Seed germination responses to seasonal temperature and drought stress are species‐specific but not related to seed size in a desert steppe: Implications for effect of climate change on community structure

Investigating how seed germination of multiple species in an ecosystem responds to environmental conditions is crucial for understanding the mechanisms for community structure and biodiversity maintenance. However, knowledge of seed germination response of species to environmental conditions is still scarce at the community level. We hypothesized that responses of seed germination to environmental conditions differ among species at the community level, and that germination response is not correlated with seed size. To test this hypothesis, we determined the response of seed germination of 20 common species in the Siziwang Desert Steppe, China, to seasonal temperature regimes (representing April, May, June, and July) and drought stress (0, ?0.003, ?0.027, ?0.155, and ?0.87 MPa). Seed germination percentage increased with increasing temperature regime, but Allium ramosum, Allium tenuissimum, Artemisia annua, Artemisia mongolica, Artemisia scoparia, Artemisia sieversiana, Bassia dasyphylla, Kochia prastrata, and Neopallasia pectinata germinated to >60% in the lowest temperature regime (April). Germination decreased with increasing water stress, but Allium ramosum, Artemisia annua, Artemisia scoparia, Bassia dasyphylla, Heteropappus altaicus, Kochia prastrata, Neopallasia pectinata, and Potentilla tanacetifolia germinated to near 60% at ?0.87 MPa. Among these eight species, germination of six was tolerant to both temperature and water stress. Mean germination percentage in the four temperature regimes and the five water potentials was not significantly correlated with seed mass or seed area, which were highly correlated. Our results suggest that the species‐specific germination responses to environmental conditions are important in structuring the desert steppe community and have implications for predicting community structure under climate change. Thus, the predicted warmer and dryer climate will favor germination of drought‐tolerant species, resulting in altered proportions of germinants of different species and subsequently change in community composition of the desert steppe.     

Nurse‐plant effects on the seed biology and germination of desert annuals

Nurse‐plants generally have positive effects on understorey species by creating more suitable conditions for stress‐intolerant plants relative to open micro‐habitats. However, long‐term effects of this plant–plant facilitation system have been rarely examined. Seeds of five desert annual species from Atiquipa coastal desert in Southern Peru were used to examine whether different microenvironmental conditions under the nurse‐plants Caesalpinia spinosa Molina (Kuntze) lead to differences in seed biology and germinability of annual plants relative to open, canopy‐free conditions. Seeds collected from plants associated with nurse‐plants were predicted to be (i) larger due to more favourable growing conditions, (ii) more viable and with greater germination rates, (iii) less variable in size and viability due to reduced environmental heterogeneity, and (iv) to germinate faster to avoid apparent competition with other annuals. Seed attribute measurements and germination trials in growth chambers were used to test these predictions. Although the plant abundance of only 2 of 5 species was strongly facilitated by the nurse‐plant, no significant differences were found in seed mass, viability or relative variability between understorey and open micro‐habitats for any of the species. Contrary to our predictions, final seed germination rates of seeds from open micro‐habitats were higher, and the open micro‐habitat treatment was more favourable for germination of seeds from both open and understorey environments. Taken together, these results suggest that plant–plant facilitation does not necessarily affect seed biology traits. Further studies addressing larger distribution ranges and/or density gradients of understorey species will illuminate the potential evolutionary effects of nurse‐plants.     

Does perianth colour affect the seed germination of two desert shrubs under different storage periods and conditions?

Haloxylon salicornicum and Salsola vermiculata (Amaranthaceae) are two perennial shrubs producing fruits with differently coloured perianths (yellow vs pink) on separate individuals. To test the impact of different storage periods (four, eight and twelve months) and temperature conditions (freezing, room temperature and natural field temperatures) on the seed germination of the two seed categories, germination tests were carried out on both species. For both species, collected from wild plants in the Arabian desert (Qatar), seed germination was tested under two photoperiods (light 12/12; dark 24 hours) at 20/30°C for winged (intact) and de‐winged seeds. For each species, all the considered factors (perianth colour, wing presence, photoperiod, storage period and storage conditions) had significant effects on seed germination (p < 0.001), while the interactions among factors varied in their level of significance depending on the species. In both species, yellow seeds germinated considerably better than the pink ones, the presence of wings slowed the germination, and both storage period and temperature conditions highly influenced their germination ability. Our results indicate that seeds from different perianth colours have different germination strategies to delay their germination and distribute the species through time by escaping germination time by imposing some kind of after‐ripening or seed dormancy. The variation in storage time and storage conditions might help in extending their germination period, where one set of seeds germinate immediately, and the other set of seeds becomes dormant and thus contributes to the formation of a soil seed bank for the long‐term recruitment of seedlings. This study highlights an ecological advantageous strategy for these two species growing in the unpredictable desert habitat.     

Effect of environmental factors on seed germination and emergence of Lepidium vesicarium

Lepidium vesicarium is a weed species with a wide distribution in the rangelands and dry‐land farming in East Azarbaijan, Iran. The experiments were undertaken to assay the effects of light, temperature, pH, osmotic potential, NaCl concentration and burial depth on seed germination and emergence of L. vesicarium. Germination was maintained at high levels (> 80%) over a wide day/night temperature range (10/5 to 30/20°C), but a severe reduction in the germination rate of L. vesicarium was found below 20/10°C. Germination of L. vesicarium was influenced by different light/dark regimes, as the germination rate was highest at 16 h light for the all treatments (0, 8, 12, 16 and 24 h light). Germination was 92–95% over a wide range of pH (2‐10). Germination was >50% at a water potential of ?0.7 MPa and salinity of 21 dS/m, indicating that drought and salt conditions have a minimal impact on seed germination. With increasing burial depth from 0 to 2 cm, the number of days required for 50% emergence increased and no germination was observed at burial depths deeper than 3 cm. This suggests that L. vesicarium would become troublesome in the rangelands and for growers in reduced‐tillage cropping systems. The ability to emerge from shallow depths, coupled with tolerance of a wide pH range, drought and salinity at germination, should be taken into account when managing this weed species.     

Effects of maternal salinity on salt tolerance during germination of Suaeda aegyptiaca,a facultative halophyte in the Arab Gulf desert

Suaeda aegyptiaca is a facultative halophyte found in saline and non‐saline habitats of the Arab Gulf desert, which produces small‐sized undispersible seeds. The interactive effects of maternal salinity and other environmental conditions, such as salinity, light and temperatures, that are prevailing during seed germination have received little attention for a facultative halophyte. This study tested the effects of maternal salinity on salt tolerance during seed germination of S. aegyptiaca under different light and temperature regimes. Seeds collected from both saline and non‐saline habitats of the United Arab Emirates (UAE) were germinated in 0, 50, 100, 200 and 400 mM NaCl, and incubated at 15/25°C, 20/30°C and 25/35°C in both 12‐h light/12‐h dark regimes and continuous darkness. Generally, seeds of the non‐saline habitat were 56% heavier and attained greater germination at the lower temperatures than seeds of the saline habitat. Seeds of the saline habitat germinated better in saline solutions at higher temperatures and in light. Germination was faster for seeds of the saline habitat than for seeds of non‐saline habitats. Germination recovery after transfer to distilled water was significantly greater for seeds from the non‐saline habitat, compared with seeds from saline habitats. Recovery was greater at lower and/or moderate temperatures, compared with at higher temperatures. Germination was significantly faster during recovery, compared with in the saline solutions. The study indicates that the maternal effect of salinity was confounded with the seed‐size effect and it cannot be conclusively confirmed.     

Phylogeny,habitat together with biological and ecological factors can influence germination of 36 subalpine Rhododendron species from the eastern Tibetan Plateau

The reproductive stages of the life cycle are crucial in explaining the distribution patterns of plant species because of their extreme vulnerability to environmental conditions. Despite reported evidence that seed germination is related to habitat macroclimatic characteristics, such as mean annual temperature, the effect of this trait in controlling plant species distribution has not yet been systematically and quantitatively evaluated. To learn whether seed germination can predict species distribution along altitude gradients, we examined germination data of 36 Rhododendron species in southeastern Tibet originating from contrasting altitudes, habitats, plant heights, seed masses, and phylogenies. Germination varied significantly with altitude, habitat, plant height, and phylogeny and was higher in the light than in the dark. Germination percentage was highest at 10:20°C in the light and 15:25°C in the dark. As altitude increased, germination percentages first rose and then decreased, being highest at 3,500–4,000 m. Germination percentage and rate were highest on rocky slopes, increasing as seed mass and plant height rose. Variations in germination percentage and rate were not significant at subgenera, section, and subsection levels, but they were significant at species level. The results suggested that the relationship between germination and altitude may provide insights into species distribution patterns. Further, germination patterns are a result of long‐term evolution as well as taxonomic constraints.     

Responses of Primula vulgaris to light quality in the maternal and germination environments

• In the model species Arabidopsis thaliana phytochromes mediate dormancy and germination responses to seasonal cues experienced during seed maturation on the maternal plants. However, the effect of the maternal light environment on seed germination in native wild species has not been well studied. This is particularly important given its practical application in the context of environmental restoration, when there can be marked changes in the canopy.
• Plants of Primula vulgaris were grown in the field over two vegetative seasons under four shading treatments from low to high ratio of red to far‐red light (R:FR). Leaf and seed traits were assessed in response to the light treatments. The germination of seeds from these four maternal environments (pre‐dispersal) was investigated at seven light and five temperature treatments (post‐dispersal).
• Thinner leaves, larger leaf area and greater chlorophyll content were found in plants growing in reduced R:FR. Shading in the maternal environment led to increased seed size and yield, although the conditions experienced by the maternal plants had no effect on seed germination. Seeds responded strongly to the cues experienced in their immediate germination environment. Germination was always enhanced under higher R:FR conditions.
• The observed phenotypic trait variation plays a major role in the ability of P. vulgaris to grow in a wide range of light conditions. However, the increased germination capacity in response to a higher R:FR for all maternal environments suggests potential for seedling establishment under vegetative shade only in the presence of canopy gaps.

     

Evaluating asymbiotic seed culture methods and establishing Disa (Orchidaceae) germinability in vitro: relationships, requirements and first-time reports

Many Disa species (80%) have not been germinated in vitro, fuelling conservation fears. Winter-rainfall species are typically germinable, whilst summer-rainfall species are exclusively intractable in vitro. We aimed to establish the in vitro germination requirements for previously ungerminated Disa species. Conventional asymbiotic seed culture protocols were reviewed and their efficacy in summer-rainfall Disa established. Mature seed was subjected to manipulations of media composition and viscosity, incubation temperature and illumination. Immature cultures were also established across four seed maturity classes. Four first-time germinations (D. cooperi, D. nervosa, D. pulchra and D. woodii) resulted from media with increased water availability. Germination rate (>12 weeks), percentage (<30%) and synchrony were not in accordance with values reported for winter-rainfall species (>80% germination in 8 weeks). Germination of immature seed under control conditions was similar to mature seed germination under modified conditions, but neither percentage approached the calculated germination potential (∼viability). Germination control in Disa is proposed as a trade-off between water availability and the presence of phyto-inhibitors in the environment of the embryo—two features typical of seeds exhibiting water-impermeable dormancy.     

The role of seed germination in the invasion process of Honey locust (Gleditsia triacanthos L., Fabaceae): comparison with a native confamilial

Identifying plant traits that promote invasiveness has been a major goal in invasion ecology. Germination plays a central role in the life cycle of plants and therefore could be a key trait in determining species invasiveness. In this study, seed germination of two confamilial, co‐occurring species that share ecological characteristics, the exotic invasive Gleditsia triacanthos L and the native Acacia aroma Gillies ex. Hook. & Arn., was compared. Seeds were obtained from individuals of three localities in the Chaco Serrano region of Córdoba, Argentina. Percent of seed germination and mean germination time were recorded in chemically and mechanically scarified seeds, and the former variable was also recorded in seeds subjected to: passage through the digestive tract of dispersers, fire simulations, fire simulation plus mechanical scarification, seed longevity, and dormancy break over time. In general, both species showed similar germination percentage. However, non‐scarified seeds of the exotic species lost physical dormancy when subjected to experiments of dormancy break over time, whereas, the native species had shorter mean germination time. The greater percentage of seed germination over time of the exotic species than of the native one might be triggering the spread of the former, whereas the shorter mean germination time might be hindering its expansion to more arid regions. The study of different mechanisms for achieving seed germination, particularly in hard seed species, could provide important information on the expansion of invasive species as well as useful knowledge for their management.     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Seed germination of three species of <Emphasis Type="Italic">Vallisneria</Emphasis> (Hydrocharitaceae), and the effects of freshwater microalgae

Two experiments were conducted to investigate seed germination under natural temperature and light regimes and to evaluate the influence of freshwater microalgae on seed germination of threeVallisneria species. Seeds exposed to natural seasonal temperature and light changes for 24 months germinated only in spring, perhaps indicating an annual dormancy/non-dormancy cycle. The ecological background of the natural habitat seems to play predominant roles in seed germinability. Mean cumulative seed germination percentages of Vallisneria natans, V. denseserrulata, andV.spinulosa of the first year were 35.2, 19.2, and 11.2%, respectively, and of the second year were 80, 72, and 32.4%, respectively. Germination rates differed significantly among the three Vallisneria species exposed to natural seasonal temperature and light changes in the first year, but differed only between Vallisneria spinulosa and the other two species in the second year. The differences of germination may influence the geographical distribution and thus be related to different survival strategies of the species. Seed germination rates in culture solution inoculated with algae were significantly higher than in solution with no algae. A strong correlation occurred between total biomass of freshwater microalgae and seed germination of the three studied species. Apparently, the extracellular products of freshwater microalgae may play an important role in seed germination. These results support the hypothesis that Vallisneria colonization may be mediated by algae facilitation.     

Seed moisture content affects afterripening and smoke responsiveness in three sympatric Australian native species from fire‐prone environments

Germination of freshly collected seeds of three sympatric herbaceous species native to fire‐prone environments in south‐western Australia was significantly improved through the application of novel combinations of dry heat, gibberellic acid, smoke water and dry afterripening. For fresh seeds, combinations of dry heat, gibberellic acid and/or smoke water resulted in >80% germination in Austrostipa elegantissima (Poaceae) and Stylidium affine (Stylidaceae) seeds and >60% germination in Conostylis candicans (Haemodoraceae) seeds, compared with <10% germination of control seeds. For fresh seeds, two broad germination patterns were observed in response to smoke water: nil – low germination for both control and smoke water‐treated seeds (A. elegantissima and S. affine); and a significant smoke response (35%) compared with control seeds (1%) (C. candicans). During afterripening, high germination for A. elegantissima seeds was achieved following 3 months storage of seeds at equilibrium relative humidities of 23–75%, but seeds stored at 5–13% equilibrium relative humidities took 6–36 months to achieve similar levels of germination. Germination of C. candicans seeds also increased after 3 months storage, to >60% at each equilibrium relative humidity and further increases over time were slight. For S. affine seeds >60% germination was achieved only after 36 months storage at 50% equilibrium relative humidity. Seeds from all three species were smoke‐responsive at some point, but the interaction/effects of afterripening on the smoke response varied significantly between species. This study highlights an apparent effect of seed dormancy status on response to smoke and a surprisingly high level of ecological variation in pre‐germination requirements (cues) for these co‐occurring species that may relate to variation(s) in microsite selection forces operating on the soil seed bank of the different species.     

Host influence on germination and reproduction of the facultative hemi‐parasitic weed Rhamphicarpa fistulosa

Rice Vampireweed, Rhamphicarpa fistulosa, was a minor parasitic weed until recently when rice cultivation in sub‐Saharan Africa was expanded into marginal wetlands, that are the parasite's natural habitat. Unlike most of the parasitic weeds, R. fistulosa is facultative, meaning that the parasite is able to complete its life cycle without a host. However, when not connected to a host plant, its biomass and seed production is lower. Because very little is known regarding the germination ecology of the parasite, the main objective of our study was to identify the cues that favour germination. We hypothesised that, first, being a wetland species, germination of R. fistulosa is stimulated by light and high soil moisture. Second, we hypothesised that if host plant presence increases its reproductive output then a germination stimulatory effect from host presence is likely to have developed. A Petri‐dish and pot experiment showed that light and completely saturated soils were a requirement for germination, demonstrating that germination requirements of R. fistulosa are typical of species that grow in environments with fluctuating water levels. A pot experiment in which five infestation levels of R. fistulosa were installed in the absence and presence of a rice plant, showed that host plant presence resulted in a 3.7 times higher seed production rate and a 15% larger average seed size. Despite this reproductive advantage, a pot experiment with three rice cultivars, selected because of their difference in strigolactone production, showed that host plant presence, regardless of the development stage, did not influence the emergence rate of R. fistulosa. In a follow‐up study, the germination stimulation effect of root exudates collected from the same three rice cultivars and a treatment consisting of an artificial germination stimulant (GR24) was compared with a treatment consisting of plain water. In these treatments, seeds of R. fistulosa were compared with seeds of the obligate parasite Striga hermonthica. Germination of S. hermonthica was strongly advanced by the presence of root exudates and GR24 but was completely absent in water, whereas germination of R. fistulosa in all treatments was similar to that in plain water. The absence of a host recognition mechanism at the germination stage suggests that the regulation of germination through light and soil moisture is near optimal. Our finding might also indicate that for this facultative parasitic plant species, a more opportunistic germination strategy is superior. Implications of the findings for management of R. fistulosa in rice cultivation are discussed.     

Seed characteristics and soil surface patch type interact to affect germination of semi-arid woodland species

Biological soil crusts are common in many arid and semi-arid regions and they can alter microenvironments which are likely to directly and indirectly influence vascular plant establishment. The effect of biological soil crusts on germination is also influenced by the biological characteristics of the seeds themselves but rarely have the effects of both crust type and seed morphology on germination been examined in the same study. In this study, seed of five semi-arid woodland species with contrasting seed morphology were sown on top of patch types that commonly occur in natural woodlands (foliose lichen, short-turf moss, tree leaf litter, disturbed crust) and their emergence was followed. Percent germination varied between patch types and, for the largest-seeded species (Maireana excavata), final germination was significantly lower on the biological soil crust and litter patch types because they strongly acted as a physical barrier to seed penetration into the soil. The remaining four species showed no significant differences in final percent germination with patch type because most seeds either completely or partially penetrated the surface layer. Germination time courses, however, showed that biological soil crusts delayed the timing of germination of these species. Hence, soil crusts might differentially affect the spatial patterning of species in semi-arid woodlands by their subtle influence on seedling emergence that is determined by differences in seed morphology and subsequent positioning within crusts.     

Soil seed bank,fire season,and temporal patterns of germination in a seeder-dominated Mediterranean shrubland

Soil seed banks play a major role in the post-fire regeneration of Mediterranean shrublands. They vary throughout the year in species composition, abundance, and readiness to germinate. After fire, germination occurs mainly during the following fall to spring. Time of germination can determine recruitment success. It is unclear what factors control post-fire germination and its timing. We tested the effects of season and fire on the readily germinable soil seed bank of a seeder-dominated shrubland. Plots were burned early and late in the summer season (ES, LS). Soil samples were collected before and after fire, and germinated in a chamber simulating successively autumn, winter, and spring conditions. Samples were kept moistened at all times. Fire intensity was similar between ES and LS. Several species of Cistus and herbs, mostly annuals, were dominant. Most germination occurred during the simulated-autumn period, with little subsequent germination during the following two periods. Germination speed (T ₅₀) during simulated-autumn was similar for shrubs and herbs, and independent of season or fire. Germination was lower for two shrubs (Rosmarinus officinalis, Cistus salvifolius) and higher for herbaceous dicots in LS than in ES soils. Fire reduced monocots and enhanced Cistus. Germination period significantly interacted with fire and season in some groups or species, altering the simulated-autumn germination peak. We demonstrate that the seed bank can germinate swiftly under simulated-autumn conditions. Hence, water availability is the main controlling factor of germination. Fire season differentially affected some species or groups, and could affect the post-fire regeneration.     

Changes in Germination Responses of Silene secundiflora in Relation to the Climate of its Habitat

Silene secundiflora is a Mediterranean species restricted in range to the southern coasts of Spain and the Balearic Islands. Germination tests were made in incubators and on thermo-gradient bars to evaluate changes in temperature-responses at intervals after seed harvest. Results were compared to average temperatures occurring in the region and interpreted in relation to the climatic characteristics of the area. Germination of freshly harvested seed was restricted to a narrow temperature band between ca. 7 and 16°C. After-ripening during the first four months resulted in a progressive widening of the temperature band favouring germination, but this remained below average summer temperatures. Exposure of imbibed seed to high temperatures (31°C) reduced the subsequent temperature maximum for germination by about 4°C, but did not induce a fully effective secondary dormancy. The results appeared to be well correlated with a situation in which germination normally occurs in September at the start of the winter growing season.     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

A field study of seed germination in the endangered Trillium reliquum Freeman (Trilliaceae)

Previous studies examining the seeds of most Trillium species have reported double dormancy, a type of seed dormancy where two cold periods and one warm period are needed for complete germination. In the present paper, we describe a field study examining the federally endangered Trillium reliquum Freeman (Trilliaceae) in which moderate to high numbers of seeds germinated after one winter following seed production. Sixteen baskets with seeds were placed in four T. reliquum populations (four baskets in each population) in Georgia, USA, in June 2005. In spring 2006, all seed baskets contained seedlings. Germination percentages ranged from 33.3 to 83.3% across sites with a mean of 56.9 ± 3.9%. Trillium reliquum had higher germination percentages compared with other field‐based germination studies with other Trillium species. Our findings will inform future demographic studies of T. reliquum and suggest that double dormancy in seeds may not be as widespread as previously reported within the genus Trillium.     

Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.