Patterns of seed mass variation and their effects on seedling traits inAlliaria petiolata (Brassicaceae)

Seed mass is considered to be the least plastic component of reproductive yield. Yet, in invasive populations of garlic mustard, Alliaria petiolata, seed mass was highly variable (eightfold among populations, 2.5-7.5 fold within populations, two-threefold within individuals, and 1.4-1.8 fold within fruits). Variation in seed mass among populations explained nearly half of the total variance. Variation among seeds within fruits accounted for a further 25% of variance. Individual seed mass within a plant decreased with increased distance from the main stem, suggesting that access to parental resources limits seed size in a predictable manner. MANOVAs and Roy-Bargmann stepdown analyses revealed significant effects of seed mass, but not seed position (within a fruit or within an infructescence), on an array of subsequent seedling traits. Smaller seeds germinated significantly earlier, and seedlings from small seeds produced their first primary leaves significantly later and grew significantly taller. After accounting for seed mass as a covariate, only one seedling trait, date of first leaf emergence, was affected by seed position in a fruit. Differences in seed mass may therefore affect seedling recruitment via effects on early seedling growth in this weedy species.     

Rapid Development of Adaptive,Climate-Driven Clinal Variation in Seed Mass in the Invasive Annual Forb Echium plantagineum L.

We examined adaptive clinal variation in seed mass among populations of an invasive annual species, Echium plantagineum, in response to climatic selection. We collected seeds from 34 field populations from a 1,000 km long temperature and rainfall gradient across the species'' introduced range in south-eastern Australia. Seeds were germinated, grown to reproductive age under common glasshouse conditions, and progeny seeds were harvested and weighed. Analyses showed that seed mass was significantly related to climatic factors, with populations sourced from hotter, more arid sites producing heavier seeds than populations from cooler and wetter sites. Seed mass was not related to edaphic factors. We also found that seed mass was significantly related to both longitude and latitude with each degree of longitude west and latitude north increasing seed mass by around 2.5% and 4% on average. There was little evidence that within-population or between-population variation in seed mass varied in a systematic manner across the study region. Our findings provide compelling evidence for development of a strong cline in seed mass across the geographic range of a widespread and highly successful invasive annual forb. Since large seed mass is known to provide reproductive assurance for plants in arid environments, our results support the hypothesis that the fitness and range potential of invasive species can increase as a result of genetic divergence of populations along broad climatic gradients. In E. plantagineum population-level differentiation has occurred in 150 years or less, indicating that the adaptation process can be rapid.     

Implications of a long-term, pollinator-mediated selection on floral traits in a generalist herb

Background and Aims

The phenotypic selection of a diverse insect assemblage was studied on a generalist plant species (Paeonia broteroi) in ten flowering seasons, with tests for whether visitor preferences for plants with larger flowers eventually translated into significant differences among plants in visitation rates, seed production, seed mass, seed germination and seedling survival.

Methods

Selection gradients were used to assess if selection on flower size contributed to explain differences in visitation rates, seed production and seed mass. First, independent analyses were carried out for each season; then for the ten season as a whole. Seedling emergence and survival were assessed by generalized linear models.

Key Results

Directional selection was found on flower size through visitation rates and seed production, and stabilizing selection through seed mass. Thus, larger flowers were more visited, and produced more, but lighter seeds, than smaller flowers. The results suggest a conflicting selection on flower size through seed number and size. Floral integration found in the study populations was larger than that in populations of a distant region. Finally, seed size did not influence seedling emergence and survival; thus, any advantages of seed size may be constrained under natural conditions before plants become reproductive individuals.

Conclusions

Plants with larger flowers may be benefited by producing more lighter seeds than fewer heavier ones, as they may contribute disproportionately to the seed bank, and have better chances that any descendant could eventually recruit. However, it seems unlikely that differences in flower size and integration found among populations in different regions could have been originated by rapid evolutionary change. First, because of the conflicting selection described; second, because of the remarkably low seedling survival found under natural conditions. Consequently, the influence of pollinator selection alone does not seem to explain differences in flower size and integration.Key words: Paeonia broteroi, long-term selection, conflicting selection, flower size, seed production, generalist pollination     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Intrinsic and extrinsic drivers of recruitment across the distribution range of a seed-dimorphic herb

Recruitment of new individuals, through germination and seedling survival, is a key process for short-lived plants. Here, we analyzed intraspecific variation in recruitment across the latitudinal range of Plantago coronopus, a widespread herb that produces both large basal seeds with a mucilaginous coat and small apical seeds without coat. We experimentally tested the effects of seed traits and water availability on recruitment, by using seeds from a wide environmental stress gradient from N Africa to N Europe. Our experiments were carried out in controlled environmental conditions and in dunes where the species naturally occurs. Water shortage decreased seed germination and seedling survival for all populations, showing the importance of water supply for P. coronopus. Basal seeds showed higher and faster germination rates than apical seeds. Since among-population variation in seed mass was not related to potential germination rate, it is the mucilaginous coat rather than size difference that likely drives the differential success between seed morphs. Seed mass positively affected seedling survival instead, but only in controlled conditions with regular water supply. An experiment in a dune showed indeed that the highest survival corresponded to the local population and not the one with the largest seeds. Our results demonstrate that both intrinsic and extrinsic factors drive inter-population variation in the early life stages of this short-lived plant, allowing it to adapt across the environmentally heterogeneous distribution range. Gathering information on intraspecific variation in recruitment-related traits will help us to understand and predict plant responses in a context of climatic change.     

Negative evidence of local adaptation to the establishment conditions in a perennial herb

The differential adaptation of populations of the same species to their local environmental conditions through divergent selection, known as local adaptation, is a key step in the process of diversification of species. Here, we explore the local adaptation of the perennial mountain herb Helleborus foetidus to variable environmental conditions of seedling emergence and establishment at two different spatial scales (habitats and regions) with special attention to the role of physical and chemical soil properties. The possibility of local adaptation was evaluated under the ??local versus foreign?? and the ??home versus away?? criteria. Reciprocal sowing experiments were carried out by cross-sowing seeds among habitats and regions, controlling for maternal effects by means of seed mass, and recording seedling emergence and survival. Several topsoil properties were measured linked to each sowing point. Only partial patterns of local adaptation were found, which were insufficient to eventually state the existence of local adaptation at any spatial scale or under any criteria assessed. Here, we discuss how soil properties and selection on seed size may be related to the non-achievement of local adaptation criteria. Negative evidence of local adaptation seems to be due to a congruency in the selective pressures exerted by the different soil environments on seedling emergence and survival.     

Propagule pressure governs establishment of an invasive herb

The success of plant invasions may be limited by the availability of propagules and/or of suitable microsites, with microsite availability being affected by, for example, disturbance and interspecific competition. A mechanistic understanding of the contributions of propagule pressure and microsite limitation to plant invasions is therefore required to minimise future invasions. Here, we investigated the relative roles of propagule pressure, the availability of microsites, and their interaction on the establishment of an invasive herb, Lupinus polyphyllus, in two geographic regions representing different climate and growth conditions in Finland (a more productive southern region and a harsher central region). We carried out a field experiment in 14 L. polyphyllus populations, in which we manipulated both propagule pressure and disturbance. In a complementary greenhouse experiment, we manipulated propagule pressure and interspecific competition. Seedling establishment of L. polyphyllus was higher in the more productive southern region than in the harsher central region. The number of L. polyphyllus seedlings increased with increasing propagule pressure regardless of disturbance or interspecific competition. However, the number of L. polyphyllus seedlings per sown seed (relative establishment) tended to decrease with increasing propagule pressure, indicating that the positive effect of propagule pressure on early invasion is partially counteracted by density-dependent mortality at high seed densities. Our results highlight the dominant role of propagule pressure over disturbance and interspecific competition in the establishment of L. polyphyllus, suggesting that the early stage of invasion is limited by the availability of propagules rather than the availability of suitable microsites.     

Effects of seed mass on seedling height and competition in European white oaks

The relationship between seed size and fitness in plants may depend on offspring density, especially in cases where seed size affects the outcome of competition. We investigated the relationship between seed mass, germination, intraspecific competition and seedling height in a glasshouse experiment on three European white oak species (Quercus robur, Q. petraea, Q. pubescens). Within offspring families, seed mass showed a moderate, but statistically significant effect on seedling height, i.e. seedlings from heavier seeds were slightly taller. In contrast, competition caused pronounced inequality in seedling height in pairs of competing seedlings, but in only 55.2% of all pairs the dominant competitor arose from the heavier seed. It is thus possible that a positive effect of seed mass on seedling growth can be mediated through the density of conspecific seedlings and that heterogeneity in offspring density will contribute to the maintenance of seed mass variation in oak populations.     

Introduced plants of Lupinus polyphyllus are larger but flower less frequently than conspecifics from the native range: Results of the first year

Introduced species, which establish in novel environments, provide an opportunity to explore trait evolution and how it may contribute to the distribution and spread of species. Here, we explore trait changes of the perennial herb Lupinus polyphyllus based on 11 native populations in the western USA and 17 introduced populations in Finland. More specifically, we investigated whether introduced populations outperformed native populations in traits measured in situ (seed mass) and under common garden conditions during their first year (plant size, flowering probability, and number of flowering shoots). We also explored whether climate of origin (temperature) influenced plant traits and quantified the degree to which trait variability was explained collectively by country and temperature as compared to other population‐level differences. Three out of four plant traits differed between the native and introduced populations; only seed mass was similar between countries, with most of its variation attributed to other sources of intraspecific variation not accounted for by country and temperature. Under common garden conditions, plants originating from introduced populations were larger than those originating from native populations. However, plants from the introduced range flowered less frequently and had fewer flowering shoots than their native‐range counterparts. Temperature of a population''s origin influenced plant size in the common garden, with plant size increasing with increasing mean annual temperature in both native and introduced populations. Our results of the first year reveal genetic basis for phenotypic differences in some fitness‐related traits between the native and introduced populations of L. polyphyllus. However, not all of these trait differences necessarily contribute to the invasion success of the species and thus may not be adaptive, which raises a question how persistent the trait differences observed in the first year are later in individuals’ life for perennial herbs.     

Does the spatial variation in selective pressures explain among-site differences in seed mass? A test with Buxus balearica

Positive effects of seed size on germination and survival can be offset by a greater probability of predation or a poorer dispersal of larger seeds. We hypothesized that spatial variation in local selective pressures acting on seed mass may lead to differences in both optimal and observed seed mass among discrete populations. We first examined the variation in seed mass across a total of 14 populations of Buxus balearica; in six of such populations, we further examined the temporal variation by measuring seed mass during three consecutive years. Second, we evaluated seed mass effects on different recruitment phases (seed rain to seedling establishment) in three populations. Lastly, for these three populations, we estimated the resulting overall phenotypic selection on seed mass during recruitment and compared the observed and the predicted optimal seed masses. Most variation (c. 70%) in seed mass occurred among populations, and although we found inter-annual variation in seed mass, the differences in seed mass among populations were consistent over time. Conflicting selective pressures on seed mass appeared during recruitment, and their direction and strength varied among populations, depending on the relative local importance of seed predation vs. germination and establishment. Observed seed mass matched predicted optimal seed mass in two of the three examined populations, suggesting local adaptative responses to the spatial mosaic of selective pressures.     

EFFECT OF SEED SIZE ON SEEDLING SUCCESS IN THREE SPECIES OF SESBANIA (FABACEAE)

Variation in seed size may produce variation in seedling fitness, but the relationship is not simple. Differences in seed size within and among species may not have the same effects. We examined effects of differences in seed size within and among three species of Sesbania, S. macrocarpa, S. drummondii, and S. vesicaria, on seedling emergence and growth in the greenhouse and the field. Of the three species, the largest-seeded species, S. vesicaria, produced the largest, longest-lived seedlings in both the greenhouse and the field. Even though plant size differed, annual S. macrocarpa produced the same seed mass as annual S. vesicaria in the greenhouse. Within-species effects were less clear. In the greenhouse, S. vesicaria seedlings grown from large seeds remained largest until maturity, but the other species did not exhibit this effect. Some persistent within-species effects of seed size differences on height were observed in the field in 1981, but not in 1980, suggesting that field conditions increase the importance of seed size differences. Unscarified S. drummondii seeds germinated before seeds of the two annual species. Within species, larger seeds of the annuals and smaller seeds of the perennial germinated first. Differences among the species in the importance of seed size to seedling fitness may allow the species to have different patterns of regulation of reproduction in response to stress. Sesbania vesicaria showed the largest within-species effects of seed size and has the lowest plasticity in seed size, suggesting that patterns of plasticity have been selected such that the most important component of yield varies least.     

Latitudinal variation in seeds characteristics of Acer platanoides and A. pseudoplatanus

Climate change will likely affect population dynamics of numerous plant species by modifying several aspects of the life cycle. Because plant regeneration from seeds may be particularly vulnerable, here we assess the possible effects of climate change on seed characteristics and present an integrated analysis of seven seed traits (nutrient concentrations, samara mass, seed mass, wing length, seed viability, germination percentage, and seedling biomass) of Acer platanoides and A. pseudoplatanus seeds collected along a wide latitudinal gradient from Italy to Norway. Seed traits were analyzed in relation to the environmental conditions experienced by the mother trees along the latitudinal gradient. We found that seed traits of A. platanoides were more influenced by the climatic conditions than those of A. pseudoplatanus. Additionally, seed viability, germination percentage, and seedling biomass of A. platanoides were strongly related to the seed mass and nutrient concentration. While A. platanoides seeds were more influenced by the environmental conditions (generally negatively affected by rising temperatures), compared to A. pseudoplatanus, A. platanoides still showed higher germination percentage and seedling biomass than A. pseudoplatanus. Thus, further research on subsequent life-history stages of both species is needed. The variation in seed quality observed along the climatic gradient highlights the importance of studying the possible impact of climate change on seed production and species demography.     

Sabal palmetto seed size: causes of variation,choices of predators,and consequences for seedlings

High variation in seed size, as is common among angiosperms, may be maintained in a plant species when several factors select for seed size. Variation may also result from differences among adult plants, such as nutrient and water availability or the amount of photosynthetic tissue. In a study of Sabal palmetto seed ecology I found high seed size variation both within- and among-palms, and investigated possible factors maintaining this variation. Seed size was positively correlated with the number of leaves on parent palms. Larger seeds produced more vigorous seedlings that had greater leaf length, area, and mass, and greater root mass. Caryobruchus gleditsiae (Bruchidae: Coleoptera), whose larvae develop within palm seeds, preferentially oviposited on larger seeds, which in turn produced larger beetle offspring. By choosing the largest seeds available, ovipositing beetles thus affect both the quantity and the quality of seeds available for recruitment. I conclude that because beetle predation selects against large seeds, while larger seeds promote seedling vigor, the maintenance of seed size variation may be an adaptation of S. palmetto promoting both seed escape from predators and seedling vigor.     

Effects of seed traits variation on seedling performance of the invasive weed,Ambrosia artemisiifolia L.

Seedling performance can determine the survival of a juvenile plant and impact adult plant performance. Understanding the factors that may impact seedling performance is thus critical, especially for annuals, opportunists or invasive plant species. Seedling performance can vary among mothers or populations in response to environmental conditions or under the influence of seed traits. However, very few studies have investigated seed traits variations and their consequences on seedling performance. Specifically, the following questions have been addressed by this work: 1) How the seed traits of the invasive Ambrosia artemisiifolia L. vary among mothers and populations, as well as along the latitude; 2) How do seed traits influence seedling performance; 3) Is the influence on seedlings temperature dependent. With seeds from nine Western Europe ruderal populations, seed traits that can influence seedling development were measured. The seeds were sown into growth chambers with warmer or colder temperature treatments. During seedling growth, performance-related traits were measured. A high variability in seed traits was highlighted. Variation was determined by the mother identity and population, but not latitude. Together, the temperature, population and the identity of the mother had an effect on seedling performance. Seed traits had a relative impact on seedling performance, but this did not appear to be temperature dependent. Seedling performance exhibited a strong plastic response to the temperature, was shaped by the identity of the mother and the population, and was influenced by a number of seed traits.     

Propagule size,dispersal ability,and seedling performance in Asclepias syriaca

Summary Using the propagules of common milkweed (Asclepias syriaca L.) we tested the predictions that decreasing size may increase dispersal ability, but also decrease the probability of seedling success, of wind-dispersed seeds. In 1982 and 1983 we released seeds from four milkweed clones at two heights in an open field and measured their dispersal distances. In the laboratory we measured falling times in a dead air space, seed mass and area, and coma mass and length of the same seeds. The seeds were later planted in a greenhouse and germination, mortality, and seedling dry mass were recorded.Seed mass was negatively correlated with dispersal distance in 1982, but not in 1983 under highly variable wind conditions. Coma mass/seed mass ratio was positively correlated with dispersal distance in 1982. During both years seed mass and coma mass/seed mass ratio were highly significantly correlated with falling time in dead air space. However, heavy seeds had superior germination, survivorship, and seedling mass at harvest, and seeds that germinated fell faster in dead air than those that did not. We found substantial variation in seed morphology both among clones and among pods within clones. Propagule falling times in dead air and (in 1982) dispersal distances in the field also differed significantly among clones and pods. The potential therefore exists for differential dispersal and establishment of milkweed genotypes.     

Interaction Between “Safe Sites” and “Safe Sides” for Germination of Neurada procumbens (Neuradaceae) in the Middle East

The seed bank of Neurada procumbens, a prostrate annual common in sandy desert habitats, is vulnerable to surface disturbances. Moreover, its seeds are highly heteromorphic, having both spinose and non-spinose sides, which affects both the vertical distribution of seeds and diaspores, and the precise position in which they lie in the soil. Here we explore the ecological implications of two factors, “safe sites” within habitats and seed orientation or “safe sides”, on seedling establishment and the seed bank. In three natural populations, the vertical distribution of buried, germinable diaspores decreased with soil depth. Most diaspores occurred at the surface (0–1 cm below the surface), while none were observed below 10 cm. Seedling emergence decreased with depth and reached zero in diaspores below 5 cm. Seedling emergence also proved highly sensitive to the orientation of diaspores in soil. Diaspores lying on their smooth side with the spinose upper side directed upwards reached the highest seedling emergence of 61 %. Root length ranged from 10 to 16 cm, and the root to shoot ratio varied between 22.8 and 30.3 cm, depending on diaspore size. Diaspores near the surface may constitute a more transient component of the seed bank, as few of these remained viable for more than a year. Persistence of Neurada seed banks seems to depend not only on vertical movement of diaspores among habitat “safe sites” but also on different seed position alternatives.     

濒危树种格木天然群体自由授粉子代苗期生长变异

格木是我国热带和南亚热带地区主要的珍稀濒危树种之一,具有重要的经济和生态价值。本文应用从格木主要天然分布区收集的6个群体的种子进行育苗试验,揭示格木天然群体子代芽苗和幼苗生长变异状况及其与种子性状、亲本环境因子间的相关性。结果表明,格木天然群体子代芽苗和幼苗表型性状在群体间和群体内均存在丰富的变异,苗高的变异高于地径,苗期选择以高生长为主。格木苗期生长与种子的大小、千粒重相关显著,胚根长度与经度、地径与年均气温、高径比与海拔均存在显著的相关性(p＜0.05)。研究结果为深入开展格木良种选育奠定了基础,亦将有助于其天然资源的保护经营和人工种植业的发展。     

Large population sizes mitigate negative effects of variable weather conditions on fruit set in two spring woodland orchids

Global circulation models predict increased climatic variability, which could increase variability in demographic rates and affect long-term population viability. In animal-pollinated species, pollination services, and thus fruit and seed set, may be highly variable among years and sites, and depend on both local environmental conditions and climatic variables. Orchid species may be particularly vulnerable to disruption of their pollination services, as most species depend on pollinators for successful fruit set and because seed germination and seedling recruitment are to some extent dependent on the amount of fruits and seeds produced. Better insights into the factors determining fruit and seed set are therefore indispensable for a better understanding of population dynamics and viability of orchid populations under changing climatic conditions. However, very few studies have investigated spatio-temporal variation in fruit set in orchids. Here, we quantified fruit production in eight populations of the orchid Orchis purpurea that does not reward pollinators and 13 populations of the rewarding Neottia (Listera) ovata during five consecutive years (2002–2006). Fruit production in large populations showed much higher stability than that in small populations and was less affected by extreme weather conditions. Our results highlight the potential vulnerability of small orchid populations to an increasingly variable climate through highly unpredictable fruit-set patterns.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Lantana camara and the fruit- and seed-damaging fly,Ophiomyia lantanae (Agromyzidae): Seed predator,recruitment promoter or dispersal disrupter?

The seed damaging fly, Ophiomyia lantanae is a widespread agent that infests the fruits and seeds of Lantana camara. Its effectiveness in reducing seed dispersal or seedling recruitment has not been comprehensively tested. We determined the effects of O. lantanae damage on fruit removal rates of L. camara in a field study. We also investigated recruitment responses (seed germination in growth cabinets and seedling emergence in pot trials) of two common L. camara biotypes, pink and pink-edged red (PER) to O. lantanae damage. We found that fewer damaged fruits were removed than undamaged fruits, suggesting that frugivores select against damaged fruit. In all recruitment experiments, the responses to damage were biotype dependent. In germination experiments, these differential responses were primarily due to significantly reduced germination of damaged PER seeds. Seedling emergence from pots of the PER biotype was also lower in damaged seeds (27.5%) than undamaged seeds (56.1%). In contrast, emergence increased with seed damage in the pink biotype, from 28.2 to 41.1%. O. lantanae damage significantly reduced seed mass only in the smaller seeded PER biotype. In the larger seeded pink biotype, it may have promoted seedling emergence by interfering with dormancy mechanisms. We also examined in situ seed banks at L. camara populations prior to peak seed production. Mean viable seed densities ranged from 78.6 to 402.8 seeds m⁻². Of these, non-dormant seeds comprised 24.6 to 98.2 seeds m⁻², suggesting that recruitment is unlikely to be seed limited. We conclude that while O. lantanae influences recruitment and dispersal processes, the magnitude of the responses measured are unlikely to greatly influence plant densities in south-east Queensland infestations.