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1. Trees commonly reproduce via masting cycles, which involves synchronized inter-annual variability in fruit crop size. A few individuals in a population will commonly produce much more fruit than others. If these trees produce fruit more frequently, as indicated by a lower inter-annual variability in fruit production, they may dominate fruit production over time. 2. By measuring fruit production of 1635 individuals of 10 temperate tree species across 4 years in northern lower Michigan, we estimated the inter-annual variability and synchrony in each species. We compared fruit production estimates with measurements of tree size, soil nutrient availability and neighbourhood crowding to investigate the source of inter-individual variation in number of fruit produced. 3. We found that trees' fruit production increased with tree size. The trees that accounted for the largest proportion of total fruit production had lower inter-annual variability and higher synchrony in fruit production. These 'super-producer' trees tended to have high nutrient availability and few neighbouring trees, but there were no effects of nutrient availability or neighbourhood crowding on fruit production in the population as a whole. 4. Synthesis. Masting is a population-level phenomenon, and is typically studied at this level. However, when we apply individual tree observations of fruit production to this phenomenon, it reveals super-producers which produce fruit more consistently than the rest of the population. By reducing inter-annual variability in fruit production, but increasing synchrony and making large numbers of fruit, super-producers may be able to reap the benefits of masting while governing population fruit production over time.

In tropical forest communities, seedling recruitment can be limited by the number of fruit produced by adults. Fruit production tends to be highly unequal among trees of the same species, which may be due to environmental factors. We observed fruit production for ~2,000 trees of 17 species across 3 years in a wet tropical forest in Costa Rica. Fruit production was modeled as a function of tree size, nutrient availability, and neighborhood crowding. Following model selection, tree size and neighborhood crowding predicted both the probability of reproduction and the number of fruit produced. Nutrient availability only predicted only the probability of reproduction. In all species, larger trees were more likely to be reproductive and produce more fruit. In addition, number of fruit was strongly negatively related to presence of larger neighboring trees in 13 species; presence of all neighboring trees had a weak‐to‐moderate negative influence on reproductive status in 16 species. Among various metrics of soil nutrient availability, only sum of base cations was positively associated with reproductive status, and for only four species. Synthesis Overall, these results suggest that direct influences on fruit production tend to be mediated through tree size and crowding from neighboring trees, rather than soil nutrients. However, we found variation in the effects of neighbors and nutrients among species; mechanistic studies of allocation to fruit production are needed to explain these differences. We observed fruit production for ~2,000 trees of 17 species across three years in a wet tropical forest in Costa Rica. Fruit production was negatively related to presence of larger neighboring trees in 13 species, but relationships between fruit production and soil nutrient availability were less common. These results suggest that influences on fruit production tend to be mediated through tree size and crowding rather than soil nutrients.

1. Though not often examined together, both plant-soil feedbacks (PSFs) and functional traits have important influences on plant community dynamics and could interact. For example, seedling functional traits could impact seedling survivorship responses to soils cultured by conspecific versus heterospecific adults. Furthermore, levels of functional traits could vary with soil culturing source. In addition, these relationships might shift with light availability, which can affect trait values, microbe abundance, and whether mycorrhizal colonization is mutualistic or parasitic to seedlings. 2. To determine the extent to which functional traits mediate PSFs via seedling survival, we conducted a field experiment. We planted seedlings of four temperate tree species across a gradient of light availability and into soil cores collected beneath conspecific (sterilized and live) and heterospecific adults. We monitored seedling survival twice per week over one growing season, and we randomly selected subsets of seedlings to measure mycorrhizal colonization and phenolics, lignin, and NSC levels at three weeks. 3 . Though evidence for PSFs was limited, Acer saccharum seedlings exhibited positive PSFs (i.e., higher survival in conspecific than heterospecific soils). In addition, soil microbes had a negative effect on A . saccharum and Prunus serotina seedling survival, with reduced survival in live versus sterilized conspecific soil. In general, we found higher trait values (measured amounts of a given trait) in conspecific than heterospecific soils and higher light availability. Additionally, A . saccharum survival increased with higher levels of phenolics, which were higher in conspecific soils and high light. Quercus alba survival decreased with higher AMF colonization. 4. We demonstrate that functional trait values in seedlings as young as three weeks vary in response to soil source and light availability. Moreover, seedling survivorship was associated with trait values for two species, despite both drought and heavy rainfall during the growing season that may have obscured survivorship-trait relationships. These results suggest that seedling traits could have an important role in mediating the effects of local soil source and light levels on seedling survivorship and thus plant traits could have an important role in PSFs.

1. The Janzen-Connell (J-C) Model proposes that host-specific enemies maintain high tree species diversity by reducing seedling performance near conspecific adults and promoting replacement by heterospecific seedlings. Support for this model often comes from decreased performance for a species at near versus far distances from conspecific adults. However, the relative success of conspecific versus heterospecific seedlings recruiting under a given tree species is a critical, but untested, component of the J-C Model. 2. In a shade-house experiment, we tested plant-soil feedbacks as a J-C mechanism in six tropical tree species. We assessed effects of conspecific versus heterospecific cultured soil extracts on seedling performance for each species, and we compared performance of conspecific versus heterospecific seedlings grown with soil extract cultured by a particular tree species. Additionally, we tested whether soil microbes were creating these plant-soil feedbacks and whether low light increased species vulnerability to pathogens. 3. Among 30 potential comparisons of survival and mass for seedlings grown in conspecific versus heterospecific soil extracts, survival decreased in seven and increased in two, whereas mass decreased in 13 and increased in 1. To integrate survival and growth, we also examined seedling performance [(mean total mass x mean survival time)/(days of experiment)], which was lower in 16 and higher in 2 of 30 comparisons between seedlings grown with soil extract cultured by conspecific versus heterospecific individuals. Based on performance within a soil extract, conspecific seedlings were disadvantaged in 15 and favoured in 7 of 30 cases relative to heterospecific seedlings. 4. Species pairwise interactions of soil modification and seedling performance occurred regardless of sterilization, suggesting chemical mediation. Microbes lacked host-specificity and reduced performance regardless of extract source and irradiance. 5. Synthesis. These results, along with parallel research in temperate forests, suggest that plant-soil feedbacks are an important component of seedling dynamics in both ecosystems. However, negative conspecific feedbacks were more prevalent in tropical than temperate species. Thus, negative plant-soil feedbacks appear to facilitate species coexistence via negative distance-dependent processes in tropical but not temperate forests, but the feedbacks were mediated through chemical effects rather than through natural enemies as expected under the J-C Model.

Plant growth responses to resources may be an important mechanism that influences species' distributions, coexistence, and community structure. Irradiance is considered the most important resource for seedling growth in the understory of wet tropical forests, but multiple soil nutrients and species have yet to be examined simultaneously with irradiance under field conditions. To identify potentially limiting resources, we modeled tree seedling growth as a function of irradiance and soil nutrients across five sites, spanning a soil fertility gradient in old-growth, wet tropical forests at La Selva Biological Station, Costa Rica. We measured an array of soil nutrients including total nitrogen (total N), inorganic N (nitrate [NO 3 − ] and ammonium [NH 4 + ]), phosphate (PO 4 − ), and sum of base cations (SBC; potassium, magnesium, and calcium). Shade in the forest understory did not preclude seedling growth correlations with soil nutrients. Irradiance was a significant predictor of growth in 52% of the species, inorganic N in 54% (NO 3 − in 32%; NH 4 + in 34%), total N in 47%, SBC in 39%, and PO 4 − in 29%. Overall, growth was correlated with both irradiance and soil nutrients in 45% of species and with soil nutrients only in an additional 48%; rarely was irradiance alone correlated with growth. Contrary to expectations, the magnitudes of growth effects, assessed as the maximum growth response to significant resources for each species, were similar for irradiance and most soil nutrients. Among species whose growth correlated with soil nutrients, the rank importance of nutrient effects was SBC, followed by N (total N, NO 3 − , and/or NH 4 + ) and PO 4 − . Species' growth responsiveness (i.e., magnitudes of effect) to irradiance and soil nutrients was negatively correlated with species' shade tolerance (survival under 1% full sun). In this broad survey of species and resources, the nearly ubiquitous effects of soil nutrients on seedling growth challenge the idea that soil nutrients are less important than irradiance in the light-limited understory of wet tropical forests.

1. The study of invasiveness typically emphasizes early successional life-history traits in exotic plants, which enable the capture of high resources in disturbed environments and rapid growth. A key issue in invasion dynamics is whether such behaviours come at the expense of traits such as low-light survivorship, which allow species to become more dominant later in succession. 2. We used maximum-likelihood analysis to compare the growth and survivorship of two exotic trees, Ailanthus altissima and Acer platanoides, with nine dominant native tree species in closed-canopy forests in Connecticut, USA. Growth was modelled as a function of light and survivorship as a function of recent growth; combining models yielded estimates of light-dependent mortality. 3. The exotic species had strikingly high growth rates, exceeding all native species at light levels ≥ 10% full sun, and growing 2.6 times faster than the fastest-growing native species at 80% full sun. At low-light levels (< 3% full sun), however, growth rates of five native species exceeded both exotics. Exotic species survivorship (as a function of light-driven growth) was strongly dependent on the degree of shading: at 1% full sun, the annual mortality rate of A. platanoides was 10% and A. altissima was 17%; only two native species had higher mortalities. However, at 5% full sun, A. platanoides' mortality was < 1%, superior to all but three native species. Mortality of all species dropped to < 1% by 10% full sun, except A. altissima whose mortality remained high at c. 10%. 4. A life-history trade-off analysis (based on radial growth, height allometry and low-light survivorship) shows a nearly linear trade-off for most species. The native species and Ailanthus follow the common life-history trade-off of low-light survivorship vs. high-light growth. However, A. platanoides diverges from this trade-off pattern by combining very high growth rates with moderately high shade tolerance. 5. Simulations with SORTIE-ND (a forest dynamics model) indicate that poor survivorship of A. altissima will limit it to disturbed sites, whereas A. platanoides' unusual combination of traits makes it invasive in both disturbed and undisturbed forests. Overall, native shade-tolerant trees and slow stand dynamics make undisturbed forests highly resistant to invasion by exotic trees that are intolerant of shade. 6. Synthesis. This study showcases the importance of rapid growth in invasive plants, holding even for exotic tree species known to invade established forests. For A. altissima, high growth rates were accompanied by poor low-light survivorship. A. platanoides departs from the general trade-off pattern that exists among native species and A. altissima, and consequently it can be highly invasive in closed-canopy forests.

1. The Janzen-Connell (J-C) Model proposes that host-specific enemies could maintain high tree species diversity by reducing seedling performance near conspecific adults. An implicit, but untested assumption of the J-C Model is that negative conspecific feedbacks would promote replacement by heterospecific seedlings. 2. In a glasshouse experiment, we tested plant-soil feedbacks as a J-C mechanism in four temperate tree species. We assessed effects of conspecific- relative to heterospecific-cultured soil extracts on seedling survival, total mass and performance for each focal species. To test the implicit assumption of replacement by heterospecific seedlings, we also compared relative performance of conspecific versus heterospecific seedlings grown with soil extract cultured by a particular tree species. We also tested whether soil microbes caused these plant-soil feedbacks and whether low irradiance increased seedling vulnerability to pathogens. 3. When grown with conspecific versus heterospecific soil extract, Acer rubrum mass decreased, Quercus rubra mass increased and Fraxinus americana increased survival. Conspecific extract reduced Acer saccharum mass in low light but increased it in high light. To integrate survival and growth, we examined seedling performance [(mean total mass x mean survival time)/(days of experiment)] at low and high light. In conspecific versus heterospecific soil extract, seedling performance was lower in two, higher in four and neutral in 18 of 24 cases, suggesting no advantage to dispersing away from conspecifics. Based on relative seedling performance within a soil extract, conspecific seedlings were disadvantaged in two, favoured in three and neutral in 19 of 24 cases relative to heterospecific seedlings. 4. Species pairwise interactions of soil modification and seedling performance were chemically mediated, occurring regardless of sterilization. Microbes lacked host specificity and reduced performance regardless of extract source. Additionally, microbial factors reduced seedling performance for Q. rubra regardless of light availability, and for A. rubrum and F. americana only in high light. 5. Synthesis. These chemical-mediated plant-soil feedbacks probably influence community dynamics, but are inconsistent with the J-C Model. Even when a species' seedlings responded more negatively to conspecific than heterospecific soil, heterospecific seedlings were not necessarily favoured in that species' soil, precluding heterospecific replacement as an explanation for coexistence.

In closed-canopy forests, gap formation and closure are thought to be major drivers of forest dynamics. Crown defoliation by insects, however, may also influence understory resource levels and thus forest dynamics. We evaluate the effect of a forest tent caterpillar outbreak on understory light availability, soil nutrient levels and tree seedling height growth in six sites with contrasting levels of canopy defoliation in a hardwood forest in northern lower Michigan. We compared resource levels and seedling growth of six hardwood species before, during and in the three years after the outbreak (2008-2012). Canopy openness increased strongly during the forest tent caterpillar outbreak in the four moderately and severely defoliated sites, but not in lightly defoliated sites. Total inorganic soil nitrogen concentrations increased in response to the outbreak in moderately and severely defoliated sites. The increase in total inorganic soil nitrogen was driven by a strong increase in soil nitrate, and tended to become stronger with increasing site defoliation. Seedling height growth increased for all species in the moderately and severely defoliated sites, but not in lightly defoliated sites, either during the outbreak year or in the year after the outbreak. Growth increases did not become stronger with increasing site defoliation, but were strongest in a moderately defoliated site with high soil nutrient levels. Growth increases tended to be strongest for the shade intolerant species Fraxinus americana and Prunus serotina, and the shade tolerant species Ostrya virginiana. The strong growth response of F. americana and P. serotina suggests that recurring forest tent caterpillar outbreaks may facilitate the persistence of shade intolerant species in the understory in the absence of canopy gaps. Overall, our results suggest that recurrent canopy defoliation resulting from cyclical forest insect outbreaks may be an additional driver of dynamics in temperate closed-canopy forests.

Plant–soil feedbacks (PSFs) are often involved in fundamental ecological processes such as plant succession and species coexistence. After a plant initiating PSFs dies, legacies of PSFs occurring as soil signatures that influence subsequent plants could persist for an unknown duration. Altered resource environments following plant death (especially light availability) could affect whether legacy effects manifest and persist. To evaluate PSFs and their legacies, we obtained soils from a chronosequence of Prunus serotina harvests. In a greenhouse experiment, we planted conspecific seedlings under two light levels in these soils of varying time since the influence of live Prunus serotina, and compared seed/seedling survival in soils from live trees, stumps, and surrounding forest matrix within each site and across the chronosequence. PSF legacies were measured as the difference between seedling performance in live tree and stump soils within a site. Negative PSF legacies of P. serotina were short-lived, lasting up to 0.5 years after tree removal. These effects occurred under 5% but not 30% full sun. PSFs and their legacies manifested in seed/seedling survival, but not biomass. Though restricted to low light, short-lived legacies of P. serotina PSFs could have lasting impacts on plant community dynamics during post-disturbance regeneration by disfavoring P. serotina regeneration in small tree-fall gaps.

1 Traditional rankings of shade tolerance of trees make little reference to individual size. However, greater respiratory loads with increasing sapling size imply that larger individuals will be less able to tolerate shade than smaller individuals of the same species and that there may be shifts among species in shade tolerance with size. 2 We tested this hypothesis using maximum likelihood estimation to develop individual-tree-based models of the probability of mortality as a function of recent growth rate for seven species: trembling aspen, paper birch, yellow birch, mountain maple, white spruce, balsam fir and eastern white cedar. 3 Shade tolerance of small individuals, as quantified by risk of mortality at low growth, was mostly consistent with traditional shade tolerance rankings such that cedar > balsam fir > white spruce > yellow birch > mountain maple = paper birch > aspen. 4 Differences in growth-dependent mortality were greatest between species in the smallest size classes. With increasing size, a reduced tolerance to shade was observed for all species except trembling aspen and thus species tended to converge in shade tolerance with size. At a given level of radial growth larger trees, apart from aspen, had a higher probability of mortality than smaller trees. 5 Successional processes associated with shade tolerance may thus be most important in the seedling stage and decrease with ontogeny.