Current and lagged climate affects phenology across diverse taxonomic groups.

The timing of life events (phenology) can be influenced by climate. Studies from around the world tell us that climate cues and species' responses can vary greatly. If variation in climate effects on phenology is strong within a single ecosystem, climate change could lead to ecological disruption, but detailed data from diverse taxa within a single ecosystem are rare. We collated first sighting and median activity within a high-elevation environment for plants, insects, birds, mammals and an amphibian across 45 years (1975-2020). We related 10 812 phenological events to climate data to determine the relative importance of climate effects on species' phenologies. We demonstrate significant variation in climate-phenology linkage across taxa in a single ecosystem. Both current and prior climate predicted changes in phenology. Taxa responded to some cues similarly, such as snowmelt date and spring temperatures; other cues affected phenology differently. For example, prior summer precipitation had no effect on most plants, delayed first activity of some insects, but advanced activity of the amphibian, some mammals, and birds. Comparing phenological responses of taxa at a single location, we find that important cues often differ among taxa, suggesting that changes to climate may disrupt synchrony of timing among taxa.

Feeding friend and foe: ample pollen mitigates the effects of pollen theft for a gynodioecious plant, Polemonium foliosissimum (Polemoniaceae).

BACKGROUND AND AIMS: Most angiosperms rely on pollinators to transport pollen and effect fertilization. While some floral visitors are effective pollinators, others act as thieves, consuming pollen but effecting little pollination in return. The importance of pollen theft in male and female reproductive success has received little attention. Here, we examined if pollen consumption by flies altered pollen receipt and exacerbated pollen limitation for a bumblebee-pollinated plant, Polemonium foliosissimum (Polemoniaceae). METHODS: To examine the effect of pollen-thieving flies, we took a three-pronged approach. First, we used single-visit observations to quantify pollen removal and pollen deposition by flies and bumblebees. Second, we manipulated pollen in the neighbourhood around focal plants in two years to test whether pollen reduction reduced pollen receipt. Third, we combined pollen reduction with hand-pollination to test whether pollen thieving exacerbated pollen limitation. Polemonium foliosissimum is gynodioecious in most populations in the Elk Mountains of central Colorado, USA. Thus, we also tested whether pollen theft affected hermaphrodites and females differently. RESULTS: Flies removed significantly more pollen and deposited less pollen per visit than did bumblebees. Reduction of pollen in the neighbourhood around focal plants reduced pollen receipt in both years but only nearly significantly so in 2015. In 2016, plants were significantly pollen-limited; hand-pollination significantly increased seeds per fruit for both hermaphrodites and females. However, the reduction of pollen around focal plants did not exacerbate pollen limitation for either hermaphrodites or females. CONCLUSIONS: Our results suggest that plants tolerate significant consumption of pollen by thieves and pollinators by producing ample pollen to feed both and fertilize available ovules. Our results demonstrate that pollen limitation in P. foliosissimum is driven by lack of effective pollinators rather than lack of pollen. Teasing out these effects elucidates the relative importance of drivers of reproductive success and thus the expected response to selection by different floral visitors.

Genotype-specific effects of ericoid mycorrhizae on floral traits and reproduction in Vaccinium corymbosum.

PREMISE: Most plants interact with mycorrhizal fungi and animal pollinators simultaneously. Yet, whether mycorrhizae affect traits important to pollination remains poorly understood and may depend on the match between host and fungal genotypes. Here, we examined how ericoid mycorrhizal fungi affected flowering phenology, floral traits, and reproductive success, among eight genotypes of highbush blueberry, Vaccinium corymbosum (Ericaceae). We asked three overarching questions: (1) Do genotypes differ in response to inoculation? (2) How does inoculation affect floral and flowering traits? (3) Are inoculated plants more attractive to pollinators and less pollen limited than non-inoculated plants of the same genotype? METHODS: To examine these questions, we experimentally inoculated plants with ericoid mycorrhizal fungi, grew the plants in the field, and measured flowering and floral traits over 2 years. In year 2, we conducted a hand-pollination experiment to test whether plants differed in pollen limitation. RESULTS: Inoculated plants had significantly higher levels of colonization for some genotypes, and there were significant floral trait changes in inoculated plants for some genotypes as well. On average, inoculated plants produced significantly larger floral displays, more fruits per inflorescence, and heavier fruits with lower sugar content, than non-inoculated, control plants. Hand pollination enhanced the production of fruits, and fruit mass, for non-inoculated plants but not for those that were inoculated. CONCLUSIONS: Our results demonstrate that inoculation with ericoid mycorrhizal fungi enhanced flowering and altered investment in reproduction in genotype-specific ways. These findings underscore the importance of examining belowground symbionts and genotype-specific responses in their hosts to fully understand the drivers of aboveground interactions.

Herbivore removal reduces influence of arbuscular mycorrhizal fungi on plant growth and tolerance in an East African savanna.

The functional relationship between arbuscular mycorrhizal fungi (AMF) and their hosts is variable on small spatial scales. Here, we hypothesized that herbivore exclusion changes the AMF community and alters the ability of AMF to enhance plant tolerance to grazing. We grew the perennial bunchgrass, Themeda triandra Forssk in inoculum from soils collected in the Kenya Long-term Exclosure Experiment where treatments representing different levels of herbivory have been in place since 1995. We assessed AMF diversity in the field, using terminal restriction fragment length polymorphism and compared fungal diversity among treatments. We conducted clipping experiments in the greenhouse and field and assessed regrowth. Plants inoculated with AMF from areas accessed by wild herbivores and cattle had greater biomass than non-inoculated controls, while plants inoculated with AMF from where large herbivores were excluded did not benefit from AMF in terms of biomass production. However, only the inoculation with AMF from areas with wild herbivores and no cattle had a positive effect on regrowth, relative to clipped plants grown without AMF. Similarly, in the field, regrowth of plants after clipping in areas with only native herbivores was higher than other treatments. Functional differences in AMF were evident despite little difference in AMF species richness or community composition. Our findings suggest that differences in large herbivore communities over nearly two decades has resulted in localized, functional changes in AMF communities. Our results add to the accumulating evidence that mycorrhizae are locally adapted and that functional differences can evolve within small geographical areas.

Is Plant Fitness Proportional to Seed Set? An Experiment and a Spatial Model.

Individual differences in fecundity often serve as proxies for differences in overall fitness, especially when it is difficult to track the fate of an individual's offspring to reproductive maturity. Using fecundity may be biased, however, if density-dependent interactions between siblings affect survival and reproduction of offspring from high- and low-fecundity parents differently. To test for such density-dependent effects in plants, we sowed seeds of the wildflower Ipomopsis aggregata (scarlet gilia) to mimic partially overlapping seed shadows of pairs of plants, one of which produced twice as many seeds. We tested for differences in offspring success using a genetic marker to track offspring to flowering multiple years later. Without density dependence, the high-fecundity parent should produce twice as many surviving offspring. We also developed a model that considered the geometry of seed shadows and assumed limited survivors so that the number of juvenile recruits is proportional to the area. Rather than a ratio of 2∶1 offspring success from high- versus low-fecundity parents, our model predicted a ratio of 1.42∶1, which would translate into weaker selection. Empirical ratios of juvenile offspring and of flowers produced conformed well to the model's prediction. Extending the model shows how spatial relationships of parents and seed dispersal patterns modify inferences about relative fitness based solely on fecundity.

Gender inequality in predispersal seed predation contributes to female seed set advantage in a gynodioecious species.

Most flowering plants are hermaphrodites. However, in gynodioecious species, some members of the population are male-sterile and reproduce only by setting seed, while others gain fitness through both male and female function. How females compensate for the loss of male function remains unresolved for most gynodioecious species. Here, as with many plants, fitness differences may be influenced by interactions with multiple species. However, whether multiple species interactions result in gender-specific fitness differences remains unknown. Using observational data from 2009-2010, we quantified seed set of the two sex morphs of Polemonium foliosissimu and asked how it is affected by pollination, and seed predation from a dipteran predispersal seed predator (Anthomyiidae: Hylemya sp.). We assessed seed production and losses to predation in 27 populations for one year and in six populations for a second year. Females set significantly more seed than did hermaphrodites in both years. Of the fitness components we assessed, including the number of flowers per plant, fruit set, seeds/fruit, and proportion of fruits destroyed by Hylemya, only fruit destruction differed significantly between the sexes. In one year, seeds/fruit and predation had a stronger effect on seed set for hermaphrodites than for females. Because predispersal seed predators do not pollinate flowers, their effects may depend on successful pollination of flowers on which they oviposit. To examine if genders differed in pollen limitation and seed predation and/or their interactive effects, in 2011 we hand-pollinated flowers and removed seed predator eggs in a fully factorial design. Both sexes were pollen limited, but their degree of pollen limitation did not differ. However, predation reduced.seed set more for hermaphrodites than for females. We found no significant interaction between hand pollen and seed predation, and no interaction between hand pollination and gender. Our results suggest that while interactions with both pollinators and seed predators affect reproductive success, floral enemies can cause inequality in seed set between genders. The next step is to understand how the seed set advantage affects long-term fitness and persistence of females in gynodioecious populations.

Enough is enough: the effects of symbiotic ant abundance on herbivory, growth, and reproduction in an African acacia.

Understanding how cooperative interactions evolve and persist remains a central challenge in biology. Many mutualisms are thought to be maintained by "partner fidelity feedback," in which each partner bases their investment on the benefits they receive. Yet, we know little about how benefits change as mutualists vary their investment, which is critical to understanding the balance between mutualism and antagonism in any given partnership. Using an obligate ant-plant mutualism, we manipulated the density of symbiotic acacia ants (Crematogaster mimosae) and examined how the costs and benefits to Acacia drepanolobium trees scaled with ant abundance. Benefits of ants to plants saturated with increasing ant abundance for protection from branch browsing by elephants and attack by branch galling midges, while varying linearly for protection from cerambycid beetles. In addition, the risk of catastrophic whole-tree herbivory by elephants was highest for trees with very low ant abundance. However, there was no relationship between ant abundance and herbivory by leaf-feeding invertebrates, nor by vertebrate browsers such as giraffe, steinbuck, and Grant's gazelle. Ant abundance did not significantly influence rates of branch growth on acacias, but there was a significant negative relationship between ant abundance and the number of fruits produced by host plants, suggesting that maintaining high-density ant colonies is costly. Because benefits to plants largely saturated with increasing colony size, while costs to plant reproduction increased, we suggest that ant colonies may achieve abundances that are higher than optimal for host plants. Our results highlight the conflicts of interest inherent in many mutualisms, and demonstrate the value of examining the shape of curves relating costs and benefits within these globally important interactions.

Spatial pattern enhances ecosystem functioning in an African savanna.

The finding that regular spatial patterns can emerge in nature from local interactions between organisms has prompted a search for the ecological importance of these patterns. Theoretical models have predicted that patterning may have positive emergent effects on fundamental ecosystem functions, such as productivity. We provide empirical support for this prediction. In dryland ecosystems, termite mounds are often hotspots of plant growth (primary productivity). Using detailed observations and manipulative experiments in an African savanna, we show that these mounds are also local hotspots of animal abundance (secondary and tertiary productivity): insect abundance and biomass decreased with distance from the nearest termite mound, as did the abundance, biomass, and reproductive output of insect-eating predators. Null-model analyses indicated that at the landscape scale, the evenly spaced distribution of termite mounds produced dramatically greater abundance, biomass, and reproductive output of consumers across trophic levels than would be obtained in landscapes with randomly distributed mounds. These emergent properties of spatial pattern arose because the average distance from an arbitrarily chosen point to the nearest feature in a landscape is minimized in landscapes where the features are hyper-dispersed (i.e., uniformly spaced). This suggests that the linkage between patterning and ecosystem functioning will be common to systems spanning the range of human management intensities. The centrality of spatial pattern to system-wide biomass accumulation underscores the need to conserve pattern-generating organisms and mechanisms, and to incorporate landscape patterning in efforts to restore degraded habitats and maximize the delivery of ecosystem services.

Inoculum source dependent effects of ericoid, mycorrhizal fungi on flowering and reproductive success in highbush blueberry (Vaccinium corymbosum).

Most terrestrial angiosperms form mutualisms with both mycorrhizal fungi and animal pollinators. Yet, the effects of mycorrhizae on pollinator behavior and plant reproduction are unknown for most species, and whether the source or type of mycorrhizal fungi affects reproductive success has rarely been examined. We examined whether inoculating highbush blueberry (Vaccinium corymbosum; Ericaceae) with ericoid mycorrhizal fungi enhanced investment in flowering and attractiveness to pollinators, and thus reduced their levels of pollen limitation over that of non-inoculated plants. We also examined the degree to which pollen limitation was dependent on inoculation source and the surrounding pollinator community context. Three-year-old saplings of Vaccinium corymbosum 'Bluecrop' or highbush blueberry (Ericaceae) were inoculated with a) ericoid mycorrhizal fungi within soil of the rhizosphere of plants growing at a local blueberry farm, b) a commercially available ericoid inoculant, c) both the local soils and commercial inoculum, or d) were not inoculated and served as controls. They were grown for one year in pots in a common garden and, in the following year, were moved to six farms in central Vermont that were known from prior studies to differ in pollinator abundance and diversity. We conducted a hand pollination experiment at each farm to examine if inoculation or pollinator abundance (i.e., farm context) affected reproductive success. Plants treated with all types of inoculums were more likely to flower, and produced more inflorescence buds than non-inoculated plants in 2018. However, in 2019, plants in the combination inoculum treatment, alone, produced more inflorescence buds than those in the other treatments. Neither the source of inoculum nor hand pollination affected fruit set (the proportion of flowers setting fruit), or fruit sugar content. Hand pollination, but not inoculation, increased berry mass and the average number of seeds produced/berry. Our results add to the growing body of evidence that mycorrhizal fungi can affect reproductive traits of their hosts but that the effects of mycorrhizal fungi depend on the mycorrhizal symbionts.

Additive effects of herbivory, nectar robbing and seed predation on male and female fitness estimates of the host plant Ipomopsis aggregata.

Many antagonistic species attack plants and consume specific plant parts. Understanding how these antagonists affect plant fitness individually and in combination is an important research focus in ecology and evolution. We examined the individual and combined effects of herbivory, nectar robbing, and pre-dispersal seed predation on male and female estimates of fitness in the host plant Ipomopsis aggregata. By examining the effects of antagonists on plant traits, we were able to tease apart the direct consumptive effects of antagonists versus the indirect effects mediated through changes in traits important to pollination. In a three-way factorial field experiment, we manipulated herbivory, nectar robbing, and seed predation. Herbivory and seed predation reduced some male and female fitness estimates, whereas plants tolerated the effects of robbing. The effects of herbivory, robbing, and seed predation were primarily additive, and we found little evidence for non-additive effects of multiple antagonists on plant reproduction. Herbivory affected plant reproduction through both direct consumptive effects and indirectly through changes in traits important to pollination (i.e., nectar and phenological traits). Conversely, seed predators primarily had direct consumptive effects on plants. Our results suggest that the effects of multiple antagonists on estimates of plant fitness can be additive, and investigating which traits respond to damage can provide insight into how antagonists shape plant performance.

Termites create spatial structure and govern ecosystem function by affecting N2 fixation in an East African savanna.

The mechanisms by which even the clearest of keystone or dominant species exert community-wide effects are only partially understood in most ecosystems. This is especially true when a species or guild influences community-wide interactions via changes in the abiotic landscape. Using stable isotope analyses, we show that subterranean termites in an East African savanna strongly influence a key ecosystem process: atmospheric nitrogen fixation by a monodominant tree species and its bacterial symbionts. Specifically, we applied the 15N natural abundance method in combination with other biogeochemical analyses to assess levels of nitrogen fixation by Acacia drepanolobium and its effects on co-occurring grasses and forbs in areas near and far from mounds and where ungulates were or were not excluded. We find that termites exert far stronger effects than do herbivores on nitrogen fixation. The percentage of nitrogen derived from fixation in Acacia drepanolobium trees is higher (55-80%) away from mounds vs. near mounds (40-50%). Mound soils have higher levels of plant available nitrogen, and Acacia drepanolobium may preferentially utilize soil-based nitrogen sources in lieu of fixed nitrogen when these sources are readily available near termite mounds. At the scale of the landscape, our models predict that termite/soil derived nitrogen sources influence >50% of the Acacia drepanolobium trees in our system. Further, the spatial extent of these effects combine with the spacing of termite mounds to create highly regular patterning in nitrogen fixation rates, resulting in marked habitat heterogeneity in an otherwise uniform landscape. In summary, we show that termite-associated effects on nitrogen processes are not only stronger than those of more apparent large herbivores in the same system, but also occur in a highly regular spatial pattern, potentially adding to their importance as drivers of community and ecosystem structure.

Termites, vertebrate herbivores, and the fruiting success of Acacia drepanolobium.

In African savannas, vertebrate herbivores are often identified as key determinants of plant growth, survivorship, and reproduction. However, plant reproduction is likely to be the product of responses to a suite of abiotic and biotic factors, including nutrient availability and interactions with antagonists and mutualists. In a relatively simple system, we examined the role of termites (which act as ecosystem engineers--modifying physical habitat and creating islands of high soil fertility), vertebrate herbivores, and symbiotic ants, on the fruiting success of a dominant plant, Acacia drepanolobium, in East African savannas. Using observational data, large-scale experimental manipulations, and analysis of foliar N, we found that Acacia drepanolobium trees growing at the edge of termite mounds were more likely to reproduce than those growing farther away, in off-mound soils. Although vertebrate herbivores preferentially used termite mounds as demonstrated by dung deposits, long-term exclusion of mammalian grazers did not significantly reduce A. drepanolobium fruit production. Leaf N was significantly greater in trees growing next to mounds than in those growing farther away, and this pattern was unaffected by exclusion of vertebrates. Thus, soil enrichment by termites, rather than through dung and urine deposition by large herbivores, is of primary importance to fruit production near mounds. Across all mound-herbivore treatment combinations, trees that harbored Crematogaster sjostedti were more likely to fruit than those that harbored one of the other three ant species. Although C. sjostedti is less aggressive than the other ants, it tends to inhabit large, old trees near termite mounds which are more likely to fruit than smaller ones. Termites play a key role in generating patches of nutrient-rich habitat important to the reproductive success of A. drepanolobium in East African savannas. Enhanced nutrient acquisition from termite mounds appears to allow plants to tolerate herbivory and the reduced defense by a relatively ineffective ant partner. Our results underscore the importance of simultaneously examining top-down and bottom-up effects to understand those factors most important to plant reproductive success.

Flowers as viral hot spots: Honey bees (Apis mellifera) unevenly deposit viruses across plant species.

RNA viruses, once considered specific to honey bees, are suspected of spilling over from managed bees into wild pollinators; however, transmission routes are largely unknown. A widely accepted yet untested hypothesis states that flowers serve as bridges in the transmission of viruses between bees. Here, using a series of controlled experiments with captive bee colonies, we examined the role of flowers in bee virus transmission. We first examined if honey bees deposit viruses on flowers and whether bumble bees become infected after visiting contaminated flowers. We then examined whether plant species differ in their propensity to harbor viruses and if bee visitation rates increase the likelihood of virus deposition on flowers. Our experiment demonstrated, for the first time, that honey bees deposit viruses on flowers. However, the two viruses we examined, black queen cell virus (BQCV) and deformed wing virus (DWV), were not equally distributed across plant species, suggesting that differences in floral traits, virus ecology and/or foraging behavior may mediate the likelihood of deposition. Bumble bees did not become infected after visiting flowers previously visited by honey bees suggesting that transmission via flowers may be a rare occurrence and contingent on multiplicative factors and probabilities such as infectivity of virus strain across bee species, immunocompetence, virus virulence, virus load, and the probability a bumble bee will contact a virus particle on a flower. Our study is among the first to experimentally examine the role of flowers in bee virus transmission and uncovers promising avenues for future research.

RNA virus spillover from managed honeybees (Apis mellifera) to wild bumblebees (Bombus spp.).

The decline of many bumblebee species (Bombus spp.) has been linked to an increased prevalence of pathogens caused by spillover from managed bees. Although poorly understood, RNA viruses are suspected of moving from managed honeybees (Apis mellifera) into wild bumblebees through shared floral resources. We examined if RNA viruses spillover from managed honeybees, the extent to which viruses are replicating within bumblebees, and the role of flowers in transmission. Prevalence and active infections of deformed wing virus (DWV) were higher in bumblebees collected near apiaries and when neighboring honeybees had high infection levels. We found no DWV in bumblebees where honeybee foragers and honeybee apiaries were absent. The prevalence of black queen cell virus (BQCV) was also higher in bumblebees collected near apiaries. Furthermore, we detected viruses on 19% of flowers, all of which were collected within apiaries. Our results corroborate the hypothesis that viruses are spilling over from managed honeybees to wild bumblebees and that flowers may be an important route for transmission.

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography.

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.

Home sick: impacts of migratory beekeeping on honey bee (Apis mellifera) pests, pathogens, and colony size.

Honey bees are important pollinators of agricultural crops and the dramatic losses of honey bee colonies have risen to a level of international concern. Potential contributors to such losses include pesticide exposure, lack of floral resources and parasites and pathogens. The damaging effects of all of these may be exacerbated by apicultural practices. To meet the pollination demand of US crops, bees are transported to areas of high pollination demand throughout the year. Compared to stationary colonies, risk of parasitism and infectious disease may be greater for migratory bees than those that remain in a single location, although this has not been experimentally established. Here, we conducted a manipulative experiment to test whether viral pathogen and parasite loads increase as a result of colonies being transported for pollination of a major US crop, California almonds. We also tested if they subsequently transmit those diseases to stationary colonies upon return to their home apiaries. Colonies started with equivalent numbers of bees, however migratory colonies returned with fewer bees compared to stationary colonies and this difference remained one month later. Migratory colonies returned with higher black queen cell virus loads than stationary colonies, but loads were similar between groups one month later. Colonies exposed to migratory bees experienced a greater increase of deformed wing virus prevalence and load compared to the isolated group. The three groups had similar infestations of Varroa mites upon return of the migratory colonies. However, one month later, mite loads in migratory colonies were significantly lower compared to the other groups, possibly because of lower number of host bees. Our study demonstrates that migratory pollination practices has varying health effects for honey bee colonies. Further research is necessary to clarify how migratory pollination practices influence the disease dynamics of honey bee diseases we describe here.

Mutualism as reciprocal exploitation: African plant-ants defend foliar but not reproductive structures.

The foundation of many plant-ant mutualisms is ant protection of plants from herbivores in exchange for food and/or shelter. While the role of symbiotic ants in protecting plants from stem- and leaf-feeding herbivores has been intensively studied, the relationship between ant defense and measures of plant fitness has seldom been quantified. We studied ant aggression, damage by herbivores and seed predators, and fruit production among Acacia drepanolobium trees occupied by four different acacia-ant species in an East African savanna. Levels of ant aggression in response to experimental disturbance differed strongly among the four species. All four ant species recruited more strongly to new leaf growth on host plants following disturbance, while recruitment to developing fruits was on average an order of magnitude lower. Host plants occupied by more aggressive ant species suffered significantly less vegetative damage from leaf-feeding insects, stem-boring beetles, and vertebrate browsers than host plants occupied by less aggressive ant species. However, there were no differences among fruiting host plants occupied by different ant species in levels of seed predation by bruchid seed predators. Fruit production on host trees was significantly correlated with tree stem diameter but not with the identity of resident ants. Our results demonstrate that defense of host plants may differ substantially among ant species and between vegetative and reproductive structures and that fruit production is not necessarily correlated with high levels of aggression by resident ants.

Oviposition choices by a pre-dispersal seed predator (Hylemya sp.) : I. Correspondence with hummingbird pollinators, and the role of plant size, density and floral morphology.

Although the importance of pollinators has most often been examined in the evolution of floral characters, seed predators may also play a role in shaping floral evolution. In this study, I examined the role of interplant distance, plant size, and flower morphology on Ipomopsis aggregatás (Polemoniaceae) attractiveness to a pre-dispersal seed predator, Hylemya sp. (Anthomyiidae) and to hummingbird pollinators. The attractiveness of I. aggregata individuals to Hylemya was nonlinearly related to interplant distance in experimental arrays. Clumped and highly dispersed plants were preyed upon more frequently than those at intermediate distances. I found no relationship between interplant distance and visitation rates by hummingbird pollinators in these experimental arrays. However, in natural populations studied, clumped plants were more frequently approached by hummingbirds than those growing more widely dispersed. Display size was unrelated to visitation by Hylemya on inflorescences I clipped and maintained as "large", "small" and "control". Display size was also unrelated to the total number of visits by hummingbird pollinators to each of these experimental plants, however "large" display plants were more likely to be visited first in any given visitation sequence. Of various morphological measurements, corolla length showed the strongest positive correlation with Hylemya egg presence. To the extent that plant spacing and morphology is correlated with pollinator visits and ultimate seed set, Hylemya could be choosing flowers optimally, and playing a role in the evolution of floral traits.

Oviposition choices by a pre-dispersal seed predator (Hylemya sp.) : II. A positive association between female choice and fruit set.

Pre-dispersal seed predators must choose proper hosts and flowers on those hosts that have a high probability of setting seed. In this study, I documented that flowers of Ipomopsis aggregata (Polemoniaceae) chosen by a pre-dispersal seed predator (Anthomyiidae: Hylemya sp.) had a higher probability of seed set than flowers the fly did not choose. I experimentally tested the hypothesis that Hylemya uses a post-fertilization cue in making oviposition choices. I found no evidence for this hypothesis.

Nectar robbing in Ipomopsis aggregata : effects on pollinator behavior and plant fitness.

Hummingbirds foraging in alpine meadows of central Colorado, United States, face a heterogeneous distribution of nectar rewards. This study investigated how variability in nectar resources caused by nectar-robbing bumblebees affected the foraging behavior of hummingbird pollinators and, subsequently, the reproductive success of a host plant (Ipomopsis aggregata). We presented hummingbirds with experimental arrays of I. aggregata and measured hummingbird foraging behavior as a function of known levels of nectar robbing. Hummingbirds visited significantly fewer plants with heavy nectar robbing (over 80% of available flowers robbed) and visited fewer flowers on those plants. These changes in hummingbird foraging behavior resulted in decreased percent fruit set as well as decreased total seed set in heavily robbed plants. These results indicate that hummingbird avoidance of nectar-robbed plants and flowers reduces plant fitness components. In addition, our results suggest that the mutualisms between pollinators and host plants may be affected by other species, such as nectar robbers.