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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'.

The encapsulation of seeds in hard coats and fruit walls (pericarp layers) fulfils protective and dispersal functions in many plant families. In angiosperms, packaging structures possess a remarkable range of different morphologies and functionalities, as illustrated by thermo and hygro-responsive seed pods and appendages, as well as mechanically strong and water-impermeable shells. Key to these different functionalities are characteristic structural arrangements and chemical modifications of the underlying sclerenchymatous tissues. Although many ecological aspects of hard seed encapsulation have been well documented, a detailed understanding of the relationship between tissue structure and function only recently started to emerge, especially in the context of environmentally driven fruit opening and seed dispersal (responsive encapsulations) and the outstanding durability of some seed coats and indehiscent fruits (static encapsulations). In this review, we focus on the tissue properties of these two systems, with particular consideration of water interactions, mechanical resistance, and force generation. Common principles, as well as unique adaptations, are discussed in different plant species. Understanding how plants integrate a broad range of functions and properties for seed protection during storage and dispersal plays a central role for seed conservation, population dynamics, and plant-based material developments.

1. Changing disturbance–climate interactions will drive shifts in plant communities: these effects are not adequately quantified by environmental niche models used to predict future species distributions. We quantified the effects of more frequent fire and lower rainfall – as projected to occur under a warming and drying climate – on population responses of shrub species in biodiverse Mediterranean-climate type shrublands near Eneabba, southwestern Australia. 2. Using experimental fires, we measured the density of all shrub species for four dominant plant functional groups (resprouter/non-sprouter × serotinous/soil seed bank) before and after fire in 33 shrubland sites, covering four post-fire rainfall years and fire intervals from 3–24 years. 3. Generalized linear mixed effects models were used to test our a priori hypotheses of rainfall, fire interval and plant functional type effects on post-fire survival and recruitment. 4. At shortened fire intervals, species solely dependent on seedling recruitment for persistence were more vulnerable to local extinction than were species with both seedling recruitment and vegetative regrowth. Nevertheless, seedling recruitment was essential for population maintenance of resprouting species. Serotinous species were less resilient than soil seed storage species regardless of regeneration mode. Critically, in relation to changing climate, a 20% reduction in post-fire winter rainfall (essential for seedling recruitment) is predicted to increase the minimum inter-fire interval required for self-replacement by 50%, placing many species at risk of decline. 5. Synthesis. Our results highlight the potentially deleterious biodiversity impacts of climate and fire regime change, and underscore weaknesses inherent in studies considering single impact factors in isolation. In fire-prone ecosystems characterized by a projected warming and drying climate, and increasing fire hazard, adaptive approaches to fire management may need to include heightened wildfire suppression and lengthened intervals for prescribed fire to best support the in situ persistence of perennial plant species and of plant biodiversity. This conclusion is at odds with the view that more managed fire may be needed to mitigate wildfire risk as climate warms.

The mapping of functional traits onto chronograms is an emerging approach for the identification of how agents of natural selection have shaped the evolution of organisms. Recent research has reported fire-dependent traits appearing among flowering plants from 60 million yr ago (Ma). Although there are many records of fossil charcoal in the Cretaceous (65–145 Ma), evidence of fire-dependent traits evolving in that period is lacking. We link the evolutionary trajectories for five fire-adapted traits in Pinaceae with paleoatmospheric conditions over the last 250 million yr to determine the time at which fire originated as a selective force in trait evolution among seed plants. Fire-protective thick bark originated in Pinus c. 126 Ma in association with low-intensity surface fires. More intense crown fires emerged c. 89 Ma coincident with thicker bark and branch shedding, or serotiny with branch retention as an alternative strategy. These innovations appeared at the same time as the Earth's paleoatmosphere experienced elevated oxygen levels that led to high burn probabilities during the mid-Cretaceous. The fiery environments of the Cretaceous strongly influenced trait evolution in Pinus. Our evidence for a strong correlation between the evolution of fire-response strategies and changes in fire regime 90–125 Ma greatly backdates the key role that fire has played in the evolution of seed plants.

Climatic change is anticipated to alter disturbance regimes for many ecosystems. Among the most important effects are changes in the frequency, size, and intensity of wildfires. Serotiny (long-term canopy storage and the heat-induced release of seeds) is a fire-resilience mechanism found in many globally important terrestrial ecosystems. Life-history traits and physiographic differences in ecosystems lead to variation in serotiny; therefore, some systems may exhibit greater resilience to shifting disturbances than others do. We present a conceptual framework to explore the consequences of changing disturbance regimes (such as mean and variance in fire severity or return intervals) to serotinous species and ecosystems and implications of altered serotinous resilience at local and regional scales. Four case studies are presented, and areas needing further research are highlighted. These studies illustrate that, despite the reputed fire resilience of serotiny, more fire does not necessarily mean more serotinous species across all systems in which they occur.

BackgroundMediterranean-type ecosystems (MTEs) are experiencing declining rainfall, increasing temperature, and shifting fire regimes as climate changes. While changes in fire regimes and post-fire recruitment are widely reported, evidence for changing plant demographic rates is limited.AimsWe hypothesised increased time to maturity and decreased serotinous seed stores available for post-fire recruitment due to declining rainfall over recent decades for two fire-killed serotinous shrubs of south-east Australian MTEs, Hakea decurrens and Banksia ornata.MethodsFruit and cone production for populations across time since fire chronosequences were measured in the same regions in the 1990s and in 2017.Key resultsEstimated time to 50% maturity increased from 3–15 years and 6–15 years for H. decurrens and B. ornata, respectively, while estimated canopy seed stores were 90% and 50% lower in 2017 than in the 1990s.ConclusionsDelayed reproductive maturity and decreased total seed stores were significantly related to decreasing rainfall received by 2017 populations over their lifetimes (5–17% less than for stands in the 1990s).ImplicationsShifts in inter-fire rates of seed production and storage, combined with changes to fire regimes and post-fire recruitment conditions due to climate change may already threaten the persistence of some species.

Current phylogenetic evidence shows that fire began shaping the evolution of land plants 125 Ma, although the fossil charcoal record indicates that fire has a much longer history (>350 Ma). Serotiny (on‐plant seed storage) is generally accepted as an adaptation to fire among woody plants. We developed a conceptual model of the requirements for the evolution of serotiny, and propose that serotiny is only expressed in the presence of a woody rachis as supporting structure, compact scales covering seeds as protective structure, seed wing as dispersal structure, and crown fire as the agent of selection and mechanism for seed release. This model is strongly supported by empirical data for modern ecosystems. We reconstructed the evolutionary history of intrinsic structural states required for the expression of serotiny in conifers, and show that these were diagnostic for early (‘transitional’) conifers from 332 Ma (late‐Carboniferous). We assessed the likely flammable characteristics of early conifers and found that scale‐leaved conifers burn rapidly and with high intensity, supporting the idea that crown fire regimes may have dominated early conifer ecosystems. Synthesis. Coupled with strong evidence for frequent fire throughout the Permian‐Carboniferous and fossil evidence for other fire‐related traits, we conclude that many early conifers were serotinous in response to intense crown fires, indicating that fire may have had a major impact on the evolution of plant traits as far back as 350 Ma.

Widespread tree mortality caused by outbreaks of native bark beetles (Circulionidae: Scolytinae) in recent decades has raised concern among scientists and forest managers about whether beetle outbreaks fuel more ecologically severe forest fires and impair postfire resilience. To investigate this question, we collected extensive field data following multiple fires that burned subalpine forests in 2011 throughout the Northern Rocky Mountains across a spectrum of prefire beetle outbreak severity, primarily from mountain pine beetle ( Dendroctonus ponderosae ). We found that recent (2001–2010) beetle outbreak severity was unrelated to most field measures of subsequent fire severity, which was instead driven primarily by extreme burning conditions (weather) and topography. In the red stage (0–2 y following beetle outbreak), fire severity was largely unaffected by prefire outbreak severity with few effects detected only under extreme burning conditions. In the gray stage (3–10 y following beetle outbreak), fire severity was largely unaffected by prefire outbreak severity under moderate conditions, but several measures related to surface fire severity increased with outbreak severity under extreme conditions. Initial postfire tree regeneration of the primary beetle host tree [lodgepole pine ( Pinus contorta var. latifolia )] was not directly affected by prefire outbreak severity but was instead driven by the presence of a canopy seedbank and by fire severity. Recent beetle outbreaks in subalpine forests affected few measures of wildfire severity and did not hinder the ability of lodgepole pine forests to regenerate after fire, suggesting that resilience in subalpine forests is not necessarily impaired by recent mountain pine beetle outbreaks. Significance Understanding how multiple disturbances may interact to affect ecosystems is important for ecosystem management as climate-driven disturbance activity increases. Recent severe bark beetle (Circulionidae: Scolytinae) outbreaks have led to widespread concern about the potential for increased wildfire severity and decreased postfire forest resilience throughout the northern hemisphere. Using extensive field data collected in multiple recent (occurring in 2011) wildfires throughout the Northern Rocky Mountains (United States), we found that recent (2001–2010) prefire mountain pine beetle (Dendroctonus ponderosae ) outbreak severity affected few measures of wildfire severity and was not directly related to postfire tree seedling establishment, suggesting that subalpine forests dominated by serotinous lodgepole pine ( Pinus contorta var. latifolia ) may be resilient to these two combined disturbances.

Impermeable seed/fruit coats, i.e., physical dormancy (PY), occurs only in several genera belonging to 19 angiosperm families. However, recent reviews on fire ecology have proposed including serotiny under PY and dormancy-broken seeds waiting for germination as ‘imposed secondary dormancy’. Serotiny occurs in 12 angiosperm and gymnosperm families, mutually exclusive from PY families, suggesting no linkage between these two traits. Imposed secondary dormancy essentially refers to the non-dormant seeds waiting for germination, as no dormancy-breaking treatments are required for germination. Understanding the evolutionary and ecological significance of PY is imperative, one of the first steps in this direction is to correctly define the seed traits. In contemporary species, the ability of PY to develop impermeable or permeable coats is under the influence of climate.