Seed Addition Experiments Research Articles

Seed density significantly affects species richness and composition in experimental plant communities.

Studies on the importance of seed arrival for community richness and composition have not considered the number of seeds arriving and its effect on species richness and composition of natural communities is thus unknown. A series of experimental dry grassland communities were established. All communities were composed of the same 44 species in exactly the same proportions on two substrates using three different seed densities.The results showed that seed density had an effect on species richness only at the beginning of the experiment. In contrast, the effects on species composition persisted across the entire study period. The results do not support the prediction that due to higher competition for light in nutrient-rich soil, species richness will be the highest in the treatment with the lowest seed density. However, the prevalence of small plants in the lowest seed density supported the expectation that low seed density guarantees low competition under high soil nutrients. In the nutrient-poor soil, species richness was the highest at the medium seed density, indicating that species richness reflects the balance between competition and limitations caused by the availability of propagules or their ability to establish themselves. This medium seed density treatment also contained the smallest plants.The results demonstrate that future seed addition experiments need to consider the amount of seed added so that it reflects the amount of seed that is naturally found in the field. Differences in seed density, mimicking different intensity of the seed rain may also explain differences in the composition of natural communities that cannot be attributed to habitat conditions. The results also have important implications for studies regarding the consequences of habitat fragmentation suggesting that increasing fragmentation may change species compositions not only due to different dispersal abilities but also due to differential response of plants to overall seed density.

Urbanized landscapes favored by fig‐eating birds increase invasive but not native juvenile strangler fig abundance

Propagule pressure can determine the success or failure of invasive plant range expansion. Range expansion takes place at large spatial scales, often encompassing many types of land cover, yet the effect of landscape context on propagule pressure remains largely unknown. Many studies have reported a positive correlation between invasive plant abundance and human land use; increased propagule pressure in these landscapes may be responsible for this correlation. We tested the hypothesis that increased rates of seed dispersal by fig-eating birds, which are more common in urban habitats, result in an increase in invasive strangler fig abundance in landscapes dominated by human land use. We quantified abundance of an invasive species (Ficus microcarpa) and a native species (F. aurea) of strangler fig in plots spanning the entire range of human land use in South Florida, USA, from urban parking lots to native forest. We then compared models that predicted juvenile fig abundance based on distance to adult fig seed sources and fig-eating bird habitat quality with models that lacked one or both of these terms. The best model for juvenile invasive fig abundance included both distance to adult and fig-eating bird habitat terms, suggesting that landscape effects on invasive fig abundance are mediated by seed-dispersing birds. In contrast, the best model for juvenile native fig abundance included only presence/absence of adults, suggesting that distance from individual adult trees may have less effect on seed limitation for a native species compared to an invasive species undergoing range expansion. However, models for both species included significant effects of adult seed sources, implying that juvenile abundance is limited by seed arrival. This result was corroborated by a seed addition experiment that indicated that both native and invasive strangler figs were strongly seed limited. Understanding how landscape context affects the mechanisms of plant invasion may lead to better management techniques. Our results suggest that prioritizing removal of adult trees in sites with high fig-eating bird habitat may be the most effective method to control F. microcarpa abundance.

Vertebrate herbivory impacts seedling recruitment more than niche partitioning or density-dependent mortality

In tropical forests, resource-based niches and density-dependent mortality are mutually compatible mechanisms that can act simultaneously to limit seedling populations. Differences in the strengths of these mechanisms will determine their roles in maintaining species coexistence. In the first assessment of these mechanisms in a Congo Basin forest, we quantified their relative strengths and tested the extent to which density-dependent mortality is driven by the distance-dependent behavior of seed and seedling predators predicted by the Janzen-Connell hypothesis. We conducted a large-scale seed addition experiment for five randomly selected tropical tree species, caging a subset of seed addition quadrats against vertebrate predators. We then developed models to assess the mechanisms that determine seedling emergence (three months after seed addition) and survival (two years after seed addition). As predicted, both niche differentiation and density-dependent mortality limited seedling recruitment, but predation had the strongest effects on seedling emergence and survival. Seedling species responded differently to naturally occurring environmental variation among sites, including variation in light levels and soil characteristics, supporting predictions of niche-based theories of tropical tree species coexistence. The addition of higher densities of seeds into quadrats initially led to greater seedling emergence, but survival to two years decreased with seed density. Seed and seedling predation reduced recruitment below levels maintained by density-dependent mortality, an indication that predators largely determine the population size of tree seedlings. Seedling recruitment was unrelated to the distance to or density of conspecific adult trees, suggesting that recruitment patterns are generated by generalist vertebrate herbivores rather than the specialized predators predicted by the Janzen-Connell hypothesis. If the role of seed and seedling predation in limiting seedling recruitment is a general phenomenon, then the relative abundances of tree species might largely depend on species-specific adaptations to avoid, survive, and recover from damage induced by vertebrate herbivores. Likewise, population declines of herbivorous vertebrate species (many of which are large and hunted) may trigger shifts in species composition of tropical forests.

Plant Fitness Assessment for Wild Relatives of Insect Resistant Bt-Crops

When field tests of transgenic plants are precluded by practical containment concerns, manipulative experiments can detect potential consequences of crop-wild gene flow. Using topical sprays of bacterial Bacillus thuringiensis larvicide (Bt) and larval additions, we measured fitness effects of reduced herbivory on Brassica rapa (wild mustard) and Raphanus sativus (wild radish). These species represent different life histories among the potential recipients of Bt transgenes from Bt cole crops in the US and Asia, for which rare spontaneous crosses are expected under high exposure. Protected wild radish and wild mustard seedlings had approximately half the herbivore damage of exposed plants and 55% lower seedling mortality, resulting in 27% greater reproductive success, 14-day longer life-spans, and 118% more seeds, on average. Seed addition experiments in microcosms and in situ indicated that wild radish was more likely to spread than wild mustard in coastal grasslands.

Regeneration of native vascular plants facilitated by Ischaemum aristatum var. glaucum tussocks: an experimental demonstration

AbstractFacilitation of a perennial tussock grass, Ischaemum aristatum var. glaucum, was hypothesized to be the provision of germination safe sites against inundation. We conducted field census and seed addition experiments to test this hypothesis. Facilitation effects of I. aristatum var. glaucum tussocks were obvious; in contrast to 280 seedlings/m2 of ten native species observed on tussocks, seedlings hardly emerged on bare ground, even after seed addition. Although effects of moss occurrence at tussocks were not significant on the total number or species richness of emerged seedlings, significantly positive effects were observed on the seedling survival of some species, including endangered species. Conservation of facilitators will efficiently ensure the regeneration success of native vascular plants.

Invasive competitor and native seed predators contribute to rarity of the narrow endemicAstragalus sinuatusPiper

The conservation of rare plant species hinges on our ability to identify the underlying mechanisms that limit rare plant populations. Theory on rarity suggests that both predispersal seed predation and competition can be important mechanisms influencing abundance and/or distribution of rare plant populations. Yet few studies have tested these interactions, and those that have evaluated each mechanism independently. Astragalus sinuatus Piper (Whited's milkvetch) is a narrow endemic plant species restricted to eight populations within a 10-km2 area in eastern Washington. We used experimental and observational methods to test the effects of native insect predispersal seed predators and an invasive grass (Bromus tectorum L. [cheatgrass]) on seed set and population density of A. sinuatus. We quantified per capita seed production and pod predation rates across four sites and among four years. Seed predation rates were high across four sites (66-82%) and all years (65-82%). Experimental reduction of predispersal seed predators significantly increased per capita seed set of A. sinuatus (164-345%) at two experimental sites. Concurrently, two seed addition experiments demonstrated the effect of seed loss and presence of B. tectorum on seedling recruitment and establishment of A. sinuatus over four growing seasons. In the first seed addition experiment, we found no difference in recruitment and establishment between low (40) and high (120) seed addition levels. In the second addition experiment (one level of addition; 40 seeds), we found that recruitment and survivorship increased 200% in plots where B. tectorum was removed compared to plots where B. tectorum was present. Thus, seed addition had no impact in the presence of B. tectorum; conversely, in the absence of B. tectorum, seed addition was highly effective at increasing population numbers. Results suggest that, in areas where B. tectorum is present, recruitment is site limited, and it is seed limited when B. tectorum is absent. We recommend that managers reduce B. tectorum in an effort to increase population growth of A. sinuatus; in areas where B. tectorum is absent, short-term reduction of insect predators should be considered as a strategy to increase population growth of this rare species.

Seedling germination success and survival of the invasive shrub Scotch broom ( Cytisus scoparius) in response to fire and experimental clipping in the montane grasslands of the Nilgiris, south India

The spread of the exotic shrub Scotch broom in the montane grasslands of the Nilgiris is one of the major threats to biodiversity and ecosystem function there. It is likely that fire suppression over the past few decades is the proximate cause of expansion of broom populations. This study capitalizes on a wildfire event to examine fire effects on mature broom populations and soil seedbanks. Fire resulted in widespread death of mature broom stands but also stimulated broom soil seedbanks. However, this initial difference in seedling densities in burned and unburned plots was lost over time due to continuous recruitment in unburned plots. In a seed addition experiment, plots which were clipped prior to the fire showed higher germination success, possibly because fire temperature was moderated by biomass removal. In another experiment, non-dormant broom seeds were added to burned plots, which then received clipping treatments; there were no differences in broom seedling survival in clipped vs. unclipped plots. Overall, these results suggest that prescribed burning might contribute to the control of Scotch broom invasion by helping eliminate mature stands without significantly increasing regeneration from seed.

Biotic resistance via granivory: establishment by invasive, naturalized, and native asters reflects generalist preference

Escape from specialist natural enemies is frequently invoked to explain exotic plant invasions, but little attention has been paid to how generalist consumers in the recipient range may influence invasion. We examined how seed preferences of the widespread generalist granivore Peromyscus maniculatus related to recruitment of the strongly invasive exotic Centaurea stoebe and several weakly invasive exotics and natives by conducting laboratory feeding trials and seed addition experiments in the field. Laboratory feeding trials showed that P. maniculatus avoided consuming seeds of C. stoebe relative to the 12 other species tested, even when seeds of alternative species were 53-94% smaller than those of C. stoebe. Seed addition experiments conducted in and out of rodent exclosures revealed that weakly invasive exotics experienced relatively greater release from seed predation than C. stoebe, although this was not the case for natives. Seed mass explained 81% of the variation in recruitment associated with rodent exclusion for natives and weak invaders, with larger-seeded species benefiting most from protection from granivores. However, recruitment of C. stoebe was unaffected by rodent exclusion, even though the regression model predicted seeds of correspondingly large mass should experience substantial predation. These combined laboratory and field results suggest that generalist granivores can be an important biological filter in plant communities and that species-specific seed attributes that determine seed predation may help to explain variation in native plant recruitment and the success of exotic species invasions.

Integral projection model of insect herbivore effects onCirsium altissimumpopulations along productivity gradients

Prediction of the role of native insect herbivores in the population growth and spatial distribution of native plants within the environment remains limited. We developed an integral projection model (IPM) to explore the effect of native insect herbivores on the population dynamics and distribution of the native plant Cirsium altissimum (tall thistle), in two different productivity zones in tallgrass prairie in Nebraska USA. Model parameters were extracted from two field experiments: a seed addition experiment that demonstrated seed limitation by insect herbivores on adult recruitment and an insect herbivory exclusion experiment that demonstrated significant herbivore impacts on both rosette performance and adult fecundity. Zones differed in elevation, soil moisture and biomass. Using our model, we asked: (1) does insect herbivory reduce plant population growth rate (λ), (2) does the effect on (λ) differ between productivity zones, (3) does it primarily operate by limiting growth or fecundity and (4) is there evidence for density dependence in mediating impact on (λ)? We found that insect herbivory suppressed tall thistle population growth rate, but the magnitude of this effect did not vary with ecosystem productivity. Insect herbivores' effect on λ arose primarily through suppression of fecundity. We found no evidence of negative density dependence countering seed limitation effects of insect herbivory on tall thistle population growth. The similar magnitudes of these effects in different productivity zones eliminate variation in ecosystem productivity as a majority condition in determining insect herbivores' impact in shaping local distribution of this native plant. Overall, the results show that native insect herbivores suppress population growth of this native plant, mediated through seed limitation that functionally determines adult density, without differences related to ecosystem productivity.

Simplifying methods to assess site suitability for plant recruitment

Few studies link seed dispersal with its demographic consequences, or provide reliable estimates of seed dispersal effectiveness. One reason is the complexity of measuring the suitability for plant recruitment of seed arrival sites. In this study, I compare three methods that differ in the effort required to measure site suitability for seedling recruitment. All are based on the proportion of seeds that become seedlings (seedling-to-seed ratios). Method I is the most detailed and labour intensive. The fate of seeds was followed throughout the different steps of the recruitment process, from fruit removal until seedling emergence, including both seeds dispersed by different animals and undispersed seeds. Method II is based on seed addition experiments. Seeds were sowed in plots, and seedlings emerging were counted in the following two seasons. In Method III, average seed input during dispersal was measured with soil seed bank samples taken in pre- and post-dispersal periods, and seedling emergence estimated with samples of three seasons. Method II provided results similar to those of Method I, which, conversely, provides more insight in the actual processes driving recruitment. Method III, however, systematically underestimated site suitability (seedling-to-seed ratios) by about 50% as compared to the other methods in all microhabitats studied. Relative instead of absolute indices of site suitability were, however, reliable with this method. Method II and III are significantly less costly and could be good alternatives to Method I for some purposes, simplifying future studies on the demographic consequences of seed dispersal and the effectiveness of dispersers.

Long‐distance Dispersal of Invasive Grasses by Logging Vehicles in a Tropical Dry Forest

ABSTRACTPredicting responses of vegetation to environmental factors in human‐altered tropical ecosystems requires an understanding of the controls on plant population expansion across landscapes (i.e., long‐distance dispersal) as well as of factors affecting recruitment at local scales (i.e., microsite conditions). We studied the roles of light availability, habitat type, soil disturbance, and seed dispersal in a selectively logged forest in lowland Bolivia where the exotic forage grass Urochloa (Panicum) maxima is abundant on roads and log landings but does not invade felling gaps or unlogged forest. Shade‐house trials and seed addition experiments with U. maxima revealed that this C4 grass thrives in high light but also grows in partial shade (10% full sun, but not 1% full sun), and that felling gaps, but not undisturbed forest, are suitable for grass establishment. To determine if seed dispersal by logging vehicles explains the discrepancy between actual and potential grass recruitment sites, we collected grass seeds that fell from trucks onto log landings located long distances (>500 m) from established grass populations. Trucks dispersed an estimated 1800 alien grass seeds per log landing during the early dry season; automobiles also transported seeds of grass (135 seeds/vehicle). The seeds collected (and relative abundances) were the exotics U. (Panicum) maxima (97%) and Urochloa (Brachiaria) brizantha (2%), and the pan‐tropical weeds Sorghum halapense (1%) and Rottboellia cochinchinensis (0.2%). Grasses invade this forest where disturbance coincides with seed dispersal by motor vehicles, while dispersal limitation apparently prevents invasion of many sites otherwise suitable for grass recruitment (i.e., felling and natural gaps).

Herb establishment in a young Pseudotsuga menziesii forest: a seven-year seed addition experiment1

The abundance of old-growth associated herbs in young Pseudotsuga menziesii forests is limited in part by the low availability of propagules of these species, but it remains unclear whether the slow post-logging development of old-growth species in younger forests also reflects a lack of microhabitats suitable for seed germination. This study attempts to fill an almost total lack of knowledge about germination of understory species in second-growth Douglas-fir forests, and in young coniferous forests in general, while providing a preliminary exploration of the relative importance of these two limiting factors. The ability of six common forest herbs to germinate in a 40-year-old Pseudotsuga menziesii stand was tested, and survival and growth were monitored for seven years. Five of the six species germinated well (8–50% germination), while one open-site species failed to germinate. After seven years, survival of four old-growth associated species was ∼10% (of the original seed number), while only 1% of seeds of a disturbance-adapted species were still present as seedlings. For all species, seedlings grew slowly and did not reach reproductive size, forming a seedling bank analogous to that of shade-tolerant tree seedlings. All old-growth associated species showed higher overall seedling numbers or greater survival in gaps than in non-gap microsites, and two species showed higher survival in the more mesic lower part of the watershed. Naturally occurring seedlings were rare and included only easily dispersed species that are forest generalists or characteristic of mature, but not of old-growth forests. I conclude that germination of late-seral herb species in young Pseudotsuga menziesii forests is not strictly microsite limited, but that these species are instead limited by a combination of low propagule availability and low resource levels with concomitant slow growth rates.

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.

Fire and mice: Seed predation moderates fire's influence on conifer recruitment

In fire-adapted ecosystems, fire is presumed to be the dominant ecological force, and little is known about how consumer interactions influence forest regeneration. Here, we investigated seed predation by deer mice (Peromyscus maniculatus) and its effects on recruitment of ponderosa pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) seedlings in unburned and recently burned fire-adapted montane forests in west-central Montana, USA. Deer mice were almost twice as abundant in burned than unburned stands. Deer mouse removal of seeds from petri dishes was two times higher in burned than in unburned stands, and seed removal levels were 8% higher for ponderosa pine than for the smaller Douglas-fir seeds. In seed-addition experiments, emergence of seedlings in deer mouse-exclusion cages was almost six times higher in burned compared to unburned forest. In both burned and unburned forest, emergence was lower for ponderosa pine than for Douglas-fir. Seedling survival to establishment did not differ between conifer species but was considerably higher in burned than in unburned forest. However, effects of seed predation on recruitment prevailed over fire effects: in cages allowing access by deer mice, emergence and establishment were extremely rare for both conifer species in both burned and unburned forest. This research suggests that consumer interactions can substantially influence recruitment even in fire-adapted forest ecosystems.

Seed limitation restricts population growth in shaded populations of a perennial woodland orchid

Seed production and seedling recruitment are thought to be of minor importance in determining population dynamics and long-term viability in long-lived perennial plants. Seed addition experiments, on the other hand, have amply shown that supplemental addition of seeds almost always, irrespective of longevity, results in increased seedling recruitment. Any change in the environment that affects fruit and seed production can thus be expected to affect seedling recruitment, but the extent to which increased fruit and seed production affect overall population dynamics remains relatively unknown. In this paper, we present demographic data of six populations of the long-lived woodland orchid Orchis purpurea that were monitored for seven consecutive years (2002-2008) occurring in two contrasting light environments. We use a nested life table response experiment (LTRE) at the vital rate level to disentangle the relative contributions of each of six annual transitions, six sites, and two light environments on the population dynamics of this species and to determine vital rate variations that contributed most to variation in population growth rate. Population growth rates (lamda) were significantly higher in the light environment than in the shaded environment (average lamda = 0.9930 and 1.0492 in the shaded and light environment, respectively). The LTRE analysis showed that variation in fecundity and, to a lesser extent, variation in growth made the largest total contributions to variation in lamda, whereas the contributions of variation in survival were almost negligible. Fruit production was two times larger and the net reproductive rate (R0) was approximately six times higher in the light environment than in shaded areas, suggesting that variables related to reproduction are the key drivers of population dynamics of this long-lived orchid species in different light environments. Our results indicate that light is an important factor affecting population dynamics of Orchis purpurea and illustrate that, even in long-lived species, flower and seed production can have important effects on the population dynamics.

Regeneration processes of biological soil crusts, macro-cryptogams and vascular plant species after fine-scale disturbance in a temperate region: Recolonization or successional replacement?

Open base-rich sandy habitats are often characterised by an extensive cover of biological soil crusts (BSCs). We studied the impact of mechanical disturbances on cyanobacteria/algae-dominated BSCs, macro-cryptogams (bryophytes, lichens), phanerogams and their regeneration processes in a 3-year field experiment in a temperate region. We compared controls with two fine-scale disturbance regimes, each based on 50 plots (20 cm×20 cm): 1. BSCs were raked (weak disturbance), 2. BSCs were completely removed (strong disturbance). We used a Sileno conicae-Cerastion semidecandri habitat as a model area for the field study and ask if the crusts are able to regenerate or will be replaced by macro-cryptogams and/or phanerogams. To clarify whether there is an autochthonous recolonization by vascular plant species out of the seed bank, we analysed the upper soil layer with the seedling-emergence method. Furthermore, we considered the possibility of seed limitation in the harsh environment and added seeds of four habitat-typical species. To assess the impact of disturbance on the soil nutrient content we analysed ammonium, nitrate, total N and phosphate contents. There was a recolonization by macro-cryptogams 2 years after strong disturbance. The cyanobacteria/algae crusts resettled only to 5% in comparison to the previous 40% cover. Therefore, the strong disturbance impact results in a replacement of the crust. Two years after disturbance the ruderalisation indices are higher on strongly disturbed plots than on weakly disturbed and intact plots. Total species numbers of phanerogams were lower on strongly disturbed plots, even in the last year of the experiment. A few habitat-typical phanerogams showed significantly lower numbers of individuals on strongly disturbed plots ( Koeleria glauca, Myosotis stricta; seedlings: Erodium cicutarium, Saxifraga tridactylites). The intermediate-disturbance hypothesis implies that weak disturbance should lead to higher species numbers. This was not the case in our system. Similarity in species composition between seed bank and above-ground vegetation was 70%. The seed-addition experiment revealed that emergence, survival and establishment were species specific; two added species emerged in intact and weakly disturbed crusts, demonstrating seed limitation of the system. Nutrient contents showed variability in time; ammonium and total N revealed treatment effects (strong disturbance resulted in lower contents). There was no recolonization of BSCs to the pre-disturbance stage in a 3-year period, but a successional replacement occurred; especially acrocarpic bryophytes and phanerogams replaced the cyanobacteria/algae stages in the case of our fine-scale disturbance regime.

Seed arrival, ecological filters, and plant species richness: a meta‐analysis

Theoretical models predict that effects of dispersal on local biodiversity are influenced by the size and composition of the species pool, as well as ecological filters that limit local species membership. We tested these predictions by conducting a meta-analysis of 28 studies encompassing 62 experiments examining effects of propagule supply (seed arrival) on plant species richness under contrasting intensities of ecological filters (owing to disturbance and resource availability). Seed arrival increased local species richness in a wide range of communities (forest, grassland, montane, savanna, wetland), resulting in a positive mean effect size across experiments. Mean effect size was 70% higher in disturbed relative to undisturbed communities, suggesting that disturbance increases recruitment opportunities for immigrating species. In contrast, effect size was not significantly influenced by nutrient or water availability. Among seed-addition experiments, effect size was positively correlated with species and functional diversity within the pool of added seeds (species evenness and seed-size diversity), primarily in disturbed communities. Our analysis provides experimental support for the general hypothesis that species pools and local environmental heterogeneity interactively structure plant communities. We highlight empirical gaps that can be addressed by future experiments and discuss implications for community assembly, species coexistence, and the maintenance of biodiversity.

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.

Importance of eelgrass early life history stages in response to oyster aquaculture disturbance

Seagrasses are a critical element in many estuaries and act as drivers of abiotic and biotic processes. One species, Zostera marina L., has been declining globally. A potential contributor to this decline is shellfish aquaculture, although we know little about its impacts. On the US west coast, shellfish aquaculture co-occurs with protected eelgrass habitats. Many aquaculture practices constitute a periodic disturbance, and a key concern is eelgrass recovery. We used observations and experiments to understand how oyster aquaculture practices (i.e. dredging (oysters grown on the bot- tom and harvested mechanically via dredging) and off-bottom longline culture (oysters suspended off the bottom on rope and harvested by hand)) influence eelgrass recovery. Studies of natural recruit- ment showed highest seedling densities in dredged beds (7 seedlings m -2 ) and lowest under long- lines (0.1 seedlings m -2 ). Seed production was highest in dredged beds (295 seeds m -2 ) and lowest in longline beds (52 seeds m -2 ). Seed addition experiments were conducted to understand the effect of oyster aquaculture and adult eelgrass neighbors on seedling germination, growth, and survival. In March 2005, seedling germination was 146% higher in eelgrass removal treatments compared to control plots, with no difference among aquaculture and reference areas. By April 2005, there were no effects of neighbors, but reference areas had greater seed densities (11 seedlings m -2 ) compared to longline areas (3.2 seedlings m -2 ). By August 2005, seedling mortality in longline and reference control plots was 100%. In dredged areas, seedlings in removal plots had greater biomass (0.38 g) than seedlings in control plots (0.14 g). We propose that if eelgrass is to be disturbed by aquaculture, dredge beds may recover more successfully than longline beds.

Are Plant Populations Seed Limited? A Critique and Meta‐Analysis of Seed Addition Experiments

We examine the relative importance of processes that underlie plant population abundance and distribution. Two opposing views dominate the field. One posits that the ability to establish at a site is determined by the availability of suitable microsites (establishment limitation), while the second asserts that recruitment is limited by the availability of seeds (seed limitation). An underlying problem is that establishment and seed limitation are typically viewed as mutually exclusive. We conducted a meta-analysis of seed addition experiments to assess the relative strength of establishment and seed limitation to seedling recruitment. We asked (1) To what degree are populations seed and establishment limited? (2) Under what conditions (e.g., habitats and life-history traits) are species more or less limited by each? (3) How can seed addition studies be better designed to enhance our understanding of plant recruitment? We found that, in keeping with previous studies, most species are seed limited. However, the effects of seed addition are typically small, and most added seeds fail to recruit to the seedling stage. As a result, establishment limitation is stronger than seed limitation. Seed limitation was greater for large-seeded species, species in disturbed microsites, and species with relatively short-lived seed banks. Most seed addition experiments cannot assess the relationship between number of seeds added and number of subsequent recruits. This shortcoming can be overcome by increasing the number and range of seed addition treatments.