Greenhouse Common Garden Research Articles

Teosinte populations exhibit weak local adaptation to their rhizosphere biota despite strong effects of biota source on teosinte fitness and traits.

While biotic interactions often impose selection, species and popula- tions vary in whether they are locally adapted to biotic interactions. Evo- lutionary theory predicts that environmental conditions drive this variable local adaptation by altering the fitness impacts of species interactions. To investigate the influence of an environmental gradient on adaptation be- tween a plant and its associated rhizosphere biota, we cross-combined teosinte (Zea mays ssp. mexicana) and rhizosphere biota collected across a gradient of decreasing temperature, precipitation, and nutrients in a greenhouse common garden experiment. We measured both fitness and phenotypes expected to be influenced by biota, including concentrations of nutrients in leaves. Independent, main effects of teosinte and biota source explained most variation in teosinte fitness and traits. For example, biota from warmer sites provided population-independent fitness benefits across teosinte hosts. Effects of biota that depended on teosinte genotype were often not specific to their local hosts, and most traits had similar relation- ships to fitness across biota treatments. However, we found weak patterns of local adaptation between teosinte and biota from colder sites, suggest- ing environmental gradients may alter the importance of local adaptation in teosinte-biota interactions, as evolutionary theory predicts.

Honest floral signalling traits vary across and within populations in an insect‐pollinated plant

Abstract In flowering plants that produce concealed rewards, pollinator foraging preferences may select for floral advertisement traits that are correlated with rewards. To date, studies have not focused on the potential for honest signals to vary across populations, which could occur due to differences in pollinator communities or plant mating system. We tested for variation in honest signals across and within populations and mating systems in Arabis alpina, a broadly distributed arctic‐alpine perennial herb that is visited by a variable community of insects. In a greenhouse common garden, we tested for correlations between corolla area, floral scent and nectar volume in 29 populations. In 12 field populations, we examined variation in pollen limitation and corolla area. Across and within populations and mating systems, larger flowers generally produced more nectar. Total scent emission was not correlated with nectar production, but two compounds—phenylacetaldehyde and benzyl alcohol—may be honest signals in some populations. Corolla area was correlated with pollen limitation only across populations. Our results suggest that honest signals may be similar across populations but may not result from contemporary direct selection on floral advertisements. Read the free Plain Language Summary for this article on the Journal blog.

Reforestation of high elevation pines: Direct seeding success depends on seed source and sowing environment.

Forest persistence in regions impacted by increasing water and temperature stress will depend upon species' ability to either rapidly adjust to novel conditions or migrate to track ecological niches. Predicted, rapid climate change is likely to outpace the adaptive and migratory capacity of long-lived isolated tree species, and reforestation may be critical to species' persistence. Facilitating persistence both within and beyond a species' range requires identification of seed lots best adapted to the current and future conditions predicted with rapid climate change. We evaluate variation in emergent seedling performance that leads to differential survival among species and populations for three high elevation five-needle pines. We paired a fully reciprocal field common garden experiment with a greenhouse common garden study to (1) quantify variation in seedling emergence and functional traits, (2) ask how functional traits affect performance under different establishment conditions, and (3) evaluate whether trait and performance variation demonstrates local adaptation and plasticity. Among study species-limber, Great Basin bristlecone, and whitebark pines-we found divergence in emergence and functional traits, though soil moisture was the strongest driver of seedling emergence and abundance across all species. Generalist limber pine had a clear emergence advantage as well as traits associated with drought adaptation, while edaphic specialist bristlecone pine was characterized by low emergence yet high early survival once established. Despite evidence for edaphic specialization, soil characteristics alone did not explain bristlecone success. Across species, trait-environment relationships provided some evidence for local adaptation in drought-adapted traits, but we found no evidence of local adaptation in emergence or survival at this early life stage. For managers looking to promote persistence, sourcing seed from drier environments is likely to impart greater drought resistance into reforestation efforts through strategies such as greater root investment, increasing the probability of early seedling survival. This research demonstrates, through a rigorous reciprocal transplant experimental design, that it may be possible to select climate- and soil-appropriate seed sources for reforestation. However, planting success will ultimately rely on a suitable establishment environment, requiring careful consideration of interannual climate variability for management interventions in these climate and disturbance-impacted tree species.

Phenotypic clines in herbivore resistance and reproductive traits in wild plants along an agricultural gradient.

The conversion of natural landscapes to agriculture is a leading cause of biodiversity loss worldwide. While many studies examine how landscape modification affects species diversity, a trait-based approach can provide new insights into species responses to environmental change. Wild plants persisting in heavily modified landscapes provide a unique opportunity to examine species' responses to land use change. Trait expression within a community plays an important role in structuring species interactions, highlighting the potential implications of landscape mediated trait changes on ecosystem functioning. Here we test the effect of increasing agricultural landscape modification on defensive and reproductive traits in three commonly occurring Brassicaceae species to evaluate plant responses to landscape change. We collected seeds from populations at spatially separated sites with variation in surrounding agricultural land cover and grew them in a greenhouse common garden, measuring defensive traits through an herbivore no-choice bioassay as well as reproductive traits such as flower size and seed set. In two of the three species, plants originating from agriculturally dominant landscapes expressed a consistent reduction in flower size and herbivore leaf consumption. One species also showed reduced fitness associated with increasingly agricultural landscapes. These findings demonstrate that wild plants are responding to landscape modification, suggesting that the conversion of natural landscapes to agriculture has consequences for wild plant evolution.

Responses of floral traits to water limitation across soil types and along climatic gradients

Understanding how environmental changes affect plant-pollinator interactions is of increasing importance given that new environmental extremes continue to be reached. Drought may be a particularly destabilizing force for plant-pollinator interactions because moisture availability can affect floral trait expression. Past work has found variability in how some floral traits respond to moisture limitation, motivating work on the potential drivers of the variability in responses. Here we asked how flower size and nectar provisioning to pollinators change with moisture limitation, and if seed provenance mediates trait responses. Specifically, we conducted a greenhouse common garden using seeds of the serpentine-tolerant annual Antirrhinum vexillocalyculatum collected from pairs of populations on and off drought-prone serpentine soil and along moisture and temperature gradients. We experimentally quantified responses to two moisture treatments, and we found no evidence that seeds from different soil types vary in drought responses. However, plants from seeds from different temperature regimes varied in the way that water limitation affected nectar volume, and those from different precipitation regimes varied in the way that water limitation affected flower size. Water limitation reduced floral display and reward for plants from some populations but increased trait means in other populations. This suggests that drought will reduce floral resources for pollinators overall, but plants from some environments may continue to invest in pollination mutualisms under moisture limitation.

Climate-driven convergent evolution in riparian ecosystems on sky islands

Climate-induced evolution will determine population persistence in a changing world. However, finding natural systems in which to study these responses has been a barrier to estimating the impact of global change on a broad scale. We propose that isolated sky islands (SI) and adjacent mountain chains (MC) are natural laboratories for studying long-term and contemporary climatic pressures on natural populations. We used greenhouse common garden trees to test whether populations on SI exposed to hot and dry climates since the end of the Pleistocene have phenotypically diverged from populations on MC, and if SI populations have converged in these traits. We show: (1) populations of Populus angustifolia from SI have diverged from MC, and converged across SI, in reproductive and productivity traits, (2) these traits (cloning and aboveground biomass, respectively) are significantly correlated, suggesting a genetic linkage between them, and (3) the trait variation is driven by both natural selection and genetic drift. These shifts represent potentially beneficial phenotypes for population persistence in a changing world. These results suggest that the SI–MC comparison is a natural laboratory, as well as a predictive framework, for studying long-term responses to climate change across the globe.

Greater flowering and response to flooding inLythrum virgatumthanL. salicaria(purple loosestrife)

Newly introduced trait diversity can spur rapid evolution and facilitate local adaptation in the introduced plant Lythrum salicaria. The horticultural plant L. virgatum might further introduce meaningful trait variation by escaping into established L. salicaria populations or by hybridizing with L. salicaria. Although many experiments have focused on L. salicaria genotypes, relatively little is known about L. virgatum ecology. We used a greenhouse common garden to compare traits and flood response of L. salicaria and L. virgatum collected from two sources each in their native range. We tested the hypotheses that these two wetland taxa have comparable responses to flooding (inundation), and that flood tolerance correlated to higher fitness. Flooding produced stronger stress responses in L. virgatum. Compared to L. salicaria, L. virgatum shifted more aboveground allocation away from reproduction, decreased inflorescence biomass by 40% more, and produced 7% more stem aerenchymatous phellum, a specialized tissue that maintains aeration. Despite these more pronounced responses to flooding stress, L. virgatum had higher fitness (inflorescence biomass and reproductive allocation) than L. salicaria. Overall, L. virgatum differed from L. salicaria in functionally important ways. Lythrum virgatum persisted under flooding and produced more reproductive biomass than L. salicaria under both flooded and non-flooded conditions. However, inundation stressed L. virgatum more than L. salicaria. Lythrum virgatum is likely able to establish into the wetland habitats in which L. salicaria prevails but may possess broader habitat tolerances.

Climate and the biotic community structure plant resistance across biogeographic groups of yellow monkeyflower.

Characterizing correlates of phytochemical resistance trait variation across a landscape can provide insight into the ecological factors that have shaped the evolution of resistance arsenals. Using field-collected data and a greenhouse common garden experiment, we assessed the relative influences of abiotic and biotic drivers of genetic-based defense trait variation across 41 yellow monkeyflower populations from western and eastern North America and the United Kingdom. Populations experience different climates, herbivore communities, and neighboring vegetative communities, and have distinct phytochemical resistance arsenals. Similarities in climate as well as herbivore and vegetative communities decline with increasing physical distance separating populations, and phytochemical resistance arsenal composition shows a similarly decreasing trend. Of the abiotic and biotic factors examined, temperature and the neighboring vegetation community had the strongest relative effects on resistance arsenal differentiation, whereas herbivore community composition and precipitation have relatively small effects. Rather than simply controlling for geographic proximity, we jointly assessed the relative strengths of both geographic and ecological variables on phytochemical arsenal compositional dissimilarity. Overall, our results illustrate how abiotic conditions and biotic interactions shape plant defense traits in natural populations.

Local adaptation of seed and seedling traits along a natural aridity gradient may both predict and constrain adaptive responses to climate change.

Variation in seed and seedling traits underlies how plants interact with their environment during establishment, a crucial life history stage. We quantified genetic-based variation in seed and seedling traits in populations of the annual plant Plantago patagonica across a natural aridity gradient, leveraging natural intraspecific variation to predict how populations might evolve in response to increasing aridity associated with climate change in the Southwestern U.S. We quantified seed size, seed size variation, germination timing, and specific leaf area in a greenhouse common garden, and related these traits to the climates of source populations. We then conducted a terminal drought experiment to determine which traits were most predictive of survival under early-season drought. All traits showed evidence of clinal variation-seed size decreased, germination timing accelerated, and specific leaf area increased with increasing aridity. Populations with more variable historical precipitation regimes showed greater variation in seed size, suggestive of past selection shaping a diversified bet-hedging strategy mediated by seed size. Seedling height, achieved via larger seeds or earlier germination, was a significant predictor of survival under drought. We documented substantial interspecific trait variation as well as clinal variation in several important seed and seedling traits, yet these slopes were often opposite to predictions for how individual traits might confer drought tolerance. This work shows that plant populations may adapt to increasing aridity via correlated trait responses associated with alternative life history strategies, but that trade-offs might constrain adaptive responses in individual traits.

Among- and within-population variation in morphology, rewards, and scent in a hawkmoth-pollinated plant.

Floral scent is a complex trait that mediates many plant-insect interactions, but our understanding of how floral scent variation evolves, either independently or in concert with other traits, remains limited. Assessing variation in floral scent at multiple levels of biological organization and comparing patterns of variation in scent to variation in other floral traits can contribute to our understanding of how scent variation evolves in nature. We used a greenhouse common garden experiment to investigate variation in floral scent at three scales-within plants, among plants, and among populations-and to determine whether scent, alone or in combination with morphology and rewards, contributes to population differentiation in Oenothera cespitosa subsp. marginata. Its range spans most of the biomes in the western United States, such that variation in both the abiotic and biotic environment could contribute to trait variation. Multiple analytical approaches demonstrated substantial variation among and within populations in compound-specific and total floral scent measures. Overall, populations were differentiated in morphology and reward traits and in scent. Across populations, coupled patterns of variation in linalool, leucine-derived compounds, and hypanthium length are consistent with a long-tongued moth pollination syndrome. The considerable variation in floral scent detected within populations suggests that, similar to other floral traits, variation in floral scent may have a heritable genetic component. Differences in patterns of population differentiation in floral scent and in morphology and rewards indicate that these traits may be shaped by different selective pressures.

Flexible drought deciduousness in a neotropical understory herb.

PremiseAdaptive divergence across environmental gradients is a key driver of speciation. Precipitation seasonality gradients are common in the tropics, yet drought adaptation is nearly unexplored in neotropical understory herbs. Here, we examined two recently diverged neotropical spiral gingers, one adapted to seasonal drought and one reliant on perennial water, to uncover the basis of drought adaptation.MethodsWe combined ecophysiological trait measurements in the field and greenhouse with experimental and observational assessments of real‐time drought response to determine how Costus villosissimus (Costaceae) differs from C. allenii to achieve drought adaptation.ResultsWe found that drought‐adapted C. villosissimus has several characteristics indicating flexible dehydration avoidance via semi‐drought‐deciduousness and a fast economic strategy. Although the two species do not differ in water‐use efficiency, C. villosissimus has a more rapid growth rate, lower leaf mass per area, lower stem density, higher leaf nitrogen, and a strong trend of greater light‐saturated photosynthetic rates. These fast economic strategy traits align with both field‐based observations and experimental dry‐down results. During drought, C. villosissimus displays facultative drought‐deciduousness, losing lower leaves during the dry season and rapidly growing new leaves in the wet season.ConclusionsWe revealed a drought adaptation strategy that has not, to our knowledge, previously been documented in tropical herbs. This divergent drought adaptation evolved recently and is an important component of reproductive isolation between C. villosissimus and C. allenii, indicating that adaptive shifts to survive seasonal drought may be an underappreciated axis of neotropical understory plant diversification.

Emission Rates of Species-Specific Volatiles Vary across Communities of Clarkia Species: Evidence for Multimodal Character Displacement.

AbstractA current frontier of character displacement research is to determine whether displacement occurs via multiple phenotypic pathways and varies across communities with different species compositions. Here, we conducted the first test for context-dependent character displacement in multimodal floral signals by analyzing variation in floral scent in a system that exhibits character displacement in flower size and that has multiple types of sympatric communities. In a greenhouse common garden experiment, we measured quantitative variation in volatile emission rates of the progeny of two species of Clarkia from replicated parental communities that contain one, two, or four Clarkia species. The first two axes of a constrained correspondence analysis, which explained 24% of the total variation in floral scent, separated the species and community types. Of the 23 compounds that were significantly correlated with these axes, nine showed patterns consistent with character displacement. Two compounds produced primarily by C. unguiculata and two compounds produced primarily by C. cylindrica were emitted in higher amounts in sympatry. Character displacement in some volatiles varied across sympatric parental communities and occurred in parallel with displacement in flower size, demonstrating that this evolutionary process can be context dependent and may occur through multiple pathways.

Extensive population-level sampling reveals clinal variation in (R)-(−)-linalool produced by the flowers of an endemic evening primrose, Oenothera harringtonii

The study of floral trait diversity has a long history due to its role in angiosperm diversification. While many studies have focused on visual traits including morphology and color, few have included floral scent despite its importance in pollination. Of the studies that have included floral scent, sampling has been limited and rarely explores variation at the population level. We studied geographic variation in the flowers of Oenothera harringtonii, a rare plant endemic to a vulnerable shortgrass prairie habitat, whose population structure and conservation status are well studied. The self-incompatible flowers of O. harringtonii open at dusk, produce nectar and a strong fragrance, and are pollinated by hawkmoths. We collected floral trait (morphology, scent chemistry and emission rates) data from 650 individuals from 19 wild populations to survey floral variation across the entire range of this species. Similarly, we collected floral data from 49 individuals grown in a greenhouse common garden, to assess whether variation observed in the field is consistent when environment factors (temperature, watering regime, soil) are standardized. We identified 35 floral volatiles representing 5 biosynthetic classes. Population differentiation was stronger for floral scent chemistry than floral morphology. (R)-(−)-linalool was the most important floral trait differentiating populations, exhibiting clinal variation across the distribution of O. harringtonii without any correlated shifts in floral morphology. Populations in the north and west produced (R)-(−)-linalool consistently, those in the east and south largely lacked it, and populations at the center of the distribution were polymorphic. Floral scent emissions in wild populations varied across four years but chemical composition was largely consistent over time. Similarly, volatile emission rates and chemical composition in greenhouse-grown plants were consistent with those of wild populations of origin. Our data set, which represents the most extensive population-level survey of floral scent to date, indicates that such sampling may be needed to capture potentially adaptive geographic variation in wild populations.

Genetically based adaptive trait shifts at an expanding mangrove range margin

Neotropical black mangrove (Avicennia germinans) is expanding poleward into temperate salt marsh along Atlantic Florida, USA, with field evidence of trait shifts consistent with greater cold tolerance within range margin populations. However, whether these shifts have a genetic basis remains unanswered. To address this gap, we measured multiple phenotypic traits of twenty A. germinans maternal cohorts from areas in both the Atlantic Florida range core and margin in a 2-year greenhouse common garden with annual temperatures analogous to range margin conditions. Compared to those from the range core, range margin cohorts survived in greater numbers, established (i.e., produced first true leaves) more quickly, and were less stressed under winter temperatures. Range margin cohorts were not taller, but invested more into lateral growth and biomass accumulation that presumably reflects adaptation to their colder and open-canopy environment. Range margin cohorts also exhibited leaf traits consistent with greater resource acquisition that may compensate for a shorter growing season and reduced light quality at higher latitude. Our results suggest that genetically based phenotypic differences better enable these range margin mangroves to thrive within their stressful environment and may facilitate further poleward expansion. An improved understanding of adaptive trait variation within ecologically important mangrove foundation species should also help inform coastal restoration initiatives.

Genomic and common garden approaches yield complementary results for quantifying environmental drivers of local adaptation in rubber rabbitbrush, a foundational Great Basin shrub.

The spatial structure of genomic and phenotypic variation across populations reflects historical and demographic processes as well as evolution via natural selection. Characterizing such variation can provide an important perspective for understanding the evolutionary consequences of changing climate and for guiding ecological restoration. While evidence for local adaptation has been traditionally evaluated using phenotypic data, modern methods for generating and analyzing landscape genomic data can directly quantify local adaptation by associating allelic variation with environmental variation. Here, we analyze both genomic and phenotypic variation of rubber rabbitbrush (Ericameria nauseosa), a foundational shrub species of western North America. To quantify landscape genomic structure and provide perspective on patterns of local adaptation, we generated reduced representation sequencing data for 17 wild populations (222 individuals; 38,615 loci) spanning a range of environmental conditions. Population genetic analyses illustrated pronounced landscape genomic structure jointly shaped by geography and environment. Genetic‐environment association (GEA) analyses using both redundancy analysis (RDA) and a machine‐learning approach (Gradient Forest) indicated environmental variables (precipitation seasonality, slope, aspect, elevation, and annual precipitation) influenced spatial genomic structure and were correlated with allele frequency shifts indicative of local adaptation at a consistent set of genomic regions. We compared our GEA‐based inference of local adaptation with phenotypic data collected by growing seeds from each population in a greenhouse common garden. Population differentiation in seed weight, emergence, and seedling traits was associated with environmental variables (e.g., precipitation seasonality) that were also implicated in GEA analyses, suggesting complementary conclusions about the drivers of local adaptation across different methods and data sources. Our results provide a baseline understanding of spatial genomic structure for E. nauseosa across the western Great Basin and illustrate the utility of GEA analyses for detecting the environmental causes and genetic signatures of local adaptation in a widely distributed plant species of restoration significance.

Short‐ and long‐term effects of nutrient enrichment on salt marsh plant production and microbial community structure

Abstract Nutrient enrichment impacts ecosystems globally. Population history, especially past resource environments, of numerically dominant plant species may affect their responses to subsequent changes in nutrient availability. Eutrophication can also alter plant–microbe interactions via direct effects on associated microbial communities or indirect effects on dominant species’ biomass production/allocation as a result of modified plant–soil interactions. We combined a greenhouse common garden and a field reciprocal transplant of a salt marsh foundation species (Spartina alterniflora) within a long‐term, whole‐ecosystem, nutrient‐enrichment study to determine whether enrichment affects plant production and microbial community structure differently depending on plant population history. For the greenhouse portion, we collected 20 S. alterniflora genotypes—10 from an enriched creek that had received elevated nutrient inputs for 10 years and 10 from an unenriched reference creek—and reared them in a common garden for 1 year. For the field portion, we conducted a 2‐year, fully crossed reciprocal transplant experiment with two gardens each at the enriched and unenriched sites; we examined the effects of source site (i.e. population history), garden site and plant genotype. After 2 years, plants in enriched gardens had higher above‐ground biomass and altered below‐ground allocation compared to plants in unenriched gardens. However, performance also depended on plant population history: plants from the enriched site had decreased above‐ground and rhizome production compared to plants from the unenriched site, most notably in unenriched gardens. In addition, almost all above‐ and below‐ground traits varied depending on plant genotypic identity. Effects of nutrient enrichment on the associated microbial community were also pronounced. Following 1 year in common garden, microbial community structure varied by plant population history and S. alterniflora genotypic identity. However, at the end of the reciprocal transplant, microbial communities differed primarily between enriched and unenriched gardens. Synthesis. Nutrient enrichment can impact plant foundation species and associated soil microbes in the short term. Most importantly, nutrient enrichment can also have long‐lasting effects on plant populations and associated microbial communities that potentially compromise their ability to respond to changing resource conditions in the future.

Submerged Aquatic Vegetation Species and Populations Within Species Respond Differently to Environmental Stressors Common in Restorations.

Submerged aquatic vegetation (SAV) improves environmental conditions by acting as a sediment stabilizer and nutrient retention tool; therefore, reintroduction of SAV is a common freshwater restoration goal. Initial plant establishment is often difficult in suboptimal conditions, and planting material with specific traits may increase establishment rates. Here we evaluate the variability in plant traits based on collection location. We find consistent differences in traits of plants collected from different natural water bodies, and those differences persist in plants grown from seeds under common garden greenhouse conditions-presumably because of genetic differentiation. In three separate mesocosm experiments, we tested the interactive impacts of collection location and environmental condition (control conditions, reduced light, elevated nutrients, or a combination of reduced light and elevated nutrients) on plant reproduction and on traits that might indicate future restoration success (plant height, number of leaves, and rhizome diameter). In most cases, plant traits at the end of the experiments varied by collection location, environmental condition, and an interaction between the two. The best performing plants also depended on response variable (e.g., plant height or number of new shoots produced). Together these results suggest that unpredictable environmental conditions at restoration sites will make selection of a single high-performing plant source difficult, so we suggest incorporating a diverse set of collection locations to increase the probability of incorporating desirable traits.

A Test of the Inadvertent Uptake Hypothesis Using Plant Species Adapted to Serpentine Soil

Serpentine soils are a stressful growing environment for plants, largely due to nutrient deficiencies and high concentrations of toxic heavy metals (e.g., Ni). Plants have evolved various adaptations for tolerating these extreme environments, including metal hyperaccumulation into above-ground tissues. However, the adaptive significance of metal hyperaccumulation is a topic of debate, with several non-mutually-exclusive hypotheses under study. For example, the inadvertent uptake hypothesis (IUH) states that heavy metal accumulation is a consequence of an efficient nutrient-scavenging mechanism for plants growing in nutrient-deficient soils. Thus, it is possible that metal hyperaccumulation is simply a byproduct of non-specific ion transport mechanisms allowing plants to grow in nutrient-deficient soils, such as serpentine soils, while simultaneously tolerating other potentially toxic heavy metals. Furthermore, some nutrient needs are tissue-specific, and heavy metal toxicity can be more pronounced in reproductive tissues; thus, studies are needed that document nutrient and metal uptake into vegetative and reproductive plant tissues across species of plants that vary in the degree to which they accumulate soil metals. To test these ideas, we grew nine plant species that are variously adapted to serpentine soils (i.e., Ni-hyperaccumulating endemic, non-hyperaccumulating endemic, indicator, or indifferent) in a common garden greenhouse experiment. All species were grown in control soils, as well as those that were amended with the heavy metal Ni, and then analyzed for macronutrient (Ca, Mg, K, and P), micronutrient (Cu, Fe, Zn, Mn, and Mo), and heavy metal (Cr and Co) concentrations in their vegetative and reproductive organs (leaves, anthers, and pistils). In accordance with the IUH, we found that hyperaccumulators often accumulated higher concentrations of nutrients and metals compared to non-hyperaccumulating species, although these differences were often organ-specific. Specifically, while hyperaccumulators accumulated significantly more K and Co across all organs, Cu was higher in leaves only, while Mn and Zn were higher in anthers only. Furthermore, hyperaccumulators accumulated significantly more Co and Mo across all organs when Ni was added to the soil environment. Our work provides additional evidence in support of the IUH, and contributes to our understanding of serpentine adaptation in plants.

Phenotypic Trait Variation in Populations of a Global Invader Mayweed Chamomile (Anthemis cotula): Implications for Weed Management

Mayweed chamomile (Anthemis cotula L.) is an annual crop weed that has become a substantial impediment to diversify traditional wheat-based cropping systems such as in the Pacific Northwest (PNW), USA. Some of the broadleaf rotational crops are vulnerable to the weed as they are less competitive or lack compatible herbicides for A. cotula management. Although A. cotula has been present in the PNW for more than a century, traits that contribute to invasiveness and how these vary among the weed populations have not been investigated. We assessed trait variation with a common garden greenhouse experiment by comparing 19 A. cotula populations from the PNW and Kashmir Valley, India. Seeds from individual plants from each population were used to grow plants through their senescence. We measured phenological (e.g., flowering duration), morphological (e.g., plant biomass), reproductive fitness (e.g., number of flower heads), and physiological (floral scent VOCs: volatile organic compounds) traits on individual plants and analyzed the data using mixed-effects models. We found high inter-population variation in most of the traits measured, but the variation was not associated with the geographical distance. Seedling emergence within 30 days of planting ranged from 2 to 49% for PNW populations to 55 to 72% for Indian populations. Flowering duration ranged from 2 to 3 months among populations. Other traits such as initial and final flowering dates, the total number of flower heads, and floral scent VOCs also differed among populations. These trait patterns may indicate a localized adaptation of A. cotula populations to environmental or management regimes, variation arising from anthropogenic or natural dispersal, multiple introductions, genetic drift, or combinations of these. Regardless of the processes, the trait variation suggests that weed management plan for A. cotula may need to be tailored to specific locations instead of a uniformly adopted plan across the region.

An invading annual plant benefits less from soil biota and has reduced competitive power with a resident grass

Abstract Aims Interactions between plants and their soil biota, arbuscular mycorrhizal fungi (AMF) in particular, may play a vital role in the establishment and the range expansion of exotic plants in new environments. However, whether there are post-introduction shifts in dependence on AMF and how dependency interacts with competition remains poorly understood. Methods We conducted a common garden greenhouse experiment to examine how native (USA) and invasive (China) populations of the plant species Plantago virginica, respond to soil biota, and whether these responses change in the presence of a competitor. Important Findings We found that while native populations consistently had a higher AMF colonization rate and benefited from AMF in both biomass and seed production, invasive populations received less benefit from AMF, and even showed reduced biomass with AMF in the presence of a competitor. This low mycorrhizal dependency in invasive populations correlated with greater suppression by an indigenous competitor for the invader. The different responses of the invasive and native populations to AMF suggest that alteration of mycorrhizal dependency has occurred during the invasion of P. virginica into China. Our findings suggest that this reduced dependency incurs a cost during interspecific competition.