Cone allometry and seed protection from fire are similar in serotinous and nonserotinous conifers.

Serotiny is an adaptive trait that allows certain woody plants to persist in stand-replacing fire regimes. However, the mechanisms by which serotinous cones avoid seed necrosis and nonserotinous species persist in landscapes with short fire cycles and serotinous competitors remain poorly understood. To investigate whether ovulate cone traits that enhance seed survival differ between serotinous and nonserotinous species, we examined cone traits in 24 species within Pinaceae and Cupressaceae based on physical measurements and cone heating simulations using a computational fluid dynamics model. Fire-relevant cone traits were largely similar between cone types; those that differed (e.g. density and moisture) conferred little seed survival advantage under simulated fire. The most important traits influencing seed survival were cone size and seed depth within the cone, which was found to be an allometric function of cone mass for both cone types. Thus, nonserotinous cones should not suffer significantly greater seed necrosis than serotinous cones of equal size. Closed nonserotinous cones containing mature seeds may achieve substantial regeneration after fire if they are sufficiently large relative to fire duration and temperature. To our knowledge, this is the most comprehensive study of the effects of fire-relevant cone traits on conifer regeneration supported by physics-based fire simulation.

MASTREE+: Time-series of plant reproductive effort from six continents.

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics.

Aún existen importantes vacíos en la comprensión de la respuesta reproductiva de las plantas al cambio medioambiental, en parte, porque su monitoreo en especies de plantas longevas requiere una observación directa durante muchos años, y estos conjuntos de datos rara vez han estado disponibles. Aquí presentamos a MASTREE +, una base de datos que recopila series de tiempo de la reproducción de las plantas de todo el planeta, poniendo a disposición estos datos de libre acceso para la comunidad científica. MASTREE + incluye 73.828 puntos de observación de la reproducción anual georreferenciados (ej. conteos de semillas y frutos) en poblaciones de plantas perennes en todo el mundo. Estas observaciones consisten en 5971 series temporales a nivel de población provenientes de 974 especies en 66 países. La mediana de la duración de las series de tiempo es de 10 años (media = 12.4 años) y el conjunto de datos incluye 1.122 series de al menos dos décadas (≥20 años de observaciones). Para un subconjunto de especies bien estudiadas, MASTREE +incluye un amplio conjunto de series temporales replicadas en gradientes geográficos y climáticos. Describimos el conjunto de datos de acceso abierto disponible como un archivo.csv y presentamos una aplicación web asociada para la exploración de datos. MASTREE+ proporcionará la base para mejorar la comprensión sobre la respuesta reproductiva de plantas longevas al cambio medioambiental. Además, MASTREE+ facilitará los avances en la investigación de la ecología y la evolución de las estrategias reproductivas en plantas perennes y el papel de la reproducción vegetal como determinante de la dinámica de ecosistemas.

Strong dispersal limitation in postfire regeneration of Baker cypress, a rare serotinous conifer.

PREMISE: Dispersal capacity primarily determines the spatial establishment patterns that drive range expansions and contractions in tree species. Seedling establishment in Baker cypress (Hesperocyparis bakeri [(Jeps.) Bartel]) relies predominantly on fire events, due to its cone serotiny, shade intolerance, and small seeds that require the optimal conditions of fire-exposed, mineral soil seedbeds. METHODS: We quantified the density and spatial distribution of post-disturbance seedlings following the 2014 Eiler Fire in northern California and compared the observed recruitment to predictions from a mechanistic seed dispersal model. RESULTS: Postfire recruitment was dense, averaging 11 seedlings/m2 , and occurred primarily in the first year after fire. We estimated the mean descent velocity of the wingless seeds as ~4 m/s, the highest value reported for any putatively wind-dispersed tree species. The rapid seed descent contributed to markedly spatially constrained recruitment. Most seedlings (~81%) established within 5 m of the parent tree, and 94% established within 10 m. The maximum observed dispersal distance was 48.5 m; dispersal distance scaled linearly with canopy height. Distributions of modeled seed dispersal distance and observed seedling establishment in Baker cypress did not differ, demonstrating that wind disperses seeds a short distance because of the lack of a wing, and secondary dispersal appeared to be minimal at this recently burned site. CONCLUSIONS: If seed dispersal is solely reliant on wind, migration in response to rapid climate change will be impeded and potentially present difficulties in sustaining populations of this and other obligate seeder species with equally constrained dispersal.

Linking ice accretion and crown structure: towards a model of the effect of freezing rain on tree canopies.

BACKGROUND AND AIMS: Despite a longstanding interest in variation in tree species vulnerability to ice storm damage, quantitative analyses of the influence of crown structure on within-crown variation in ice accretion are rare. In particular, the effect of prior interception by higher branches on lower branch accumulation remains unstudied. The aim of this study was to test the hypothesis that intra-crown ice accretion can be predicted by a measure of the degree of sheltering by neighbouring branches. METHODS: Freezing rain was artificially applied to Acer platanoides L., and in situ branch-ice thickness was measured directly and from LiDAR point clouds. Two models of freezing rain interception were developed: 'IceCube', which uses point clouds to relate ice accretion to a voxel-based index (sheltering factor; SF) of the sheltering effect of branch elements above a measurement point; and 'IceTree', a simulation model for in silico evaluation of the interception pattern of freezing rain in virtual tree crowns. KEY RESULTS: Intra-crown radial ice accretion varied strongly, declining from the tips to the bases of branches and from the top to the base of the crown. SF for branches varied strongly within the crown, and differences among branches were consistent for a range of model parameters. Intra-crown variation in ice accretion on branches was related to SF (R(2) = 0·46), with in silico results from IceTree supporting empirical relationships from IceCube. CONCLUSIONS: Empirical results and simulations confirmed a key role for crown architecture in determining intra-crown patterns of ice accretion. As suspected, the concentration of freezing rain droplets is attenuated by passage through the upper crown, and thus higher branches accumulate more ice than lower branches. This is the first step in developing a model that can provide a quantitative basis for investigating intra-crown and inter-specific variation in freezing rain damage.

A field test of inverse modeling of seed dispersal.

PREMISE OF THE STUDY: Seed dispersal distance-a key process in plant population dynamics-remains poorly understood because of the difficulty of finding a source plant so well isolated from conspecifics that seeds or seedlings can be unambiguously attributed to it. Inverse modeling (IM) of seed dispersal, a simple statistical technique for parameterizing dispersal kernels, has been widely used since 1992; surprisingly, however, this approach has never been verified in the field. METHODS: We released from 20 nearby trees the winged seeds of a liana species, Entada polystachya, near the coast in a tropical, dry forest in Jalisco, Mexico. KEY RESULTS: With a two-parameter log-normal function, we found that IM predicted both the shape and scale parameters well as long as we used the entire data set. When, however, we subsampled (thus simulating the use of transects for seedlings or an array of seed traps), the estimates of the scale and shape parameters were often more than double the real values. The problem was due to the marked anisotropy (directional bias; in this case, in the direction of the diurnal sea breeze) of the individual dispersal curves. When we randomized the direction of dispersal of individual seeds from the trees (keeping dispersal distances unchanged), predictions of parameter values were excellent. CONCLUSIONS: Inverse modeling must include directional parameters when dealing with areas where strong anisotropy is to be expected, e.g., for wind dispersal of seeds near coasts or pollination by any vector where a plant species is limited to a strongly linear habitat such as river banks.

An assessment of temporal variability in mast seeding of North American Pinaceae.

Our overall objective is to synthesize mast-seeding data on North American Pinaceae to detect characteristic features of reproduction (i.e. development cycle length, serotiny, dispersal agents), and test for patterns in temporal variation based on weather variables. We use a large dataset (n = 286 time series; mean length = 18.9 years) on crop sizes in four conifer genera (Abies, Picea, Pinus, Tsuga) collected between 1960 and 2014. Temporal variability in mast seeding (CVp) for 2 year genera (Abies, Picea, Tsuga) was higher than for Pinus (3 year), and serotinous species had lower CVp than non-serotinous species; there were no relationships of CVp with elevation or latitude. There was no difference in family-wide CVp across four tree regions of North America. Across all genera, July temperature differences between bud initiation and the prior year (ΔT) was more strongly associated with reproduction than absolute temperature. Both CVp and ΔT remained steady over time, while absolute temperature increased by 0.09°C per decade. Our use of the ΔT model included a modification for Pinus, which initiates cone primordia 2 years before seedfall, as opposed to 1 year. These findings have implications for how mast-seeding patterns may change with future increases in temperature, and the adaptive benefits of mast seeding. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Emergence of morel (Morchella) and pixie cup (geopyxis carbonaria) ascocarps in response to the intensity of forest floor combustion during a wildfire.

We studied the density of ascocarps (mushrooms) of morels (Morchella) and pixie cups (Geo-pyxis carbonaria) as a function of postfire duff (forest floor organic layer) depth in the first 4 y after a wildfire. The great majority of ascocarps of both species appeared in the first summer (2004) after an Aug 2003 fire in predominantly pine-spruce montane stands in Kootenay National Park, British Columbia. The spatial distribution of the ascocarps of both species was strongly biased toward (i) microsites with thin postfire duff and (ii) proximity to standing burned tree trunks. The bases of ascocarps of both species invariably were found just below the surface of the mineral soil. A field experiment in nearby intact forest showed that complete or partial duff removal in the absence of damage to the roots or crown did not lead to ascocarps of either species. We conclude that for both fungal species an unusually large abundance of ascocarps simultaneously requires damage to the associated trees and major duff reduction.

Mast seeding patterns are asynchronous at a continental scale.

Resource pulses are rare events with a short duration and high magnitude that drive the dynamics of both plant and animal populations and communities1. Mast seeding is perhaps the most common type of resource pulse that occurs in terrestrial ecosystems2, is characterized by the synchronous and highly variable production of seed crops by a population of perennial plants3,4, is widespread both taxonomically and geographically5, and is often associated with nutrient scarcity6. The rare production of abundant seed crops (mast events) that are orders of magnitude greater than crops during low seed years leads to high reproductive success in seed consumers and has cascading impacts in ecosystems2,7. Although it has been suggested that mast seeding is potentially synchronized at continental scales8, studies are largely constrained to local areas covering tens to hundreds of kilometres. Furthermore, summer temperature, which acts as a cue for mast seeding9, shows patterns at continental scales manifested as a juxtaposition of positive and negative anomalies that have been linked to irruptive movements of boreal seed-eating birds10,11. Here, we show a breakdown in synchrony of mast seeding patterns across space, leading to asynchrony at the continental scale. In an analysis of synchrony for a transcontinental North America tree species spanning distances of greater than 5,200 km, we found that mast seeding patterns were significantly asynchronous at distances of greater than 2,000 km apart (all P < 0.05). Other studies have shown declines in synchrony across distance, but not asynchrony. Spatiotemporal variation in summer temperatures at the continental scale drives patterns of synchrony in mast seeding, and we anticipate that this affects the spatial dynamics of numerous seed-eating communities, from insects to small mammals to the large-scale migration patterns of boreal seed-eating birds.

Non-serotinous woody plants behave as aerial seed bank species when a late-summer wildfire coincides with a mast year.

ABSTRACT: Trees which lack obvious fire-adaptive traits such as serotinous seed-bearing structures or vegetative resprouting are assumed to be at a dramatic disadvantage in recolonization via sexual recruitment after fire, because seed dispersal is invariably quite constrained. We propose an alternative strategy in masting tree species with woody cones or cone-like structures: that the large clusters of woody tissue in a mast year will sufficiently impede heat transfer that a small fraction of seeds can survive the flaming front passage; in a mast year, this small fraction would be a very large absolute number.In Kootenay National Park in British Columbia, we examined regeneration by Engelmann spruce (Picea engelmannii), a non-serotinous conifer, after two fires, both of which coincided with mast years. Coupling models of seed survivorship within cones and seed maturation schedule to a spatially realistic recruitment model, we show that (1) the spatial pattern of seedlings on a 630 m transect from the forest edge into the burn was best explained if there was in situ seed dissemination by burnt trees; (2) in areas several hundred meters from any living trees, recruitment density was well correlated with local prefire cone density; and (3) spruce was responding exactly like its serotinous codominant, lodgepole pine (Pinus contorta).We conclude that non-serotinous species can indeed behave like aerial seed bank species in mast years if the fire takes place late in the seed maturation period. Using the example of the circumpolar boreal forest, while the joint probability of a mast year and a late-season fire will make this type of event rare (we estimate P = 0.1), nonetheless, it would permit a species lacking obvious fire-adapted traits to occasionally establish a widespread and abundant cohort on a large part of the landscape.

Turbulence-induced resonance vibrations cause pollen release in wind-pollinated Plantago lanceolata L. (Plantaginaceae).

In wind pollination, the release of pollen from anthers into airflows determines the quantity and timing of pollen available for pollination. Despite the ecological and evolutionary importance of pollen release, wind-stamen interactions are poorly understood, as are the specific forces that deliver pollen grains into airflows. We present empirical evidence that atmospheric turbulence acts directly on stamens in the cosmopolitan, wind-pollinated weed, Plantago lanceolata, causing resonant vibrations that release episodic bursts of pollen grains. In laboratory experiments, we show that stamens have mechanical properties corresponding to theoretically predicted ranges for turbulence-driven resonant vibrations. The mechanical excitation of stamens at their characteristic resonance frequency caused them to resonate, shedding pollen vigorously. The characteristic natural frequency of the stamens increased over time with each shedding episode due to the reduction in anther mass, which increased the mechanical energy required to trigger subsequent episodes. Field observations of a natural population under turbulent wind conditions were consistent with these laboratory results and demonstrated that pollen is released from resonating stamens excited by small eddies whose turnover periods are similar to the characteristic resonance frequency measured in the laboratory. Turbulence-driven vibration of stamens at resonance may be a primary mechanism for pollen shedding in wind-pollinated angiosperms. The capacity to release pollen in wind can be viewed as a primary factor distinguishing animal- from wind-pollinated plants, and selection on traits such as the damping ratio and flexural rigidity may be of consequence in evolutionary transitions between pollination systems.

Seed size, dispersal, and aerodynamic constraints within the Bombacaceae.

The aerodynamic constraints operating on the wind-dispersed, drag-producing diaspores of several species of the tropical family Bombacaceae were examined. Kapok (the drag-promoting appendage) was best characterized as a moderately flattened hemisphere impervious to air movement. The kapok shape was not isometric: kapok planform area was proportional to the kapok mass raised to the power 0.52 rather than to the 0.67 expected from isometry. Thus, necessarily, terminal velocity rises with seed mass much faster in this group than among taxa with winged seeds. Further, we derived the optimality argument to show that the kapok mass ought to be about 50% of the total diaspore mass (seed plus kapok). While seven of eight species had a lower kapok investment than this, and none were especially close to the theoretically optimal value, nonetheless the kapok investment values were hardly draws from a random distribution. Finally, the kapok fibers of these Bombacaceae species begin to bend at a drag of about 0.005 N, and this sets an upper limit on the efficient diaspore size of about 250 mg for the seed mass. This latter value is similar to the mass of the largest seed we know of in this family.