American Journal of Botany

Gene flow to marginal populations at a species range edge can facilitate rapid adaptation by increasing genetic diversity, reducing inbreeding depression, and introducing novel alleles. In highly inbred populations, hybrid vigor is often observed in the first generation (F1), but hybrid breakdown may diminish fitness in subsequent generations. Thus, benefits of gene flow may be overestimated when only F1 performance is assessed. We tested whether gene flow among populations of the annual plant Erythranthe laciniata (A. Gray) G.L. Nesom, from similar and contrasting environments, confers persistent fitness advantages across F1 and F2 generations at the high-elevation edge of its range in the California Sierra Nevada. Gene flow was experimentally introduced through pollen transfer between cold-edge populations, between cold edge and central populations, and within local cold edge populations, and compared to self-fertilized offspring, the predominant mating strategy of E. laciniata. For F1 progeny, we measured morphological, phenological, and fitness traits in a common garden located near the cold-climate range limit during 2008-2009, a relatively average year, and for F2 progeny in 2009-2010, a relatively wet year. Although F1 crosses showed no initial performance advantage measured in the previous year, F2 progeny from center-to-edge and edge-to-edge crosses significantly outperformed selfed and locally outcrossed lines in fruit mass, total pedicels, biomass, and height. Our findings demonstrate that gene flow can confer long-term fitness benefits, especially among populations adapted to similar selective pressures, and highlight the potential of assisted gene flow to bolster or rescue peripheral populations facing climate change. SIGNIFICANCE STATEMENTSpecies living at the edges of their geographic ranges often have small, isolated populations with limited genetic diversity, which can restrict their ability to adapt to environmental change. Gene flow from other populations may increase adaptive potential, but its long-term consequences remain uncertain because most studies evaluate only first-generation hybrids. Using experimental crosses in the mountain wildflower Erythranthe laciniata, we show that gene flow can produce stronger fitness benefits in second-generation hybrids than in the first generation at a high-elevation range edge. These results suggest that recombination among populations can generate advantageous genetic combinations that emerge over multiple generations. Our findings highlight the potential for assisted gene flow to enhance adaptation and persistence of range-edge populations under climate change.

Lack of sexual reproduction limits the fitness and long-term viability of many plant populations. This may pose a particular problem for populations at the edges of species ranges, which are often small and isolated and therefore may be less likely to attract pollinators. But despite the fact that many range-edge populations are of significant conservation concern and value, there is often little information about which visitors are effective pollinators, and few explicit tests of whether range-edge populations experience reduced pollination. Here, we assess which visitors are effective pollinators of sundial lupine (Lupinus perennis), a legume that is threatened in much of its range, and whether pollination success varies between populations in the range core and those at the species northern range edge. Across six populations in the northern USA and southern Canada (Ontario), sundial lupine was visited almost exclusively by bees, but only large bees (Bombus, Xylocopa) could be confirmed as effective pollinators in single-visit experiments. While seed production varied significantly among populations, visitation rates did not. Neither pollinator visitation, pollen receipt, nor seed production declined at sundial lupines northern range edge. We therefore found no evidence that pollination success constrains either performance of at-risk populations of sundial lupine or the species northern range limit.

PremiseHerbarium specimens are increasingly used to extract morphological traits for ecological and evolutionary studies, yet the effects of tissue desiccation on trait measurements remain poorly understood. Here, we tested whether higher tissue water content leads to greater measurement changes after herborization (H1) and whether fresh trait values can be reliably predicted from herbarium measurements (H2). MethodsWe evaluated the reliability of herbarium-based measurements by comparing fresh and dried traits of leaves, flowers, fleshy fruits, and seeds across 262 individuals representing 133 Neotropical Myrtaceae species. Phylogenetic least square models and machine-learning regressions were used to test H1 and H2. ResultsLeaves and flowers generally shrank after herborization, fruits size metrics tended to increase, and seeds were largely unaffected. Water content was significantly associated with the magnitude of herborization effects in flowers and some leaf and seed traits. Fresh trait values were accurately predicted from herbarium measurements. Prediction errors were lowest for leaf traits, followed by fruits, flowers, and seeds. DiscussionThese results partially support H1 and support H2, indicating that herbarium specimens can be reliably used for trait analyses when organ-specific responses are considered, providing a practical framework to account for potential desiccation bias in functional trait research.

Long-term field observations typically are the "gold-standard" for inferences of phenological sensitivities in montane systems but are spatially limited. Herbarium specimens provide broader spatial coverage, but their utility to accurately capture montane phenology remains poorly known. We compared flowering phenology of 45 species inferred from herbarium specimens with comparable data from nearly 50 years of direct observations at the Rocky Mountain Biological Laboratory. Estimates of flowering time and phenological sensitivity to snow density were consistent between herbarium specimens and observations, but observations revealed secondary flowering peaks. Herbarium specimens additionally yielded shallower estimates of phenological sensitivity to spring temperature than did field observations. Across co-occurring species, "early" flowering individuals inferred from herbarium specimens, rather than the mean response across all individuals, may better approximate community-level phenological responses to temperature changes. We conclude that herbarium specimens are reliable resources for closing gaps in understanding phenological variation along elevational gradients of montane systems.

The temperate rainforests of the Pacific Northwest of North America harbor many endemic taxa whose evolutionary histories have been shaped by major climatic and geologic events. The enigmatic taildropper slugs (genus Prophysaon) are one example, notable for their ability to autonomize their tails to escape predators. Despite extensive work uncovering the evolutionary history of individual lineages, relationships among the nine recognized species of Prophysaon remain poorly understood due to insufficient molecular data. To address this, we collected transcriptomes for six of the nine currently accepted species of Prophysaon. Using these data, we were able to resolve species relationships, calling into question the existing subgeneric classification based on morphology. We also detected undescribed phenotypic diversity within the P. andersonii--P. foliolatum species complex, with molecular data supporting the distinctness of two phenotypically distinct populations from Washington. Finally, our transcriptomic data suggest a moderate role of introgression in shaping the evolutionary history of Prophysaon. Here, we synonymize the subgenus Mimetarion with nominotypical Prophysaon. Future work should further investigate whether the undescribed diversity detected here represents species level differentiation.

Background and aimsThis study combines morphological and environmental data to better understand a Brunfelsia (Solanaceae) species complex, aiming to clarify patterns of variation and identify ecological factors that shape morphotype boundaries. Such an approach provides a broader perspective on how organisms respond to environmental gradients and contributes to a more comprehensive understanding of biodiversity. MethodsWe analyzed 273 herbarium specimens for 13 morphological traits using univariate and ordination analyses, namely PCA and CVA. Climatic and edaphic variables were extracted for 147 specimens with georeferenced records. To assess habitat suitability and the ecological niche of each predefined morphotype, niche models under present conditions and niche overlap tests were conducted. A redundancy analysis (RDA) was applied to evaluate how environmental predictors explain variation in vegetative and floral traits. Finally, DAPC was used to estimate membership probabilities based on morphological and environmental data. Key ResultsTwo well-differentiated groups were recovered: the capitata-hydrangeiformis morphotype, allegedly composing a cline, and the ecologically and morphologically distinct "bahia" morphotype. Variation in floral traits was better explained by environmental predictors than variation in vegetative traits; moreover, floral traits were able to delineate morphotypes more robustly when plotted in isolation. However, when analyzing the results of ecological niche overlap, a significant ecological separation of the "bahia" morphotype from the others was observed. Therefore, key morphological characters for the taxonomy of Brunfelsia covary in part with environmental variables. ConclusionsThese findings support the recognition of "bahia" morphotype as a distinct species to be formally described. This integrative approach contributes to understanding diversification processes in biodiversity hotspots and highlights hidden taxonomic diversity within Brunfelsia, where many rare and narrow-endemic taxa lie.

Premise of studyUnder increasingly frequent pollinator-limited environments, plants need to rely on modes of reproductive assurance such as selfing and cloning. However, few studies investigate the interplay between selfing and cloning in plants that can do both. Here, we explore mechanisms determining the relative expression of selfing and cloning throughout the European distribution of the wild woodland strawberry (Fragaria vesca) under a pollinator-free environment. MethodsWe established an outdoor common garden with 121 woodland strawberry genotypes from across Europe and excluded them from pollinators. For each genotype, we recorded reproductive traits and performed hand-pollination treatments. Key ResultsWe found a weak trade-off between cloning and selfing, driven by increased seed and fruit provisioning rather than flower production. The capacity to autonomously self-fertilize was determined by the lateral proximity of the anthers to the pistils (lateral herkogamy), but not by early inbreeding depression. Genotypes sampled at lower latitudes and altitudes were better at self-fertilizing and had smaller petals. The propensity to clone increased towards the east, where genotypes also had smaller petals, particularly at higher latitudes. ConclusionAt the species level, we detected a trade-off between the propensity for clonal reproduction and the capacity for self-fertilization. At a continental scale, the capacity to self-fertilize varied along a north-south gradient, whereas clonal propensity varied along an east-west gradient. Our results suggest that these two modes of reproductive assurance may compensate for reduced pollinator attractiveness (smaller petals) in regions where each mode is most strongly expressed.

Ongoing climate change will negatively impact tree populations unless they are able to acclimate to the changes in their local environment. Effective planning for climate adaptation management requires an understanding of the current state of local adaptation and physiological performance to assess whether populations are at risk of local extinction, to determine if seed movement is appropriate, and to select appropriate seed sources if intervention is needed. We established a new reciprocal transplant experiment (ACE, Adaptation to Climate and Environment) across a latitudinal gradient in North America to examine the impacts of warming on three bur oak (Quercus macrocarpa) populations across much of the species range. We established common gardens in Minnesota, Illinois, and Oklahoma with seedlings grown from seeds collected within 50 km of each of those locations from a total of sixty maternal families. We aimed to 1) assess local adaptation in each of the populations using survival and size as fitness metrics, and 2) evaluate physiological responses to different environments along the latitudinal gradient. We found that northern populations are maladapted to hotter climates as evidenced by their low survival, growth, and photosynthetic rates in the warmest common garden. The southernmost population had the highest survival rate, growth rate, and fitness of the three populations in the southernmost garden, providing evidence for local adaptation to the warmest site. However, conditions in the middle garden resulted in the highest fitness and best physiological performance for all populations. Growth and survival were correlated in the middle garden but were decoupled in the northern and southern gardens. This decoupling is likely due to stress associated with more extreme climates at the ends of the gradient that led to greater resource allocation to survival than to growth. Our results suggest that southern seed sources may perform well in warmer conditions in the north brought on by climate change, which has important implications for managers assisting broadly ranged tree species in adapting to climate change.

O_LIWhen species hybridize, resistance to introgression is presumably due to selection against hybridizing alleles. While many studies have characterized direct selection at these sites, alleles may resist introgression through correlational selection. Here we investigate the role of direct and correlational selection in reducing introgression at the color locus in Ipomoea cordatotriloba. C_LIO_LIWe used recombinant inbred lines that varied in limb color, flower size and sugar concentration to estimate the fitness advantage of the flower color allele via direct and correlational selection. To assess the effect of correlational selection on fitness, we ask if floral size or nectar sugar concentration is correlated with fecundity in pink- but not white-limbed lines. C_LIO_LIWe find no evidence for direct selection on flower color across four fitness components - germination, survival, fecundity, and siring success. Instead, both flower size and sugar concentration significantly correlate with fecundity in pink, but not white limbed lines. As a result, correlational selection on the color allele opposes introgression when recurrent migration is low (<3%). C_LIO_LIThese results demonstrate that correlational, rather than direct, selection is sufficient to resist introgression via hybridization and suggest that correlational selection is an underexplored mechanism to generate resistance to introgression across multiple loci. C_LI

Background and AimsGroundwater-dependent ecosystems support disproportionate biodiversity in arid regions, yet the population genetics of spring-specialist plants remains poorly understood. Here, we present the first species-wide genetic dataset for crimson monkeyflower ( Mimulus verbenaceus, Phrymaceae), a spring-specialist plant distributed in seeps, springs, and associated riparian areas across desert regions of North America. MethodsUsing genome-wide reduced representation sequencing data consisting of 10,760 SNPs from 175 individuals across 17 populations, we characterized the patterns of genetic diversity using FST and Neis D. Population structure was assessed using ADMIXTURE and PCA. We examined the contributions of climate to range-wide genetic variation in crimson monkeyflower using redundancy analysis. Key ResultsPatterns of genetic differentiation were more consistent with those of spring-specialist animal taxa than those of upland plants or generalist riparian plants. We found strong population structure at both broad regional scales and at fine local scales. While riparian connectivity influenced local patterns of diversity, adaptation to local climatic variation was more influential at regional scales, with temperature, relative humidity, and a monsoon-driven climate gradient structuring genetic differentiation. ConclusionsOur findings highlight the distinctive influence of isolated perennial groundwater sources, as well as adaptation to climate, in shaping genetic variation in this spring-specialist plant. These findings suggest that spring-specialist plants deserve special consideration in ecological theory, management, and conservation.

Background and AimsNon-native species are now ubiquitous members of regional floras. The factors that lead to establishment and dominance of non-native species are continuously debated. Fundamental hypotheses about drivers of invasion success include the role of phylogeny, polyploidy, genome size, and rapid niche evolution. These hypotheses have been tested in the seed plants, but ferns, the second largest group of vascular plants, have rarely been considered in these analyses, despite making up a non-trivial portion of non-native floras. MethodsWe compiled a dataset of global non-native ferns and categorized them along the invasion spectrum using descriptions from the literature and natural history collections. Using this dataset, we assessed I) the taxonomic diversity and phylogenetic clustering of non-native ferns, II) the geographic distribution of fern introductions, testing for shifts in climatic niches, and III) test for the association of invader traits across the invasion continuum, including smaller genome sizes and higher ploidal levels. Key ResultsWe generated a dataset that includes 83 taxa; of these, we classified 18 as casual, 35 as naturalized (but not invasive), and 30 as invasive. Using this dataset, we found I) weak or no phylogenetic clustering of non-native ferns, II) some regions are overrepresented as sources and recipients of introductions, III) climatic niches are often conserved between native and introduced ranges, but can differ between introductions, IV) naturalized ferns have smaller genomes, and V) invaders have higher ploidal levels. ConclusionsWe integrated regional floras, occurrence and climate data, phylogeny, and cytology to test fundamental hypotheses regarding the colonization success of ferns. This study provides insights into the ecological, genomic, and phylogenetic features associated with the colonization of new habitats by non-native ferns, a largely overlooked portion of non-native plant taxa.

Reduced competition or facilitation between kin relative to nonkin can improve plant performance, particularly under resource-limited conditions. Understanding whether kin interactions differ between invasive and native species may provide insights into the mechanisms underlying the persistence and spread of invasive species, particularly for species that spread clonally. To explore this, we conducted a greenhouse experiment using the invasive Alternanthera philoxeroides and its native congener A. sessilis in China. For both species, we grew central plants without or with neighbors, and for the latter we had three intraspecific neighbor kinship treatments (kin only, nonkin only, and both kin and nonkin [mixed] neighbors). To test whether kinship effects are affected by resource limitation, we grew the plants under two watering conditions (well-watered and drought-stressed). Our findings revealed that at both the group (i.e., pot-level) and individual levels, invasive plants had a higher biomass production and experienced a less negative relative neighbor effect in kin groups than in nonkin groups, while these patterns were reversed in the native species. Although aboveground architecture of central plants did not differ significantly between kin and nonkin neighbors in either species, neighbor plants of the invasive species produced fewer nodes in kin groups than in nonkin groups, while the reverse was true for the native species. These patterns were not affected by the watering treatment. Together, these results indicate that while the native plants has stronger kin competition, the invasive species has reduced kin competition. Such reduced competition among kin in the invasive Alternanthera philoxeroides may enhance its population dominance and facilitate its spread.

Montane plant populations are experiencing novel conditions due to climate change. Furthermore, climate change is causing increased climate perturbations, such as the 2012-2016 drought in the western US, remarkable in its aridity, longevity, and warmer temperatures. This drought provided an opportunity to understand how montane populations respond to extreme perturbations, including at range limits. We resurrected seeds of the endemic annual plant Erythranthe laciniata, collected in 2008 or earlier (before the drought) and in 2014 (the height of the drought), in a common garden experiment to understand how drought influenced evolution in contemporary field conditions. The study included nine populations across the species range, including range edges. Over 2,100 replicates were sown in three common gardens at natural populations at low, central, and high elevations. We recorded phenology and flower production to estimate lifetime fitness. This experiment took place in 2021, a year with low precipitation and high temperatures. We found higher fitness in the drought generation at the high garden, while both generations showed similar fitness at the central and low gardens. We detected climate adaptation at the low and high gardens, and rapidly evolved faster phenology at the high garden. Lifetime fitness was substantially lower at lower gardens overall, even for low-elevation populations. Low-elevation populations outperformed central populations at the central garden, suggesting adaptive mismatch. Together, these results indicate rapid contemporary adaptation that is beneficial at the leading edge of the species range. Nevertheless, low fitness at lower elevations may foreshadow range contraction under continued climate change.

Background and AimsComplex genomic histories driven by hybridization and polyploidy can shape key plant traits such as defense, stress tolerance, and toxicity, particularly in Amaran-thaceae, which includes crops such as quinoa and spinach. Within this family, white goosefoot (Chenopodium album) is both a widespread agricultural weed and a traditional food resource. However, its evolutionary history is complicated by discordant signals among genomic markers within the C. album complex, comprising diploid, tetraploid, and hexaploid taxa. Here, we tested whether reticulate evolution underlies this genome-wide discordance. MethodsUsing genome-scale phylogenomic data, we analysed 2,298 conserved nuclear loci (BUSCO genes) across 27 Amaranthaceae genomes. Both single- and multicopy gene families were included to capture signals of gene duplication, incomplete lineage sorting, and hybridization. Complementary phylogenomic approaches were used to evaluate whether the evolutionary history is best supported by strictly bifurcating relationships or by reticulate evolution. Key ResultsA consistent C. album lineage was recovered, comprising tetraploid and hexaploid C. album cytotypes together with C. suecicum, C. strictum, C. formosanum, C. acuminatum, and C. opulifolium. Phylogenetic discordance was concentrated within Chenopodium, particularly around the C. album and C. quinoa lineages. Models incorporating hybridization fit better than strictly bifurcating relationships, supporting at least two reticulation events. Hybridization signals were detected in 271 loci in tetraploid and 270 in hexaploid C. album, of which 232 were shared, indicating a shared hybrid origin rather than independent lineages. ConclusionsThe evolutionary history of the C. album lineage is best explained by reticulate processes involving hybridization and polyploidy. Conserved nuclear loci retain persistent signatures of these events, helping to resolve complex evolutionary histories in polyploid plant systems.

Forest trees pose numerous potential challenges to phylogenomic inference. Their large effective population sizes and relatively long generation times lead to deep allele coalescence and consequently incomplete lineage sorting (ILS), which biases inferences of divergence times toward older ages and introduces gene tree discordance. Deep phylogenetic divergences, reaching back into the Paleocene, introduce reference-mapping biases. Introgression--the movement of genes between lineages--may result in different phylogenies being inferred depending on which individuals are included in analysis, even if the plurality of the genome favors the divergence history unaffected by introgression. These factors influence phylogenetic inference across the Tree of Life but are particularly prevalent in forest trees. Oaks (Quercus) are notable for all three influences. In addition, our knowledge of the oak phylogeny is currently based strongly on restriction site associated DNA sequencing (RADseq) datasets published over the past decade, which may introduce additional sources of uncertainty. In this chapter, we analyze a 322-species RADseq dataset and genome resequencing data from across the genus to address sources of uncertainty in our understanding of the global oak phylogeny, which we hope will serve as a model for other research groups working on comparable woody plant groups.

Understanding evolutionary relationships in hyperdiverse plant groups remains a major challenge in systematics. The orchid genus Bulbophyllum, the second largest genus of flowering plants, represents an exceptional example of phylogenetic and morphological complexity. Relationships, particularly within the species-rich Asian clade, have remained poorly resolved due to extensive morphological variation and limited resolution in previous phylogenetic studies. Here, we reconstructed phylogenetic relationships using 63 plastid genes from 355 specimens representing 322 species and 65 of the 97 recognised sections of Bulbophyllum. Our analyses confirmed that the genus comprises five major evolutionary lineages comprised of species predominantly from Australasia, Madagascar, Continental Africa, Neotropics, and Asia. We provide the first robust phylogenetic evidence for a dichotomous split within the Asian clade into two well-supported lineages: the Asian-Malesian clade and the Malesian-Papuasian clade, with the latter containing a strongly supported Papuasian subclade. Additionally, this study supports the monophyly of several currently recognised sections while clarifying relationships in previously problematic groups. This study provides the most comprehensive plastid-based phylogenomic framework for Bulbophyllum to date and establishes a foundation for future taxonomic revision and integrative analyses of diversification and trait evolution within this hyperdiverse genus.

Most plants require animal pollination to reproduce, prompting concern that pollinator declines immediately threaten plant populations. This concern is warranted if pollinator-mediated seed losses cause declines in plant population growth rates ({lambda}). However, demographic trade-offs might reduce the risk of population decline if seed loss improves performance elsewhere in the life cycle. We conducted a multi-year pollination manipulation on four species and measured how demographic vital rates and {lambda} responded. Seed responses did not predict net changes in {lambda}. Reduced pollination decreased seed production, but only caused a net decrease in {lambda} in one species; in the others, improved survival buffered {lambda}. Increased pollination boosted seed production, but at a cost to survival that caused a net reduction in {lambda} in three species. Our results highlight the importance of demographic trade-offs for understanding the impacts of pollinator declines on plant biodiversity and, more broadly, the population-level impacts of changing mutualisms.

BackgroundPlants are exposed to various environmental challenges. With ongoing climate change, droughts and insect outbreaks are expected to become more frequent. Thus, a better understanding is needed of how different plant species respond to such single and combined challenges. This study investigated common versus species-specific responses to environmental challenges in three perennial plant species of different growth forms and whether responses differ intraspecifically among accessions. Clones of different accessions of the herbaceous species Tanacetum vulgare, the woody vine Solanum dulcamara, and the tree Populus nigra were subjected to similar control, herbivory, drought, and combined (drought and herbivory) treatments for the same periods. After the exposure, concentrations of foliar phytohormones and various morphological traits were measured. ResultsAcross all species, several foliar phytohormones and one of ten morphological traits responded consistently to the environmental challenges. Jasmonoyl-isoleucine was induced by herbivory and the combined treatment, abscisic acid (ABA) by drought and the combined treatment, and indole acetic acid by the combined treatment in all species. Root mass remained unchanged in all species. However, structural equation models (SEMs) revealed a shared regulatory pathway across species in which ABA connected treatment and root mass, indicating a common hormonal response potentially linking challenges to growth responses. Despite these common patterns, species-specific responses were pronounced. In P. nigra, a unique induction of salicylic acid was found under the combined treatment, while aboveground mass and root-shoot ratio remained unaffected by any treatment, in contrast to the other two species. Species-specific SEMs further indicated distinct phytohormone-mediated pathways underlying morphological variation. Phenotypic plasticity reflected these species-specific patterns, with none of the phytohormones or morphological traits exhibiting uniform plasticity across species. Intraspecific variation further shaped responses, as phytohormone and morphological trait plasticity depended on accession, indicating substantial accession-specific plant responses. ConclusionsOur results indicate that some responses to comparable challenges may be conserved across species, while others are species-specific. The combined treatment elicited the most pronounced responses, and such complex responses may become more frequent under current global change. Our study highlights that comprehensive understanding of plant responses requires systematic comparisons at both interspecific and intraspecific scales.

Timing of flowering is shifting with climate change. Although climate-driven shifts in phenology sometimes affect seed production, whether changing phenology will scale up to affect population dynamics of long-lived plants remains largely unknown, particularly under changing precipitation. Understanding how phenology affects persistence and extinction risk is a pressing need given contemporary biodiversity loss. We combined nearly a decade of demographic censuses and a four-year phenological survey in a rainfall manipulation experiment to examine the effects of experimental drought and irrigation on flowering phenology, vital rates (e.g., survival and individual growth), and population growth in the perennial herb Lomatium utriculatum. We found that drought advanced flowering by 3.3 days on average, and that earlier-flowering plants produced more seeds regardless of treatment. However, both rainfall treatments reduced seed production compared to controls. We quantified the phenology-mediated and direct, non-phenological effects of rainfall manipulation on population growth rates using integral projection models and a life table response experiment. Drought and irrigation increased {lambda} through increased individual growth, but these effects were partially negated by treatment-driven declines in seed output. In contrast, changes to seed production resulting from shifting flowering times had negligible effects on population growth. Our results suggest that climate-driven phenological shifts may only marginally impact population dynamics in perennial plants and highlight that assessing phenologys consequences for persistence under climate change must also account for direct demographic effects of the climate driver(s) themselves. SignificanceWill changing flowering times under climate change increase extinction risk in plant populations? Despite well-documented earlier flowering and its influence on the number of offspring produced, how changing flowering times will affect population growth or decline is still mostly unknown. We study this in a perennial wildflower subject to changes in rainfall. While we found that drought meant earlier flowering and that, all else equal, early flowering meant more seeds, these effects only marginally affected population growth. Instead, population growth was influenced mostly by rainfall-driven changes to individual plant growth. While shifting flowering times remain an important indicator of climate change, assessing extirpation in plants requires considering flowering times as only one of many life cycle processes changing with climate.

Background and AimsClimate gradients influence seed morphology, emergence, and early life-history traits with cumulative impacts to individual fitness. For ex situ seed collections, which represent an invaluable repository of potential trait information for species management and conservation, climate data can guide preservation of adaptive variation and inform deployment strategies for restoration. Here we leverage a range-wide ex situ seed collection of critically endangered black ash seeds (Fraxinus nigra) to evaluate how climatic gradients shape variation in morphology and early life-history. MethodsTo test how climate of origin, seed morphology, and early life-history interact to impact first year fitness, high-throughput X-ray imaging and neural network-based segmentation were used to quantify variation in seed morphology for 701 maternal lineages spanning 76 populations across the range of F. nigra. Following this, a subset of seeds were used to establish a common garden experiment and quantify variation in emergence, early life-history transitions, and their cumulative impact to first-year survival and growth. ResultsOn average, differences within-population explained [~]43% of the variability in seed morphology, while among-population differences explained [~]14%. This suggests that substantial genetic variation exists within populations for natural selection to act upon and differences have evolved among populations. Climate associations indicated warmer and drier environments predicted heavier seeds with faster developmental transitions and increased first-year height. Together, climate of origin, seed mass, and timing of developmental transitions best predicted cumulative fitness, with populations from more continental environments exhibiting greater survival and first-year height accumulation on average. ConclusionsOverall, these results highlight the importance of climate of origin, seed traits, and early developmental transitions to first-year fitness in a perennial tree species. This work demonstrates how ex situ collections can be used to identify climatically structured trait variation and guide conservation strategies aimed at maintaining adaptive potential under environmental change.