Intraspecific variations in seedling emergence and survival of Potentilla matsumurae (Rosaceae) between alpine fellfield and snowbed habitats

Potentilla matsumurae has a wide distribution from wind-blown fellfields to snowbeds in alpine regions of Japan. The environmental factors influencing seedling establishment differ between the fellfield and snowbed habitats; plants growing in each habitat may therefore have different germination strategies. Using a reciprocal sowing experiment, patterns of seedling emergence and survivorship were examined in both habitat types in the Taisetsu Mountains, Japan. Seeds derived from a fellfield population germinated earlier than did those derived from a snowbed population at both habitats, and the germination of fellfield seeds continued throughout the growing season. The timing of seedling emergence greatly affected subsequent survival at the fellfield. Seedlings that emerged in the first half of the growing season had low survivorship during the first year because of frost and drought damage, but the remaining seedlings had high survivorship during the winter; seedlings that emerged in the latter half of the growing season showed the opposite trend. At the snowbed, seedling survival was high throughout the growing season. Germination experiments in the laboratory highlighted a difference in the sensitivity of seeds from the fellfield and snowbed populations to fluctuating temperatures. These results indicate that intraspecific variation in emergence and survivorship may occur over a small scale in an alpine environment.     

Morphological and genetic variations of Potentilla matsumurae (Rosaceae) between fellfield and snowbed populations

Identifying ecological factors associated with local differentiation of populations is important for understanding microevolutionary processes. Alpine environments offer a unique opportunity to investigate the effects of habitat-specific selective forces and gene flow limitations among populations at a microscale on local adaptation because the heterogeneous snowmelt patterns in alpine ecosystems provide steep environmental changes. We investigated the variation in morphological traits and enzyme loci between fellfield and snowbed populations of Potentilla matsumurae, a common alpine herb with a wide distribution along snowmelt gradients in northern Japan. We found significant differences in morphological traits between fellfield and snowbed habitats in a northern distribution region. These differences were maintained when plants were grown under uniform conditions in a greenhouse. Allozyme variations among 15 populations from geographically separated regions with different historical backgrounds showed that the populations are more genetically differentiated between the fellfield and snowbed habitats within a region than between populations occupying the same habitat type in different regions. These results suggest that variation in snowmelt regimes could be a driving force creating local adaptation and genetic differentiation of alpine plant populations.     

Comparisons of germination traits of alpine plants between fellfield and snowbed habitats

We examined the seed-germination responses of 27 alpine species with reference to habitat type (fellfield and snowbed), temperature (five regimes), and light requirement. About 70% of species showed >40% germination at warm temperatures without cold stratification. However, a moist-chilling treatment markedly improved the germination percentages in most species, especially under cool conditions. Thus, cold stratification effectively reduced the temperature requirement for germination. Patterns of germination response within species were not consistent between the fellfield and snowbed habitats for species inhabiting both habitats. For interspecific comparisons, there were no significant differences in germination responses to the temperature regimes and the cold stratification between the fellfield and snowbed species. Also, germination speed and the length of germinating period did not differ between fellfield and snowbed species. Most species (86%) showed a requirement for light for germination without cold stratification. Although the extent of the light requirement was reduced after cold stratification in some species, the light requirement of most small-seeded species remained. The combination of cold stratification and the light requirement is a major factor determining the seedling emergence and formation of seed banks in alpine plants. However, habitat-specific patterns of germination traits were less clear, suggesting similar germination traits in fellfield and snowbed plants, at least under controlled conditions in the laboratory.     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Habitat-specific responses in the flowering phenology and seed set of alpine plants to climate variation: implications for global-change impacts

The timing of the snowmelt is a crucial factor in determining the phenological schedule of alpine plants. A long-term monitoring of snowmelt regimes in a Japanese alpine area revealed that the onset of the snowmelt season has been accelerated during the last 17 years in early snowmelt sites but that such a trend has not been detected in late snowmelt sites. This indicates that the global warming effect on the snowmelt pattern may be site-specific. The flowering phenology of fellfield plants in an exposed wind-blown habitat was consistent between an unusually warm year (1998) and a normal year (2001). In contrast, the flowering occurrence of snowbed plants varied greatly between the years depending on the snowmelt time. There was a large number of flowering species in the fellfield community from mid- to late to late June and from mid- to late July. The flowering peak of an early-melt snowbed plant community was in the middle of the flowering season and that of a late-melt snowbed community was in the early flowering season. These habitat-specific phenological patterns were consistent between 1998 and 2001. The effects of the variation in flowering timing on seed-set success were evaluated for an entomophilous snowbed herb, Peucedanum multivittatum, along the snowmelt gradient during a 5-year period. When flowering occurred prior to early August, mean temperature during the flowering season positively influenced the seed set. When flowering occurred later than early August, however, the plants enjoyed high seed-set success irrespective of temperature conditions if frost damage was absent. These observations are probably explained based on the availability of pollinators, which depends not only on ambient temperature but also on seasonal progress. These results suggest that the effects of climate change on biological interaction may vary depending on the specific habitat in the alpine ecosystem in which diverse snowmelt patterns create complicated seasonality for plants within a very localized area.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

不同干扰程度下沙生植物异翅独尾草的种群结构与动态特征

不断加剧的人类活动导致古尔班通古特沙漠南缘异翅独尾草(Eremurus anisopterus(Kar.et Kir.)Regel)生存生境片段化,形成许多大小不一的斑块种群。为深刻理解在不同程度破碎化斑块中异翅独尾草种群的生存现状,共选取19个样点,分析其龄级结构,编制静态生命表,绘制存活曲线和死亡曲线,并引入4个生存分析函数。结果表明:古尔班通古特沙漠南缘不同样地中异翅独尾草种群动态因人类干扰与生境破碎化程度的差异,呈现为不同的结构特征及变化趋势,各样地异翅独尾草种群龄级完整性均不同,破碎化程度高的样地中种群的龄级有残缺或断代现象;人类干扰程度中、弱的b类型、c类型斑块的种群年龄结构分别属于稳定至衰退型和增长型,而受干扰最强的a类型斑块中的种群结构表现出较强的波动性,种群趋于衰退的风险较高;存活曲线与4个生存函数曲线表明,a类型种群前、中期稳定,后期衰退;b类型种群前期衰退,中、后期稳定;c类型种群稳定增长。说明异翅独尾草种群的衰退可能是其生境破碎化引起的,因此,对于人类干扰程度强的衰退型种群应亟需减少人为干扰,依据不同生境中的干扰因素及种群生存现状,制定科学与切实可行的保护、恢复策略。     

Mast-seeding in the cycad genus Encephalartos: a test of the predator satiation hypothesis

Mast-seeding behaviour was monitored in 18 populations of eight species of the African cycad genus Encephalartos between 1988 and 1991. The coefficient of variation (V) in annual cone production for each population ranged between 88 and 200, indicating large fluctuations in reproductive effort between years. Data were collected to determine whether mast-seeding reduced levels of predispersal seed predation by satiating seed predators in mast years and whether it resulted in a reproductive advantage over plants which reproduced more frequently. Masting intensity was greatest in those populations in which individual plants suffered the highest levels of predispersal seed predation in years when only a few plants produced seeds. The principal seed predators were two congeneric weevil species, Antliarhinus zamiae and A. signatus, which develop exclusively on cycad seeds. The lowest intensity of mast-seeding was recorded for cycad populations with low levels of seed predation and in which A. zamiae and A. signatus occurred only in low numbers or were entirely absent. Larger seed crops appeared to result in lower levels of seed predation by A. zamiae and A. signatus in four populations of E. altensteinii, and differences in seed crop size accounted for 48–66% of variation in levels of seed predation in populations of five cycad species. In one population of E. altensteinii, lower levels of seed predation in plants reproducing periodically resulted in a reproductive advantage over plants reproducing more frequently. These results are consistent with the predator satiation hypothesis. However, in most cycad populations, numbers of seed predators did not appear to decrease significantly after a period of 2–8 years between reproductive episodes and, in two of three populations examined, periodic reproduction did not increase the number of seeds surviving to dispersal over a 4-year period. These results are interpreted to mean that periodic reproduction has not evolved in response to selection imposed by seed predators, but that selection may favour those plants which experience lower levels of seed predation by coning in synchrony with the majority of plants in the population.     

Heterosis in crosses between lines of Drosophila melanogaster selected for adaptation to different environments

Summary Experiments were designed to examine whether heterosis would occur in crosses of Drosophila melanogaster populations adapted to 18 °C or 28 °C environments. Crosses were examined in parental environments, an intermediate environment (23 °C) and a mixed environment (alternating 18°/28°C). Parental populations did not show divergence for larval viability, cold shock or high temperature mortalities when tested in a common environment. However, the 28 °C population was less fecund than the 18 °C population, but had higher larval competitive ability and higher adult longevity. Heterosis for viability, cold shock mortality and high temperature mortality occurred in crosses between a population adapted to 18 °C and another adapted to 28 °C, but not in crosses between two populations adapted to the same temperature. The results suggest that, in the absence of drift, heterosis is expected in crosses between lines or populations with different histories of selection but not between lines with the same selection histories.     

Impact of simulated herbivory on Alliaria petiolata survival, growth, and reproduction

In weed biological control, insect damage to target weeds can be simulated in invaded habitats to study potential responses of the plant to introduced natural enemies. In the present study, we investigated the impact of two levels of manual flower-shoot damage (shoots cut at tip or base) on Alliaria petiolata (garlic mustard) survival, size, and reproduction. Experiments were conducted in 2002 and 2003 using invasive field populations of A. petiolata under naturally varying plant densities. Plant survival was recorded, and size and reproduction parameters were measured. Manual flower-shoot damage had a significant effect on plant survival. In both years, fewer plants survived in the basal-cut treatment than in either the control (un-cut) or tip-cut treatment. Plant size and reproductive output were likewise reduced in the basal-cut treatment. In both years, total seed production was significantly lower in the basal-cut treatment than either the control or tip-cut treatment. When combined, increased mortality and reduced seed production of basal-cut plants greatly reduced the contribution these plants made to the seed bank. Plant density did not affect reproduction or plant size. The impacts of cutting were consistent across years and sites with distinct biotic and abiotic conditions, and A. petiolata densities. We anticipate that herbivore damage to A. petiolata populations by introduced biological control agents will likewise remain consistent under varying biotic and abiotic conditions if the agents are equally adapted to these.     

Estimation of genetic variability and heritability for biofuel feedstock yield in several populations of switchgrass

Information on heritability and predicted gains from selection for increased biomass yield for ethanol production in switchgrass is limited and may vary among breeding populations. The purpose of this study was to estimate heritability and predicted gains from selection for higher biomass yield within a lowland ecotype switchgrass population, Southern Lowland 93 (SL‐93), and two upland ecotype switchgrass populations, Southern Upland Northern Upland Early Maturing (SNU‐EM) and Southern Upland Northern Upland Late Maturing (SNU‐LM). Narrow‐sense heritabilities (h_n²) for biomass yield in each of the three populations were estimated via progeny–parent regression analysis. Half‐sib (HS) progeny families from 130 randomly selected plants from the SL‐93 population were evaluated for biomass yield in replicated trials in 2002 and 2003. Clonal parent plants were evaluated for biomass yield in separate environments to provide unbiased h_n² estimates from progeny–parent regression. Yield differences were highly significant among SL‐93 HS progenies within and over years. For the SL‐93 population, h_n² estimates were 0.13 and 0.12 based on individual plant and phenotypic family mean (PFM) selection, respectively. Predicted genetic gains (ΔG) per selection cycle were 0.15 kg dry matter (dm) plant^?1 and 0.10 kg dm plant^?1 for PFM and individual plant selection methods, respectively. For the SNU‐EM and SNU‐LM populations, year and year × HS family effects were highly significant (P < 0.01) and the HS family effect over years was nonsignificant (P < 0.05). However, HS family effects were highly significant within respective years (P < 0.01). Estimates of h_n² for the SNU‐EM and SNU‐LM populations based on PFM and individual plant selection were similar, ranging from 0.44 to 0.47; ΔG per selection cycle ranged from 0.22 to 0.33 kg dm plant^?1. The magnitudes of the estimates of additive genetic variation suggest that selection for higher biomass yield should be possible. The substantial effect of environment on biomass yields in the upland populations and the failure of families to respond similarly over years stress the importance of adequately testing biomass yield over years to assess yield.     

Pollinator diversity affects plant reproduction and recruitment: the tradeoffs of generalization

One outstanding and unsolved challenge in ecology and conservation biology is to understand how pollinator diversity affects plant performance. Here, we provide evidence of the functional role of pollination diversity in a plant species, Erysimum mediohispanicum (Brassicaceae). Pollinator abundance, richness and diversity as well as plant reproduction and recruitment were determined in eight plant populations. We found that E. mediohispanicum was generalized both at the regional and local (population) scale, since its flowers were visited by more than 100 species of insects with very different morphology, size and behaviour. However, populations differed in the degree of generalization. Generalization correlated with pollinator abundance and plant population size, but not with habitat, ungulate damage intensity, altitude or spatial location. More importantly, the degree of generalization had significant consequences for plant reproduction and recruitment. Plants from populations with intermediate generalization produced more seeds than plants from populations with low or high degrees of generalization. These differences were not the result of differences in number of flowers produced per plant. In addition, seedling emergence in a common garden was highest in plants from populations with intermediate degree of generalization. This outcome suggests the existence of an optimal level of generalizations even for generalized plant species.     

Habitat-dependent Ecotype Micro-distribution at the Mid-shore in Natural Populations of <Emphasis Type="Italic">Littorina saxatilis</Emphasis>

On the NW coast of Spain, there is a hybrid zone in natural populations of L. saxatilis, where the largest ecotype is adapted to the desiccation and heat stress of the upper shore, whereas the smaller one is adapted to the wave impacts of the lower shore. The two ecotypes meet and hybridise in a mid-shore area producing intermediate individuals or hybrids. It has been postulated that this hybrid zone is maintained by a selection-gradient model that assumes habitat-dependent selection for the different environments of upper and lower shore areas. In this study, we focus on the hybrid (mid-shore) area, where a transition occurs between upper-shore and lower-shore environments, in order to ascertain whether the habitat-dependent selection is maintained at the micro-habitat scale. We present data on snail density at several levels of the mid-shore (varying accordingly in the relative frequency of mussels and barnacles) obtained for three seasons in three consecutive years and three localities. In the mid-shore, the abundance of the RB ecotype increases with increasing tidal height, whereas the abundance of the SU ecotype increases with decreasing tidal height, suggesting that habitat-dependent selection is maintaining the ecotype micro-distribution.     

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.     

Are gardens effective in butterfly conservation? A case study with the pipevine swallowtail, <Emphasis Type="Italic">Battus philenor</Emphasis>

Gardens with nectar sources and larval host plants have been proposed to stem the decline in butterfly abundance caused by habitat loss. However, no study has provided evidence that gardens benefit butterflies. We examined the use of natural sites and gardens in the San Francisco bay area by the butterfly, Battus philenor. We found that natural sites were more likely to attract adult B. philenor, received more oviposition, and had higher juvenile survival than gardens sites. Butterflies were more likely to be present in gardens with established populations of the host plant, Aristolochia californica, growing in the sun. Battus philenor are unlikely to visit gardens with host plants planted within the past 7 years. Gardens between the ages of 8–40 years received oviposition, but did not always support completion of larval development of B. philenor. In gardens with host plants over 40 years of age, B. philenor consistently survived from egg to the adult stage. Natural enemy induced mortality of eggs did not differ between garden and natural sites, but overall egg survival was lower in gardens than at natural sites. It is unlikely that gardens serve as 'refugia' for B. philenor in years when populations in natural sites experience low survival or low fecundity. Even in gardens capable of supporting larvae to maturity, the density of eggs and survival rates were lower than in natural populations of the host plant suggesting that gardens were not optimal habitats. Therefore, without evidence that juvenile abundance and survival rates in gardens matches or exceeds that in natural sites, it is most likely that gardens act as population sinks for B. philenor.     

Variation in life-history traits among Urtica dioica populations with different history in parasitism by the holoparasitic plant Cuscuta europaea

Several studies demonstrate that natural enemies (e.g. parasites) have profound negative effects on the life-history traits of their hosts. If the host can compensate for the negative effects of parasitic infection by altering its life history, these modifications may partly form the basis of resistance or tolerance against parasites. Thus, parasites may be of considerable importance in shaping the evolution of life-history traits of their hosts. To examine if previous parasitism is associated with differences in life-history traits of the host, I conducted a common garden experiment with Urtica dioica plants originating from eight populations of which four were unparasitized, and four parasitized by the holoparasitic plant, Cuscuta europaea. A field survey indicated no differences between unparasitized and parasitized populations in, for example, the number of plant species and nutrient levels in the soil. Thus, it seems reasonable to assume that differences in life-history traits between the two population types in the common garden would reflect the effects of previous selection by the parasite. In the common garden, plants from parasitized populations started to flower later and allocated less biomass to asexual reproduction (measured as the production of stolons, i.e. clonal propagation) compared to plants from unparasitized populations. These results thus indicate that selection by the parasite may have favoured later onset of flowering, and may have selected against asexual reproduction.     

Resource allocation to sexual and vegetative reproduction in a forest herbSyneilesis palmata (Compositae)

The balance between sexual and vegetative reproduction inSyneilesis palmata was examined in relation to environmental conditions and the amount of reproductive resources, which is defined here as the total quantity of dry matter invested in both modes of reproduction. The allocation balance was measured for individual plants of two populations, with different densities in an open habitat, for 2 years, and those of two other populations, under different light intensities in a plantation forest (forest floor and edge), for 3 years. Relative allocation to sexual reproduction decreased with increasing reproductive resources in all populations except for the forest edge, which showed a constant allocation balance. The high density population showed lower relative allocation to sexual reproduction than the low density population, irrespective of the amount of reproductive resources. However the between-year comparison of the high density population suggested that under extremely high density, plants with a small amount of reproductive resources enhanced sexual reproduction, while plants with a large amount of reproductive resources reproduced vegetatively. On the forest floor, plants with small amounts of reproductive resources had higher relative allocation to sexual reproduction than the forest edge population, while plants with large amounts of reproductive resources had a somewhat lower one. The adaptive significance of such allocation patterns are discussed, based on qualitative data on the characteristics of both types of offspring.     

<Emphasis Type="BoldItalic">r</Emphasis>-Selected Traits in an Invasive Population

I compared life-history traits and self-fertilization rates in greenhouse culture of native Spartina alterniflora with an invasive population from a low-density Pacific estuary to see whether plants in the novel r-selective regime exhibit early reproduction, greater self-compatibility and high reproductive effort putting individuals at greater risk of death. Plants were grown from seed collected from the Atlantic and Gulf coasts of North America and a population introduced approximately 100 years ago to Willapa Bay, Washington, USA that has expanded to cover over ca. 6000 hectares. Pacific marshes have almost no native emergent vascular plants in the intertidal habitat, offering a virtually empty niche for invasive S. alterniflora. The low-density plants at the leading edge of this rapidly expanding population suffer a severe Allee effect of greatly reduced fecundity, caused by pollen limitation, compared to high-density areas. I found 99% of the invasive plants initiated reproduction in the first year of this study, while only 52% of the native range plants did so in the first year, followed by 34% in the second and 14% of native plants had not initiated reproduction after three years. The invasive plants had more than twice the reproductive effort of the native plants. Plants with the highest reproductive effort in the first year of growth died regardless of coastal site of origin, although nearly all of those that died over two years were invasive Pacific plants (27%). The invaders set two times the seed as the natives under forced selfing, suggesting greater self-compatibility or reproductive effort. These results suggest either a founding and bottleneck event and/or the invasive population has evolved from the long-lived, predominantly self-incompatible, K-selected state of the probable invasive propagules originating in dense, competitive native marshes.Co-ordinating editor: I. Olivieri     

Ecotypic differentiation determines the outcome of positive interactions in a dryland annual plant species

Positive interactions among plants have rarely been investigated with respect to their evolutionary consequences and vice versa. The outcome of facilitative interactions depends on the competitive ability and stress tolerance of the species. We tested whether this also applies to populations of conspecifics that are locally adapted to different environments and thereby differ in these traits. We hypothesised that ecotypes from less stressful environments experience a greater effect of facilitation when grown in stressful environments compared to populations adapted to these conditions.Seeds of two ecotypes of the annual grass species, Brachypodium distachyon, were collected from Mediterranean and arid origins and transplanted at an arid environment within the species’ distribution range. To examine the effect of biotic interactions on these ecotypes, we transplanted the individuals with and without the presence of the shrub Gymnocarpos decander (underneath or away from the shrub), and with and without the presence of annual vegetation (removal experiment). We examined the effect of these interactions on the two B. distachyon ecotypes by comparison of emergence success, biomass, and survival to reproduction.The presence of shrubs had a positive effect on all three variables in both ecotypes. Facilitation by shrubs enabled individuals from Mediterranean origin to grow and reproduce in arid conditions. Unlike the locals, they failed to survive to reproduction away from the shrubs, because of the markedly shorter growing season in open areas. The annual vegetation did not affect emergence or survival to reproduction in either ecotype; however, the positive effect of shrubs on biomass was reduced in the presence annual vegetation in the Mediterranean ecotype.This demonstrates that ecotypes adapted to arid conditions respond differently to these biotic interactions compared to Mediterranean populations. We argue that facilitation may have important evolutionary consequences by enabling maladapted ecotypes to invade and colonize stressful habitats.     

Immigrant and extrinsic hybrid seed inviability contribute to reproductive isolation between forest and dune ecotypes of Epipactis helleborine (Orchidaceae)

Reproductive isolation caused by divergent natural selection arising from differences between ecological environments or ecological interactions represents a major mechanism contributing to speciation, but its relative importance is poorly known. In this study, controlled reciprocal crossings and seed germination experiments were combined with genetic and morphometric analyses to test the hypothesis that previously described differences in mycorrhizal communities between forest and dune ecotypes of Epipactis helleborine were sufficiently strong to create complete reproductive isolation between the two ecotypes. Molecular analyses using 770 SNP markers showed that the two ecotypes were genetically distinct and that populations of the dune ecotype were genetically impoverished compared to populations of the forest ecotype. Morphologically, the two ecotypes were also significantly different, with plants of the dune ecotype generally being smaller than plants of the forest ecotype. The results further showed that immigrant seeds had a significantly lower probability of protocorm formation than native seeds, indicating strong immigrant inviability. Although both ecotypes were able to cross easily and to produce a large number of viable seeds, hybrid seeds showed significantly lower protocorm formation than pure seeds, further contributing to reproductive isolation. Overall, these results indicate that interfertile populations of a widespread orchid adapting to contrasting environments diverge as a consequence of concurrent selection acting against immigrants and hybrids and suggest that mycorrhizal fungi can play a role in the early stages of plant speciation.