Flowering phenology influences seed production and outcrossing rate in populations of an alpine snowbed shrub, Phyllodoce aleutica: effects of pollinators and self-incompatibility

Background and Aims

Because of differences in snowmelt time, the reproductive phenologies of alpine plants are highly variable among local populations, and there is large variation in seed set across populations. Temporal variation in pollinator availability during the season may be a major factor affecting not only seed production but also outcrossing rate of alpine plants.

Methods

Among local populations of Phyllodoce aleutica that experience different snowmelt regimes, flowering phenology, pollinator availability, seed-set rate, and outcrossing rate were compared with reference to the mating system (self-compatibility or heterospecific compatibility with a co-occurring congeneric species).

Key Results

Flowering occurred sequentially among populations reflecting snowmelt time from mid-July to late August. The visit frequency of bumble-bees increased substantially in late July when workers appeared. Both seed set and outcrossing rate increased as flowering season progressed. Although flowers were self-compatible and heterospecific compatible, the mixed-pollination experiment revealed that fertilization with conspecific, outcrossing pollen took priority over selfing and hybridization, indicating a cryptic self-incompatibility. In early snowmelt populations, seed production was pollen-limited and autogamous selfing was common. However, genetic analyses revealed that selfed progenies did not contribute to the maintenance of populations due to late-acting inbreeding depression.

Conclusions

Large variations in seed-set and outcrossing rates among populations were caused by the timing of pollinator availability during the season and the cryptic self-incompatibility of this species. Despite the intensive pollen limitation in part of the early season, reproductive assurance by autogamous selfing was not evident. Under fluctuating conditions of pollinator availability and flowering structures, P. aleutica maintained the genetic composition by conspecific outcrossing.Key words: Alpine snowbed, autogamy, bumble-bee, cryptic self-incompatibility, flowering phenology, mixed pollination, outcrossing rate, Phyllodoce aleutica, pollination success, seasonality, self-pollination     

Apomixis is not prevalent in subnival to nival plants of the European Alps

Background and Aims

High alpine environments are characterized by short growing seasons, stochastic climatic conditions and fluctuating pollinator visits. These conditions are rather unfavourable for sexual reproduction of flowering plants. Apomixis, asexual reproduction via seed, provides reproductive assurance without the need of pollinators and potentially accelerates seed development. Therefore, apomixis is expected to provide selective advantages in high-alpine biota. Indeed, apomictic species occur frequently in the subalpine to alpine grassland zone of the European Alps, but the mode of reproduction of the subnival to nival flora was largely unknown.

Methods

The mode of reproduction in 14 species belonging to seven families was investigated via flow cytometric seed screen. The sampling comprised 12 species typical for nival to subnival plant communities of the European Alps without any previous information on apomixis (Achillea atrata, Androsace alpina, Arabis caerulea, Erigeron uniflorus, Gnaphalium hoppeanum, Leucanthemopsis alpina, Oxyria digyna, Potentilla frigida, Ranunculus alpestris, R. glacialis, R. pygmaeus and Saxifraga bryoides), and two high-alpine species with apomixis reported from other geographical areas (Leontopodium alpinum and Potentilla crantzii).

Key Results

Flow cytometric data were clearly interpretable for all 46 population samples, confirming the utility of the method for broad screenings on non-model organisms. Formation of endosperm in all species of Asteraceae was documented. Ratios of endosperm : embryo showed pseudogamous apomixis for Potentilla crantzii (ratio approx. 3), but sexual reproduction for all other species (ratios approx. 1·5).

Conclusions

The occurrence of apomixis is not correlated to high altitudes, and cannot be readily explained by selective forces due to environmental conditions. The investigated species have probably other adaptations to high altitudes to maintain reproductive assurance via sexuality. We hypothesize that shifts to apomixis are rather connected to frequencies of polyploidization than to ecological conditions.     

Seeds of alpine plants are short lived: implications for long-term conservation

Background and Aims

Alpine plants are considered one of the groups of species most sensitive to the direct and indirect threats to ecosystems caused by land use and climate change. Collecting and banking seeds of plant species is recognized as an effective tool for providing propagating material to re-establish wild plant populations and for habitat repair. However, seeds from cold wet environments have been shown to be relatively short lived in storage, and therefore successful long-term seed conservation for alpine plants may be difficult. Here, the life spans of 69 seed lots representing 63 related species from alpine and lowland locations from northern Italy are compared.

Methods

Seeds were placed into experimental storage at 45 °C and 60 % relative humidity (RH) and regularly sampled for germination. The time taken in storage for viability to fall to 50 % (p₅₀) was determined using probit analysis and used as a measure of relative seed longevity between seed lots.

Key Results

Across species, p₅₀ at 45 °C and 60 % RH varied from 4·7 to 95·5 d. Seed lots from alpine populations/species had significantly lower p₅₀ values compared with those from lowland populations/species; the lowland seed lots showed a slower rate of loss of germinability, higher initial seed viability, or both. Seeds were progressively longer lived with increased temperature and decreased rainfall at the collecting site.

Conclusions

Seeds of alpine plants are short lived in storage compared with those from lowland populations/related taxa. The lower resistance to ageing in seeds of alpine plants may arise from low selection pressure for seed resistance to ageing and/or damage incurred during seed development due to the cool wet conditions of the alpine climate. Long-term seed conservation of several alpine species using conventional seed banking methods will be problematic.     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.     

The acceptability of meadow plants to the slug Deroceras reticulatum and implications for grassland restoration

Background and Aims

Despite the selective pressure slugs may exert on seedling recruitment there is a lack of information in this context within grassland restoration studies. Selective grazing is influenced by interspecific differences in acceptability. As part of a larger study of how slug–seedling interactions may influence upland hay meadow restoration, an assessment of relative acceptability is made for seedlings of meadow plants to the slug, Deroceras reticulatum.

Methods

Slug feeding damage to seedling monocultures of 23 meadow species and Brassica napus was assessed in microcosms over 14 d. The severity and rate of damage incurred by each plant species was analysed with a generalized additive mixed model. Plant species were then ranked for their relative acceptability.

Key Results

Interspecific variation in relative acceptability suggested seedlings of meadow species form a hierarchy of acceptability to D. reticulatum. The four most acceptable species were Achillea millefolium and the grasses Holcus lanatus, Poa trivialis and Festuca rubra. Trifolium pratense was acceptable to D. reticulatum and was the second highest ranking forb species. The most unacceptable species were mainly forbs associated with the target grassland, and included Geranium sylvaticum, Rumex acetosa, Leontodon hispidus and the grass Anthoxanthum odoratum. A strong positive correlation was found for mean cumulative feeding damage and cumulative seedling mortality at day 14.

Conclusions

Highly unacceptable species to D. reticulatum are unlikely to be selectively grazed by slugs during the seedling recruitment phase, and were predominantly target restoration species. Seedlings of highly acceptable species may be less likely to survive slug herbivory and contribute to seedling recruitment at restoration sites. Selective slug herbivory, influenced by acceptability, may influence community-level processes if seedling recruitment and establishment of key functional species, such as T. pratense is reduced.     

Effect of altitude on the genetic structure of an Alpine grass, Poa hiemata

Background and Aims

The persistence of plants inhabiting restricted alpine areas under climate change will depend upon many factors including levels of genetic variation in adaptive traits, population structure, and breeding system.

Methods

Using microsatellite markers, the genetic structure of populations of a relatively common alpine grass, Poa hiemata, is examined across three altitudinal gradients within the restricted Australian alpine zone where this species has previously been shown to exhibit local adaptation across a narrow altitudinal gradient.

Key Results

Genetic variation across six microsatellite markers revealed genetic structuring along altitudinal transects, and a reduction in genetic variation at high and low altitude extremes relative to sites central within transects. There was less genetic variation among transect sites compared with altitudinal gradients within transects, even though distances among transects were relatively larger. Central sites within transects were less differentiated than those at extremes.

Conclusions

These patterns suggest higher rates of gene flow among sites at similar altitudes than along transects, a process that could assist altitudinal adaptation. Patterns of spatial autocorrelation and isolation by distance changed with altitude and may reflect altered patterns of dispersal via pollen and/or seed. There was evidence for selfing and clonality in neighbouring plants. Levels of gene flow along transects were insufficient to prevent adaptive changes in morphological traits, given previously measured levels of selection.Key words: Poa hiemata, genetic structure, altitudinal gradient, microsatellite, gene flow, climate change     

Climate warming could shift the timing of seed germination in alpine plants

Background and Aims

Despite the considerable number of studies on the impacts of climate change on alpine plants, there have been few attempts to investigate its effect on regeneration. Recruitment from seeds is a key event in the life-history of plants, affecting their spread and evolution and seasonal changes in climate will inevitably affect recruitment success. Here, an investigation was made of how climate change will affect the timing and the level of germination in eight alpine species of the glacier foreland.

Methods

Using a novel approach which considered the altitudinal variation of temperature as a surrogate for future climate scenarios, seeds were exposed to 12 different cycles of simulated seasonal temperatures in the laboratory, derived from measurements at the soil surface at the study site.

Key Results

Under present climatic conditions, germination occurred in spring, in all but one species, after seeds had experienced autumn and winter seasons. However, autumn warming resulted in a significant increase in germination in all but two species. In contrast, seed germination was less sensitive to changes in spring and/or winter temperatures, which affected only three species.

Conclusions

Climate warming will lead to a shift from spring to autumn emergence but the extent of this change across species will be driven by seed dormancy status. Ungerminated seeds at the end of autumn will be exposed to shorter winter seasons and lower spring temperatures in a future, warmer climate, but these changes will only have a minor impact on germination. The extent to which climate change will be detrimental to regeneration from seed is less likely to be due to a significant negative effect on germination per se, but rather to seedling emergence in seasons that the species are not adapted to experience. Emergence in autumn could have major implications for species currently adapted to emerge in spring.     

Hygrochastic capsule dehiscence supports safe site strategies in New Zealand alpine Veronica (Plantaginaceae)

Background and Aims

Hygrochasy is a capsule-opening mechanism predominantly associated with plants in arid habitats, where it facilitates spatially and temporally restricted dispersal. Recently, hygrochastic capsules were described in detail for the first time in alpine Veronica in New Zealand. The aim of the present study was to investigate whether hygrochastic capsules are an adaptation of alpine Veronica to achieve directed dispersal to safe sites. We expect that by limiting dispersal to rainfall events, distances travelled by seeds are short and confine them to small habitat patches where both seedlings and adults have a greater chance of survival.

Methods

Dispersal distances of five hygrochastic Veronica were measured under laboratory and field conditions and the seed shadow was analysed. Habitat patch size of hygrochastic Veronica and related non-hygrochastic species were estimated and compared.

Key Results

Dispersal distances achieved by dispersal with raindrops did not exceed 1 m but weather conditions could influence the even distribution of seeds around the parent plant. Compared with related Veronica species, hygrochastic Veronica mostly grow in small, restricted habitat patches surrounded by distinctly different habitats. These habitat patches provide safe sites for seeds due to their microtopography and occurrence of adult cushion plants. Non-hygrochastic Veronica can be predominantly found in large habitats without clearly defined borders and can be spread over long distances along rivers.

Conclusions

The results suggest that hygrochasy is a very effective mechanism of restricting seed dispersal to rainfall events and ensuring short-distance dispersal within a small habitat patch. It appears that it is an adaptation for directed dispersal to safe sites that only exist within the parent habitat.     

Among-individual variation in pollen limitation and inbreeding depression in a mixed-mating shrub

Background and Aims

Variation in inbreeding depression (δ) among individual plants is considered to play a central role in mating system evolution and population genetics. Moreover, such variation could be linked to individual susceptibility to pollen limitation (PL) because those individuals strongly affected by δ for seed production will require more outcross pollen for setting a given number of fruits or seeds. However, no study has tested explicitly for associations between PL and δ at the individual plant level. This study assesses the extent of among-individual variation in PL and δ, the consistency of δ across life stages, and the relationships between individual PL and δ in the mixed-mating shrub Myrtus communis.

Methods

Controlled hand-pollinations were performed in a natural M. communis population. Marked flowers were monitored until fruit production and a greenhouse experiment was conducted with the seeds produced.

Key Results

Compared with selfing, outcross-pollination enhanced seed number per fruit, germination rate and seedling growth, but did not enhance fruit-set. Only seed number per fruit was pollen limited and, thus, cumulative pollen limitation depended more on pollen quality (outcross pollen) than on quantity. The effects of δ varied considerably across life stages and individual plants. Cumulative δ was high across individuals (mean δ = 0·65), although there were no positive correlations between δ values at different life stages. Interestingly, maternal plants showing stronger δ for seed production were more pollen limited, but they were also less affected by δ for seedling growth because of a seed size/number trade-off.

Conclusions

Results show a general inconsistency in δ across life stages and individuals, suggesting that different deleterious loci are acting at different stages. The association between δ and PL at the individual level corroborates the idea that pollen limitation may be ‘genotype-dependent’ regardless of other factors.     

Seed reserve composition in 19 tree species of a tropical deciduous forest in Mexico and its relationship to seed germination and seedling growth

Background and Aims

The size and composition of seed reserves may reflect the ecological strategy and evolutionary history of a species and also temporal variation in resource availability. The seed mass and composition of seed reserves of 19 co-existing tree species were studied, and we examined how they varied among species in relation to germination and seedling growth rates, as well as between two years with contrasting precipitation (652 and 384 mm).

Methods

Seeds were collected from a tropical deciduous forest in the northwest of Mexico (Chamela Biological Station). The seed dry mass, with and without the seed coat, and the concentrations of lipids, nitrogen and non-structural carbohydrates for the seed minus seed coat were determined. The anatomical localization of these reserves was examined using histochemical analysis. The germination capacity, rate and lag time were determined. The correlations among these variables, and their relationship to previously reported seedling relative growth rates, were evaluated with and without phylogenetic consideration.

Key Results

There were interannual differences in seed mass and reserve composition. Seed was significantly heavier after the drier year in five species. Nitrogen concentration was positively correlated with seed coat fraction, and was significantly higher after the drier year in 12 species. The rate and lag time of germination were negatively correlated with each other. These trait correlations were also supported for phylogenetic independent contrasts. Principal component analysis supported these correlations, and indicated a negative association of seedling relative growth rate with seed size, and a positive association of germination rate with nitrogen and lipid concentrations.

Conclusions

Nitrogen concentration tended to be higher after the drier year and, while interannual variations in seed size and reserve composition were not sufficient to affect interspecific correlations among seed and seedling traits, some of the reserves were related to germination variables and seedling relative growth rate.     

High outcrossing in the annual colonizing species Ambrosia artemisiifolia (Asteraceae)

Background and Aims

Variation in mating patterns may be particularly evident in colonizing species because they commonly experience wide variation in plant density. Here, the role of density for the mating system of Ambrosia artemisiifolia (common ragweed), a wind-pollinated annual colonizing species previously reported as self-compatible, is explored.

Methods

The effect of population density on the proportion of self- and cross-fertilized seeds was examined using allozyme markers and experimental arrays conducted over two seasons in the field. Also the reproductive success of isolated plants located in diverse habitats was measured. The potential occurrence of a physiological mechanism preventing self-fertilization, i.e. self-incompatibility, following controlled self- and cross-pollinations in the glasshouse was examined.

Key Results

Outcrossing rates estimated using allozyme markers were uniformly high, regardless of the spacing between plants. However, when single plants were isolated from congeners they set few seeds. Observations of pollen-tube growth and seed set following controlled pollinations demonstrated that plants of A. artemisiifolia possess a strong self-incompatibility mechanism, contrary to earlier reports and assumptions.

Conclusions

The maintenance of high outcrossing rates in colonizing populations of A. artemisiifolia is likely to be facilitated by the prodigious production of wind-borne pollen, high seed production and extended seed dormancy.Key words: Self-incompatibility, outcrossing rate, density dependence, colonization, wind-pollination, Ambrosia artemisiifolia (ragweed), Asteraceae     

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     

Effects of Water Level on Three Wetlands Soil Seed Banks on the Tibetan Plateau

Background

Although the effect of water level on germination in soil seed banks has been documented in many ecosystems, the mechanism is not fully understood, and to date no empirical studies on this subject exist. Further, no work has been done on the effect of water level on seed banks of drying and saline-alkaline wetlands in alpine areas on the Tibetan Plateau.

Methodology

We examined the effects of water level (0 cm, 5 cm and 10 cm) on seed germination and seedling establishment from soil seed banks at 0–5 cm and 5–10 cm depths in typical, drying, and saline-alkaline wetlands. We also explore the potential role of soil seed bank in restoration of drying and saline-alkaline wetlands.

Principal Findings

Species richness decreased with increase in water level, but there almost no change in seed density. A huge difference exists in species composition of the seed bank among different water levels in all three wetlands, especially between 0 cm and 5 cm and 0 cm and 10 cm. Similarity of species composition between seed bank and plant community was higher in 0 cm water level in drying wetland than in the other two wetlands. The similarity was much higher in 0 cm water level than in 5 cm and 10 cm water levels in all three wetlands. Species composition of the alpine wetland plant community changed significantly after drying and salinization, however, species composition of the seed bank was unchanged regardless of the environment change.

Conclusions/Significance

Water level greatly affects seed bank recruitment and plant community establishment. Further, different water levels in restored habitats are likely to determine its species composition of the plant community. The seed bank is important in restoration of degraded wetlands. Successful restoration of drying and salinization wetlands could depend on the seed bank.     

Mycorrhizal associations and reproductive isolation in three closely related Orchis species

Background and Aims

The maintenance of species boundaries in sympatric populations of closely related species requires some kind of reproductive isolation that limits gene flow among species and/or prevents the production of viable progeny. Because in orchids mycorrhizal fungi are needed for seed germination and subsequent seedling establishment, orchid–mycorrhizal associations may be involved in acting as a post-mating barrier.

Methods

We investigated the strength of post-mating barriers up to the seed germination stage acting between three closely related Orchis species (Orchis anthropophora, O. militaris and O. purpurea) and studied the role of mycorrhizal fungi in hybridization by burying seed packets of pure and hybrid seeds. After retrieval and assessment of seed germination, the fungi associating with protocorms originating from hybrid and pure seeds were determined and compared with those associating with adult individuals using DNA array technology.

Results

Whereas pre-zygotic post-mating barriers were rather weak in most crosses, post-zygotic post-mating barriers were stronger, particularly when O. purpurea was crossed with O. anthropophora. Germination trials in the field showed that seed germination percentages of hybrid seeds were in most cases lower than those originating from pure crosses. In all species pair combinations, total post-mating reproductive isolation was asymmetric. Protocorms associated with a smaller range of fungal symbionts than adult plants, but there was considerable overlap in mycorrhizal associations between protocorms and their respective parents.

Conclusions

Our results suggest that mycorrhizal associations contribute little to reproductive isolation. Pre-mating barriers are probably the main factors determining hybridization rates between the investigated species.     

A world-wide perspective on crucifer speciation and evolution: phylogenetics,biogeography and trait evolution in tribe Arabideae

Background and Aims

Tribe Arabideae are the most species-rich monophyletic lineage in Brassicaceae. More than 500 species are distributed in the majority of mountain and alpine regions worldwide. This study provides the first comprehensive phylogenetic analysis for the species assemblage and tests for association of trait and characters, providing the first explanations for the enormous species radiation since the mid Miocene.

Methods

Phylogenetic analyses of DNA sequence variation of nuclear encoded loci and plastid DNA are used to unravel a reliable phylogenetic tree. Trait and ancestral area reconstructions were performed and lineage-specific diversification rates were calculated to explain various radiations in the last 15 Myr in space and time.

Key Results

A well-resolved phylogenetic tree demonstrates the paraphyly of the genus Arabis and a new systematic concept is established. Initially, multiple radiations involved a split between lowland annuals and mountain/alpine perennial sister species. Subsequently, increased speciation rates occur in the perennial lineages. The centre of origin of tribe Arabideae is most likely the Irano-Turanian region from which the various clades colonized the temperate mountain and alpine regions of the world.

Conclusions

Mid Miocene early diversification started with increased speciation rates due to the emergence of various annual lineages. Subsequent radiations were mostly driven by diversification within perennial species during the Pliocene, but increased speciation rates also occurred during that epoch. Taxonomic concepts in Arabis are still in need of a major taxonomic revision to define monophyletic groups.     

Limitations to reproductive output and genetic rescue in populations of the rare shrub Grevillea repens (Proteaceae)

Background and Aims

When conserving rare plant species, managers are often faced with small and/or isolated populations displaying low levels of sexual reproduction and genetic variation. One option for reinvigorating these populations is the introduction of genetic material from other sites, but in some cases fitness may be reduced as a result of outbreeding depression. Here the pollination biology of the rare shrub Grevillea repens is studied across its natural range and reproductive responses following cross-pollination among populations are examined to determine factors that may be limiting sexual reproduction and the potential for genetic rescue.

Methods

Pollen manipulation treatments (self-, autogamous self-, cross- and open pollination) were applied to flowers to examine the breeding system and fruit and seed production in five populations of G. repens. Pollen production, presentation and viability were investigated and interpopulation crosses of increasing genetic distance performed among the populations.

Key Results

The study species is self-incompatible and displayed very low natural seed set over two seasons, due partly to low pollen viability in one of the populations. Within-population crossing increased fruit and seed production at some sites, indicating pollinator limitation. Interpopulation crosses further increased reproductive output in one population, suggesting mate limitation, and for this site there was a positive relationship between genetic distance among populations and the size of genetic rescue benefits. However, in other populations there was a decrease in fruit and seed set with increasing genetic distance.

Conclusions

The results highlight that management strategies involving interpopulation crosses can improve reproductive output in small, isolated populations of rare plants, but guidelines need to be developed on a population by population basis.Key words: Grevillea repens, Proteaceae, genetic rescue, pollination ecology, self-incompatibility, breeding system, interpopulation cross, outbreeding depression, pollinator limitation, mate limitation, resource limitation     

Light limitation and litter of an invasive clonal plant,Wedelia trilobata,inhibit its seedling recruitment

Background and Aims

Invasive clonal plants have two reproduction patterns, namely sexual and vegetative propagation. However, seedling recruitment of invasive clonal plants can decline as the invasion process proceeds. For example, although the invasive clonal Wedelia trilobata (Asteraceae) produces numerous seeds, few seedlings emerge under its dense population canopy in the field. In this study it is hypothesized that light limitation and the presence of a thick layer of its own litter may be the primary factors causing the failure of seedling recruitment for this invasive weed in the field.

Methods

A field survey was conducted to determine the allocation of resources to sexual reproduction and seedling recruitment in W. trilobata. Seed germination was also determined in the field. Effects of light and W. trilobata leaf extracts on seed germination and seedling growth were tested in the laboratory.

Key Results

Wedelia trilobata blooms profusely and produces copious viable seeds in the field. However, seedlings of W. trilobata were not detected under mother ramets and few emerged seedlings were found in the bare ground near to populations. In laboratory experiments, low light significantly inhibited seed germination. Leaf extracts also decreased seed germination and inhibited seedling growth, and significant interactions were found between low light and leaf extracts on seed germination. However, seeds were found to germinate in an invaded field after removal of the W. trilobata plant canopy.

Conclusions

The results indicate that lack of light and the presence of its own litter might be two major factors responsible for the low numbers of W. trilobata seedlings found in the field. New populations will establish from seeds once the limiting factors are eliminated, and seeds can be the agents of long-distance dispersal; therefore, prevention of seed production remains an important component in controlling the spread of this invasive clonal plant.     

Kinship between parents reduces offspring fitness in a natural population of Rhododendron brachycarpum

Background and Aims

A reduction in offspring fitness resulting from mating between neighbours is interpreted as biparental inbreeding depression. However, little is known about the relationship between the parents'' genetic relatedness and biparental inbreeding depression in their progeny in natural populations. This study assesses the effect of kinship between parents on the fitness of their progeny and the extent of spatial genetic structure in a natural population of Rhododendron brachycarpum.

Methods

Kinship coefficients between 11 858 pairs of plants among a natural population of 154 R. brachycarpum plants were estimated a priori using six microsatellite markers. Plants were genotyped, and pairs were selected from among 60 plants to vary the kinship from full-sib to unrelated. After a hand-pollination experiment among the 60 plants, offspring fitness was measured at the stages of seed maturation (i.e. ripening) under natural conditions, and seed germination and seedling survival under greenhouse conditions. In addition, spatial autocorrelation was used to assess the population''s genetic structure.

Key Results

Offspring fitness decreased significantly with increasing kinship between parents. However, the magnitude and timing of this effect differed among the life-cycle stages. Measures of inbreeding depression were 0·891 at seed maturation, 0·122 (but not significant) at seed germination and 0·506 at seedling survival. The local population spatial structure was significant, and the physical distance between parents mediated the level of inbreeding between them.

Conclusions

The level of inbreeding between individuals determines offspring fitness in R. brachycarpum, especially during seed maturation. Genetic relatedness between parents caused inbreeding depression in their progeny. Therefore, biparental inbreeding contributes little to reproduction and instead acts as a selection force that promotes outcrossing, as offspring of more distant (less related) parents survive better.     

Cushions of Thylacospermum caespitosum (Caryophyllaceae) do not facilitate other plants under extreme altitude and dry conditions in the north-west Himalayas

Background

Cushion plants are commonly considered as keystone nurse species that ameliorate the harsh conditions they inhabit in alpine ecosystems, thus facilitating other species and increasing alpine plant biodiversity. A literature search resulted in 25 key studies showing overwhelming facilitative effects of different cushion plants and hypothesizing greater facilitation with increased environmental severity (i.e. higher altitude and/or lower rainfall). At the same time, emerging ecological theory alongside the cushion-specific literature suggests that facilitation might not always occur under extreme environmental conditions, and especially under high altitude and dryness.

Methods

To assess these hypotheses, possible nursing effects of Thylacospermum caespitosum (Caryophyllaceae) were examined at extremely high altitude (5900 m a.s.l.) and in dry conditions (precipitation <100 mm year⁻¹) in Eastern Ladakh, Trans-Himalaya. This is, by far, the highest site, and the second driest, at which the effects of cushions have been studied so far.

Key Results

In accordance with the theoretical predictions, no nursing effects of T. caespitosum on other alpine plants were detected. The number and abundance of species were greater outside cushions than within and on the edge of cushions. None of the 13 species detected was positively associated with cushions, while nine of them were negatively associated. Plant diversity increased with the size of the area sampled outside cushions, but no species–area relationship was found within cushions.

Conclusions

The results support the emerging theoretical prediction of restricted facilitative effects under extreme combinations of cold and dryness, integrating these ideas in the context of the ecology of cushion plants. This evidence suggests that cases of missing strong facilitation are likely to be found in other extreme alpine conditions.