Seed rain,seedling establishment and clonal growth strategies on a glacier foreland

Abstract. Seed production, composition of the seed rain, germination, and seedling mortality, as well as vegetative growth characteristics of common pioneer plant species were studied on the foreland of the retreating Morteratsch glacier in the Swiss Alps. The frequency of diaspores trapped in different successional stages was related to their dispersal mode and was highly skewed towards a few species. Plenty of diaspores well adapted for dispersal by wind are a precondition for the most important pioneer species. Seeds from all pioneer species investigated had a good germination success, provided that the moisture content of the soil was high enough. However, requirement for seedling establishment differed among sites of increasing terrain age and among species. Only specialized pioneers such as Cerastium pedunulatum, Linaria alpina, Oxyria digyna and Saxifraga aizoides tolerate the cold and moist conditions near the glacier. However, these species are restricted to early successional stages. Seedlings of Epilobium fleischeri are affected not only by the cold and moist conditions near the glacier but also by moderately dry conditions on older sites. Availability of safe sites becomes crucial for most species with increasing age of sites and with drier conditions. Most species playing a dominant role during early succession and persisting during later successional stages have a distinctive ability to spread clonally and have a growth form with more or less widely spaced ramets: Achillea moschata. Cerastium pedunculatum. Epilobium fleischeri and Hieraium staticifolium. The growth strategy and demography of the clonal E. fleischeri is presented as an example. The life cycle of this species is characterized in succession by (1) the colonization of safe sites by small seeds adapted for wind dispersal, (2) horizontal spread by clonal growth, and (3) the persistence through phenotypic morphological plasticity in later successional stages. Seedling establishment and clonal growth are thus complementary mechanisms in plant succession on recently deglaciated terrain.     

Alpine species seed longevity in the soil in relation to seed size and shape – A 5-year burial experiment in the Central Alps

Long-term persistent seeds are generally expected to be small and spherically shaped. In contrast, transient seeds are often large, flattened and elongated. We hypothesized that alpine species follow these trends, and that most species of unstable alpine habitats have the potential to form long-term persistent seed banks. The latter would enable them to delay germination until favourable conditions are present in their environment thereby increasing the likelihood for establishment of seedlings. We selected nine alpine species of more or less unstable habitats (i.e. Achillea moschata, Artemisia genipi, Anthyllis vulneraria ssp. alpicola, Geum reptans, Linaria alpina, Oxyria digyna, Saxifraga aizoides, Saxifraga oppositifolia, and Trifolium pallescens) for this study. Seed longevity was estimated by a 5-year burial experiment in the field with seed excavations after 1 winter, 2 years and 5 years. Germination experiments in the growth chamber and viability tests were performed after each excavation. We detected longevity patterns ranging from transient to long-term persistent. Two species were classified as transient. All other species have the potential to form persistent seed banks, with almost half of the investigated species showing a trend for long-term persistence. Despite contradictory reports, long-term persistent seed banks are an important life history trait of species of unstable alpine habitats. We found that seed shape and hard-seededness are good predictors of seed persistence of alpine species. Seed size seems to be less important. The data from this study support the general hypotheses describing the relationship among seed size, seed shape and seed persistence in the soil.     

Differences in the location of subcotyledonary buds among Epilobium angustifolium L., E. dodonaei Vill. and E. fleischeri Hochst. (Onagraceae) and effects on architecture and population structure

STÖCKLIN, J., 1992. Differences in the location of subcotyledonary buds among Epilobium angustifolium L., E. dodonaei Vill. and E. fleischeri Hochst. (Onagraceae) and effects on architecture and population structure . Morphological and architectural features in the two closely related pioneer plant species Epilobium dodonaei and E. fleischeri are examined in cultivation and in the field and compared with E. angustifolium. In E. angustifolium , the aerial shoot system is renewed every year from buds on horizontal roots and results in a horizontal spread and a clonal growth form. In E. dodonaei , bud formation is restricted to the hypocotyl and in larger plants to the transitional region between root and shoot. Consequently this species shows no vegetative mobility and develops a shrub-like habit. The alpine E. fleischeri combines the habit of E. angustifolium and E. dodonaei and may either develop successive generations of shoots from the transitional region between root and shoot and/or exploit new areas by horizontal roots and the formation of shoots from root buds. The simple difference in the location of renewal buds is accentuated by cumulative growth. The study shows that E. dodonaei and E. fleischeri , which often are considered as subspecies, are separated by fundamental differences in their architectural models. The ecological and demographic implications of these differences are discussed.     

Seed weight increases with altitude in the Swiss Alps between related species but not among populations of individual species

Seed weight is a crucial plant life history trait, determining establishment success and dispersal ability. Especially in stressful environments, larger seeds may be selected at the expense of seed number, because larger seeds have a better chance of giving rise to an established offspring. We tested the hypotheses that between related species-pairs and among populations of single species a similar trend for increasing seed weight with increasing altitude should be present. Firstly, we measured seed weights from 29 species-pairs, with one species occurring in lowland areas and a congeneric species from high altitudes. Seeds of the alpine species were 28±8% larger than seeds from lowland species (P<0.01). Compared to the related lowland species, 55% of the alpine species had heavier seeds, 3% (one species) had lighter, and 41% had seeds of approximately equal weight. Secondly, we compared seed weights among populations of four species from different habitats and with different life histories. Seeds from between 11 and 34 populations per species were sampled along altitudinal gradients of 800–1,500 m (ca. 800 m in Scabiosa lucida, ca. 1,000 m in Saxifraga oppositifolia, ca. 1,000 m in Epilobium fleischeri, and ca. 1,500 m in Carex flacca). In all the four species, we found no indication for heavier seeds at higher altitudes. Our results indicate a selection pressure for species with heavier seeds at higher altitude, but the trend does not seem to operate across all cases. Phylogenetic constraints may limit the correlation among altitude and seed weight, operating particularly against selection for larger seed size, the closer populations and species are related to each other.     

Colonization dynamics of a clonal pioneer plant on a glacier foreland inferred from spatially explicit and size-structured matrix models

The regional distribution of a plant species is a result of the dynamics of extinctions and colonizations in suitable habitats, especially in strongly fragmented landscapes. Here, we studied the role of spatial dynamics of the long-lived, clonal pioneer plant Geum reptans occurring on glacier forelands in the European Alps. We used demographic data from several years and sites in the Swiss Alps in combination with dispersal data to parametrize a matrix model for G. reptans to simulate extinctions, colonizations and spatial spread of established populations on glacial forelands. We used different scenarios with varying germination rates, wind and animal dispersal capabilities, and modes of spatial spread (seed-only vs clonal spread), resulting in population growth rates (λ) ranging from 1.04 to 1.20. Our results suggest that due to the low germination rate (~1%) and the very limited wind dispersal distances (99.8% of seeds are dispersed < 5 m), G. reptans has a low probability of establishing new populations and a very low spatial spread by seed dispersal alone. In contrast to the low rate of establishment, the persistence of established populations is high and even populations of only a few individuals have an extinction probability of less than 25% within 100 years. This high persistency is partly due to clonal reproduction via aboveground stolons. Clonal reproduction increases the population size and contributes considerably to the spatial spread of established populations. Our simulation results together with the known pattern of molecular diversity of G. reptans indicate that the occurrence of populations of this species in the Alps is unlikely to be a result of recent colonizations by long-distance dispersal, but rather a result of post-glacial colonizations by large migrating populations that were fragmented when glaciers retreated. Additionally, our simulations suggest that the currently observed high rates of glacial retreat might be too fast for pioneer plants, such as G. reptans, to keep up with the retreating ice and therefore might threaten existing populations.     

Experimental evaluation of seed limitation in alpine snowbed plants

Background

The distribution and abundance of plants is controlled by the availability of seeds and of sites suitable for establishment. The relative importance of these two constraints is still contentious and possibly varies among species and ecosystems. In alpine landscapes, the role of seed limitation has traditionally been neglected, and the role of abiotic gradients emphasized.

Methodology/Principal Findings

We evaluated the importance of seed limitation for the incidence of four alpine snowbed species (Achillea atrata L., Achillea clusiana Tausch, Arabis caerulea L., Gnaphalium hoppeanum W. D. J. Koch) in local plant communities by comparing seedling emergence, seedling, juvenile and adult survival, juvenile and adult growth, flowering frequency as well as population growth rates λ of experimental plants transplanted into snowbed patches which were either occupied or unoccupied by the focal species. In addition, we accounted for possible effects of competition or facilitation on these rates by including a measure of neighbourhood biomass into the analysis. We found that only A. caerulea had significantly lower seedling and adult survival as well as a lower population growth rate in unoccupied sites whereas the vital rates of the other three species did not differ among occupied and unoccupied sites. By contrast, all species were sensitive to competitive effects of the surrounding vegetation in terms of at least one of the studied rates.

Conclusions/Significance

We conclude that seed and site limitation jointly determine the species composition of these snowbed plant communities and that constraining site factors include both abiotic conditions and biotic interactions. The traditional focus on abiotic gradients for explaining alpine plant distribution hence appears lopsided. The influence of seed limitation on the current distribution of these plants casts doubt on their ability to readily track shifting habitats under climate change unless seed production is considerably enhanced under a warmer climate.     

Seed dormancy and germination behaviour in two Euphrasia species (Orobanchaceae) occurring in the Swiss Alps

The optimal time for germination of a seed depends on environmental conditions of its habitat, the life cycle of the germinating plant, and the conditions for successful establishment, growing and reproduction. We studied the germination behaviour of the alpine annual Euphrasia minima and an alpine ecotype of E. salisburgensis in a lowland garden experiment. Seeds of both species and their hybrids germinated at constant (5 °C) and at varying temperatures (3–10 °C), and never before spring after seed ripening. Germination was spread over 3 years, which suggests that the seeds formed a persistent seed bank. The two species together with E. minima and E. minima hybrids differed significantly in the germination rate in the first and second spring.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 649–656.     

Mass effects,clonality, and phenology but not seed traits predict species success in colonizing restored grasslands

Preserved grasslands commonly host species‐rich plant communities but a large part of the grasslands were plowed up in the past. Their restoration often requires a long time and initial restoration measures might trigger ecosystem recovery, which is then followed by spontaneous colonization. We evaluate the establishment success of target grassland species, which were not sown but established spontaneously in the restored grasslands of Bílé Karpaty Mts., SE Czech Republic. According to their key functional traits and incidence in the landscape (mass effect; acquired from the results of a grid mapping project in the region), we examined the frequency of species and their mean cover in 82 restored grasslands. The best predictor of species frequency in the grasslands was their mass effect, followed by a high capacity for clonal growth and late phenology. Seed dispersal traits (seed mass, terminal velocity, epizoochory ranking index) and plant height had no significant effect. Specific leaf area was positively correlated with mass effect. Species having a high cover in the restored grasslands had a high capacity for clonal growth. In the preparation of seed mixtures, we should therefore consider that nonclonal species relying on regeneration from seeds will be generally less able to reproduce and should be promoted by artificial sowing. At the same time, species common in the landscape, which spread well clonally, and those with a late phenology, might be expected to colonize restored meadows on their own, so that sowing them is not necessary.     

Effects of lead contamination on the clonal propagative ability of Phragmites australis (common reed) grown in wet and dry environments

Clonal propagation is important for the survival and maintenance of the common reed Phragmites australis. Pot culture experiments were conducted to investigate the effects of lead (Pb) concentration (0, 500, 1500, 3000, 4500 mg·kg^?1) and water stress on the clonal reproductive ability of this species. The Pb concentration found in plant organs, in decreasing order, was roots >shoots >rhizomes. There was a negative relationship between the growth of clonal propagative modules (excluding axillary shoot buds) and Pb concentrations, which caused a decrease in biomass, rhizome growth and number of axillary and apical rhizome buds. Daughter axillary shoots exhibited a tolerance strategy, with no significant change in their number; the axillary and apical rhizome buds, daughter apical rhizome shoots and rhizomes exhibited compensatory growth during the late stage of Pb (excluding 4500 mg·kg^?1) treatment in a wet environment. Pb applications above 500 mg·kg^?1 reduced these parameters significantly in the drought treatment, except for the number of axillary shoot buds, which did not change. Our results indicate that clonal propagative resistance to Pb contamination can occur via tolerance strategies, compensatory growth and a Pb allocation strategy, enabling these reeds to maintain population stability in wet environments. However, clonal modular growth and reproductive ability were inhibited significantly by the interaction between drought and Pb, which would cause a decline in P. australis populations in a dry environment. Lead concentrations of 4500 and 500 mg·kg^?1 in soils might meet or exceed the Pb tolerance threshold of clonally propagated reeds in wet and dry environments, respectively.     

Timing of seed dispersal generates a bimodal seed bank depth distribution

The density of soil seed banks is normally highest at the soil surface and declines monotonically with depth. Sometimes, for a variety of reasons, peak density occurs below the surface but, except in severely disturbed soils, it is generally true that deeper seeds are older. In seasonally dry habitats that develop deep soil cracks during the dry season, it is possible that some seeds fall down cracks and rapidly become deeply buried. We investigated this possibility for three dominant clonal perennials (Scirpus maritimus, S. litoralis, and Juncus subulatus) in the Do?ana salt marsh, a nontidal marsh with a Mediterranean climate located in southwest Spain. Two species, which shed most of their seed during the dry season and have seeds with low buoyancy, had bimodal viable seed depth distributions, with peak densities at the surface and at 16-20 cm. A third species, which shed most seeds after soil cracks had closed and had seeds with high buoyancy, had viable seeds only in surface soil. Bimodal seed bank depth distributions may be relatively common in seasonally dry habitats with fine-textured soils, but their ecological significance has not been investigated.     

Effect of yak rumen content treatments on seed germination of 11 alpine meadow species on the Qinghai-Tibetan Plateau

The Qinghai-Tibetan Plateau is located in the ‘Third Pole’ of the world, characterized by a harsh environment. Despite this, the alpine meadow ecosystem have developed over a wide area but serious grassland degradation is threatening the ecological environment on the Plateau. Recruitment of new plants to the population, via germination and establishment, is vital to plant community survival. Previous work on the seeds in this area has mainly focused on community-wide germination strategies, seed germination characteristics and their correlations with seed size and seed mass. However, there have been no studies on the effects of soaking in rumen contents on the plant seed germination characteristics of alpine meadow species. The present study had two main objectives: (i) to determine the effect of fresh rumen content from yaks on seed germination characteristics and seedling growth of species common to the eastern Tibetan Plateau alpine meadow, and (ii) to develop an effective method to enhance seed germination. Seeds of 11 common species were collected together with fresh rumen content from three yaks that grazed there. Seed germination tests were conducted after they had been soaked in rumen content for one of six soaking periods (12, 24, 36, 48, 60 or 72 h). The seeds were incubated under natural light conditions of 8 h light at 25 °C and 16 h darkness at 15 °C, for the germination period of 34 days. The results showed that seed germination and seedling growth were affected by soaking time, seed coat completeness and seed type. After soaking in rumen content, the germination percentages of scarified (peeled or with the seed coat cut through) seeds of some species (true seeds Oxytropis ochrocephala and Medicago ruthenia var inschanica, nutlet Carex enervis, achenes Anemone rivularis and Polygonum sibiricum) and complete seeds of C. enervis, and A. rivularis were improved but the duration of soaking was also important. Seed germination of caryopsis Achnatherum inebrians (a toxic grass) was significantly inhibited by any exposure to rumen fluids. Scarified seeds generally had higher germination percentages than complete ones after treatment, but with the increase in soaking time, germination percentages declined and scarified seeds were more sensitive to the treatment than the complete seeds. After soaking in yak rumen content, the germination indices of scarified M. ruthenia at 12 h treatment, O. ochrocephala and achene Rumex acetosa at 12–24 h treatment, nutlet Kobresia humilis at 24 h treatment, P. sibiricum at 24–48 h treatment, C. enervis at 12–48 h treatment and A. rivularis at 12–60 h treatment were significantly higher than the control (P < 0.05), while the germination indices of complete C. enervis seeds at 12 h and 36 h treatment, and A. rivularis at 12–60 h treatment were significantly higher compared with the control. The germination indices of other species gradually decreased with the increase in soaking time. We concluded that yak rumen digestion could enhance, inhibit or not affect seed germination and seedling growth of the alpine meadow species, which might influence seedling recruitment, interspecific competition, and the plant community structure of the eastern Tibetan Plateau alpine meadow. Overall, yak digestion has a positive effect on alpine meadow seed germination and seed dispersal.     

Seed bank,seed size and dispersal in moisture gradients of temporary pools in Southern France

Plants reduce risk of extinction due to unpredictable rainfall by soil seed banks, dispersal or large seeds. However, seed size also increases independently in dry habitats, and since seed size is in a trade-off with seed number, size of seed banks is expected to increase in moister habitats. Therefore, we wanted to test if seed abundance in soil increases in wet habitats, if seed size increases in dry habitats, and if spread of seeds along the gradient is higher for plants of intermediate habitats in local moisture gradients.We studied 15 temporary pools in three biogeographically separated wetlands in Southern France. For each pool we studied five different moisture levels, totalling 75 local plant communities. We quantified soil seed bank by the seedling emergence method, seed size and an index of spatial spread of seeds in the soil for every species. We also quantified water levels for each plot.We found increasing abundance of seeds in the soil with increasing water levels but lower seed size and higher spread at intermediate water levels. When we controlled for niche position, we found no trade-off between seed size, spread and abundance in the soil seed bank.Type and importance of risk reduction strategies thus appeared to be strongly driven by the plant species’ moisture niche and the spatial arrangement of water levels.     

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.     

Genetic variation and reproduction strategy of Gentiana pannonica in different habitats

Gentiana pannonicascop. (Gentianaceae) is a long-lived perennial mountain species. It is a typical east−Alpine element. The centre of its distribution is situated in the eastern Alps, where the species occurs in alpine and subalpine areas. Besides the Alps, G. pannonica occurs also in the Bohemian Forest, Czech Republic. The reproduction strategy and genetic variation of this species endangered in the Czech Republic were studied. Genetic variation within and among six populations of G. pannonica in the Bohemian Forest and two populations in the Alps was studied using random amplified polymorphic DNAs (RAPD) markers. The variation found between populations from primary habitats and populations situated in the secondary mountain meadows accounted for 5% of the total genetic variation, whereas no significant genetic diversity was found between two different regions (Bohemian Forest, the Alps). Most of the variation is distributed among individuals within populations in the primary habitats, specifically 77% in the Bohemian Forest and 79% in the Alps. The pattern of among-population variation was substantially different between primary and secondary habitats. While among-population variation in primary habitats was low in both of the regions (21–23%), the proportion of RAPD variation among populations in secondary habitats was distinctly higher (65%). Within-population genetic variation was higher in primary habitats than in the secondary ones. This may be a consequence of processes such as genetic drift or bottleneck, and founder effects caused by changes in their distribution during the Holocene period. The differences of vegetation composition, seedlings recruitment and seed set were investigated in the field. Vegetation composition differed between primary and secondary habitats of G. pannonica and seedlings recruitment was recorded only in primary habitats. The reproduction output of the species under study is probably influenced by the availability of pollinators.     

Does pre-dispersal seed predation limit reproduction and population growth in the alpine clonal plant Geum reptans?

We studied the impact of the seed damaging gall midge larva Geomyia alpina on its perennial alpine host plant Geum reptans. We analysed the effect of seed predation on reproduction by seeds, i.e. seed number, seed mass, and seed viability and on growth and clonal propagation of non-protected plants in comparison to plants protected from predation by an insecticide. Additionally, we assessed the consequences of seed predation for population growth using matrix projection modelling. Seed predation resulted in a decrease in total seed mass per flower head by 23.8% in non-protected plants (P < 0.05). Individual seed mass decreased with increasing infestation intensity (P < 0.05). Seed number remained unaffected because the sucking feeding behaviour by gall midge larvae does not evoke seed abortion. Percent germination of seeds from non-protected plants was reduced by 97.9% compared to seeds from protected plants. According to reduced seed viability, modelling revealed a decrease in population growth rate from λ = 1.055 to λ = 1.041. Predation did neither influence total plant biomass nor biomass fractions. But stolon dry-weight of non-protected plants increased by 24.1% (P < 0.05), which may indicate a trade-off between sexual reproduction and clonal propagation. Our results demonstrate that despite substantial reduction of viable seeds, predation by gall midge larvae only slightly affected population growth of G. reptans suggesting that in this alpine species, persistence by longevity and clonal propagation can balance potential seed losses by predation, at least for local population growth.     

First report of seed dispersal by ants in Dicentra peregrina (Papaveraceae), an alpine plant in the Japanese Alps

Seed dispersal by ants, known as myrmecochory, is commonly observed among various plant taxa. The seeds of these plants have an elaiosome to attract ants. In Japan, myrmecochory has been well studied in several lowland plant species, but not in highland plant species that grow above the tree line. We investigated whether the seeds of Dicentra peregrina, known as the “queen of the alpine plants” in Japan, are carried by Formica gagatoides ants at 2510 m a.s.l. on Mt Norikura, Kita‐Alps, Japan. We observed F. gagatoides workers picking up D. peregrina seeds by grasping the elaiosome and carrying the seeds into their nests. We inferred from the observed ant behavior and the seed morphology that D. peregrina is a myrmecochorous species.     

Effect of temperature and light on seed germination of Erysimum naxense and Erysimum krendlii

Seed germination of two local Greek endemics was studied (Erysimum naxense, Erysimum krendlii). Seed viability was determined by using the tetrazolium method and germination was studied in synchronized cycles of five and four alternating temperatures [10/5 (for E. naxense only) and 15/10, 20/15, 25/20, and 30/25°C for both species, in cycles of 16 h day/8 h night], and in five light regimes (red, blue, green, white, and dark). Germination of E. naxense and E. krendlii seeds was determined daily for six and five weeks, respectively, with the data analyzed as viability adjusted accumulative seed germination at the end of each week. E. naxense’s seed viability was higher (90%) than that of E. krendlii (64%); seed germination (%) of both increased at low alternating temperatures (10/5°C, 15/10°C, 20/15°C). Germination of E. naxense seeds at low temperatures was light-independent, whereas at high temperatures it was increased with red light. Germination of E. krendlii seeds was inconsistently affected by light at the temperatures studied. Percentages of seed germination of both species were higher in experimental conditions similar to the ones of their natural habitats during autumn and/or spring (facilitated with Geographic Information Systems). These conclusions provide guidelines for species-specific propagation protocols and ex situ conservation.     

Effect of Light and Moisture Conditions and Seed Age on Germination of Three Closely Related Myosotis Species

Germination of three closely related species from the Myosotis palustris group (M. nemorosa, M. palustris subsp. laxiflora, M. caespitosa) differing in their habitats and capacity for clonal growth, was compared in two greenhouse experiments. To evaluate both inter- and intraspecific variation, each species was represented by seeds from several populations. Final germination percentage and germination rates T₅₀ were compared both between species and populations within species. In the first experiment, we studied the influence of two external factors, moisture and light. Four moisture levels (dry, wet, periodically flooded and permanently flooded soil) and three types of shading (without shading, shaded with green foil, shaded with solid paper sheet) were combined in a complete factorial design. In all three species, total germination percentage was the same in the three wettest treatments, and decreased in the dry treatment. Germination in the treatments shaded with green foil (simulating vegetation cover, which changed light quality) was significantly slower than in treatments without shading and treatments shaded with a solid paper sheet. There were significant differences among species, but we also found very pronounced differences among populations within a species. M. caespitosa had the uniformly highest germination percentage (reaching in some cases 100%) and also fastest germination. Germination of M. palustris subsp. laxiflora populations was slower and reached lower final proportions, and medium variability among populations. Inter-population variability in the final germination percentage was highest, and the final germination the lowest in M. nemorosa. In addition, M. nemorosa, a species typical for permanent meadow communities was delayed by permanent flooding. In the second experiment, we studied the effects of seed age and storage conditions. Three combinations of seed age and storage were used: younger seeds (half year old) with no-chilling, younger seeds with chilling and old seeds (three years old) with chilling. M. caespitosa had again the highest final germination percentage and fastest germination rates T₅₀. In addition, final germination percentage of this species slightly increased with seed age, whereas it decreased in the other two species. The germination behaviour corresponded well to expectation based on species life histories and habitat preferences. Remarkably stable and high germination percentages and fastest germination rates T₅₀ were ascertained in M. caespitosa, a species of disturbed habitats, with lowest capacity for clonal spread. M. palustris subsp. laxiflora (species with highest clonal capacity) and M. nemorosa (species with medium clonal potential) achieved lower, but still very high final germination percentage. In addition, M. nemorosa showed the highest inter-populations variability in our experiments.     

Buried seed population in the herbaceous lomas on Loma Ancon in the coastal desert of central Peru

Herbaceous lomas in the Peruvian coastal desert, of South America establish in spring, and its habitat is limited to the southern or southwestern slopes along the coast that are affected by thick fog. The time of appearance, the duration and the thickness of the fog vary greatly from year to year, so the lomas can grow only in habitats with enough water to, sustain seed germination and plant growth. This paper studies the species composition and density of the buried seed population, of the herbaceous lomas of Loma Ancon in order to clarify the mechanisms of the lomas' establishment. The mean number of species with viable seeds was about, 12 spp. m⁻² and that of dead seeds was about 22 spp. m⁻². The dominant species wereSolanum tuberiferum, S. pinnatifidum andNolana humifusa, both in viable and dead seeds. Viable seed density was about 5000–8000 seeds m⁻², which is comparable with the seed densities of other herbaceous communities. Dead seed density was about 15000–27000 seeds m⁻², or nearly three times the viable seed density, because the rate of decomposition was slow in the extremely dry conditions. The net increase of viable seeds by seed production was estimated at about 5000 seeds m⁻² in 1980, and the increase in the number of dead seeds was 2200 seeds m⁻².     

The role of fleshy pericarp in seed germination and dispersal under flooded conditions in three wetland forest species

In flooded habitats, inundations affect important forest regeneration processes, such as seed dispersal and germination. The main seed dispersal mechanism used by species in Austral South American temperate swamp and riparian forests is endozoochory, which releases seeds from the fleshy pericarp. Endozoochory could be favorable or unfavorable in wetland habitats, since this mechanism exposes seeds directly to water and can, in some cases, be detrimental to germination. In this study, we studied whether or not the fleshy pericarp favors germination after the flooding period's end. Furthermore, we quantified if the number of days which the fruit was found to be floating related to its germination success. We used the seeds of three common fleshy fruit species of flooded habitats from southern Chile, the trees Luma apiculata and Rhaphithamnus spinosus, and the vine Luzuriaga radicans. We simulated flooding periods of 7, 15, 30 and 45 days submerging seeds, with and without pericarps, in water. We found that the pericarp's presence significantly increased Luma's germination success and significantly decreased that of Luzuriaga. The germination of Rhaphithamnus was low after periods of flooding in both seed treatments, with no significant differences found between them. The fruits could float for an average of one to four weeks, depending on the species, which did not relate to their germination success. These results show that germination was affected by the presence of fleshy pericarps in flooded conditions and furthermore, that flotation allows for hydrochory from one week to one month. We suggest that in swamp forests multiple seed dispersal mechanisms are advantageous, especially for fleshy-fruited species.