Germination Phenology Research Articles

Macroclimatic Convergence and Habitat Specialisation Shape the Mediterranean Seed Germination Syndrome.

Ecological theory predicts that plant reproductive phenology in the Mediterranean regions is shaped by evolutionary processes driven by strong seasonality in precipitation-evaporation patterns. Thus, it can be expected that seed germination phenology has adapted to maximise recruitment during the season of highest water availability. Cold-cued and slow germination (i.e., the 'Mediterranean seed germination syndrome') has been hypothesised to be an adaptation to ensure that seedling emergence occurs in autumn/early winter, extending the growing season before the subsequent unfavourable summer drought. However, this hypothesis has been tested on individual species or local studies, without a proper synthesis for the whole Mediterranean region. Here we tested, for the first time, the Mediterranean seed germination syndrome using experimental data for 459 species (11,363 records, 59 families) occurring in the Mediterranean Basin. We performed a phylogenetically informed Bayesian meta-analysis to model the effect on germination proportions of seven key experimental cues: mean incubation temperature, alternating temperature regime, light and dormancy-breaking treatments (scarification, warm stratification and cold stratification) and the modulating role of seed mass on seed germination. We show that species from lowland zonal habitats of the Mediterranean align with the Mediterranean germination syndrome hypothesis, with their seeds responding positively to cool, constant temperatures and negatively to light. Yet, habitat specialists (i.e., species restricted to mountains, coasts and wetlands) deviate from the syndrome, showing nearly opposite germination requirements. Seed mass further influences the germination niche and phylogenetically related species exhibit similar germination responses. Cumulatively, these results suggest that evolutionary pressures from local habitat-related conditions override the macroclimatically imposed Mediterranean seed germination syndrome.

Germination phenology alters species coexistence outcomes

Abstract Species‐specific phenological responses to changing climate are reshuffling the timing of species interactions, however we do not fully understand the consequences of these changes for species' population dynamics and community composition. In this study, we experimentally manipulated the timing of germination for five annual plant species from southern California and used pairwise competition experiments and coexistence theory to quantify how phenological shifts may impact species interactions and coexistence. We found that phenological shifts may help promote coexistence when they confer an advantage for competitively inferior species, but in other cases promote dominance by competitively superior species. Earlier germination generally increased species' performance relative to competitors, but the relative changes in intra‐and inter‐specific interactions caused more complex effects on niche and fitness differences. Phenological differences tended to reduce stabilising niche differences for many species pairs and reduced overall coexistence probabilities. Synthesis. While phenological differences among species have typically been considered a form of niche partitioning, it seems increasingly likely that phenological offsets could destabilise species coexistence. The net effects of changing phenology on species coexistence will depend on the complex combinations of effects on intra‐ and inter‐specific interactions, which remain challenging to predict.

Time-dependent interaction modification generated from plant-soil feedback.

Pairwise interactions between species can be modified by other community members, leading to emergent dynamics contingent on community composition. Despite the prevalence of such higher-order interactions, little is known about how they are linked to the timing and order of species' arrival. We generate population dynamics from a mechanistic plant-soil feedback model, then apply a general theoretical framework to show that the modification of a pairwise interaction by a third plant depends on its germination phenology. These time-dependent interaction modifications emerge from concurrent changes in plant and microbe populations and are strengthened by higher overlap between plants' associated microbiomes. The interaction between this overlap and the specificity of microbiomes further determines plant coexistence. Our framework is widely applicable to mechanisms in other systems from which similar time-dependent interaction modifications can emerge, highlighting the need to integrate temporal shifts of species interactions to predict the emergent dynamics of natural communities.

Functional Attributes of Seeds as Indicators of Germination Sensitivity to Global Warming

Climate change has profound impacts on ecosystems, and one critical aspect is its effect on seed germination, a crucial stage in plant life cycles. Various studies have explored the responses of plant species to rising temperatures, and there is now a pressing need to integrate this wealth of information into a coherent framework. The aim of this study was to survey literature about seed traits and thence to evaluate germination responses to climate change and to propose functional groups for germination. Increased temperature affects seed traits, particularly germinability. Many species show increased germination percentages in warmer temperatures, although the extent varies among species and temperature ranges. Some maintain a consistent percentage, whereas others reduce it to retain seeds in the seed bank. Temperature changes also affect the timing and season of germination, with some species accelerating germination, others delaying it, thereby influencing competition and exposure to adverse conditions. Shifts in temperature can alter seed requirements, affecting responses to temperature, humidity, light, chemical stimuli, and dormancy. Modifications in germination have profound effects on seed bank and seedling bank dynamics, affecting plant populations and ecological community resilience. Changes in germination can disrupt competitive dynamics, favoring some over others, altering community composition and potentially impairing ecosystem functionality. Germination Niche, Germination Potential, and Germination Phenology are fundamental concepts in the evaluation of climate change's implications for germination.

A long-term experiment reveals no trade-off between seed persistence and seedling emergence.

Information on seed persistence and seedling emergence from the soil seed bank is critical for understanding species coexistence and predicting community dynamics. However, quantifying seed persistence in the soil is challenging; thus, its association with other life-history traits is poorly known on a broad scale. Using germination phenology for 349 species in a 42-yr experiment, we quantified the persistence-emergence correlations and their associations with intrinsic regeneration traits using Bayesian phylogenetic multilevel models. We showed no trade-off between seed persistence and seedling emergence. Physically dormant seeds were more persistent but exhibited lower emergence than nondormant seeds. Monocarpic species had both higher persistence and emergence than polycarpic species. Seed mass posed a marginal proxy for persistence, while emergence almost doubled from the smallest to the largest seeds. This study challenges the traditional assumption and is the first demonstration of noncorrelation between persistence and emergence, probably owing to the complexity of regenerative strategies. Species with short persistence and low emergence would be the most vulnerable for insitu conservation. Our analyses of this unique, long-term dataset provide a strong incentive for further experimental studies and a rich data resource for future syntheses.

Precipitation timing and soil substrate drive phenology and fitness of Arabidopsis thaliana in a Mediterranean environment

Abstract In Mediterranean climates, the timing of seasonal rains determines germination, flowering phenology and fitness. As climate change alters seasonal precipitation patterns, it is important to ask how these changes will affect the phenology and fitness of plant populations. We addressed this question experimentally with the annual plant species Arabidopsis thaliana. In a first experiment, we manipulated the date of rainfall onset and recorded germination phenology on sand and soil substrates. In a second experiment, we manipulated germination date, growing season length and mid‐season drought to measure their effects on flowering time and fitness. Within each experiment, we manipulated seed dormancy and flowering time using multilocus near‐isogenic lines segregating strong and weak alleles of the seed dormancy gene DOG1 and the flowering time gene FRI. We synthesized germination phenology data from the first experiment with fitness functions from the second experiment to project population fitness under different seasonal rainfall scenarios. Germination phenology tracked rainfall onset but was slower and more variable on sand than on soil. Many seeds dispersed on sand in spring and summer delayed germination until the cooler temperatures of autumn. The high‐dormancy DOG1 allele also prevented immediate germination in spring and summer. Germination timing strongly affected plant fitness. Fecundity was highest in the October germination cohort and declined in spring germinants. The late flowering FRI allele had lower fecundity, especially in early fall and spring cohorts. Projections of population fitness revealed that: (1) Later onset of autumn rains will negatively affect population fitness. (2) Slow, variable germination on sand buffers populations against fitness impacts of variable spring and summer rainfall. (3) Seasonal selection favours high dormancy and early flowering genotypes in a Mediterranean climate with hot dry summers. The high‐dormancy DOG1 allele delayed germination of spring‐dispersed fresh seeds until more favourable early fall conditions, resulting in higher projected population fitness. These findings suggest that Mediterranean annual plant populations are vulnerable to changes in seasonal precipitation, especially in California where rainfall onset is already occurring later. The fitness advantage of highly dormant, early flowering genotypes helps explain the prevalence of this strategy in Mediterranean populations. Read the free Plain Language Summary for this article on the Journal blog.

Contrasting responses to climate variability generate seasonal priority effects between native and invasive forest herbs

Abstract Invasive plants are often characterized by rapid germination and precocious phenology. Theory suggests that early germination may provide invaders with a competitive advantage over slower germinating natives, but the relative contribution of rapid germination versus other intrinsic competitive traits (e.g. rapid growth, high fecundity, broad environmental tolerance) to the success of invaders is poorly understood. Depending on the relationship between germination and competition, shifts in germination phenology due to climate change may increase the dominance of invaders or buffer communities against their impacts. We investigated the link between temperature variation, germination phenology and competitive interactions with a sequence of controlled environment experiments. First, we evaluated the relationships between temperature variation and germination phenology for two herbaceous species found in North American woodlands, the non‐native Hesperis matronalis and native Cryptotaenia canadensis. We then leveraged temperature‐response differences to manipulate the relative germination phenology of these taxa and quantified the effects of their phenological differences on competition. Seeds of the invasive H. matronalis germinated rapidly, reaching 50% germination in under 10 days in all treatment combinations. C. canadensis did not reach 50% germination with less than 7 weeks of cold stratification. However, with more than 10 weeks of cold stratification and low (20/10°C) incubation temperatures, the germination phenology of C. canadensis was well‐matched to that of H. matronalis. When grown together, we found that precocious germination phenology doubled the competitive impact of H. matronalis relative to its other intrinsic competitive traits. Phenological advantage of just 2–3 days relative to C. canadensis was enough to secure competitive dominance at the seedling stage. Synthesis. This study revealed that the mechanistic link between the germination phenology and competitive success of an invasive plant can be strongly mediated by climate sensitivity differences between introduced and native species. Climate change will likely exacerbate these differences, especially in regions where warming reduces cold stratification. Our findings suggest that phenological diversity in native plant communities is an important property of invasion resistance. The relationship between environmental variation, germination dynamics and competition provides a path forward for forecasting climate change impacts on seasonal community assembly and highlights the need to incorporate phenological diversity in restorations.

Seed Dormancy and Germination Requirements of Torilis scabra (Apiaceae)

The timing of seed germination significantly affects the fitness and life cycle of plants. Torilis scabra is a perennial medicinal herb occurring in mixed forests but the increasing use and modification of forestlands in recent decades has led to the degeneration of its natural habitat. Nonetheless, the requirements for germination in T. scabra remain unclear. The present study focused on identifying conditions necessary to break T. scabra seed dormancy and describing its seed dormancy type. By periodically collecting seeds that were sown in the field, germination phenology was studied. The impact of light, temperature, and warm/cold stratification on breaking seed dormancy and promoting germination was also determined through incubating seeds in laboratory conditions. Additionally, the effect of GA3 was explored to more accurately identify the type of dormancy present. The results demonstrated that the seeds of T. scabra possessed small, undeveloped embryos with physiological dormancy at the time of maturity. In the field, embryo growth initiated in early spring and the embryo–seed length ratio increased by ~300% before the radical emerged. In the laboratory, the embryo–seed length ratio increased from 0.24 to 0.82 when seeds were subjected to cold stratification at 4 °C and then transferred to 15/25 °C. Germination was observed across a broad temperature range after cold stratification. GA3 also helped to break dormancy but after-ripening did not. Taken together, the results suggest that seeds of T. scabra have non-deep simple morphophysiological dormancy.

Changes in phenology can alter patterns of natural selection: the joint evolution of germination time and postgermination traits.

The timing of a developmental transition (phenology) can influence the environment experienced by subsequent life stages. When phenology causes an organism to occupy a particular habitat as a consequence of the developmental cues used, it can act as a form of habitat tracking. Evolutionary theory predicts that habitat tracking can alter the strength, direction, and mode of natural selection on subsequently expressed traits. To test whether germination phenology altered natural selection on postgermination traits, we manipulated germination time by planting seedlings in seven germination cohorts spanning 2 yr. We measured selection on postgermination traits relating to drought, freezing, and heat tolerance using a diverse combination of Arabidopsis thaliana mutants and naturally occurring ecotypes. Germination cohorts experienced variable selection: when dry, cold, and hot environments were experienced by seedlings, selection was intensified for drought, freezing, and heat tolerance, respectively. Reciprocally, postgermination traits modified the optimal germination time; genotypes had maximum fitness after germinating in environments that matched their physiological tolerances. Our results support the theoretical predictions of feedbacks between habitat tracking and traits expressed after habitat selection. In natural populations, whether phenological shifts alter selection on subsequently expressed traits will depend on the effectiveness of habitat tracking through phenology.

Cold stratification in winter is more than enough for seed dormancy-break of summer annuals in eastern North America: implications for climate change

AbstractGermination of seeds of some summer annuals in Kentucky (eastern USA) in late-winter lead to the hypothesis that under present climate conditions the whole length of the winter cold stratification (CS) period is not required for dormancy-break of seeds of summer annuals with physiological dormancy (PD). We evaluated our data from germination phenology studies of 45 species (69 datasets) and buried-seed studies of 33 species (44 datasets). We determined time and temperature of germination after CS and percentage of the total number of hours of CS during winter (% of winter CS) seeds received prior to start of germination. In the phenology studies, mean temperature during the week of first germination for C3 and C4 species was 11.1 and 14.4°C, respectively, and % of winter CS was 80.8 and 87.4, respectively. In the buried-seed studies, % of CS for C3 and C4 species was 40.8 and 48.1, respectively, when they germinated to 25% at 20/10°C. For 32 of 33 species in the buried-seed studies, the minimum temperature at which seeds germinated decreased with increased CS; thus, seeds had Type 2 non-deep PD. The time of germination is controlled by a number of hours of CS, a decrease in minimum temperature at which seeds can germinate and a temperature increase in early spring. Seeds can germinate at relatively high temperatures as early as December and January, but they continue to be CS until spring. Temperature increases in eastern North America due to global warming are not likely to inhibit the germination of summer annuals with PD in spring.

From Cone to Seed and Seedling—Characterization of Three Portuguese Pinus pinaster Aiton Populations

Seed production in Pinus pinaster Aiton is not usually considered a limiting factor for natural regeneration; instead, seed weight is a more limiting factor in successful pine recruitment. Divergent relationships between seed weight and germination rate were previously observed amongst maritime pine populations of central coastal Portugal. The present study followed cone-to-seed and seed-to-seedling approaches to decrease intra-specific variability and clarify the impact of cone size and seed mass on seedling recruitment. The main objectives of this study were (1) to determine and compare the mass of cones and seeds of three maritime pine populations located along a geographic gradient along the coastal center of Portugal and (2) to clarify the relations thereof between cone and seed traits with germination phenology and initial seedling growth. Results demonstrated that heavier cones tended to generate more mature seeds, but not necessarily heavier ones, although seed weight was suggested to be an indicator of robust seedlings. The outcomes of this study reinforce the great intra-population variability of maritime pine, showing the ecology of this species and its ability to adapt to various environments successfully.

Width of the temperature range for seed germination of herbaceous plant species in temperate eastern North America: life cycles, seasons and temperature variation and implication for climate warming

AbstractTo persist (without immigration) in habitats with unpredictable environmental conditions, annuals must produce seeds each year or have a seed bank. Thus, we predicted that compared to perennials, annuals have a wider germination temperature range (GTR, the difference in temperature between the week with the highest and the week with the lowest germination during the natural germination season). We determined the GTR via germination phenology data for 350 herbaceous species in 59 families from the eastern USA: summer annuals (SA), 63; winter annuals (WA), 83; monocarpic perennials (MP), 28; and polycarpic perennials (PP), 176. There was no significant phylogenetic signal for the GTR. The width of the GTR during the first spring germination season was 9.6, 8.7 and 8.8°C for MP, PP and SA, respectively, and during the first autumn germination season 12.8, 11.8 and 12.4°C for MP, PP and WA, respectively. Annuals did not have a wider GTR than perennials in either the spring or the autumn germination season. Our data suggest that selection for early germination in either spring or autumn has resulted in only small differences in the GTR. We predict that global warming will have little or no effect on reshaping the germination phenology of herbaceous species of temperate eastern North America.

Effects of temperature on seed dormancy and germination of the coastal dune plant Viola grayi: germination phenology and responses to winter warming.

In temperate sand dunes, rising air temperature from climate change could not only further elevate surface soil temperatures during summers, but also drastically change the range of soil temperatures in other seasons. Winter warming may shift the timing of seed germination of dune species that require cold stratification for dormancy release. We assessed the effects of temperature on dormancy and germination of Viola grayi seeds and evaluated whether winter warming could affect its germination phenology by conducting germination experiments and analyzing soil temperature data in cold and warm winters. Viola grayi seeds were dormant when dispersed in spring. One-month moist-chilling treatment (4°C) effectively released dormancy, while short, intermittent lower temperatures (alternating 20°/5°C) did not. Continuous higher temperatures induced secondary dormancy in nondormant seeds. During a cold, snowy winter, the surface soil temperatures of the sand dune remained at 0°-2°C for approximately 1 month owing to the accumulated snow, while the period of such stable low soil temperatures was much shorter during a warm, less-snowy winter, and the highest soil temperature reached 20°-25°C. These results suggest that dispersed seeds germinate in the following spring after winter chilling, but they may remain dormant after warm winters. With winter warming, seed dormancy of V. grayi seeds could be prolonged and the associated germination delayed. Assessing the minimum requirements for dormancy release and the potential to form persistent soil seed banks is important for judging the necessity and urgency of conservation efforts for temperate dune species.

Epicotyl morphophysiological dormancy in seeds of Paeonia ostii (Paeoniaceae): Seasonal temperature regulation of germination phenology

Paeonia is one of the most popular ornamental plants in temperate regions, and its seeds have epicotyl dormancy. Our primary aim was to determine the sequence of temperatures required for radicle and shoot emergence in seeds of P. ostii and how these two distinct stages of dormancy-break are correlated with natural seasonal temperature changes. We tested the effects of various temperature regimes and sequences of regimes on growth of the underdeveloped embryo inside the seed, radicle emergence and shoot growth in the laboratory and germination responses to temperature under natural temperatures in an experimental garden and analyzed ABA and GA concentrations during the dormancy-breaking process. Prior to radical emergence, embryo length increased c. 330%. Embryo elongation and radicle emergence occurred in early October following seed dispersal in summer. The epicotyl-plumule did not differentiate until after embryo growth was completed and the radicle had emerged. Cold stratification was required to break epicotyl dormancy; consequently, shoot emergence did not occur until February when the temperature began to increase. Thus, seed dormancy break in P. ostii is phenologically well adapted to the seasonal cycle of the temperate zone. ABA and GA analyses suggest that root dormancy might be due to high ABA content and shoot dormancy to low GA content. • Seeds of Paeonia ostii have deep simple epicotyl morphophysiological dormancy. • The plumule does not differentiate until embryo growth is competed. • Radicle emergence occurs only at warm temperatures in autumn. • Shoots emerge only in spring in root-emerged seeds that have been cold stratified. • The timing of dormancy-break is regulated by changes in seasonal temperatures.

Reproductive Cycle of the Seagrass Zostera noltei in the Ria de Aveiro Lagoon.

Sexual reproduction in seagrasses is essential to increase their resilience towards environmental stressors, but its phenology is still unknown in some regions, limiting our knowledge about the recovery capacity of these ecosystems. In this study, the flowering effort, reproductive phenology, seed production and ability of germination of Zostera noltei was studied for the first time in the Ria de Aveiro lagoon, Portugal. Flowering of Z. noltei in the Ria de Aveiro lasts from June to November, reaching a peak between July and August. All the meadows showed similar flowering effort and phenology over time. Comparing with other European populations, the flowering effort of Z. noltei in Ria de Aveiro lasted for a longer period, which could be related with the milder temperatures in summer and autumn and the great anthropogenic stress to which the meadows are subjected in the lagoon. The number of seeds produced and their ability of germination were similar among meadows and sampling periods, reaching levels similar to those of other European regions. Nevertheless, future studies are needed to determine the fate of the produced seeds in the field to have a better understanding about the natural recovery capacity of the species.

Warming during maternal generations delays offspring germination in native and nonnative species

As environmental conditions shift due to global warming and other human‐caused environmental changes, plastic responses in phenological traits like germination or flowering time may become increasingly important. While phenological plasticity is a common response to global warming, with many populations exhibiting earlier germination or flowering in warmer years, warming may also result in transgenerational plasticity, especially on early life stages. In other words, seeds produced by mothers inhabiting warmer environments may germinate faster (or slower) than seeds produced by mothers inhabiting ambient or cooler environments. Here, we use seeds collected from a field warming experiment to examine how germination and early growth differ in response to ambient versus warmed (+3°C) temperatures experienced by both maternal and offspring generations. Because nonnative species are often more phenotypically plastic than native species and because a variety of life‐history traits and environmental factors affect the evolution of both within‐ and transgenerational plasticity, we include multiple invasive and native plant species in our study. On average, warming experienced during maternal generations delayed germination by ~0.2 days °C−1, although species varied in the magnitude of response. In contrast, warming during the offspring generation tended to advance germination by ~0.1 days °C−1. Nonnative species demonstrated higher germination success than native species, but we detected no differences in germination timing between native and nonnative species or that native and nonnative species differed in either within‐ or transgenerational plasticity, although species (independent of native status) did exhibit differing degrees of within‐ and transgenerational plasticity in germination timing and early growth. This study suggests that temperatures experienced by maternal plants can influence their offspring's germination phenology, potentially even more so than temperatures experienced in the offspring's immediate environment.

Dormancy cycles in Aquilegia oxysepala Trautv. et Mey. (Ranunculaceae), a species with non-deep simple morphophysiological dormancy

Seed dormancy and the formation of a soil seed bank is important plant regeneration strategies, especially if the environment is unpredictable. The present research explores how environmental factors control seed dormancy release, and how seed dormancy is related to the soil seed bank and regeneration of the perennial Aquilegia oxysepala. The effects of incubation temperature, light, cold and warm stratification, gibberellic acid (GA3) along with the germination phenology of A. oxysepala in the field were used to determine the type of seed dormancy. Seasonal change of seed dormancy was determined by regularly exhuming buried seeds and incubating them in laboratory conditions. A. oxysepala seeds has underdeveloped (small) embryos along with physiological dormancy at dispersal. With the increased amounts of cold stratification, the germination of A. oxysepala increased gradually. GA3 served as a substitute for cold stratification. Breaking of physiological dormancy under natural temperatures in the field occurred in winter, while growth of embryos and germination of seeds occurred in early spring. Viable seeds that had not germinated in early spring were induced into secondary dormancy by high soil temperatures. A. oxysepala provides one of a few examples of dormancy cycling in seeds with morphophysiological dormancy. Freshly matured seeds of A. oxysepala seeds have non-deep simple morphophysiological dormancy. The annual dormancy–non-dormancy cycle maintains the coordination between timing of seedling emergence with favorable seasons, thus increasing the survival chances of seedlings in environments with seasonal changes.

Germination responses to winter warm spells and warming vary widely among woody plants in a temperate forest.

Winter underpins key ecological processes, such as dormancy loss and seedling emergence. Enhanced warm spells, together with warming are occurring and will continue in the future. The consequences of these climate phenomena on germination were investigated among co-occurring woody plants, whose seeds are bird-dispersed in autumn and require cold stratification for spring emergence. Seeds from nine common southeastern USA plants were collected in autumn. We verified that seeds of the study species required cold stratification for dormancy loss. We then examined the following aspects in the laboratory or field: effect of warm spells during cold stratification on germination, effect of a warm spell during winter on seed survival and germination phenology, and effect of warming from autumn dispersal through winter dormancy loss on timing of germination. While no consistent effects of warm spells were found in the laboratory on quantity of germination, warm spells advanced spring field germination for several species. Some species germinated during cold stratification and during warm spells, especially extreme spells, in the laboratory. In the field, about half of Lonicera maackii seedlings that emerged with a warm spell died by late winter. With warming from autumn through spring, laboratory germination shifted from spring to predominately autumn for some species. With precocious germination during warm spells or germination phenology shifts, two scenarios are possible. Seedlings may die during winter, reducing the size of the soil seed bank and number of emergents, or they would survive in warmer winters, which would give them a competitive advantage over spring-emerging seedlings.

Variation in the seasonal germination niche across an elevational gradient: the role of germination cueing in current and future climates.

The timing of germination has profound impacts on fitness, population dynamics, and species ranges. Many plants have evolved responses to seasonal environmental cues to time germination with favorable conditions; these responses interact with temporal variation in local climate to drive the seasonal climate niche and may reflect local adaptation. Here, we examined germination responses to temperature cues in Streptanthus tortuosus populations across an elevational gradient. Using common garden experiments, we evaluated differences among populations in response to cold stratification (chilling) and germination temperature and related them to observed germination phenology in the field. We then explored how these responses relate to past climate at each site and the implications of those patterns under future climate change. Populations from high elevations had stronger stratification requirements for germination and narrower temperature ranges for germination without stratification. Differences in germination responses corresponded with elevation and variability in seasonal temperature and precipitation across populations. Further, they corresponded with germination phenology in the field; low-elevation populations germinated in the fall without chilling, whereas high-elevation populations germinated after winter chilling and snowmelt in spring and summer. Climate-change forecasts indicate increasing temperatures and decreasing snowpack, which will likely alter germination cues and timing, particularly for high-elevation populations. The seasonal germination niche for S. tortuosus is highly influenced by temperature and varies across the elevational gradient. Climate change will likely affect germination timing, which may cascade to influence trait expression, fitness, and population persistence.

Hydrothermal sensitivities of seed populations underlie fluctuations of dormancy states in an annual plant community.

Plant germination ecology involves continuous interactions between changing environmental conditions and the sensitivity of seed populations to respond to those conditions at a given time. Ecologically meaningful parameters characterizing germination capacity (or dormancy) are needed to advance our understanding of the evolution of germination strategies within plant communities. The germination traits commonly examined (e.g., maximum germination percentage under optimal conditions) may not adequately reflect the critical ecological differences in germination behavior across species, communities, and seasons. In particular, most seeds exhibit primary dormancy at dispersal that is alleviated by exposure to dry after-ripening or to hydrated chilling to enable germination in a subsequent favorable season. Population-based threshold (PBT) models of seed germination enable quantification of patterns of germination timing using parameters based on mechanistic assumptions about the underlying germination physiology. We applied the hydrothermal time (HTT) model, a type of PBT model that integrates environmental temperature and water availability, to study germination physiology in a guild of coexisting desert annual species whose seeds were after-ripened by dry storage under different conditions. We show that HTT assumptions are valid for describing germination physiology in these species, including loss of dormancy during after-ripening. Key HTT parameters, the hydrothermal time constant (θHT ) and base water potential distribution among seeds (Ψb (g)), were effective in describing changes in dormancy states and in clustering species exhibiting similar germination syndromes. θHT is an inherent species-specific trait relating to timing of germination that correlates well with long-term field germination fraction, while Ψb (g) shifts with depth of dormancy in response to after-ripening and seasonal environmental variation. Predictions based on variation among coexisting species in θHT and Ψb (g) in laboratory germination tests matched well with 25-yr observations of germination dates and fractions for the same species in natural field conditions. Seed dormancy and germination strategies, which are significant contributors to long-term species demographics under natural conditions, can be represented by readily measurable functional traits underlying variation in germination phenologies.