Large and non-spherical seeds are less likely to form a persistent soil seed bank.

There is some evidence that seed traits can affect the long-term persistence of seeds in the soil. However, findings on this topic have differed between systems. Here, we brought together a worldwide database of seed persistence data for 1474 species to test the generality of seed mass-shape-persistence relationships. We found a significant trend for low seed persistence to be associated with larger and less spherical seeds. However, the relationship varied across different clades, growth forms and species ecological preferences. Specifically, relationships of seed mass-shape-persistence were more pronounced in Poales than in other order clades. Herbaceous species that tend to be found in sites with low soil sand content and precipitation have stronger relationships between seed shape and persistence than in sites with higher soil sand content and precipitation. For the woody plants, the relationship between persistence and seed morphology was stronger in sites with high soil sand content and low precipitation than in sites with low soil sand content and higher precipitation. Improving the ability to predict the soil seed bank formation process, including burial and persistence, could benefit the utilization of seed morphology-persistence relationships in management strategies for vegetation restoration and controlling species invasion across diverse vegetation types and environments.

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 in situ 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.

Intraspecific variation in seed size is mediated by seed dispersal modes and animal dispersers - evidence from a global-scale dataset.

Seed dispersal mechanisms play a crucial role in driving evolutionary changes in seed and fruit traits. While previous studies have primarily focussed on the mean or maximum values of these traits, there is also significant intraspecific variation in them. Therefore, it is pertinent to investigate whether dispersal mechanisms can explain intraspecific variations in these traits. Taking seed size as a case study, we compiled a global dataset comprising 3424 records of intraspecific variation in seed size (IVSS), belonging to 691 plant species and 131 families. We provided the first comprehensive quantification of dispersal mechanism effects on IVSS. Biotic-dispersed species exhibited a larger IVSS than abiotic-dispersed species. Synzoochory species had a larger IVSS than endozoochory, epizoochory, and myrmecochory species. Vertebrate-dispersed species exhibited a larger IVSS than invertebrate-dispersed species, and species dispersed by birds exhibited a larger IVSS than mammal-dispersed species. Additionally, a clear negative correlation was detected between IVSS and disperser body mass. Our results prove that the IVSS is associated with the seed dispersal mechanism. This study advances our understanding of the dispersal mechanisms' crucial role in seed size evolution, encompassing not only the mean value but also the variation.

Seed traits and recruitment interact with habitats to generate patterns of local adaptation in a perennial grass.

A fundamental challenge in the field of ecology involves understanding the adaptive traits and life history stages regulating the population dynamics of species across diverse habitats. Seed traits and early seedling vigor are thought to be key functional traits in plants, with important consequences for recruitment, establishment, and population persistence. However, little is known about how diverse seed traits interact with seed and microsite availability to impact plant populations. Here, we performed a factorial experiment involving seed addition and surface soil disturbance to explore the combined effects of seed and site availability using genotypes characterized by varying seed mass and dormancy traits. Additionally, we included hybrids that exhibited recombined seed trait relationships compared with natural genotypes, allowing us to assess the impact of specific seed traits on establishment across different sites. We detected a significant three-way interaction between seed addition, site conditions, and soil surface disturbance, influencing both seedling establishment and adult recruitment in Panicum hallii, a perennial grass found in coastal mesic (lowland) and inland xeric (upland) habitats. This establishment/recruitment pattern suggests that mesic and xeric establishment at foreign sites is constrained by the interplay of seed and site limitations. Notably, soil surface disturbance facilitated establishment and recruitment of the xeric genotype while limiting the mesic genotype across all sites. Our results highlight the importance of seed size and dormancy as key factors impacting seedling establishment and adult recruitment, suggesting a potential interactive relationship between these traits.

Mother-reliant or self-reliant: the germination strategy of seeds in a species-rich alpine meadow is associated with the existence of pericarps.

BACKGROUND AND AIMS: Some plants germinate their seeds enclosed by a pericarp, whereas others lack the outer packaging. As a maternal tissue, the pericarp might impart seeds with different germination strategies. Plants in a community with different flowering times might separately disperse and germinate their seeds; therefore, flowering time can be considered as one manifestation of maternal effects on the offspring. The mass of the seed is another important factor influencing germination and represents the intrinsic resource of the seed that supports germination. Using seeds from a species-rich alpine meadow located in the Hengduan Mountains of China, a global biodiversity hotspot, we aimed to illustrate whether and how the type of seed (with or without a pericarp) modulates the interaction of flowering time and seed mass with germination. METHODS: Seeds were germinated in generally favourable conditions, and the speed of germination [estimated by mean germination time (MGT)] was calculated. We quantified the maternal conditions by separation of flowering time for 67 species in the meadow, of which 31 produced seeds with pericarps and 36 yielded seeds without pericarps. We also weighed 100 seeds of each species to assess their mass. KEY RESULTS: The MGT varied between the two types of seeds. For seeds with pericarps, MGT was associated with flowering time but not with seed mass. Plants with earlier flowering times in the meadow exhibited more rapid seed germination. For seeds without a pericarp, the MGT depended on seed mass, with smaller seeds germinating more rapidly than larger seeds. CONCLUSIONS: The distinct responses of germination to flowering time and seed mass observed in seeds with and without a pericarp suggest that germination strategies might be mother-reliant for seeds protected by pericarps but self-reliant for those without such protection. This new finding improves our understanding of seed germination by integrating ecologically mediated maternal conditions and inherent genetic properties.

The seed morphospace, a new contribution towards the multidimensional study of angiosperm sexual reproductive biology.

BACKGROUND: The evolutionary success of flowering plants is associated with the vast diversity of their reproductive structures. Despite recent progress in understanding angiosperm-wide trends of floral structure and evolution, a synthetic view of the diversity in seed form and function across angiosperms is lacking. SCOPE: Here we present a roadmap to synthesise the diversity of seed forms in extant angiosperms, relying on the morphospace concept, i.e. a mathematical representation which relates multiple traits and describes the realised morphologies. We provide recommendations on how to broaden the range of measurable traits beyond mass, by using key morphological traits representative of the embryo, endosperm, and seed coat but also fruit attributes (e.g., dehiscence, fleshiness). These key traits were used to construct and analyse a morphospace to detect evolutionary trends and gain insight into how morphological traits relate to seed functions. Finally, we outline challenges and future research directions, combining the morphospace with macroevolutionary comparative methods to underline the drivers that gave rise to the diversity of observed seed forms. CONCLUSIONS: We conclude that this multidimensional approach has the potential, although still untapped, to improve our understanding of covariation among reproductive traits, and further elucidate angiosperm reproductive biology as a whole.

Seed functional ecology in Brazilian rock outcrop vegetation: an integrative synthesis.

BACKGROUND AND AIMS: Rock outcrop vegetation is distributed worldwide and hosts a diverse and unique flora that evolved under harsh environmental conditions. Unfortunately, seed ecology in such ecosystems has received little attention, especially regarding seed traits, germination responses to abiotic factors and the potential role of phylogenetic relatedness on such features Here, we provide the first quantitative and phylogenetically-informed synthesis of the seed functional ecology of Brazilian rock outcrop vegetation, with a particular focus on quartzitic and ironstone campo rupestre. METHODS: Using a database of functional trait data, we calculated the phylogenetic signal of seven seed traits for 371 taxa and tested whether they varied among growth forms, geographic distribution, and microhabitats. We also conducted meta-analyses that included 4,252 germination records for 102 taxa to assess the effects of light, temperature, and fire-related cues on the germination of campo rupestre species and explored how the aforementioned ecological groups and seed traits modulate germination responses. KEY RESULTS: All traits and germination responses showed a moderate-to-strong phylogenetic signal. Campo rupestre species responded positively to light and had maximum germination between 20-25 ºC. The effect of temperatures beyond this range was moderated by growth form, species geographic distribution, and microhabitat. Seeds exposed to heat shocks above 80 °C lost viability, but smoke accelerated germination. We found a moderating effect of seed mass for in responses to light and heat shocks, with larger, dormant seeds tolerating heat better but less sensitive to light. Species from xeric habitats evolved phenological strategies to synchronise germination during periods of increased soil water availability. CONCLUSIONS: Phylogenetic relatedness plays a major role in shaping seed ecology of Brazilian rock outcrop vegetation. Nevertheless, seed traits and germination responses varied significantly between growth forms, species geographic distribution and microhabitats, providing support to the regeneration niche hypothesis and the role of functional traits in shaping germination in these ecosystems.

Effect of intraspecific seed trait variation on the germination of eight tropical dry forest species.

Functional traits can have intraspecific and interspecific variations essential in the structure and dynamics of natural communities. These traits may have implications in the germination and seedling establishment phases in seeds. The objective of this study was to evaluate the effect of variations in mass, volume, and nutrient content (C, N, and P) on the germination of eight species representative of the tropical dry forest (TDF). Our results showed that seed size, both in terms of mass and volume, did not predict germination rates or percentages, nor were they related to nutrient content. In contrast, N content was the most important trait in the germination phase. Larger seeds did not germinate more or faster, but they could offer better resistance against desiccation, since they had higher C/N ratios in their tissues, a characteristic of orthodox seeds. The species A. guachapele, B. arborea, H. crepitans, and V. tortuosa presented a high biological potential in terms of their regeneration capacity, particularly, because the characteristics of their seeds, as well as the nutrient content, revealed consistent implications in their reproductive success, promoting high germination percentages in less time. In general, the results obtained in this study provide basic knowledge for future research, offering starting points for further exploration of species-specific adaptations and how they may be affected by the environment.

Gradual genome size evolution and polyploidy in Allium from the Qinghai-Tibetan Plateau.

BACKGROUND AND AIMS: Genome size is an important plant trait, with substantial interspecies variation. The mechanisms and selective pressures underlying genome size evolution are important topics in evolutionary biology. There is considerable diversity in Allium from the Qinghai-Tibetan Plateau, where genome size variation and related evolutionary mechanisms are poorly understood. METHODS: We reconstructed the Allium phylogeny using DNA sequences from 71 species. We also estimated genome sizes of 62 species, and determined chromosome numbers in 65 species. We examined the phylogenetic signal associated with genome size variation, and tested how well the data fit different evolutionary models. Correlations between genome size variations and seed mass, altitude and 19 bioclimatic factors were determined. KEY RESULTS: Allium genome sizes differed substantially between species and within diploids, triploids, tetraploids, hexaploids and octaploids. Size per monoploid genome (1Cx) tended to decrease with increasing ploidy levels. Allium polyploids tended to grow at a higher altitude than diploids. The phylogenetic tree was divided into three evolutionary branches. The genomes in Clade I were mostly close to the ancestral genome (18.781 pg) while those in Clades II and III tended to expand and contract, respectively. A weak phylogenetic signal was detected for Allium genome size. Furthermore, significant positive correlations were detected between genome size and seed mass, as well as between genome size and altitude. However, genome size was not correlated with 19 bioclimatic variables. CONCLUSIONS: Allium genome size shows gradual evolution, followed by subsequent adaptive radiation. The three well-supported Allium clades are consistent with previous studies. The evolutionary patterns in different Allium clades revealed genome contraction, expansion and relative stasis. The Allium species in Clade II may follow adaptive radiation. The genome contraction in Clade III may be due to DNA loss after polyploidization. Allium genome size might be influenced by selective pressure due to the conditions on the Qinghai-Tibetan Plateau (low temperature, high UV irradiation and abundant phosphate in the soil).

Light controls alpine meadow community assembly during succession by affecting species recruitment from the seed bank.

Some research indicates that soil seed banks can promote species coexistence through storage effects. However, the seed bank mechanism that maintains plant assembly and its role in degraded grassland restoration are still not clear. We collected seed bank samples from early, mid and late secondary successional stages of an abandoned subalpine meadow on the Tibetan Plateau, and samples from each stage were exposed to full (i.e., natural), mid, and low light treatments in the field to represent light availability at the bottom/understory (soil surface) of a plant community in the early, mid and late stages of succession, respectively. Species richness, seed density, species composition, and community weighted mean values (CWMs) of seed mass of the species whose seeds germinated in soil samples were evaluated. In response to the light treatments, species richness increased significantly with increased light only for the late successional stage, seed density increased significantly with increased light only in the early and mid successional stages, and seed mass decreased significantly with increased light only in the mid and late successional stages. Species composition differed significantly among the light treatments only in the late successional stage. For the successional series, species richness and seed mass of the species that germinated increased significantly with succession only under mid and full light treatments. Seed density decreased significantly with succession in each light treatment. Species composition differed significantly between the early- and late stage and between the mid and late stage in each light treatment. Both the abiotic (light) and biotic (seed mass) factors influence seed bank recruitment to the plant community. Regeneration of small-seeded species in the seed bank was inhibited under low light in the late successional stage. The balance of stochastic and deterministic processes along a successional gradient was determined by regeneration from the seed bank depending on light intensity change. Differences in seed response to light intensity change largely determined plant community assembly. Our findings should help in the development of effective conservation and restoration strategies.

Weak evidence of trade-offs modulated by seed mass among a guild of closely related winter annuals.

Plant-plant interactions are integral to the establishment and persistence of diversity in plant communities. For annual plant species that depend on seeds to regenerate, seed characteristics that confer fitness advantages may mediate processes such as plant-plant interactions. Seed mass is known to vary widely and has been shown to associate with species' differences in stress tolerance and competitive effects. However, understanding of how seed mass influences species' responses to competition is less well understood. Using natural assemblages of six closely related annual plant species in Western Australia, we implemented a thinning study to assess how seed mass influences the outcomes of plant-plant interactions. We found relatively weak evidence for competition or facilitation among species. Our strongest results indicated that heavy-seeded species had lower survivorship than light-seeded species when interacting with heterospecifics. Seed mass was also negatively related to overall survival, counter to expectations. These findings indicate some evidence for trade-offs mediated by seed mass in this system. However, we acknowledge that other factors may have influenced our results, such as the use of natural assemblages (rather than using sowing experiments) and the presence of important small-scale environmental variation not captured with our choice of abiotic variables. Further research is required to clarify the role of seed mass in this diverse annual system, ideally including many focal species, and using sowing experiments.

Propagule size and seed development duration: high photosynthate allocation and growth allometry.

MAIN CONCLUSION: The divergences in propagule mass have been more consistently associated with divergences in seed development duration or fruit pedicel cross-sectional area than with divergences in any other biotic factors. Allometry and Corner's rule became an important theme in evolutionary biology of plant trait structure and function. Being one of the most widely noticed plant traits, propagule (seed and fruit) mass variation mechanism across species is still controversial. Here we examined correlations between propagule mass and seed development duration as well as other traits, such as cross-sectional area of fruit pedicel, life form, fruit type and leaf area over four census years, to test an important life history strategy for propagule biomass allocation. We investigated 491 species, belonging to 91 families and 320 genera, representing 95% of native wild species in Beijing Botanical Garden. The scaling correlations between propagule mass and seed development duration and the other traits were determined using phylogenetic generalized linear models. Results show a significant positive relationship among propagule mass and seed development duration, leaf area and pedicel cross-sectional area for all species and for each life form (except vines) regardless of phylogeny. The variation in seed mass has been more consistently associated with variation in seed development duration than with divergences in any other variable, such as growth form, fruit type, pedicel cross-sectional area and leaf area, whereas variation in fruit weight was found to be more associated with variation in pedicel cross-sectional area than the other. Biotic factors, such as seed development duration, pedicel cross-sectional area, growth form and leaf area, mediate propagule size variation, of which seed development duration or pedicel cross-sectional area is the most important. The results further supported a hypothesis that large-seeded species evolved owing to high photosynthate availability and growth allometry in plant body. A mechanistic mathematical model involving seed development duration was provided to expound propagule mass variations across species.

Contrasting effects of nectar yeasts on the reproduction of Mediterranean plant species.

PREMISE: Yeasts are often present in floral nectar and can influence plant fitness directly (independently of pollinators) or indirectly by influencing pollinator visitation and behavior. However, few studies have assessed the effect of nectar yeasts on plant reproductive success or compared effects across different plant species, limiting our understanding of the relative impact of direct vs. indirect effects. METHODS: We inoculated the nectar of six plant species in the field with the cosmopolitan yeast Metschnikowia reukaufii to analyze the direct and indirect effects on female reproductive success over 2 years. The pollinator assemblage for each species was recorded during both flowering years. RESULTS: Direct yeast effects on female fecundity were statistically nonsignificant for all plant species. There were significant indirect, pollinator-mediated effects on fruit production and seed mass for the two species pollinated almost exclusively by bumblebees or hawkmoths, with the direction of the effects differing for the quantity- and quality-related fitness components. There were no consistent effects of the yeast on maternal fecundity for any of the species with diverse pollinator assemblages. CONCLUSIONS: Effects of M. reukaufii on plant reproduction ranged from negative to neutral or positive depending on the plant species. The among-species variation in the indirect effects of nectar yeasts on plant pollination could reflect variation in the pollinator community, the specific microbes colonizing the nectar, and the order of microbial infection (priority effects), determining potential species interactions. Elucidating the nature of these multitrophic plant-pollinator-microbe interactions is important to understand complex processes underlying plant pollination.

The role of maternal age, growth, and environment in shaping offspring performance in an aerial conifer seed bank.

PREMISE: Maternal effects have been demonstrated to affect offspring performance in many organisms, and in plants, seeds are important mediators of these effects. Some woody plant species maintain long-lasting canopy seed banks as an adaptation to wildfires. Importantly, these seeds stored in serotinous cones are produced by the mother plant under varying ontogenetic and physiological conditions. METHODS: We sampled the canopy seed bank of a highly serotinous population of Pinus pinaster to test whether maternal age and growth and the environmental conditions during each crop year affected seed mass and ultimately germination and early survival. After determining retrospectively the year of each seed cohort, we followed germination and early survival in a semi-natural common garden. RESULTS: Seed mass was related to maternal age and growth at the time of seed production; i.e., slow-growing, older mothers had smaller seeds, and fast-growing, young mothers had larger seeds, which could be interpreted either as a proxy of senescence or as a maternal strategy. Seed mass had a positive effect on germination success, but aside from differences in seed mass, maternal age had a negative effect and diameter had a positive effect on germination timing and subsequent survival. CONCLUSIONS: The results highlight the importance of maternal conditions combined with seed mass in shaping seedling establishment. Our findings open new insights in the offspring performance deriving from long-term canopy seed banks, which may have high relevance for plant adaptation.

Seed Quantity or Quality?-Reproductive Responses of Females of Two Dioecious Woody Species to Long-Term Fertilisation.

Although seed quality and quantity, as well as reproductive performance are important life history stages of plants, little is known about the reproductive responses of trees to environmental changes such as increased anthropogenic deposition of nitrogen (N) and phosphorus (P). Dioecious plants are good models with which to test the environmental impact on female or male reproductive responses individually. We analysed effects of different long-term nutritional availability on the reproductive performance of two dioecious species (Taxus baccata L. and Juniperus communis L.) characterised by different life histories. By using pot experiments with vegetatively propagated plants grown in different fertilisation conditions, we observed an increase in plant growth and strobili production but a decrease in seed efficiency. Seeds produced by fertilised plants had greater seed mass. Fertiliser addition did not change C or N content nor the C/N ratio of T. baccata seeds, but increased N content and the N/P ratio; however, it did lower the C/N ratio in J. communis. Fertilisation did not change the metabolite profile in T. baccata but 18 metabolites were changed in J. communis. The study revealed new links between species life history, environmental changes, and reproduction. The findings imply that future environmental conditions may alter both seed productivity, and quality, as well as plant reproductive behaviour.

Persistent soil seed banks promote naturalisation and invasiveness in flowering plants.

With globalisation facilitating the movement of plants and seeds beyond the native range, preventing potentially harmful introductions requires knowledge of what drives the successful establishment and spread of alien plants. Here, we examined global-scale relationships between naturalisation success (incidence and extent) and invasiveness, soil seed bank properties (type and densities) and key species traits (seed mass, seed dormancy and life form) for 2350 species of angiosperms. Naturalisation and invasiveness were strongly associated with the ability to form persistent (vs. transient) seed banks but relatively weakly with seed bank densities and other traits. Our findings suggest that seed bank persistence is a trait that better captures the ability to become naturalised and invasive compared to seed traits more widely available in trait databases. Knowledge of seed persistence can contribute to our ability to predict global naturalisation and invasiveness and to identify potentially invasive flowering plants before they are introduced.

Exposure time is an important variable in quantifying post-dispersal seed removal.

A literature synthesis concluded that small mammals have the greatest impact on post-dispersal removal of intermediate-sized seeds (Dylewski et al. 2020). However, this study failed to consider the duration of seed exposure to predators. Re-analyses of the corrected dataset revealed only a weak effect of seed mass on seed removal.

The seed germination spectrum of alpine plants: a global meta-analysis.

Assumptions about the germination ecology of alpine plants are presently based on individual species and local studies. A current challenge is to synthesise, at the global level, the alpine seed ecological spectrum. We performed a meta-analysis of primary data from laboratory experiments conducted across four continents (excluding the tropics) and 661 species, to estimate the influence of six environmental cues on germination proportion, mean germination time and germination synchrony; accounting for seed morphology (mass, embryo : seed ratio) and phylogeny. Most alpine plants show physiological seed dormancy, a strong need for cold stratification, warm-cued germination and positive germination responses to light and alternating temperatures. Species restricted to the alpine belt have a higher preference for warm temperatures and a stronger response to cold stratification than species whose distribution extends also below the treeline. Seed mass, embryo size and phylogeny have strong constraining effects on germination responses to the environment. Globally, overwintering and warm temperatures are key drivers of germination in alpine habitats. The interplay between germination physiology and seed morphological traits further reflects pressures to avoid frost or drought stress. Our results indicate the convergence, at the global level, of the seed germination patterns of alpine species.

Selection patterns on early-life phenotypic traits in Pinus sylvestris are associated with precipitation and temperature along a climatic gradient in Europe.

Understanding the dynamics of selection is key to predicting the response of tree species to new environmental conditions in the current context of climate change. However, selection patterns acting on early recruitment stages and their climatic drivers remain largely unknown in most tree species, despite being a critical period of their life cycle. We measured phenotypic selection on Pinus sylvestris seed mass, emergence time and early growth rate over 2 yr in four common garden experiments established along the latitudinal gradient of the species in Europe. Significant phenotypic plasticity and among-population genetic variation were found for all measured phenotypic traits. Heat and drought negatively affected fitness in the southern sites, but heavy rainfalls also decreased early survival in middle latitudes. Climate-driven directional selection was found for higher seed mass and earlier emergence time, while the form of selection on seedling growth rates differed among sites and populations. Evidence of adaptive and maladaptive phenotypic plasticity was found for emergence time and early growth rate, respectively. Seed mass, emergence time and early growth rate have an adaptive role in the early stages of P. sylvestris and climate strongly influences the patterns of selection on these fitness-related traits.

Adaptive responses to temperature and precipitation variation at the early-life stages of Pinus sylvestris.

Early-stage fitness variation has been seldom evaluated at broad scales in forest tree species, despite the long tradition of studying climate-driven intraspecific genetic variation. In this study, we evaluated the role of climate in driving patterns of population differentiation at early-life stages in Pinus sylvestris and explored the fitness and growth consequences of seed transfer within the species range. We monitored seedling emergence, survival and growth over a 2-yr period in a multi-site common garden experiment which included 18 European populations and spanned 25° in latitude and 1700 m in elevation. Climate-fitness functions showed that populations exhibited higher seedling survival and growth at temperatures similar to their home environment, which is consistent with local adaptation. Northern populations experienced lower survival and growth at warmer sites, contrary to previous studies on later life stages. Seed mass was higher in populations from warmer areas and was positively associated with survival and growth at more southern sites. Finally, we did not detect a survival-growth trade-off; on the contrary, bigger seedlings exhibited higher survival probabilities under most climatic conditions. In conclusion, our results reveal that contrasting temperature regimes have played an important role in driving the divergent evolution of P. sylvestris populations at early-life stages.