Maternal warming affects early life stages of an invasive thistle

Maternal environment can influence plant offspring performance. Understanding maternal environmental effects will help to bridge a key gap in the knowledge of plant life cycles, and provide important insights for species’ responses under climate change. Here we show that maternal warming significantly affected the early life stages of an invasive thistle, Carduus nutans. Seeds produced by plants grown in warmed conditions had higher germination percentages and shorter mean germination times than those produced by plants under ambient conditions; this difference was most evident at suboptimal germination temperatures. Subsequent seedling emergence was also faster with maternal warming, with no cost to seedling emergence percentage and seedling growth. Our results suggest that maternal warming may accelerate the life cycle of this species via enhanced early life‐history stages. These maternal effects on offspring performance, together with the positive responses of the maternal generation, may exacerbate invasions of this species under climate change.     

Air humidity as key determinant of morphogenesis and productivity of the rare temperate woodland fern Polystichum braunii

(1) Most ferns are restricted to moist and shady habitats, but it is not known whether soil moisture or atmospheric water status are decisive limiting factors, or if both are equally important. (2) Using the rare temperate woodland fern Polystichum braunii, we conducted a three‐factorial climate chamber experiment (soil moisture (SM) × air humidity (RH) × air temperature (T)) to test the hypotheses that: (i) atmospheric water status (RH) exerts a similarly large influence on the fern's biology as soil moisture, and (ii) both a reduction in RH and an increase in air temperature reduce vigour and growth. (3) Nine of 11 morphological, physiological and growth‐related traits were significantly influenced by an increase in RH from 65% to 95%, leading to higher leaf conductance, increased above‐ and belowground productivity, higher fertility, more epidermal trichomes and fewer leaf deformities under high air humidity. In contrast, soil moisture variation (from 66% to 70% in the moist to ca. 42% in the dry treatment) influenced only one trait (specific leaf area), and temperature variation (15 °C versus 19 °C during daytime) only three traits (leaf conductance, root/shoot ratio, specific leaf area); RH was the only factor affecting productivity. (4) This study is the first experimental proof for a soil moisture‐independent air humidity effect on the growth of terrestrial woodland ferns. P. braunii appears to be an air humidity hygrophyte that, whithin the range of realistic environmental conditions set in this study, suffers more from a reduction in RH than in soil moisture. A climate warming‐related increase in summer temperatures, however, seems not to directly threaten this endangered species.     

EFFECTS OF PARENTAGE AND SIZE OF THE POLLEN LOAD ON PROGENY PERFORMANCE IN CAMPANULA AMERICANA

To examine the effects of maternal and paternal parentage and the size of the pollen load on seed size and weight and on progeny performance we conducted a controlled crossing experiment using a natural population of Campanula americana. We found that seed size was positively correlated with early seedling performance for all but one of traits we measured (days to emergence), but was not significantly correlated with any of the later vegetative measures or reproductive output. We detected significant effects due to the maternal parent for the vegetative traits days to emergence, days to first leaf, and final plant height, as well as total seed weight, and mean seed weight per fruit. Significant paternal effects were found for all of the seedling traits except number of leaves after vernalization. The progeny from fruits receiving high pollen loads significantly outperformed the progeny from fruits receiving low pollen loads for the traits days to first and second leaf, numbers of leaves after vernalization, and days to first flower. These results not only demonstrate the importance of parentage and seed weight on progeny performance, but also indicate that variations in the size of the pollen load may be important in seedling establishment in natural populations.     

First‐year seedlings and climate change: species‐specific responses of 15 North American tree species

Species will respond individually to climate change and this poses a challenge for modeling climate–vegetation dynamics using broader taxonomic or biogeographical classifications. Additionally, responses to climate and environmental conditions may shift with ontogeny, further complicating efforts to understand the likely rates and directions of vegetation change. We measured emergence, leaf‐out rate, growth, and survival of first‐year seedlings in response to warming, precipitation regime shifts, and seedbed condition (leaf litter presence/absence). We grouped species into three levels of organization (species‐specific, biome‐level and broad taxonomic group) and hypothesized that most metrics of seedling performance would be best described by species‐specific models, as even similar species may respond in vastly different ways to global change. Results showed that the species‐specific model was the best fit for emergence and development rates, whereas growth and survival could be captured through broader groupings, with the broadleaf temperate group exhibiting the greatest growth and conifers the shortest survival times. The sign and magnitude of response to climate and seedbed condition varied with treatment combinations and metric of performance. For example, seedlings grew more in response to warming, but conditions too dry or too wet limited this positive response. Also, warmer temperatures generally increased emergence, development, and growth, but decreased survival, whereas leaf litter presence decreased emergence and slowed development, but increased survival. The results presented here are for first‐year seedlings and in many cases the responses are different from other studies using older plants. Future research and climate vegetation modeling needs to assess performance at multiple development stages and determine where key bottleneck phases for population growth occur for individual species.     

Trait dimensionality and population choice alter estimates of phenotypic dissimilarity

The ecological niche is a multi‐dimensional concept including aspects of resource use, environmental tolerance, and interspecific interactions, and the degree to which niches overlap is central to many ecological questions. Plant phenotypic traits are increasingly used as surrogates of species niches, but we lack an understanding of how key sampling decisions affect our ability to capture phenotypic differences among species. Using trait data of ecologically distinct monkeyflower (Mimulus) congeners, we employed linear discriminant analysis to determine how (1) dimensionality (the number and type of traits) and (2) variation within species influence how well measured traits reflect phenotypic differences among species. We conducted analyses using vegetative and floral traits in different combinations of up to 13 traits and compared the performance of commonly used functional traits such as specific leaf area against other morphological traits. We tested the importance of intraspecific variation by assessing how population choice changed our ability to discriminate species. Neither using key functional traits nor sampling across plant functions and organs maximized species discrimination. When using few traits, vegetative traits performed better than combinations of vegetative and floral traits or floral traits alone. Overall, including more traits increased our ability to detect phenotypic differences among species. Population choice and the number of traits used had comparable impacts on discriminating species. We addressed methodological challenges that have undermined cross‐study comparability of trait‐based approaches. Our results emphasize the importance of sampling among‐population trait variation and suggest that a high‐dimensional approach may best capture phenotypic variation among species with distinct niches.     

Genetic variation in flowering phenology and reproductive performance in a Mediterranean high‐mountain specialist,Armeria caespitosa (Plumbaginaceae)

Adaptive responses to past climate change may play an important role in the persistence of high‐mountain plants, which are vulnerable to global warming. Armeria caespitosa is a high‐mountain plant, endemic to the Iberian Central Range. Differences in abiotic environment along the elevational gradient impose two opposing stress gradients (i.e. water stress and duration of the growth season) on the species. Furthermore, the species is found in two interspersed, contrasting microhabitats (rocky outcrops and dry cryophilic grasslands) that have different effects on plants depending of the elevation. As a result of this, the species shows great among‐population variation in many reproductive and vegetative traits. We used a common garden approach to determine whether this phenotypic variation has a genetic basis or is the result of plastic responses shaped by heterogeneous environmental conditions. Plants from the high‐elevation edge and dry cryophilic grasslands flowered earlier and produced more viable fruits but were smaller. These results confirm that among‐population variation in flowering phenology and reproductive performance traits in A. caespitosa is partially genetically based. The results also show that the stronger selection response in favour of early‐flowering individuals in populations at the low‐elevation edge did not correspond with the greater proportion of early‐flowering individuals. Genetic variability associated with flowering onset may be relevant in coping with the impacts of ongoing global warming. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 384–395.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Climate‐driven reduction of genetic variation in plant phenology alters soil communities and nutrient pools

We examined the hypothesis that climate‐driven evolution of plant traits will influence associated soil microbiomes and ecosystem function across the landscape. Using a foundation tree species, Populus angustifolia, observational and common garden approaches, and a base population genetic collection that spans 17 river systems in the western United States, from AZ to MT, we show that (a) as mean annual temperature (MAT) increases, genetic and phenotypic variation for bud break phenology decline; (b) soil microbiomes, soil nitrogen (N), and soil carbon (C) vary in response to MAT and conditioning by trees; and (c) with losses of genetic variation due to warming, population‐level regulation of community and ecosystem functions strengthen. These results demonstrate a relationship between the potential evolutionary response of populations and subsequent shifts in ecosystem function along a large temperature gradient.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

Taller and larger: shifts in Arctic tundra leaf traits after 16 years of experimental warming

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate‐vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open‐top chambers (OTCs) to passively warm vegetation by 1–2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf δ¹⁵N. Long‐term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect δ¹⁵N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.     

模拟增温和降水变化对北京东灵山辽东栎种子出苗和幼苗生长的影响

气候变化将增加地表平均气温、改变降水格局, 会影响到种子出苗和幼苗生长, 进而影响物种的更新动态。为探讨增温和降水变化对东灵山地区建群树种辽东栎(Quercus mongolica)种子出苗和一年生幼苗生长和适应状况的影响, 该文利用环境控制生长箱开展了温度和降水量的双因素控制实验, 温度设置3个梯度: 月平均气温(对照)、增温2 ℃和增温6 ℃; 降水量设置3个梯度: 月平均降水量(对照)、减水30%和加水30%。结果表明: 1)辽东栎的种子出苗率和一年生幼苗的生长对增温和降水变化的响应不一致, 种子出苗率主要受到降水及其与温度交互作用的影响, 幼苗生长仅受到温度和降水独立作用的影响; 2)春季增温2 ℃或降水量增加均使辽东栎种子出苗期提前; 增温6 ℃与降水量减少的水热组合延迟了种子出苗期并使其存活率和出苗率显著降低, 但在此温度下增加降水量则增加了出苗速率和出苗率。3)增温2 ℃对其生长无显著影响, 增温6 ℃则在不同水分条件下显著地增加了幼苗的比叶面积、抑制了叶的伸长生长, 同时也显著降低了各器官生物量积累, 并减少了幼苗生物量向根的分配; 降水量减少降低了幼苗根生物量, 但未影响总生物量和根冠比, 降水量增加显著促进了幼苗地上部分的生长, 特别是叶的生长。因此, 适当地增温或增加降水量将增加辽东栎幼苗的更新潜力, 但增温和降水量减少导致的干旱化将显著降低幼苗的更新潜力。     

Warming affects different components of plant–herbivore interaction in a simplified community but not net interaction strength

Global warming impacts natural communities through effects on performance of individual species and through changes in the strength of interactions between them. While there is a body of evidence of the former, we lack experimental evidence on potential changes in interaction strengths. Knowledge about multispecies interactions is fundamental to understand the regulation of biodiversity and the impact of climate change on communities. This study investigated the effect of warming on a simplified community consisting of three species: rosy apple aphid Dysaphis plantaginea feeding on plantain, Plantago lanceolata, and a heterospecific neighbouring plant species, perennial ryegrass, Lolium perenne. The aphid does not feed on L. perenne. The experimental design consisted of monocultures and mixtures of L. perenne and P. lanceolata at three temperature levels. We did not find indication for indirect temperature effects on D. plantaginea through changes in leaf nitrogen or relative water content. However, experimental warming affected the life history traits of the aphid directly, in a non‐linear manner. Aphids performed best at moderate warming, where they grew faster and had a shorter generation time. In spite of the increased population growth of the aphids under warming, the herbivory rates were not changed and consequently the plant–herbivore interaction was not altered under warming. This suggests reduced consumption rates at higher temperature. Also plant competition affected the aphids but through an interaction with temperature. We provide proof‐of‐concept that net interactions between plants and herbivores should not change under warming despite direct effects of warming on herbivores when plant–plant interaction are considered. Our study stresses the importance of indirect non–trophic interactions as an additional layer of complexity to improve our understanding of how trophic interactions will alter under climate change.     

Size‐based ecological interactions drive food web responses to climate warming

Predicting climate change impacts on animal communities requires knowledge of how physiological effects are mediated by ecological interactions. Food‐dependent growth and within‐species size variation depend on temperature and affect community dynamics through feedbacks between individual performance and population size structure. Still, we know little about how warming affects these feedbacks. Using a dynamic stage‐structured biomass model with food‐, size‐ and temperature‐dependent life history processes, we analyse how temperature affects coexistence, stability and size structure in a tri‐trophic food chain, and find that warming effects on community stability depend on ecological interactions. Predator biomass densities generally decline with warming – gradually or through collapses – depending on which consumer life stage predators feed on. Collapses occur when warming induces alternative stable states via Allee effects. This suggests that predator persistence in warmer climates may be lower than previously acknowledged and that effects of warming on food web stability largely depend on species interactions.     

The impact of climate change measured at relevant spatial scales: new hope for tropical lizards

Much attention has been given to recent predictions that widespread extinctions of tropical ectotherms, and tropical forest lizards in particular, will result from anthropogenic climate change. Most of these predictions, however, are based on environmental temperature data measured at a maximum resolution of 1 km², whereas individuals of most species experience thermal variation on a much finer scale. To address this disconnect, we combined thermal performance curves for five populations of Anolis lizard from the Bay Islands of Honduras with high‐resolution temperature distributions generated from physical models. Previous research has suggested that open‐habitat species are likely to invade forest habitat and drive forest species to extinction. We test this hypothesis, and compare the vulnerabilities of closely related, but allopatric, forest species. Our data suggest that the open‐habitat populations we studied will not invade forest habitat and may actually benefit from predicted warming for many decades. Conversely, one of the forest species we studied should experience reduced activity time as a result of warming, while two others are unlikely to experience a significant decline in performance. Our results suggest that global‐scale predictions generated using low‐resolution temperature data may overestimate the vulnerability of many tropical ectotherms to climate change.     

Climate change and plant regeneration from seed

At the core of plant regeneration, temperature and water supply are critical drivers for seed dormancy (initiation, break) and germination. Hence, global climate change is altering these environmental cues and will preclude, delay, or enhance regeneration from seeds, as already documented in some cases. Along with compromised seedling emergence and vigour, shifts in germination phenology will influence population dynamics, and thus, species composition and diversity of communities. Altered seed maturation (including consequences for dispersal) and seed mass will have ramifications on life history traits of plants. Predicted changes in temperature and precipitation, and thus in soil moisture, will affect many components of seed persistence in soil, e.g. seed longevity, dormancy release and germination, and soil pathogen activity. More/less equitable climate will alter geographic distribution for species, but restricted migratory capacity in some will greatly limit their response. Seed traits for weedy species could evolve relatively quickly to keep pace with climate change enhancing their negative environmental and economic impact. Thus, increased research in understudied ecosystems, on key issues related to seed ecology, and on evolution of seed traits in nonweedy species is needed to more fully comprehend and plan for plant responses to global warming.     

Short photoperiod reduces the temperature sensitivity of leaf‐out in saplings of Fagus sylvatica but not in horse chestnut

Leaf phenology is one of the most reliable bioindicators of ongoing global warming in temperate and boreal zones because it is highly sensitive to temperature variation. A large number of studies have reported advanced spring leaf‐out due to global warming, yet the temperature sensitivity of leaf‐out has significantly decreased in temperate deciduous tree species over the past three decades. One of the possible mechanisms is that photoperiod is limiting further advance to protect the leaves against potential damaging frosts. However, the “photoperiod limitation” hypothesis remains poorly investigated and experimentally tested. Here, we conducted a photoperiod‐ and temperature‐manipulation experiment in climate chambers on two common deciduous species in Europe: Fagus sylvatica (European beech, a typically late flushing species) and Aesculus hippocastanum (horse chestnut, a typically early flushing species). In agreement with previous studies, we found that the warming significantly advanced the leaf‐out dates by 4.3 and 3.7 days/°C for beech and horse chestnut saplings, respectively. However, shorter photoperiod significantly reduced the temperature sensitivity of beech only (3.0 days/°C) by substantially increasing the heat requirement to avoid leafing‐out too early. Interestingly, the photoperiod limitation only occurs below a certain daylength (photoperiod threshold) when the warming increased above 4°C for beech trees. In contrast, for chestnut, no photoperiod threshold was found even when the ambient air temperature was warmed by 5°C. Given the species‐specific photoperiod effect on leaf phenology, the sequence of the leaf‐out timing among forest tree species may change under future climate warming conditions. Nonphotoperiodic species may benefit from warmer springs by starting the growing season earlier than photoperiodic sensitive species, modifying forest ecosystem structure and functions, but this photoperiod limitation needs to be further investigated experimentally in numerous species.     

Critical periods for impact of climate warming on early seedling establishment in subarctic tundra

Climate warming is expected to shift bioclimatic zones and plant species distribution. Yet, few studies have explored whether seedling establishment is a possible bottleneck for future migration and population resilience. We test how warming affects the early stages of seedling establishment in 10 plant species in subarctic tundra. To zoom into the life phases where the effects of warming actually take place, we used a novel approach of breaking down the whole‐season warming effect into full factorial combination of early‐, mid‐, and late‐season warming periods. Seeds were sown in containers placed under field conditions in subarctic heath and were exposed to 3 °C elevation of surface temperature and 30% addition of summer precipitation relative to ambient. Heating was achieved with Free Air Temperature Increase systems. Whole‐season heating reduced germination and establishment, significantly in four out of 10 species. The whole‐season warming effect originated from additive effects of individual periods, although some of the periods had disproportionally stronger influence. Early‐germinating species were susceptible to warming; the critical phases were early summer for germination and mid summer for seedling survival. Graminoids, which emerged later, were less susceptible although some negative effects during late summer were observed. Some species with intermediate germination time were affected by all periods of warming. Addition of water generally could not mitigate the negative effects of whole‐season heating, but at individual species level both strengthening and amelioration of these negative effects were observed. We conclude that summer warming is likely to constrain seedling recruitment in open micro sites, which is a common seed regeneration niche in tundra ecosystem. Importantly, we described both significant temporal and species‐specific variation in the sensitivity of seedling establishment to warming which needs to be taken into consideration when modelling population dynamics and vegetation transitions in a warmer climate.     

Summer aridity rather than management shapes fitness‐related functional traits of the threatened mountain plant Arnica montana

Semi‐natural mountain grasslands are increasingly exposed to environmental stress under climate change. However, which are the environmental factors that limit plants in these grasslands? Also, is the present management effective against these changes? Fitness‐related functional traits may offer a way to detect changes in performance and provide new insights into their vulnerability to climate change. We investigated changes in performance and variability of functional traits of the mountain grassland target species Arnica montana along a climate gradient in Central German low mountain ranges. This gradient represents at its lower end climate conditions that are expected at its upper end under future climate change. We measured vegetative, generative, and physiological traits to account for multiple ways of plant responses to the environment. Using mixed effects and multivariate models, we evaluated changes in trait values among individuals as well as the variability of their populations in order to assess performance under changing summer aridity and different management regimes. Fitness‐related performance of most traits showed strongly positive associations with reduced summer aridity at higher elevations, while only specific leaf area and leaf dry matter content showed no association. This suggests a higher performance level at less arid montane sites and that the physiological traits are less sensitive to this climate change factor. The coefficient of variation of almost all traits declined steadily with decreasing site aridity. We suggest that this reduced variability indicates a lower environmental stress level for A. montana toward its environmental optimum at montane elevations, especially because the trait performance increased simultaneously. Surprisingly, management factors and habitat characteristics had only low influence on both trait performance and variability. In summary, summer aridity had a stronger effect to shape the trait performance and variability of A. montana under increased environmental stress than management and other habitat characteristics.     

Phenotypic variation in seedlings of a “keystone” tree species (Quercus douglasii): the interactive effects of acorn source and competitive environment

Blue oak (Quercus douglasii) is a deciduous tree species endemic to California that currently exhibits poor seedling survival to sapling age classes. We used common garden techniques to examine how genetic variation at regional and local scales affected phenotypic expression in traits affecting oak seedling growth and survival. Between-population variation was examined for seedlings grown from acorns collected from a northern, mesic population and a southern, xeric population. Within-population variation was examined by comparing seedlings from different maternal families within the mesic population. Acorns were planted into neighborhoods of an annual dicot (Erodium botrys), an annual grass (Bromus diandrus), and a perennial bunchgrass (Nassella pulchra). By varying the species composition of herbaceous neighborhoods into which acorns were planted, the interactive effects of competition and acorn germplasm source on phenotypic expression could also be examined. Potential maternal effects, expressed as variation in acorn size, were assessed by weighing each acorn before planting. Probability of seedling emergence increased significantly with acorn size in the xeric population but not in the mesic population. Similarly, the effect of acorn size on seedling leaf area, stem weight, and root weight was also population-dependent. At a within-population level, acorn size effects on seedling traits varied significantly among maternal families. In addition to acorn size effects, rates of oak seedling emergence were also dependent on an interaction of population source and competitive environment. Interactions between maternal family and competitive environment in the expression of seedling leaf characters suggest the possibility of genetic variation for plasticity in traits such as specific leaf area. Using carbon isotope discrimination () as an index of relative water-use efficiency (WUE), higher water use efficiency was indicated for oak seedlings grown in the annual plant neighborhoods compared to seedlings grown in the bunchgrass neighborhood. This trend may represent an adaptive plastic response because, compared to the bunchgrass neighborhood, soil water depletion was more rapid within annual plant neighborhoods.