Disruption of a belowground mutualism alters interactions between plants and their floral visitors

Plants engage in diverse and intimate interactions with unrelated taxa. For example, aboveground floral visitors provide pollination services, while belowground arbuscular mycorrhizal fungi (AMF) enhance nutrient capture. Traditionally in ecology, these processes were studied in isolation, reinforcing the prevailing assumption that these above- and belowground processes were also functionally distinct. More recently, there has been a growing realization that the soil surface is not a barrier to many ecological interactions, particularly those involving plants (who live simultaneously above and below ground). Because of the potentially large impact that mycorrhizae and floral visitors can have on plant performance and community dynamics, we designed an experiment to test whether these multi-species mutualisms were interdependent under field conditions. Using benomyl, a widely used fungicide, we suppressed AMF in a native grassland, measuring plant, fungal, and floral-visitor responses after three years of fungal suppression. AMF suppression caused a shift in the community of floral visitors from large-bodied bees to small-bodied bees and flies, and reduced the total number of floral visits per flowering stem 67% across the 23 flowering species found in the plots. Fungal suppression has species-specific effects on floral visits for the six most common flowering plants in this experiment. Exploratory analyses suggest these results were due to changes in floral-visitor behavior due to altered patch-level floral display, rather than through direct effects of AMF suppression on floral morphology. Our findings indicate that AMF are an important, and overlooked, driver of floral-visitor community structure with the potential to affect pollination services. These results support the growing body of research indicating that interactions among ecological interactions can be of meaningful effect size under natural field conditions and may influence individual performance, population dynamics, and community structure.     

Scale dependence of reproductive failure in fragmented Echinacea populations

I investigated reproduction in a three-year study of Echinacea angustifolia, purple coneflower, growing in a fragmented prairie landscape. I quantified the local abundance of flowering conspecifics at individual-based spatial scales and at a population-based spatial scale. Regression analyses revealed that pollen limitation increased while seed set and fecundity decreased with isolation of individual plants. Isolation, defined as the distance to the k(th) nearest flowering conspecific, was a good predictor of pollen limitation, for all nearest neighbors considered (k = 1-33), but the strength of the relationship, as quantified by R2, peaked at intermediate scales (k = 2-18). The relationship of isolation to seed set and fecundity was similarly strongest at intermediate scales (k = 3-4). The scale dependence of individual density effects on reproduction (density of flowering plants within x meters) resembled that of isolation. Analyses at a population-based scale showed that pollen limitation declined significantly with population size. Seed set and fecundity also declined with population size, but significantly so only in 1998. Whether quantifying local abundance with population- or individual-based measures, reproductive failure due to pollen limitation is a consistent consequence of Echinacea scarcity. However, individual-based measures of local abundance predicted pollen limitation from a wider sample of plants with a simpler model than did population size. Specifically, the largest site, a nature preserve, is composed of plants with intermediate individual isolation and, as predicted, intermediate pollen limitation, but its large population size poorly predicted population mean pollen limitation.     

Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers

The timing of life history traits is central to lifetime fitness and nowhere is this more evident or well studied as in the phenology of flowering in governing plant reproductive success. Recent changes in the timing of environmental events attributable to climate change, such as the date of snowmelt at high altitudes, which initiates the growing season, have had important repercussions for some common perennial herbaceous wildflower species. The phenology of flowering at the Rocky Mountain Biological Laboratory (Colorado, USA) is strongly influenced by date of snowmelt, which makes this site ideal for examining phenological responses to climate change. Flower buds of Delphinium barbeyi, Erigeron speciosus, and Helianthella quinquenervis are sensitive to frost, and the earlier beginning of the growing season in recent years has exposed them to more frequent mid-June frost kills. From 1992 to 1998, on average 36.1% of Helianthella buds were frosted, but for 1999-2006 the mean is 73.9%; in only one year since 1998 have plants escaped all frost damage. For all three of these perennial species, there is a significant relationship between the date of snowmelt and the abundance of flowering that summer. Greater snowpack results in later snowmelt, later beginning of the growing season, and less frost mortality of buds. Microhabitat differences in snow accumulation, snowmelt patterns, and cold air drainage during frost events can be significant; an elevation difference of only 12 m between two plots resulted in a temperature difference of almost 2 degrees C in 2006 and a difference of 37% in frost damage to buds. The loss of flowers and therefore seeds can reduce recruitment in these plant populations, and affect pollinators, herbivores, and seed predators that previously relied on them. Other plant species in this environment are similarly susceptible to frost damage so the negative effects for recruitment and for consumers dependent on flowers and seeds could be widespread. These findings point out the paradox of increased frost damage in the face of global warming, provide important insights into the adaptive significance of phenology, and have general implications for flowering plants throughout the region and anywhere climate change is having similar impacts.     

Predispersal seed herbivores, not pollinators, exert selection on floral traits via female fitness

Herbivores that oviposit in flowers of animal-pollinated plants depend on pollinators for seed production and are therefore expected to choose flowers that attract pollinators. This provides a mechanism by which seed herbivores and pollinators could impose conflicting selection on floral traits. We measured phenotypic selection on floral traits of Lobelia siphilitica (Lobeliaceae) via female fitness to determine the relative strength of selection by pollinators and a specialist predispersal seed herbivore. We were able to attribute selection on flowering phenology to the herbivores. However, no selection could be attributed to pollinators, resulting in no conflicting selection on floral traits. Unlike pollinators, whose preference for certain floral traits does not always translate into higher fitness, any discrimination by seed herbivores is likely to decrease fitness of the preferred floral phenotype. Thus predispersal seed herbivores may be a significant agent of selection on floral traits.     

Age-size plasticity for reproduction in monocarpic plants

Empirical and theoretical investigations of monocarpy have usually addressed the question of minimum or threshold sizes for reproduction. However, the range of flowering sizes observed in many monocarpic species is extraordinarily large (well beyond what can be called a "threshold"), and the sizes of flowering and nonflowering plants may overlap greatly. We attempt to explain these reproductive patterns in terms of optimal reaction norms predicted by simple deterministic life history models. We assume that individuals differ in their growth trajectories due to the heterogeneous quality of microsites and ask how the optimal age and size at flowering varies with environmental variation in growth and for different assumptions about fecundity and mortality. Under two very different growth functions (one with no age- or size-related decline in growth rate and another with such a decline as size approaches an asymptote), the optimal reaction norms imply considerable plasticity for size at reproduction, particularly when poor growth is associated with higher mortality or lower asymptotic size. Deterministic models such as these may be more applicable to long-lived than to short-lived monocarps, because fitness potential should be less affected by stochastic variability in yearly growing condition in the former than in the latter. We consider the case of a tropical monocarpic and masting tree species, Cerberiopsis candelabra (Apocynaceae), and show that our model results can account for wide ranges of reproductive size and overlap in size of flowering and nonflowering plants, in accord with observation. We suggest that empirical attention to norms of reaction across growth environments will be a more profitable approach than investigation of size thresholds per se.     

Mimics and magnets: the importance of color and ecological facilitation in floral deception

Plants that lack floral rewards can attract pollinators if they share attractive floral signals with rewarding plants. These deceptive plants should benefit from flowering in close proximity to such rewarding plants, because pollinators are locally conditioned on floral signals of the rewarding plants (mimic effect) and because pollinators are more abundant close to rewarding plants (magnet effect). We tested these ideas using the non-rewarding South African plant Eulophia zeyheriana (Orchidaceae) as a study system. Field observations revealed that E. zeyheriana is pollinated solely by solitary bees belonging to a single species of Lipotriches (Halictidae) that appears to be closely associated with the flowers of Wahlenbergia cuspidata (Campanulaceae), a rewarding plant with which the orchid is often sympatric. The pale blue color of the flowers of E. zeyheriana differs strongly from flowers of its congeners, but is very similar to that of flowers of W. cuspidata. Analysis of spectral reflectance patterns using a bee vision model showed that bees are unlikely to be able to distinguish the two species in terms of flower color. A UV-absorbing sunscreen was applied to the flowers of the orchid in order to alter their color, and this resulted in a significant decline in pollinator visits, thus indicating the importance of flower color for attraction of Lipotriches bees. Pollination success in the orchid was strongly affected by proximity to patches of W. cuspidata. This was evident from one of two surveys of natural populations of the orchid, as well as experiments in which we translocated inflorescences of the orchid either into patches of W. cuspidata or 40 m outside such patches. Flower color and location of E. zeyheriana plants relative to rewarding magnet patches are therefore key components of the exploitation by this orchid of the relationship between W. cuspidata and Lipotriches bee pollinators.     

Pollination in Dianthus deltoides (Caryophyllaceae): Effects of Habitat Fragmentation on Visitation and Seed Set

Abstract: I analyse the effects of habitat fragmentation on the pollination success of a perennial, butterfly-pollinated, caryophyllaceous herb, the maiden pink, Dianthus deltoides L. The study was conducted in July 1986 and July 1987 at two different sites in southwest Sweden, an undisturbed "mainland" site and a fragmented site consisting of "habitat islands" within a heavily utilized agricultural area The fragmented area had a lower diversity and abundance of both flowering plants and flower-visiting insects. Dianthus flowers received fewer visits in the fragmented area than in the mainland area, and the seed set was much lower. Hand pollination increased seed set up to 4.1 times in the fragmented area, but no significant differences were found between hand-pollinated and control flowers at the mainland site. There were no differences between the two sites in standing crop of nectar, ovule number per flowers, or seed set of bagged flowers, band-pollinated flowers, and hand-pollinated fertilized flowers Thus, the difference in natural seed set between the two sites can be explained by differences in pollinator service.     

Plant Reproductive Traits as a Function of Local, Regional, and Global Abundance

Most ecological generalizations stem from the study of common organisms, but most species are rare. There are a number of reasons to expect that rare and common species may differ consistently in their characteristics, with possible implications for conservation. Past analyses of this issue, however, have generally considered only a single measure of rarity and have not corrected analyses for the lack of independence in the traits of related species. We compared several reproductive characteristics of Mediterranean annual crucifers as a function of their global range, regional abundance, and local population density in Israel, making use of independent contrast methods to correct for the phylogenetic relationships of the species involved. We found plants growing at low local density to be disproportionately likely to be self-compatible. Petal length and floral depth (sepal length) were correlated with breeding system but showed significant interaction effects between breeding system and abundance. Floral longevity was inversely related to abundance and also showed significant interaction effects between breeding system and abundance. Overall, rare species tended to display more extreme values for floral traits than did common plants with the same breeding systems; they had unusually large and deep flowers if self-incompatible and unusually small, shallow ones if self-compatible.     

Phytoremediation of petroleum-contaminated soils: Pre-screening for suitable plants and rhizospheral fungi

A field study was conducted in a petroleum-contaminated site in Kermanshah refinery (Iran) to find the petroleum-resistant plant species and the rhizospheral fungi for being used in bioremediation. Results showed that the amounts of petroleum pollution in non-vegetated areas is 6.8% and in vegetated areas is 2.2%. Plant samples were collected from petroleum-polluted areas and determined using morphological characters. It was found that eight plant species were growing on the contaminated sites: Polygonum aviculare, Amaranthus retroflexus, Noea mucronata, Alhaji cameleron, Crozophora heirosolymitrana, Poa sp., Convolvulus arvensis and Trifolium repense. The rhizospheral fungi of the plants were collected and determined using microscopic studies and taxonomical keys. The results determined the presence of 21 species in the rhizosphere of the plants growing in the polluted areas; three of these species were common in all of the plants and the others have species-specific distribution within the plants. The highest number of rhizospheral fungi (11 species) were determined for P. aviculare in non-polluted areas and nine species in polluted areas. It seems that the plant is the best candidate for using phytoremediation. However, the variation of fungi in petroleum-polluted areas was more than non-polluted zones. The culture of fungi in oil-contaminated media showed that, although all the studied fungi were resistant to low petroleum pollution (1% v/v), a few species, especially Fusarium species, showed higher resistance to petroleum pollution (10% v/v) and hence they may be suitable for mycoremediation in highly polluted areas.     

Ecological and evolutionary mechanisms for low seed: ovule ratios: need for a pluralistic approach?

Central to the ecology and evolution of a broad range of plants is understanding why they routinely have submaximal reproduction manifested as low seed : ovule and fruit : flower ratios. We know much less about the processes responsible for low seed : ovule ratios than we do for fruit : flower ratios. Current hypotheses for low seed : ovule ratios are largely drawn from those for fruit : flower ratios, including proximate (ecological) causes of pollen limitation, resource limitation, and pollen quality, as well as the ultimate (evolutionary) hypothesis of "bet hedging" on stochastic pollination. Yet, such mechanisms operating on fruit : flower ratios at the whole-plant level may not best explain low seed : ovule ratios at the individual-flower level. We tested each of these proximate and ultimate causes for low seed : ovule ratios using the specialized pollination mutualism between senita cacti (Pachycereus schottii) and senita moths (Upiga virescens). Seed : ovule ratios were consistently low (approximately 0.61). Such excess ovule production by senita likely has a strong genetic component given the significant differences among plants in ovule number and the consistency in ovule production by plants within and among flowering seasons. Excess ovule production and low seed : ovule ratios could not be explained by pollen limitation, resource limitation, pollen quality, or bet hedging. Nevertheless, phenotypic selection analyses did show significant selection gradients for increased ovule number, suggesting that other evolutionary processes may be responsible for excess ovule production and low seed : ovule ratios. In contrast, low fruit : flower ratios at the whole-plant level were explained by an apparent equilibrium between pollen and resource limitation. Thus, mechanisms responsible for low fruit : flower ratios at the whole-plant level are not necessarily in accord with those of low seed : ovule ratios at the individual-flower level. This suggests that we may need to adopt a more pluralistic approach to seed : ovule ratios and consider alternative hypotheses, including a greater array of proximate and ultimate causes. Initial results of this study suggest that floral allometry, selection on correlated floral traits, stigma clogging with pollen grains, and style clogging with pollen tubes may provide promising avenues for understanding low seed : ovule ratios.     

Soil nutrient supply modulates temperature-induction cues in mast-seeding grasses

Synchronous and intermittent reproduction in long-lived plants, known as mast seeding, is induced by climatic cues, but the mechanism explaining variation in masting among neighboring but edaphically segregated species is unknown. Soil nutrients can enhance flowering, and thus, populations on nutrient-rich soils may require less-favorable growing temperatures to flower. We tested this hypothesis by predicting the probability of flowering in response to air temperature for five species of alpine Chionochloa grasses in South Island, New Zealand, over 37 years and relating our predictions to soil N supply (NH4(+) + NO3(-)). Summer air temperatures better predicted flowering than spring air temperatures, which were correlated with soil N mineralization. Species on N-rich soils required lower mean temperatures to induce flowering and/or responded more consistently across a gradient of air temperatures, contributing to the higher probability of their tillers and tussocks flowering at low summer temperatures. Our results suggest that flowering primarily occurs in response to warm summer temperatures, but species on N-rich soils require less favorable growing conditions because they invest relatively less N in seeds. Thus, predicting masting requires a consideration of the interactions among climate, the internal resources of plants, and mineral nutrient uptake.     

乐安江重金属污染对浮游植物群落结构的影响

１９９３年和１９９４年对乐安江６个采样站浮游植物的两次调查结果表明，藻类的群落结构因不同河段水质状况不同而不同：未受矿山废水污染的上游海口站，藻类多样性指数较高，密度较低，优势种多为清洁神和耐中污的种类，水质较好；接纳大量矿山及采矿化工废水和生活污水的沽口站，藻类受到明显抑制，多样性指数极低，耐污种占绝对优势，水质极差；随流程和净化时间的延长，下游各站藻类的种类、数量、多样性指数、种类数比均不断上升，优势比不断下降，体现出明显的水质净化趋势．乐安江水体Ｃｕ２＋浓度与藻类的种类、数量、种类多样性指数、种类数比呈负相关的关系，Ｃｕ２＋浓度与藻类的优势比呈正相关关系；ｐＨ值与上述指标关系正好与Ｃｕ２＋所呈现的关系相反．藻类群落结构在春季丰水期与秋季枯水期的变化表明，春季乐安江水质好于秋季．     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Effects of Population Size on Seed Production and Germinability in an Endangered, Fragmented Grassland Plant

Abstract: Fragmentation and isolation of plant populations are thought to affect demographic processes such as seed production and cause reductions in fitness. I followed seed set over a 3-year period in eight populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) that differed in population size from 13 to over 5000 flowering plants. Germinability of the resultant seed was also examined to determine whether small populations had lower fitness than large populations. Seed set was significantly associated with population size in 2 of the 3 years. Small populations (<30 flowering plants) produced significantly fewer seeds per head in 1994 and 1995 than did large populations (500 to over 5000 flowering plants), which did not differ significantly from one another. There was, however, substantial variation within populations. In 1993 seed production did not follow any simple relationship with population size, possibly because environmental stress from low rainfall had an overriding impact. Differences in seed germinability between populations were largely not evident, suggesting that this aspect of fitness has not declined substantially in small populations relative to large populations. This study suggests that nongenetic, demographic factors are of immediate importance to the persistence of small populations of R. leptorrhynchoides because of their potential impacts on seedling recruitment.     

Anther-mimicking floral guides exploit a conflict between innate preference and learning in bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Bee-pollinated plants are frequently dichogamous: e.g. each flower has a discernable male and female phase, with only the male phase offering a pollen reward. Pollen-collecting bees should therefore discriminate against female-phase flowers to maximise their rate of pollen harvest, but this behaviour would reduce plant fitness due to inferior pollination. Here, we test the hypothesis that flowers use pollen-mimicking floral guides to prevent flower-phase discrimination. Such floral guides resemble pollen in spectral reflection properties and are widespread among angiosperm flowers. In an array of artificial flowers, bumblebees learned less well to discriminate between flower variants simulating different flowering phases when both flower variants carried an additional pollen-yellow guide mark. This effect depended crucially on the pollen-yellow colour of the guide mark and on its spatial position within the artificial flower. We suggest that floral guides evolved to inhibit flower-phase learning in bees by exploiting the innate colour preferences of their pollinators.     

Effects of plant size and distribution on the numbers of invertebrate species and individuals inhabiting the brown alga Pelvetia fastigiata

The invertebrates living on specimens of the brown alga Pelvetia fastigiata, growing in the intertidal zone at La Jolla, California (USA) in November 1975 were enumerated. Within 7 collecting sites, larger plants generally shelter more animal species and individuals than smaller plants. The number of species on a given plant can be described as an equilibrium number; numbers of species and individuals can become similar on defaunated and transplanted algae of comparable sizes placed at the same experimental site. Such equilibria are site-specific because the colonization, immigration and loss rates of animals differ among the sites studied. Relationships between plant size and the number of animal species and individuals on P. fastigiata also differ among the collecting sites. The between-site differences are related to tidal level, to habitat diversity and to habitat patch-size. Small isolated plants without epiphytes shelter few species. The faunas of larger isolated plants at mid-tide levels generally include many thallus-dwelling, tubiculous, vagrant and epiphyte-dwelling species, but few such species commonly inhabit plants within aggregations of P. fastigiata. Within aggregations, the plants host relatively few epiphytes and thus lack habitat diversity, and the net emigration rates of many animals including epiphyte-dwellers are relatively high. Plants within aggregations, however, usually shelter more animal indivuduals than isolated plants. Thus, faunal diversity is reduced, not increased within the largest patches of P. fastigiata.     

Indirect Effects of Biological Control on Plant Diversity Vary across Sites in Montana Grasslands

Abstract:  Biological control with specialist, nonindigenous, herbivorous insects is an important option for controlling invasive exotic plants in wildlands and nature reserves. It is assumed that biological control agents will reduce the dominance of the target weed, thereby increasing the native diversity of the associated plant community. However, this hypothesis has rarely been tested. We introduced Aphthona nigriscutis into grassland sites infested with the invasive exotic species Euphorbia esula L. on a nature reserve in Montana (U.S.A.). Two sites with better soil had been treated previously with herbicide, whereas two other sites had not. We measured the density and biomass of Euphorbia vegetative and flowering stems and number of native and exotic shrubs, grass-like plants, and forbs in 48 microplots in Aphthona release and control macroplots at each site. After 5 years, Aphthona release was associated with a 33–39% decline in Euphorbia aboveground biomass compared with controls at all sites. Other effects of the biocontrol depended on the site. Biocontrol slowed the recovery of species diversity at the sites previously treated with herbicide but slowed the loss of diversity at sites without a history of herbicide. Biocontrol introduction was not associated with a disproportionate increase in nontarget exotic species. Release of Aphthona caused a decline in the biomass of flowering stems relative to controls at good-soil, previous-herbicide sites but was associated with a relative increase in flower stem mass at poor-soil, no-herbicide sites. Our results suggest that biocontrol reductions in weed dominance will not always be associated with increased species diversity. More emphasis should be placed on conserving desirable communities and less on simple weed control. Monitoring of community-level effects should accompany biocontrol introductions on nature reserves.     

Modeling the Persistence of Small Populations of Strongly Interdependent Species: Figs and Fig Wasps

Conservation problems are usually studied at the population or ecosystem levels. Formulating predictive theory for the domain in between has been difficult. Fig trees and their pollinating wasps, principally tropical groups of organisms, form pairs of obligate mutualists that provide unique opportunities for studying the influence of species interactions on the survival of small populations. Survival of each partner depends on that of the associated species. The pollinator population can be maintained only if figs are produced year-round. Because fig trees flower synchronously at the individual level, wasps have to locate a new individual host tree at each generation. We describe results of simulation models estimating the minimum number of trees required to maintain a wasp population using two levels of the criteria: (1) different probability of survival (50% and 99%) and (2) different time of survival (5 or 1000 years). We also examined how these different estimates are sensitive to differences in the seasonality of flowering period and in the length of the period of female receptivity in figs. Such estimates can be used to understand the potential effects of the reduction of fig population size via fragmentation. Unlike most studies on the effect of low population size on population viability, our paper focuses on maintenance of a biotic interaction, rather than on single-species dynamics. The biotic interaction on which we focus is important because figs in many tropical ecosystems may be keystone resources for frugivores that are in turn essential seed dispersal agents for other plants.     

Population Size and Reproduction in Phlox pilosa

Abstract: We analyzed the relationships between population size and reproductive characteristics in the perennial prairie forb Phlox pilosa , an obligate outcrossing butterfly-pollinated species. We examined 27 populations ranging in size from 9 to over 75,000 flowering ramets in two regions of the state of Iowa (eastcentral and northwest) in 1993 and 1994. We collected flowers from each population and scored them for pollen arrival to stigmas and number of pollen tubes. We collected fruiting ramets from each population at the end of the Phlox growing season and scored them for height, biomass, and reproductive variables, including the number of flowers initiated and opened and the number of capsules initiated and matured. In both years, population size was significantly correlated with the number of capsules matured per ramet. Differences between populations in capsule production were set primarily at the pollination stage. In 1993, pollen arrival to stigmas was significantly lower than in 1994 and was correlated with population size in eastcentral Iowa populations, indicating that lower reproduction in small populations that year was at least partially due to inadequate amounts of pollen being moved. In 1993, weather conditions likely depressed pollinator activity, but absolute capsule formation was high because of high flower production per ramet and high population densities. In 1994, when pollen arrival to stigmas was relatively high and unrelated to population size, outcross pollen movement was greater in larger populations. Increased efficacy of outcross pollen movement in 1994 may have resulted from lower flower production and less dense populations forcing greater pollinator movement between ramets or from variation between years in fine-scale spatial genetic substructuring of populations. Our results indicate that the viability of Phlox pilosa can be best ensured by protecting and creating populations of at least 1000–2000 flowering ramets.     

Inbreeding Depression, Environmental Stress, and Population Size Variation in Scarlet Gilia (Ipomopsis aggregata)

Despite a large body of theory, few studies have directly assessed the effects of variation in population size on fitness components in natural populations of plants. We conducted studies on 10 populations of scarlet gilia, Ipomopsis aggregata , to assess the effects of population size and year-to-year variation in size on the relative fitness of plants. We showed that seed size and germination success are significantly reduced in small populations (those 100 flowering plants) of scarlet gilia. Plants from small populations are also more susceptible to environmental stress. When plants from small and large populations were subjected to an imposed stress (combined effects of transplanting and experimental clipping, simulating ungulate herbivory) in a common garden experiment, plants from small populations suffered higher mortality and were ultimately of smaller size than plants from large populations. In addition, experimental evidence indicates that observed fitness reductions are genetic, due to the effects of genetic drift and/or inbreeding depression. When pollen was introduced from distant populations into two small populations, seed mass and percentage of germination were bolstered, while pollen transferred into a large population had no significant effect. Year-to-year variation in population size and its effects on plant fitness are also discussed. In one small population, for example, a substantial increase in size from within did not introduce sufficient new (archived) genetic material to fully overcome the effects of inbreeding depression.