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Abstract Synchronized episodic reproduction among long‐lived plants shapes ecological interactions, ecosystem dynamics, and evolutionary processes worldwide. Two active scientific fields investigate the causes and consequences of such synchronized reproduction: the fields of masting and fire‐stimulated flowering. While parallels between masting and fire‐stimulated flowering have been previously noted, there has been little dialogue between these historically independent fields. We predict that the synthesis of these fields will facilitate new insight into the causes and consequences of synchronized reproduction. Here we briefly review parallels between masting and fire‐stimulated flowering, using two case studies and a database of 1870 plant species to facilitate methodological, conceptual, geographical, taxonomic, and phylogenetic comparisons. We identify avenues for future research and describe three key opportunities associated with synthesis. First, the taxonomic and geographic complementarity of empirical studies from these historically independent fields highlights the potential to derive more general inferences about global patterns and consequences of synchronized reproduction in perennial plants. Second, masting's well developed conceptual framework for evaluating adaptive hypotheses can help guide empirical studies of fire‐stimulated species and enable stronger inferences about the evolutionary ecology of fire‐stimulated flowering. Third, experimental manipulation of reproductive variation in fire‐stimulated species presents unique opportunities to empirically investigate foundational questions about ecological and evolutionary processes underlying synchronized reproduction. Synthesis of these fields and their complementary insights offers a unique opportunity to advance our understanding of the evolutionary ecology of synchronized reproduction in perennial plants. 

Summary A recent study posited that fire in grasslands promotes persistence of plant species by improving mating opportunities and reproductive outcomes. We devised an investigation to test these predicted mechanisms in two widespread, long‐lived perennials. We expect fire to synchronize flowering, increase mating and boost seed set.We quantified individual flowering phenology and seed set ofLiatris asperaandSolidago speciosafor 3 yr on a preserve in Minnesota, USA. The preserve comprises two management units burned on alternating years, allowing for comparisons between plants in burned and unburned areas within the same year, and plants in the same area across years with and without burns.Fire increased flowering synchrony and increased time between start date and peak flowering. Individuals of both species that initiated flowering later in the season had higher seed set. Fire was associated with substantially higher flowering rates and seed set inL.asperabut notS.speciosa. InL.aspera, greater synchrony was associated with increased mean seed set.Although fire affected flowering phenology in both species, reproductive success improved only in the species in which fire also synchronized among‐year flowering. Our results support the hypothesis that reproduction in some grassland species benefits from fire. 

Premise Herbivores directly influence plant reproduction by damaging reproductive tissue which reduces seed production in consumed plants. Consumption of reproductive tissue may also indirectly depress reproduction in unconsumed plants by isolating unconsumed individuals from prospective mates and reducing pollination. However, empirical support for such hypothesized indirect effects remains limited. Methods To investigate potential indirect effects of herbivory on mating opportunities and pollination, we quantified spatial patterns of white-tailed deer (Odocoileus virginianus) herbivory and reproductive outcomes in Lilium philadelphicum (Liliaceae). We mapped 708 flowering plants, monitored deer herbivory, and examined how distance to prospective mates influenced rates of ovule fertilization, a proxy for pollen receipt. Results Most floweringL. philadelphicum plants (83%) failed to produce seed. Deer consumption of flowers prevented seed production in 35% of individuals. The probability of deer herbivory declined with distance to flowering conspecifics. Sixty-five percent of the remaining, unconsumed plants failed to produce seed due to apparent pollen limitation. While ovule fertilization rates declined with multiple predictors quantifying distances to flowering conspecifics, isolation metrics that excluded plants consumed by deer significantly improved predictions of ovule fertilization. Conclusions Our results offer empirical support for the hypothesis that herbivory reduces pollination and seed production in unconsumed plants by altering spatial mating opportunities. This feedback between deer herbivory and distance-dependent pollination contributes to widespread reproductive failure in L. philadelphicum, though the extent to which deer reduce seed production varies with plant isolation. Herbivory may exacerbate pollen limitation in other species by isolating unconsumed plants from prospective mates. 

Data and R code necessary to replicate figures and analyses presented in the following manuscript: Beck and Wagenius. In press. Herbivory exacerbates pollen limitation by isolating unconsumed plants from prospective mates. American Journal of Botany. 

Periodic fire enhances seedling recruitment for many plant species in historically fire-dependent ecosystems. Fire is expected to promote recruitment by generating environmental conditions that promote seedling emergence and survival. However, fire may also increase flowering and seed production. This makes it difficult to distinguish the effects of microsite conditions from seed availability in observational studies of seedling recruitment. Experiments that manipulate seed inputs across a representative range of conditions are needed to elucidate how seed availability versus microsite conditions influence post-fire seedling recruitment and plant demography. We experimentally manipulated time since fire across 36 patches of remnant tallgrass prairie distributed across 6400 ha in western Minnesota (USA). Over two years, we sowed 11,057Echinacea angustifolia(Asteraceae) seeds across 84 randomly placed transects and tracked 974 experimentally sown seedlings to evaluate how time since fire influenced seedling emergence and survival after experimentally controlling for variation in seed inputs. We also quantified six environmental variables and evaluated whether these covariates were associated with seedling emergence and survival. Fire influenced both seedling emergence and seedling survival. Seedlings emerged from approximately 1 percent of all seeds sown prior to experimental burns. Seeds sown one year after experimental burns emerged at 15 times the rate of seeds sown in the fall before burns, but emergence then declined as time since fire increased. Sowing seeds at high densities reduced rates of seedling emergence but increased overall recruitment. Increases in litter depth were associated with reduced emergence. Meanwhile, the probability that seedlings survived to late summer was greatest when they emerged 0-1 years after fire. The probability of seedling survival decreased with litter depth and increased with the local density of conspecific seedlings. Our findings experimentally support widespread predictions that fire enhances seedling recruitment by generating microsite conditions favorable for seedling emergence and survival – especially by increasing the light available to newly emerged seedlings. Nevertheless, recruitment also increased with seed inputs indicating that both seed availability and microsite conditions influence post-fire recruitment. Explicitly discriminating between seed-limitation and microsite-limitation is critical for understanding the demographic processes that influence plant population dynamics in historically fire-dependent ecosystems. 

Anthropogenically fragmented populations may have reduced fitness due to loss of genetic diversity and inbreeding. The extent of such fitness losses due to fragmentation and potential gains from conservation actions are infrequently assessed together empirically. Controlled crosses within and among populations can identify whether populations are at risk of inbreeding depression and whether inter-population crossing alleviates fitness loss. Because fitness depends on the environment and life stage, studies quantifying cumulative fitness over a large portion of the lifecycle in conditions that mimic natural environments are most informative. To assess the fitness consequences of habitat fragmentation, we leveraged controlled within-family, within-population, and between-population crosses to quantify inbreeding depression and heterosis in seven populations of Echinacea angustifolia within a 6,400-ha area. We then assessed cumulative offspring fitness after 14 yr of growth in a natural experimental plot (N = 1,136). The mean fitness of progeny from within-population crosses varied considerably, indicating genetic differentiation among source populations, even though these sites are all less than 9 km apart. The fitness consequences of within-family and between-population crosses varied in magnitude and direction. Only one of the seven populations showed inbreeding depression of high effect, while four populations showed substantial heterosis. Outbreeding depression was rare and slight. Our findings indicate that local crossings between isolated populations yield unpredictable fitness consequences ranging from slight decreases to substantial increases. Interestingly, inbreeding depression and heterosis did not relate closely to population size, suggesting that all fragmented populations could contribute to conservation goals as either pollen recipients or donors. 

Anthropogenically fragmented populations may have reduced fitness due to loss of genetic diversity and inbreeding. The extent of such fitness losses due to fragmentation and potential gains from conservation actions are infrequently assessed together empirically. Controlled crosses within and among populations can identify whether populations are at risk of inbreeding depression and whether interpopulation crossing alleviates fitness loss. Because fitness depends on environment and life stage, studies quantifying cumulative fitness over a large portion of the lifecycle in conditions that mimic natural environments are most informative. To assess fitness consequences of habitat fragmentation, we leveraged controlled within-family, within-population, and between-population crosses to quantify inbreeding depression and heterosis in seven populations of Echinacea angustifolia within a 6400-hectare area. We then assessed cumulative offspring fitness after 14 years of growth in a natural experimental plot (N = 1136). Mean fitness of progeny from within-population crosses varied considerably, indicating genetic differentiation among source populations, even though these sites are all less than 9 km apart. The fitness consequences of within-family and between-population crosses varied in magnitude and direction. Only one of the seven populations showed inbreeding depression of high effect, while four populations showed substantial heterosis. Outbreeding depression was rare and slight. Our findings indicate that local crossings between isolated populations yield unpredictable fitness consequences ranging from slight decreases to substantial increases. Interestingly, inbreeding depression and heterosis did not relate closely to population size, suggesting that all fragmented populations could contribute to conservation goals as either pollen recipients or donors. 

Many plant species in historically fire-dependent ecosystems exhibit fire-stimulated flowering. While greater reproductive effort after fire is expected to result in increased reproductive outcomes, seed production often depends on pollination, the spatial distribution of prospective mates, and the timing of their reproductive activity. Fire-stimulated flowering may thus have limited fitness benefits in small, isolated populations where mating opportunities are restricted and pollination rates are low. We conducted a 6-y study of 6,357 Echinacea angustifolia (Asteraceae) individuals across 35 remnant prairies in Minnesota (USA) to experimentally evaluate how fire effects on multiple components of reproduction vary with population size in a common species. Fire increased annual reproductive effort across populations, doubling the proportion of plants in flower and increasing the number of flower heads 65% per plant. In contrast, fire’s influence on reproductive outcomes differed between large and small populations, reflecting the density-dependent effects of fire on spatiotemporal mating potential and pollination. In populations with fewer than 20 individuals, fire did not consistently increase pollination or annual seed production. Above this threshold, fire increased mating potential, leading to a 24% increase in seed set and a 71% increase in annual seed production. Our findings suggest that density-dependent effects of fire on pollination largely determine plant reproductive outcomes and could influence population dynamics across fire-dependent systems. Failure to account for the density-dependent effects of fire on seed production may lead us to overestimate the beneficial effects of fire on plant demography and the capacity of fire to maintain plant diversity, especially in fragmented habitats.