for target plant species that are dispersed by these rodents.     

Influence of litter and weather on seedling recruitment in a mixed oak-pine woodland

The effects of regular litter removal and annual variation in temperature and precipitation on seedling recruitment of species differing in their seed size and mode of dispersal were studied in a 16-year (1984-1999) experiment in a mixed oak-pine wood in southern Poland. Litter was the most important factor in determining spatial variability in seedling recruitment, whereas differences in climatic conditions among years, especially temperature fluctuations in late winter and early spring, determined the temporal variability in seedling recruitment. Compared with control plots, significantly more new individuals of bryophytes and seedlings as well as a number of new species of vascular plants were noted in the litter-removal plots over the 16-year study. Litter strongly impeded seedling emergence of small-seeded species. The negative effect of litter on seedling recruitment of large-seeded species and the recruitment of new shoots in species growing clonally was much weaker. There was a significant positive correlation between the numbers of seedlings in the litter-removal and control plots and temperatures in January to March. In the litter-removal plots this mainly affected small-seeded species. Seedling recruitment was less consistently related to variation in precipitation. Positive relationships were found only between the number of seedlings of large-seeded species in the litter-removal plots and precipitation in July of the current year and in September of the previous year, and between the number of seedlings in the control plots and precipitation in September and November of the previous year.     

Widespread seed limitation affects plant density but not population trajectory in the invasive plant Centaurea solstitialis

In some plant populations, the availability of seeds strongly regulates recruitment. However, a scarcity of germination microsites, granivory or density-dependent mortality can reduce the number of plants that germinate or survive to flower. The relative strengths of these controls are unknown for most plant populations and for exotic invaders in particular. We conducted a seed addition experiment with a granivore exclusion treatment in a field setting to explore how these factors interact to regulate populations of the widespread invader Centaurea solstitialis (yellow starthistle) at three study sites across the plant’s range in California. We coupled the experimental approach with observational studies within established C. solstitialis populations to estimate seed rain, recruitment and mortality at natural densities. Seed limitation occurred in both experimental and observational plots in all populations. Although vertebrate granivores were active at each site, they had no effect on C. solstitialis recruitment. Density increased mortality, but the effect was variable and weak relative to its effect on fecundity. The seed limitation that was evident at the seedling stage persisted to flowering. Seed-limited populations such as these ought to be highly sensitive to losses to seed predators, and many biological control agents, including those established for C. solstitialis, are seed predators. However, flowering plant density was decoupled from seed production by a strong compensatory response in the surviving plants; seed production was nearly constant in plots across all seed addition levels. Thus, flowering plant density is reduced by the established biocontrol agents, but seed production compensates to replace the population every generation, and no long-term decline is predicted.     

Population ecology of wild sunflowers: effects of seed density and post-dispersal vertebrate seed predators

Assessing the effects of seed density on the population dynamics of wild plant species with crop relatives will be vital in determining the potential effects of introducing traits into wild populations as a result of crop-to-wild gene flow. We examined experimental sunflower (Helianthus annuus) patches in eastern Kansas to determine the effects of seed density and predation on seedling recruitment and seed production in the next generation. High seed density treatment plots had significantly more seedlings and adult plants than did low seed density treatment plots. Overwinter vertebrate seed predator exclusion treatments resulted in increases in plant density compared to plots in which vertebrates were not excluded. Control patches (no seeds added) contained virtually no plants. Head production and estimated total seed production for a patch were not statistically different among treatments (excluding control plots). Although initial seed density and vertebrate post-dispersal seed predation do appear to have effects on seedling recruitment, neither appear to be limiting seed production of competing adult plants. Therefore, variation in seed densities (over the range examined) may have limited effects on local population dynamics. It is important to note that the choice of seed densities may affect the results obtained: the seed densities used in this study may, in retrospect, be higher than in the small roadside populations typical in eastern Kansas, yet other natural sites have much larger densities. Further, the effects of increased seed density at a local site may have other important effects such as altering metapopulation dynamics through increased long-distance dispersal or increased local seed bank size.     

Seed predation limits post-fire recruitment in the waratah (<Emphasis Type="Italic">Telopea speciosissima</Emphasis>)

Seed predation may reduce recruitment in populations that are limited by the availability of seeds rather than microsites. Fires increase the availability of both seeds and microsites, but in plants that lack a soil- or canopy-stored seed bank, post-fire recruitment is often delayed compared to the majority of species. Pyrogenic flowering species, such as Telopea speciosissima, release their non-dormant seeds more than 1 year after fire, by which time seed predation and the availability of microsites may differ from that experienced by plants recruiting soon after fire. I assessed the role of post-dispersal seed predation in limiting seedling establishment after fire in T. speciosissima, in southeastern Australia. Using a seed-planting experiment, I manipulated vertebrate access to seeds and the combined cover of litter and vegetation within experimental microsites in the 2 years of natural seed fall after a fire. Losses to vertebrate and invertebrate seed predators were rapid and substantial, with 50% of seeds consumed after 2 months in exposed locations and after 5 months when vertebrates were excluded. After 7 months, only 6% of seeds or seedlings survived, even where vertebrates were excluded. Removing litter and vegetation increased the likelihood of seed predation by vertebrates, but had little influence on losses due to invertebrates. Microsites with high-density vegetation and litter cover were more likely to have seed survival or germination than microsites with low-density cover. Recruitment in pyrogenic flowering species may depend upon the release of seeds into locations where dense cover may allow them to escape from vertebrate predators. Even here, conditions suitable for germination must occur soon after seed release for seeds to escape from invertebrate predators. Seed production will also affect recruitment after any one fire, while the ability of some juvenile and most adult plants to resprout after fire buffers populations against rapid declines when there is little successful recruitment.     

Burning Influences on Wiregrass (Aristida beyrichiana) Restoration Plantings: Natural Seedling Recruitment and Survival

Regeneration and expansion of Aristida beyrichiana and Aristida stricta (wiregrass) populations in remaining fire‐maintained Pinus palustris (longleaf pine) stands of the southeastern United States has become an objective of land managers. Although growing‐season fire is required for successful wiregrass seed production, studies examining naturally occurring wiregrass seedling dynamics are few. This study investigates how seedling survivorship is affected by season of burn, seedling size, time since germination, and proximity to adult plants. Restoration at this research site was begun in 1992 with the planting of containerized longleaf pine and wiregrass seedlings. Study plots were established in November 1997 after a growing‐season prescribed fire (June 1996) that resulted in successful seed production and seedling recruitment. Burn treatment plots included (1) no burn (control), (2) fire in the dormant season of the first year after germination (March 1998), (3) fire in the growing season of the first year after germination (August 1998), and (4) fire in the growing season of the second year after germination (July 1999). Seedling mortality increased with growing season burning and close proximity to planted adults. Natural seedling recruitment continued into the second year after initial seed‐drop in all plots, which verifies that wiregrass seed banking occurs for a minimum of 2 years after seed drop. Where wiregrass management objectives include population expansion, seedling recruits should be allowed 1 to 2 years post‐germination without growing season fire for successful establishment.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

Seed and microsite limitations of recruitment and the impacts of post-dispersal seed predation at the within population level

Seed predation may cause important seed losses in plant populations, but its impact on the dynamics of populations will depend on the degree of seed or microsite limitations for recruitment. Seed losses will only affect recruitment if it is seed limited. The nature of recruitment limitation (seeds or microsites) is usually ascribed to whole plant populations but it may vary within populations among microhabitats and habitats. Thus, the potential impact of seed predation will also vary within the population, being highest where recruitment is seed limited. The impact to the whole population will depend on the spatial concordance between the intensity of seed predation and that of seed limitation. Recruitment limitations (with seed addition experiments), seed predation (with seed removal experiments), and the dynamics of seed availability in the soil (with soil samples taken both after seed dispersal and before the following dispersal event) of the shrub Corema album (Empetraceae) were investigated in dunes in NW Spain, at microhabitats ‘open ground’, ‘underneath C. album ♀’, and ‘underneath C. album ♂’ at two habitats, sparse and dense scrub. The nature of recruitment limitation (seeds vs. microsites) varied within the population. It was seed limited in the microhabitat ‘open ground’ and microsite limited under shrub cover. The spatial patterns of seedling recruitment were unrelated to seed availability but strongly affected by germination requirements. The spatial discordance between seed availability and recruitment implies a crucial constraint for processes affecting seed availability (seed predation but also e.g., dispersal) to impact recruitment. They will not affect its spatial pattern but only its quantity as long as they act in those sites selected by seeds to germinate. Seed predation was highest underneath mother plants and lowest in open ground. Thus, its potential impact is low, as it is centred where recruitment is not seed limited. This study shows that the analysis of seed predation in relation to recruitment limitations at smaller spatial scales within the population provides more insight to understand its impact.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

Plant recruitment bottlenecks in temperate forest fragments: seed limitation and insect herbivory

Numerous studies have documented declines in plant diversity in response to habitat loss in fragmented landscapes. However, determining the mechanisms that lead to species loss is challenging using solely a correlative approach. Here we link correlative assessments of plant community composition with seed additions for a focal species to test the hypothesis that distributions of forests plants within a fragmented landscape are limited by seed dispersal. Woody plant species richness of fragments declined as fragments (n=26) became more isolated by agricultural fields. We predicted that if these isolation effects were driven by poor dispersal rather than other effects associated with habitat loss, then plants should vary in their response to isolation in relation to their seed size (i.e., stronger effects for plants with larger seeds). As predicted under this dispersal limitation hypothesis, sensitivity of bird-dispersed shrubs to isolation was related to their seed mass, with species with heavy seeds (e.g., Lindera benzoin) exhibiting stronger declines in presence across isolation gradients than species with light seeds. Seed addition experiments were performed for Lindera benzoin in two high isolation forest fragments (nearest neighbor mean distance=803 m) where Lindera was naturally absent, and two low isolation fragments (nearest neighbor mean distance=218 m) with naturally occurring Lindera populations. Seed addition and control plots (n=50 1 m² plots per fragment) were monitored for 13 censuses over 3 years. Across all four fragments, seed additions resulted in significant increases in Lindera seedling recruitment with no differences in final seedling establishment among fragments. However, insect herbivory was higher on Lindera seedlings in high isolation compared to low isolation fragments and was negatively correlated with seedling survival over some years. Consistent with prior work, our results confirm that seed dispersal plays a significant role in affecting plant diversity in fragmented landscapes. However, results also suggest the need for a better understanding of how additional processes, such as herbivory, may be altered as habitat is lost and what effects such changes have for forest plants.     

Experimental Assessment of Factors Limiting Seedling Recruitment of an Amazonian Understory Herb

Understory herbs are an important and species-rich component of tropical forests, but little is known about factors limiting recruitment of these herbs. In a Peruvian flood plain forest, we studied seedling recruitment of the widespread clonal herb Heliconia metallica . We analyzed natural recruitment and experimentally added seeds, excluded predators and removed the understory vegetation at occupied and unoccupied sites to test whether seed limitation, predation, disturbance, and the presence of mature conspecific plants influence seedling recruitment. The number of naturally recruited seedlings was higher at flooded sites and close to flowering H. metallica ramets. Predation caused considerable seed loss shortly after sowing. Seed survival strongly increased with the openness of a site, but was not affected by the presence of mature conspecifics. Seed addition increased the number of seedlings at both occupied and unoccupied sites. Two years after sowing, the number of established seedlings and their size increased with light availability at a site. In experimentally disturbed plots, light availability, seedling establishment, and survival were higher. The effect of disturbance on seedling survival was stronger at occupied sites, probably due to strong light competition from mature H. metallica plants. Herbivore damage on leaves of seedlings was lower at occupied sites, but seedlings grew faster at unoccupied sites. We conclude that seedling recruitment of H. metallica depends on natural disturbance and is limited by both seed availability and the presence of mature conspecific plants.     

Influences of neighbouring plants on seedling establishment in limestone grassland

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

Variability of seedling recruitment under dominant, moss, and litter removal over four years

In a four-year removal experiment we investigated the effect of removing a dominant species (Nardus stricta), the litter layer and moss layer on species composition of established vegetation and on numbers and species composition of seedlings. The experiment was conducted in an oligotrophic wet meadow (Molinion with some features ofViolion caninae according to phytosociological classification). After four years of the experiment, the redundancy analysis (RDA) did not reveal any changes in the composition of the established plant community. Seedling numbers and their species composition, however, varied significantly both among treatments and over years. Whereas the differences between treatments were mainly in total seedling numbers, the years differed in both seedling numbers and relative representation of individual species. In particular,Myosotis nemorosa seedlings highly increased their abundance in a favourable year 1995 constituting one third of all the seedlings found, whereas in other years their abundance was between 4% and 10%. The effects of year and treatment were not additive (significant year × treatment interaction). This means that plots under different treatments are differently affected by climatic conditions of individual years. The greatest increase of seedling recruitment was observed in the plots with the moss layer removed. Microsites with high seedling recruitment persisted in the plots for no more than two years. Why the established vegetation composition does not differ among treatments even after four years, despite the relatively pronounced and immediate response of seedlings? The number of species with regular seedling recruitment is limited. They are mostly perennials, and consequently, it might take even longer than four years before recruitment limitation affects the population size of established plants.     

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.     

Alien seedling recruitment as a response to altitude and soil disturbance in the mountain grasslands of central Argentina

Climate and disturbance are considered key factors in explaining plant invasion, mainly by their effect on alien seedling recruitment. We tested whether soil disturbance by digging and altitude, as a sub-set of climate, affected the seedling recruitment of two alien species (Cirsium vulgare and Melilotus alba) in the mountain grasslands of central Argentina. We performed a seed-addition factorial experiment with altitude as a fixed factor co-variable (6 levels), time since sowing (6 levels) as an autoregressive co-variable, and soil disturbance by digging (two levels) and seed addition (two levels) as fixed factors. Seeds of the studied species were sown in replicated undisturbed and disturbed soil plots, in grassland stands located every 200 m from 1200 to 2200 m a.s.l. The number of emerging seedlings at each plot was recorded during a period of 8 months, at intervals of 30–60 days. Seedlings of both alien species emerged at all the studied altitudes, but only the recruitment of C. vulgare increased linearly with altitude. Moreover, the time window of seedling recruitment for this species increased with altitude. Soil disturbance produced a significant decrease in overall seedling emergence of both species; however, an increase in emergence was observed for C. vulgare at an altitude over 2000 m a.s.l. Natural emergence inside control plots without seed addition was only observed for Cirsium vulgare, which indicates that this species was not seed limited. Our results show that altitude and the absence of disturbance do not restrict the recruitment of C. vulgare and M. alba in these natural mountain grasslands. Moreover, these alien species seem to be tolerant to and even favoured by conditions occurring at higher altitudes and in absence of micro-site soil disturbances in the Córdoba mountain grasslands.     

Effects of microsite conditions on seedling establishment on the foreland of Coleman Glacier,Washington

Abstract. Questions: How do physical microsite conditions of microsites affect germination and seedling survival in different successional stages? Do different species germinate in similar microsites in a given successional stage? Location: Coleman Glacier foreland, Mount Baker, Washington State, USA. Methods: Two methods were used to characterize safe sites. 1. Grids of 300 10 cm × 10 cm plots were located in four different age classes on the foreland. 2.105 pairs of plots, with and without seedlings of Abies amabilis, were located in each age class. For each plot we identified all seedlings and all individuals < 1 m tall. Microsite characteristics such as topography and presence of rocks or woody debris were noted for each plot. Microsite characteristics were compared between plots with and without each species. In addition we examined the effect of distance from seed sources on the presence of Alnus viridis seeds and seedlings in a newly disturbed area. Results: In early successional sites, seedlings of several species were positively associated with depressions and presence of rocks, and negatively associated with ridges. Patterns were generally consistent among species. In later succession, seedlings were not significantly associated with any microsite characteristics. For Alnus viridis, seed density decreased with distance from seed sources but seedling density did not. Conclusions: Because of harsh conditions in early succession, physical microsites are important, and most species have similar microsite requirements. In later succession, physical microsites characteristics are not as important and are more variable. Microsites appear to be more important than seed rain in controlling the distribution of Alnus viridis in early succession.     

Effect of competition intensity on recruitment of palatable and unpalatable grasses

In this study, we made an attempt to reveal how competition intensity from established plants impacts on palatable and unpalatable grass seedlings recruitment, in a natural mesic grassland of central Argentina. Our objective was to assess the seedling recruitment of a palatable species (Chascolytrum subaristatum) and an unpalatable species (Nassella trichotoma) in microsites differing in competition intensity from established plants. Identity (C. subaristatum and N. trichotoma) and defoliation severity were used as surrogate for competition intensity. In March 2017, we permanently marked established individuals of N. trichotoma and C. subaristatum and placed two circular plots adjacent to each individual. In one plot we added seeds of N. trichotoma and in the other seeds of C. subaristatum. After seeding, established plants were randomly assigned to one of three level of defoliation: without defoliation, low defoliation severity and high defoliation severity. From April to November 2017 (i.e. over a complete annual growing cycle), we measured seedling density, recruitment and growth. Our results supported the hypothesis that seedlings of palatable grasses are more competitive than seedlings of unpalatable grasses. Seedling of the palatable grass C. subaristatum recruited successfully regardless the intensity of competition from established plants, whereas seedlings of the unpalatable grass N. trichotoma recruited better under low competitive pressure from established plants. Our results suggest that the availability of microsites with low competitive pressure from the established vegetation, created by selective grazing of palatable grasses, promotes the recruitment of unpalatable grass seedlings. This mechanism may contribute to the species replacement process commonly observed in heavy grazed grasslands.     

Factors limiting rosette recruitment in scarlet gilia, Ipomopsis aggregata: seed and disturbance limitation

We performed a factorial combination of seed additions, surface soil disturbance, and protection from potential seed predation in experimental plots to elucidate the relative importance of each factor in the recruitment of scarlet gilia seedlings. We detected a significant interaction between seed addition and surface soil disturbance on seedling recruitment over 2 years of establishment. Plots that received both supplemental seed addition and disturbance established 10.5 times as many rosettes as control plots. We did not detect main or interactive effects of caging on seedling establishment. We explored the importance of density-dependent recruitment by investigating the number of rosettes per experimental plot through time. In addition, we used planned contrasts to compare the absolute and percent changes in control and treatment plot rosette densities between the study years. We found significant downward concave curvature to the seedling recruitment curve, suggesting negative density-dependent recruitment across the experimental range of rosette densities. We found a significant difference between the control plots and the seed addition-disturbance plots in terms of absolute changes in plot density from 1997 to 1998. Greater net mortality occurred in plots that received both seed addition and disturbance. However, we found no differences among treatments in the percent change in rosette density across the study years. Our study demonstrates that seedling recruitment in natural plant populations may be limited by the interaction of seed and microsite availability, and that seedling recruitment in scarlet gilia may be negatively density dependent. Our results suggest that the fecundity effects of particular plant-animal interactions (e.g., pollination, herbivory) may affect scarlet gilia population dynamics, particularly under conditions of high soil disturbance. Received: 11 June 1999 / Accepted: 24 November 1999