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• Annually variable and synchronous seed production by plant populations, or masting, is a widespread reproductive strategy in long-lived plants. Masting is thought to be selectively beneficial because interannual variability and synchrony increase the fitness of plants through economies of scale that decrease the cost of reproduction per surviving offspring. Predator satiation is believed to be a key economy of scale, but whether it can drive phenotypic evolution for masting in plants has been rarely explored. • We used data from seven plant species (Quercus humilis, Quercus ilex, Quercus rubra, Quercus alba, Quercus montana, Sorbus aucuparia and Pinus pinea) to determine whether predispersal seed predation selects for plant phenotypes that mast. • Predation selected for interannual variability in Mediterranean oaks (Q. humilis and Q. ilex), for synchrony in Q. rubra, and for both interannual variability and reproductive synchrony in S. aucuparia and P. pinea. Predation never selected for negative temporal autocorrelation of seed production. • Predation by invertebrates appears to select for only some aspects of masting, most importantly high coefficient of variation, supporting individual-level benefits of the population-level phenomenon of mast seeding. Determining the selective benefits of masting is complex because of interactions with other seed predators, which may impose contradictory selective pressures.

• Masting is a widespread reproductive strategy in plants that helps to reduce seed predation and increase pollination. However, masting can involve costs, notably negative density-dependent (NDD) seedling survival caused by concentrating reproduction in intermittent events. Masting benefits have received widespread attention, but the costs are understudied, which precludes understanding why some plant species have evolved intense masting, while others reproduce regularly. • We followed seed production, seed predation (both 13 yr), and seedling recruitment and survival (11 yr) in Sorbus aucuparia. We tested whether NDD in seedling survival after mast years can reduce the benefits of pulsed reproduction that come through predator satiation. • Seed predation rates were extreme in our population (mean = 75%), but were reduced by masting. The commonly accepted, but untested, assertion that pulsed recruitment is associated with strong NDD was unsupported. Consequently, the proportion of seedlings that survived their first year increased with fruit production. This provides a rare test of economies of scale beyond the seed stage. • Our results provide estimation of the costs of mast seeding, and indicate that these may be lower than expected. Low masting costs, if common, may help explain why masting is such a widespread reproductive strategy throughout the plant kingdom.

Plants often face chronic seed loss from predispersal seed predation by insects. Although many studies have documented the rates of seed loss for single species in different communities, it is unclear how rates of predispersal seed predation vary among co‐occurring species within the same community. If interspecific asymmetries in seed loss are great, this common interaction could have important implications for coexistence. Species traits, such as seed size or seed nitrogen and carbon, might correlate with interspecific variation in predispersal seed predation among co‐occurring grassland forb species. We collected infructescences from 13 co‐occurring forb species from each of four western Montana grasslands over 2 years. We quantified the magnitude of seed loss due to predispersal seed predation by insects. The average level of seed loss was 15.8%. Larger seeded species suffered significantly higher levels of predispersal seed predation than smaller seeded species, and seed size predicted seed loss more than seed nitrogen or carbon. Although large‐seeded species often have greater proportional recruitment and early survival than small‐seeded species, our study suggests that these advantages are partially counterbalanced by greater predispersal seed loss for larger versus smaller seeded species. Asymmetries in predispersal seed predation may importantly affect coexistence among these species.

Floral herbivory represents a major threat to plant reproductive success, driving the importance of plant tolerance mechanisms that minimize fitness costs. However, the cumulative insect herbivory plants experience under natural conditions complicates predictions about tolerance contributions to net fitness. Apical damage can lead to compensatory seed production from late season flowering that ameliorates early season fitness losses. Yet, the compensation realized depends on successful development and herbivore escape by later season flowers. Using monocarpic perennial Cirsium canescens, we quantified seed-reproductive fitness of plants with vs. without experimental damage to the early-developing large apical flower head, with and without a 30–40% herbivory reduction on subsequent flower heads, for two flowering cohorts. Plants with reduced herbivory clearly demonstrated the release of apical dominance and compensation, not overcompensation, for apical damage via greater seed maturation by later flower heads. In contrast, plants that experienced ambient herbivory levels on subsequent heads undercompensated for early apical damage. Individuals had lower total seed set when the apical head was damaged. Compensation was, therefore, possible through a small increase in total flower heads, caused by a higher rate of floral bud survival, and a higher seed maturation rate by subsequent heads, leading to more viable seeds per matured flower head. With ambient cumulative floral herbivory, compensation for apical damage was not sufficient to improve fitness. Variation in the intensity of biological interactions played a role in the success of plant tolerance as a mechanism to maximize individual fitness.

Seed predation is an antagonistic interaction that negatively affects the performance of individual plants and can limit plant population dynamics. In animal-dispersed plants, crop size is an important determinant of plant reproductive success through its effect on seed dispersers and predators. Seed traits, such as size or chemical composition, can also increase the tolerance to seed predators or reduce their performance. We investigated the interaction between Quercus faginea and two specialized pre-dispersal insect seed predators (weevils and moths) during two years of contrasting crop size to determine the consequences of oak reproductive investment on seed production and insect performance. Crop size was 44% lower and acorns were 32% smaller in the second year, although acorn predation by insects was proportionally similar between both years at the population level. Individual trees producing larger crops showed a lower incidence of insect predators during the year of abundant acorn production, whereas trees producing bigger acorns experienced higher seed predation rates by insects, and acorns held more insect larvae in the low crop year. Competition between insects increased when acorn production was low, and higher tannin content in acorns further constrained the number of weevil larvae developing together in the same acorn. However, the abundance and size of insect larvae produced per tree were similar between the two crop years, and this was due to larvae often depleting acorn reserves when resources were low. Oak reproductive output increased nearly two-fold during the large crop year. Crop size variation, acorn production in a given year and acorn size and chemical composition seem to be important traits for reducing damage by insect predators in Quercus faginea and improve oak reproductive success.

Plants interact with both antagonistic and mutualistic animals during reproduction, with the outcomes of these interactions significantly influencing plant reproductive success, population dynamics, and the evolution of plant traits. Here, we investigated the spatial and temporal variations in the interactions between Juniperus thurifera, its seed-dispersing birds, and three specific arthropod species that attack the fleshy cones during the predispersal period. We assessed how plant traits affect levels of cone damage by arthropods and seed dispersal by birds, the occurrence of competition among arthropod species, and the impact of seed predators on the activity of frugivores. Plant traits, cone damage by arthropods, and seed dispersal by birds showed spatiotemporal variability. Fluctuation in cone abundance was the leading factor determining damage by arthropods and bird dispersal with a secondary role of cone traits. Large crops satiated predispersal seed predators, although the amount of frugivory did not increase significantly, suggesting a potential satiation of bird dispersers. Crop size and cone traits at individual trees determined preferences by seed predator species and the foraging activity of bird dispersers. Competition among arthropods increased during years of low cone production, and seed predators sometimes negatively affected bird frugivory. High supra-annual variations in cone production appear to be a key evolutionary mechanism enhancing J. thurifera reproductive success. This strategy reduces the impact of specialized seed predators during years of high seed production, despite the potential drawback of satiating seed dispersers.

Variation in selection among populations and years has important implications for evolutionary trajectories of populations. Yet, the agents of selection causing this variation have rarely been identified. Selection on the time of reproduction within a season in plants might differ both among populations and among years, and selection can be mediated by both mutualists and antagonists. We investigated if differences in the direction of phenotypic selection on flowering phenology among 20 populations of Gentiana pneumonanthe during 2 yr were related to the presence of the butterfly seed predator Phengaris alcon, and if butterfly incidence was associated with the abundance of the butterfly's second host, Myrmica ants. In plant populations without the butterfly, phenotypic selection favored earlier flowering. In populations where the butterfly was present, caterpillars preferentially attacked early-flowering individuals, shifting the direction of selection to favoring later flowering. Butterfly incidence in plant populations increased with ant abundance. Our results demonstrate that antagonistic interactions can shift the direction of selection on flowering phenology, and suggest that such shifts might be associated with differences in the community context.

Premise of the study: The processes maintaining flower color polymorphisms have long been of evolutionary interest. Mechanistic explanations include selection through pollinators, antagonists, local environments, drift, and pleiotropic effects. We examined the maintenance of inflorescence color polymorphisms in the genus Protea (Proteaceae) of South Africa, in which ∼40% of species contain different color morphs. Methods: We studied 10 populations of four bird-pollinated Protea species and compared adult performance, floral and leaf morphology, vegetative pigmentation, germination, and seedling survival between co-occurring pink and white morphs. We also tested for differences in pollination success and pre-dispersal seed predation. Key results: White morphs produced seeds 10% heavier and 3.5 times more likely to germinate, which all else being equal, should fuel positive selection on white. In one studied population per species, however, white morphs were more susceptible to seed predation by endophagous larvae. Pollinators had no morph-specific effects on female fecundity, as measured by amount or probability of seed set. Differences in stem color indicated that white morphs produced smaller quantities of pigment and associated compounds throughout, possibly explaining their higher seed palatability. Conclusions: Our findings suggest a mechanism for some white protea polymorphisms: deleterious pleiotropic effects on pink morphs are occasionally offset by reduced losses to seed-eating larvae. Because trends were repeated across species, we suggest that similar processes may also occur in other proteas, placing a new emphasis on seed predators for influencing some of South Africa's amazing floral diversity.

Seed ecologists have often stated that they expect larger-seeded species to have lower survivorship through postdispersal seed predation than smaller-seeded species. Similar predictions can be made for the relationship between survivorship through predispersal seed predation and seed mass. In order to test these predictions, we gathered data regarding survivorship through 24 hours of exposure to postdispersal seed predators for 81 Australian species, and survivorship through predispersal seed predation for 170 Australian species. These species came from an arid environment, a subalpine environment, and a temperate coastal environment. We also gathered data from the published literature (global) on survivorship through postdispersal seed predation for 280 species and survivorship through predispersal seed predation for 174 species. We found a weak positive correlation between seed mass and the percentage of seeds remaining after 24 hours of exposure to postdispersal seed predators at two of three field sites in Australia, and no significant relationship across 280 species from the global literature, or at the remaining field site. There was no significant relationship between seed mass and survivorship through predispersal seed predation either cross-species or across phylogenetic divergences in any of the vegetation types, or in the compilation of data from the literature. Postdispersal seed removal was responsible for a greater percentage of seed loss in our field studies than was predispersal seed predation. On average, 83% of diaspores remained after 24 hours of exposure to postdispersal seed removers, whereas 87% of seeds survived all predispersal seed predation that occurred between seed formation and seed maturity. Mean seed survival was higher in the field studies than in the literature compilations, and species showing 100% survival were heavily underrepresented in the literature. These differences may be due to biases in species selection or publication bias. Seed defensive tissue mass increased isometrically with seed mass, but there was no significant relationship between the amount of defensive tissue per gram of seed reserve mass and survivorship through postdispersal seed predation.

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 foliosissimum 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.