Soil salinity regulates spatial-temporal heterogeneity of seed germination and seedbank persistence of an annual diaspore-trimorphic halophyte in northern China.

BACKGROUND AND AIMS: Seed heteromorphism is a plant strategy that an individual plant produces two or more distinct types of diaspores, which have diverse morphology, dispersal ability, ecological functions and different effects on plant life history traits. The aim of this study was to test the effects of seasonal soil salinity and burial depth on the dynamics of dormancy/germination and persistence/depletion of buried trimorphic diaspores of a desert annual halophyte Atriplex centralasiatica. METHODS: We investigated the effects of salinity and seasonal fluctuations of temperature on germination, recovery of germination and mortality of types A, B, C diaspores of A. centralasiatica in the laboratory and buried diaspores in situ at four soil salinities and three depths. Diaspores were collected monthly from the seedbank from December 2016 to November 2018, and the number of viable diaspores remaining (not depleted) and their germinability were determined. RESULTS: Non-dormant type A diaspores were depleted in the low salinity "window" in the first year. Dormant diaspore types B and C germinated to high percentages at 0.3 and 0.1 mol L-1 soil salinity, respectively. High salinity and shallow burial delayed depletion of diaspore types B and C. High salinity delayed depletion time of the three diaspore types and delayed dormancy release of types B and C diaspores from autumn to spring. Soil salinity modified the response of diaspores in the seedbank by delaying seed dormancy release in autum and winter and by providing a low-salt concentration window for germination of non-dormant diaspores in spring and early summer. CONCLUSIONS: Buried trimorphic diaspores of annual desert halophyte A. centralasiatica exhibited diverse dormancy/germination behavior in respond to seasonal soil salinity fluctuation. Prolonging persistence of the seedbank and delaying depletion of diaspores under salt stress in situ primarily is due to inhibition of dormancy-break. The differences in dormancy/germination and seed persistence in the soil seedbank may be a bet-hadging strategy adapted to stressful temporal and spatial heterogeneity, and allows A. centralasiatica to persist in the unpredictable cold desert enevironment.

A long-term experiment reveals no trade-off between seed persistence and seedling emergence.

Information on seed persistence and seedling emergence from the soil seed bank is critical for understanding species coexistence and predicting community dynamics. However, quantifying seed persistence in the soil is challenging; thus, its association with other life-history traits is poorly known on a broad scale. Using germination phenology for 349 species in a 42-yr experiment, we quantified the persistence-emergence correlations and their associations with intrinsic regeneration traits using Bayesian phylogenetic multilevel models. We showed no trade-off between seed persistence and seedling emergence. Physically dormant seeds were more persistent but exhibited lower emergence than nondormant seeds. Monocarpic species had both higher persistence and emergence than polycarpic species. Seed mass posed a marginal proxy for persistence, while emergence almost doubled from the smallest to the largest seeds. This study challenges the traditional assumption and is the first demonstration of noncorrelation between persistence and emergence, probably owing to the complexity of regenerative strategies. Species with short persistence and low emergence would be the most vulnerable for in situ conservation. Our analyses of this unique, long-term dataset provide a strong incentive for further experimental studies and a rich data resource for future syntheses.

Environmental and seed-position effects on viability and germination of buried seeds of an invasive diaspore-heteromorphic annual grass.

Environmental factors, such as temperature and moisture, and plant factors, such as seed position on the mother plant, can affect seed viability and germination. However, little is known about the viability and germination of seeds in different positions on the mother plant after burial in soil under natural environmental conditions. Here, diaspores from three positions on a compound spike and seeds from two/three positions in a diaspore of the invasive diaspore-heteromorphic annual grass Aegilops tauschii were buried at four depths for more than 2 years (1-26 months) under natural conditions and viability and germination monitored monthly. Viability of seeds in each diaspore/seed position decreased as burial depth and duration increased and was associated with changes in soil temperature and moisture. Germination was highest at 2 cm and lowest at 10 cm soil depths, with peaks and valleys in autumn/spring and winter/summer, respectively. Overall, seeds in distal diaspore and distal seed positions had higher germination percentages than those in basal diaspore and basal seed positions, but basal ones lived longer than distal ones. Chemical content of fresh diaspores/seeds was related to diaspore/seed position effects on seed germination and viability during burial. We conclude that seeds in distal diaspores/seed positions have a 'high risk' strategy and those in basal positions a 'low risk' strategy. The two risk strategies may act as a bet-hedging strategy that spreads risks of germination failure in the soil seed bank over time, thereby facilitating the survival and invasiveness of A. tauschii.

Trade-offs between diaspore dispersal and dormancy within a spike of the invasive annual grass Aegilops tauschii.

MAIN CONCLUSION: Differences in dispersal and dormancy of heteromorphic diaspores of Aegilos tauschii may increase its flexibility to invade/occupy weedy unpredictable habitats by spreading risk in space and time. In plant species that produce dimorphic seeds, there often is a negative relationship between dispersal and dormancy, with high dispersal-low dormancy in one morph and low dispersal-high dormancy in the other, which may function as a bet-hedging strategy that spreads the risk of survival and ensures reproductive success. However, the relationship between dispersal and dormancy and its ecological consequences in invasive annual grasses that produce heteromorphic diaspores is not well studied. We compared dispersal and dormancy responses of diaspores from the basal (proximal) to the distal position on compound spikes of Aegilops tauschii, an invasive grass with heteromorphic diaspores. Dispersal ability increased and degree of dormancy decreased as diaspore position on a spike increased from basal to distal. There was a significant positive correlation between length of awns and dispersal ability, and awn removal significantly promoted seed germination. Germination was positively correlated with GA concentration and negatively correlated with ABA concentration, and the ABA: GA ratio was high in seeds with low germination/high dormancy. Thus, there was a continuous inverse-linear relationship between diaspore dispersal ability and degree of dormancy. This negative relationship between diaspore dispersal and degree of dormancy at different positions on a spike of Aegilops tauschii may facilitate seedling survival in space and time.

Seed dormancy in space and time: global distribution, paleoclimatic and present climatic drivers, and evolutionary adaptations.

Seed dormancy is an important life history state that increases survival and fitness of seed plants, and thus it has attracted much attention. However, global biogeography, effects of paleoenvironment, evolutionary roles of dormancy transitions, and differences in adaptations of seed dormancy between life-forms are poorly understood. We compiled global distribution records for seed dormancy of 12 743 species and their phylogeny to explore the biogeographic patterns, environmental drivers, and evolutionary transitions between seed dormancy and nondormancy. Biogeographic patterns reveal a low proportion of dormancy in tropical rainforest regions and arctic regions and a high proportion of dormancy in remaining tropical, subtropical, and temperate regions for all species and woody species. Herbaceous plants show a greater proportion of dormancy in most global regions except arctic regions. Seasonal environments have a consistent positive influence on the dormancy pattern for both life-forms, but precipitation and temperature were important driving factors for woody and herbaceous plants, respectively. Seed dormancy was the dominating state during the evolutionary history of seed plants, and dormancy transitions had a significant relationship with paleotemperatures. Dormancy and nondormancy transitions in response to fluctuating environments during long-term evolutionary history may have played important roles in the diversification of seed plants. Our results add to the current knowledge about seed dormancy from macro-adaptive perspectives and the potential adaptive mechanisms of seed plants.

A Seed Mucilage-Degrading Fungus From the Rhizosphere Strengthens the Plant-Soil-Microbe Continuum and Potentially Regulates Root Nutrients of a Cold Desert Shrub.

Seed mucilage plays important roles in the adaptation of desert plants to the stressful environment. Artemisia sphaerocephala is an important pioneer plant in the Central Asian cold desert, and it produces a large quantity of seed mucilage. Seed mucilage of A. sphaerocephala can be degraded by soil microbes, but it is unknown which microorganisms can degrade mucilage or how the mucilage-degrading microorganisms affect rhizosphere microbial communities or root nutrients. Here, mucilage-degrading microorganisms were isolated from the rhizosphere of A. sphaerocephala, were screened by incubation with mucilage stained with Congo red, and were identified by sequencing and phylogenetic analyses. Fungal-bacterial networks based on high-throughput sequencing of rhizosphere microbes were constructed to explore the seasonal dynamic of interactions between a mucilage-degrading microorganism and its closely related microorganisms. The structural equation model was used to analyze effects of the mucilage-degrading microorganism, rhizosphere fungal-bacterial communities, and soil physicochemical properties on root C and N. The fungus Phanerochaete chrysosporium was identified as a mucilage-degrading microorganism. Relative abundance of the mucilage-degrading fungus (MDF) was highest in May. Subnetworks showed that the abundance of fungi and bacteria closely related to the MDF also were highest in May. Interactions between the MDF and related fungi and bacteria were positive, which might enhance mucilage degradation. In addition, the MDF might regulate root C and N by affecting rhizosphere microbial community structure. Our results suggest that MDF from the rhizosphere strengthens the plant-soil-microbe continuum, thereby potentially regulating microbial interactions and root nutrients of A. sphaerocephala.[Formula: see text] Copyright © 2021 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license.

Seed development in Paeonia ostii (Paeoniaceae), with particular reference to embryogeny.

BACKGROUND: Seeds of Paeonia ostii have been proposed as a source of raw material for the production of edible oil; however, lack of information about the developmental biology of the seeds hampers our ability to use them. Our aim was to investigate development of the seed coat, endosperm and embryo of P. ostii in relation to timing of accumulation of nutrient reserves from pollination to seed maturity. Ovules and developing seeds of P. ostii were collected at various stages of development from zygote to maturity. Seed fresh mass, dry mass, germination, moisture, soluble sugars, starch, protein and oil content were determined. Ontogeny of seeds including embryo, endosperm and seed coat were analyzed histologically. RESULTS: The ovule of P. ostii is anatropous, crassinucellate and bitegmic. The zygote begins to divide at about 5 days after pollination (DAP), and the division is not accompanied by cell wall formation. By 25 DAP, the proembryo begins to cellularize. Thereafter, several embryo primordia appear at the surface of the cellularized proembryo, but only one matures. Endosperm development follows the typical nuclear type. The seed coat is derived from the outer integument. During seed development, soluble sugars, starch and crude fat content increased and then decreased, with maximum contents at 60, 80 and 100 DAP, respectively. Protein content was relatively low compared with soluble sugars and crude fat, but it increased throughout seed development. CONCLUSIONS: During seed development in P. ostii, the seed coat acts as a temporary storage tissue. Embryo development of P. ostii can be divided into two stages: a coenocytic proembryo from zygote (n + n) that degenerates and a somatic embryo from peripheral cells of the proembryo (2n → 2n). This pattern of embryogeny differs from that of all other angiosperms, but it is similar to that of gymnosperms.

The pleurogram, an under-investigated functional trait in seeds.

BACKGROUND: A structure called the pleurogram makes up a large part of the seed coat of some species in subfamilies Caesalpinioideae and Mimosoideae of Fabaceae, but little is known about its function. It has been hypothesized that this structure acts as a hygroscopic valve during the maturation drying of seeds. However, a new hypothesis has recently emerged that proposes a distinct function for the pleurogram. SCOPE: Here, we provide an overview of the structure and function of the pleurogram, which is diverse and complex. This large structure can be dislodged, thereby creating a pathway for water entry into water-impermeable seeds. However, the pleurogram is non-functional as a pathway of water into the seed of some species. Thus, the evolutionary history of species with a pleurogram may be related to a loss/gain in its function. A complete model for the function of the pleurogram is proposed. CONCLUSIONS: The pleurogram may act on several stages of the seed, from maturation to germination. As a hygroscopic valve, it regulates dehydration of the seed during maturation. As a pathway for water entry into the seed, the pleurogram acts as a water gap in seeds with physical dormancy, thereby regulating dormancy break/germination. The occurrence of a pleurogram in several genera of legumes and Cucurbitaceae is confirmed. Single or multiple pleurograms can serve as (the) point(s) of water entry into seeds that do not otherwise have a hilar water gap.

Seed germination of gynodioecious species: theoretical considerations and a comparison of females and hermaphrodites.

MAIN CONCLUSION: Better seed germination of females than of hermaphrodites is not a major contributor to the greater geometric lifetime fitness that females require to be maintained in a gynodioecious population. Gynodioecy is a sexually dimorphic breeding system in which females (F, male sterile) and hermaphrodites (H) coexist in the same population. For plants with nuclear (biparental) inheritance of male sterility, theory predicts that except when the product of selfing rate (s) and inbreeding depression (δ) in H is high (sδ > 0.50), F must compensate (female advantage) for the loss of gene transmission via pollen production by producing more or higher-quality offspring than H to be maintained in the population. For species with cytoplasmic (maternal) inheritance of male sterility, the female requires only a small compensation in seed production or some other offspring fitness trait to persist. Reallocation to seeds of resources saved by loss of pollen production is expected to increase the quantity (number) and/or quality (mass, germinability) of seeds produced by F, thus compensating for the lack of pollen production. The primary aim of our study was to compare seed germination of F and H via a literature review. Based on theoretical considerations, we hypothesized that seeds of F should germinate better or equally as well as those of H. We found that of 235 case studies for 47 species Fgerm > Hgerm in 48.1%, Fgerm = Hgerm in 38.3% and Fgerm < Hgerm in 13.6%. Our results are very similar to those of a previously published meta-analysis that included germination of F and H for 12 species. For 162 cases on seed size, F > H in 29.0%, F = H in 63.6% and F < H in 7.4%. Since [(Fgerm > Hgerm) < (Fgerm ≤ Hgerm)] and [(Fseedsize > Hseedsize) < (Fseedsize ≤ Hseedsize)], these results suggest that seed quality is not a major fitness component of female advantage.

Dynamics of the diaspore and germination stages of the life history of an annual diaspore-trimorphic species in a temperate salt desert.

MAIN CONCLUSION: Individuals of the annual halophyte Atriplex centralasiatica produce three kinds of diaspores that differ in dispersal, dormancy/germination response and type of seed bank formed, which likely is a bet-hedging strategy in the rainfall-unpredictable environment on the semi-arid, saline Ordos Plateau in Inner Mongolia, China. Seasonal fluctuations in environmental conditions provide germination cues for the establishment of seedlings at the right time and place to ensure plant survival and population regeneration. Diaspore heteromorphism is a phenomenon in which diaspores with stark qualitative differences in morphology and ecology are produced by the same maternal plant. Germination responses and dispersal times of the annual halophyte Atriplex centralasiatica were examined to determine the role of diaspore heteromorphism in its adaptation to salt desert conditions. A. centralasiatica is a tumbleweed that produces three types of diaspores that differ in morphology and ecophysiology. The relative potential dispersal ability and intensity of dormancy of the three diaspore types was type A (fan-shaped diaspores with yellow fruits) < type B (fan-shaped diaspores with black fruits) < type C (globular diaspores with black fruits). In the field, type A retained high germinability, but all of them were depleted from the (transient) soil seedbank in the first growing season. Types B and C cycled between dormancy and nondormancy, and 0 and > 90.0% remained in the soil seedbank 2 years after dispersal, respectively. The dormancy, dispersal and salt tolerance of type B diaspores were intermediate between those of A and C. Type A exhibited low dispersal-nondormancy, type B exhibited intermediate dispersal-intermediate dormancy and type C exhibited high dispersal ability-high dormancy. In the unpredictable salt desert habitat, the functional differences in germination and dispersal of the three diaspores act as a bet-hedging mechanism and ensure population establishment in different years by spreading germination over time and space.

Wing loading, not terminal velocity, is the best parameter to predict capacity of diaspores for secondary wind dispersal.

Lift-off velocity may be the most useful surrogate to measure the secondary dispersal capacity of diaspores. However, the most important diaspore attribute determining diaspore lift-off velocity is unclear. Furthermore, it is not known whether terminal velocity used to characterize the primary dispersal capacity of diaspores can also be used to predict their secondary wind dispersal capacity. Here, we investigate how diaspore attributes are related to lift-off velocity. Thirty-six species with diaspores differing in mass, shape index, projected area, wing loading, and terminal velocity were used in a wind tunnel to determine the relationship between diaspore attributes and lift-off velocity. We found that diaspore attributes largely explained the variation in lift-off velocity, and wing loading, not terminal velocity, was the best parameter for predicting lift-off velocity of diaspores during secondary wind dispersal. The relative importance of diaspore attributes in determining lift-off velocity was modified by both upwind and downwind slope directions and type of diaspore appendage. These findings allow us to predict diaspore dispersal behaviors using readily available diaspore functional attributes, and they indicate that wing loading is the best proxy for estimating the capacity for secondary dispersal by wind.

A review of the seed biology of Paeonia species (Paeoniaceae), with particular reference to dormancy and germination.

MAIN CONCLUSION: Most Paeonia species have epicotyl dormancy. Germination of peony seeds requires warm stratification for embryo growth and radicle protrusion followed by cold stratification for epicotyl growth. The genus Paeonia (Paeoniaceae) includes many popular ornamentals, has colorful flowers and contains several Chinese medicinal species. The germination protocol for seeds of Paeonia species is complex and impedes the breeding of new cultivars and contributes to the rarity and high cost of the plants. Although numerous reports on seed dormancy/germination in peonies are scattered throughout the literature, most of them are in Chinese. The primary aims of this paper are to provide a general overview of the available information on seed dormancy/germination in peonies and to make some suggestions regarding propagation for the peony industry and breeders. Most Paeonia species have epicotyl dormancy. The embryo is differentiated into organs, but it is underdeveloped (small) and must grow inside the seed before the radicle can emerge. Germination of peony seeds requires warm stratification for embryo growth and radicle protrusion followed by cold stratification for epicotyl growth. In addition, the epicotyl is sensitive to cold stratification only after the root has grown to a certain length. GA3 treatment enhances embryo growth and subsequent germination percentages. Further investigations on the physiology, genetics and proteomics would contribute to a better understanding of seed dormancy in Paeonia.

Seed mucilage interacts with soil microbial community and physiochemical processes to affect seedling emergence on desert sand dunes.

Seedling emergence is a critical stage in the establishment of desert plants. Soil microbes participate in plant growth and development, but information is lacking with regard to the role of microbes on seedling emergence. We applied the biocides (captan and streptomycin) to assess how seed mucilage interacts with soil microbial community and physiochemical processes to affect seedling emergence of Artemisia sphaerocephala on the desert sand dune. Fungal and bacterial community composition and diversity and fungal-bacterial interactions were changed by both captan and streptomycin. Mucilage increased soil enzyme activities and fungal-bacterial interactions. Highest seedling emergence occurred under streptomycin and mucilage treatment. Members of the phyla Firmicutes and Glomeromycota were the keystone species that improved A. sphaerocephala seedling emergence, by increasing resistance of young seedlings to drought and pathogen. Seed mucilage directly improved seedling emergence and indirectly interacted with the soil microbial community through strengthening fungal-bacterial interactions and providing favourable environment for soil enzymes to affect seedling emergence. Our study provides a comprehensive understanding of the regulatory mechanisms by which soil microbial community and seed mucilage interactively promote successful establishment of populations of desert plants on the barren and stressful sand dune.

Morphophysiological dormancy in the basal angiosperm order Nymphaeales.

Background and Aims: Substantial evidence supports the hypothesis that morphophysiological dormancy (MPD) is the basal kind of seed dormancy in the angiosperms. However, only physiological dormancy (PD) is reported in seeds of the ANA-grade genus Nymphaea. The primary aim of this study was to determine the kind of dormancy in seeds of six species of Nymphaea from the wet-dry tropics of Australia. Methods: The effects of temperature, light and germination stimulants on germination were tested on multiple collections of seeds of N. immutabilis, N. lukei, N. macrosperma, N. ondinea, N. pubescens and N. violacea. Embryo growth prior to hypocotyl emergence was monitored. Key Results: Germination was generally <10 % after 28 d in control treatments. Germination percentage was highest at 30 or 35 °C for seeds exposed to light and treated with ethylene or in anoxic conditions in sealed vials of water, and it differed significantly between collections of N. lukei, N. macrosperma and N. violacea. Seeds of N. pubescens did not germinate under any of the conditions. Embryo growth (8-37 % in length) occurred before hypocotyl emergence (germination) in seeds of the five species that germinated. Conclusions: Fresh seeds were dormant, and the amount of pregermination embryo growth in seeds of N. lukei and N. immutabilis was relatively small, while in seeds of N. macrosperma, N. ondinea and N. violacea it was relatively large. Thus, seeds of N. lukei and N. immutabilis had PD and those of N. macrosperma, N. ondinea and N. violacea had MPD. Overall, we found that seeds in the most phylogenetically derived clades within Nymphaea have MPD, suggesting that PD is the most likely basal trait within the Nymphaeales. This study also highlights the broad range of dormancy types and germination strategies in the ANA-grade angiosperms.

A function for the pleurogram in physically dormant seeds.

BACKGROUND AND AIMS: Different structures have been shown to act as a water gap in seeds with physical dormancy (PY), and in Fabaceae they are commonly located in the hilar region. However, the function of the pleurogram, a structure in the extra-hilar region that is common in legume seeds, remains unknown. Our aims were to review the literature for occurrence of the pleurogram in Fabaceae, determine if the pleurogram can open, and compare the functional morpho-anatomy of water gaps in seeds of 11 Senna species. METHODS: Imbibition tests showed that all 11 species had PY. Structural features of the hilar and extra-hilar regions of the seeds were investigated using light and scanning electron microscopy, and dye-tracking was performed to trace the pathways of water through the seed coat. KEY RESULTS: A pleurogram has been reported for 37 legume genera. Water gaps differed among Senna species, with lens, hilum, micropyle and pleurogram taking up water after PY was broken. In Senna alata seeds, only the pleurogram acted as a water gap, whereas in S. reniformis and S. silvestris water entered the seed through both the pleurogram and the hilar region. In the pleurogram of S. alata and S. reniformis, the palisade layer moved outward, exposing the hourglass cells, whereas in S. silvestris the palisade layer was broken. CONCLUSIONS: The pleurogram acts as a water gap in some of the 11 Senna species examined, but it is non-functional in others. Opening the pleurogram occurs due to the formation of a linear slit in the palisade layer. The pleurogram is of functional significance by creating a wide opening, whereby water can reach the embryo and start germination. This is the first report of the pleurogram functioning as a water gap. Because this structure is shared by at least 37 genera, it also may be a water gap in many other legume species.

Effects of climate warming and prolonged snow cover on phenology of the early life history stages of four alpine herbs on the southeastern Tibetan Plateau.

PREMISE OF THE STUDY: Much research has focused on plant responses to ongoing climate change, but there is relatively little information about how climate change will affect the early plant life history stages. Understanding how global warming and changes in winter snow pattern will affect seed germination and seedling establishment is crucial for predicting future alpine population and vegetation dynamics. METHODS: In a 2-year study, we tested how warming and alteration in the snowmelt regime, both in isolation and combination, influence seedling emergence phenology, first-year growth, biomass allocation, and survival of four native alpine perennial herbs on the southeastern Tibetan Plateau. KEY RESULTS: Warming promoted seedling emergence phenology of all four species and biomass per plant of two species but reduced seedling survival of three species. Prolonged snow cover partly mediated the affects of warming on Primula alpicola (survival and biomass), Pedicularis fletcheri (phenology, biomass, and root:shoot ratio) and Meconopsis integrifolia (survival). For the narrowly distributed species M. racemosa, seedling growth was additively decreased by warming and prolonged snow cover. CONCLUSIONS: Both warming and alteration of the snow cover regime can influence plant recruitment by affecting seedling phenology, growth, and survival, and the effects are largely species-specific. Thus, climate change is likely to affect population dynamics and community structure of the alpine ecosystem. This is the first experimental demonstration of the phenological advancement of seedling emergence in the field by simulated climate warming.

Global biogeography of seed dormancy is determined by seasonality and seed size: a case study in the legumes.

Seed dormancy is expected to provide ecological advantages by adjusting germination to the favorable growth period. However, many species produce nondormant seeds, particularly in wet tropical forests, a biogeographic pattern that is not well accounted for in current models. We hypothesized that the global distribution of dormant seeds derives from their adaptive value in predictably fluctuating (i.e. seasonal) environments. However, the advantage conferred by dormancy might ultimately depend on other seed attributes, particularly size. This general model was tested within a phylogenetically informed framework using a data set comprising > 216 000 world-wide observations of Fabaceae, spanning three orders of magnitude in seed size and including both dormant and nondormant seeds. Our results confirmed our hypothesis: nondormant seeds can only evolve in climates with long growing seasons and/or in lineages that produce larger seeds. Conversely, dormancy should be evolutionarily stable in temperate lineages with small seeds. When the favorable season is fleeting, seed dormancy is the only adaptive strategy. Based on these results, we predict that, within a given lineage, taxa producing larger, nondormant seeds will necessarily predominate in aseasonal environments, while plants bearing small, dormant seeds will be dominant under short growing seasons.

Phylogeny strongly drives seed dormancy and quality in a climatically buffered hotspot for plant endemism.

BACKGROUND AND AIMS: Models of costs and benefits of dormancy (D) predict that the evolutionarily stable strategy in long-term stable environments is for non-dormancy (ND), but this prediction remains to be tested empirically. We reviewed seed traits of species in the climatically buffered, geologically stable and nutrient-impoverished campo rupestre grasslands in Brazil to test the hypothesis that ND is favoured over D. We examined the relative importance of life-history traits and phylogeny in driving the evolution of D and assessed seed viability at the community level. METHODS: Germination and viability data were retrieved from 67 publications and ND/D was determined for 168 species in 25 angiosperm families. We also obtained the percentage of embryoless, viable and dormant seeds for 74 species. Frequencies of species with dormant and non-dormant seeds were compared with global databases of dormancy distribution. KEY RESULTS: The majority of campo rupestre taxa (62·5 %) had non-dormant seeds, and the ND/D ratio was the highest for any vegetation type on Earth. Dormancy was unrelated to other species life-history traits, suggesting that contemporary factors are poor predictors of D. We found a significant phylogenetic structure in the dormancy categorical trait. Dormancy diversity was highly skewed towards the root of the phylogenetic tree and there was a strong phylogenetic signal in the data, suggesting a major role of phylogeny in determining the evolution of D versus ND and seed viability. Quantitative analysis of the data revealed that at least half of the seeds produced by 46 % of the surveyed populations were embryoless and/or otherwise non-viable. CONCLUSIONS: Our results support the view that long-term climatic and geological stability favour ND. Seed viability data show that campo rupestre species have a markedly low investment in regeneration from seeds, highlighting the need for specific in situ and ex situ conservation strategies to avoid loss of biodiversity.

Effects of germination time on seed morph ratio in a seed-dimorphic species and possible ecological significance.

BACKGROUND AND AIMS: Diaspores of heteromorphic species may germinate at different times due to distinct dormancy-breaking and germination requirements, and this difference can influence life history traits. The primary aim of this study was to determine the effect of germination time of the two seed morphs of Suaeda corniculata subsp. mongolica on life history traits of the offspring. METHODS: Germinated brown and black seeds were sown on the 20th of each month from April to September in a simulated but near-natural habitat of the species. Phenological and vegetative traits of the maternal plants, and number, size and germination percentage of the offspring were determined. KEY RESULTS: Germinated seeds sown late in the year produced smaller plants that had a higher proportion of non-dormant brown than dormant black seeds, and these brown seeds were larger than those produced by germinated seeds sown early in the year. The length of the seedling stage for brown seeds was shorter than that for black seeds, and the root/shoot ratio and reproductive allocation of plants from brown seeds were more variable than they were for plants from black seeds. Late-germinating brown seeds produced larger plants than late-germinating black seeds. CONCLUSIONS: Altering the proportion of the two seed types in response to germination timing can help alleviate the adverse effects of delayed germination. The flexible strategy of a species, such as S. corniculata, that produces different proportions of dimorphic seeds in response to variation in germination timing may favour the maintenance and regeneration of the population in its unpredictable environment.