Polyploidy affects the seed,dormancy and seedling characteristics of a perennial grass,conferring an advantage in stressful climates

• Polyploidy (the state of having more than two genome copies) is widely distributed in flowering plants and can vary within species, with polyploid races often associated with broad ecological tolerances. Polyploidy may influence within‐species variation in seed development, germination and establishment. We hypothesized that interactions between polyploidy and the seed developmental environment would affect subsequent dormancy, germination and early growth traits, particularly in stressful environments.
• Using seeds developed in a common garden under ambient and warmed conditions, we conducted germination trials under drought and temperature stress, and monitored the subsequent growth of seedlings. The study species, Themeda triandra, is a widespread, keystone, Australian native grass and a known polyploid complex.
• Tetraploid plants produced heavier, more viable seeds than diploids. Tetraploids were significantly more dormant than diploids, regardless of seed developmental environment. Non‐dormant tetraploids were more sensitive to germination stress compared to non‐dormant diploids. Finally, tetraploid seedlings were larger and grew faster than diploids, usually when maternal plants were exposed to developmental temperatures atypical to the source environment.
• Seed and seedling traits suggest tetraploids are generally better adapted to stressful environments than diploids. Because tetraploid seeds of T. triandra are more dormant they are less likely to germinate under stress, and when they do germinate, seedling growth is rapid and independent of seed developmental environment. These novel results demonstrate that polyploidy, sometimes in interaction with developmental environment and possibly also asexuality, can have within‐species variation in seed and seedling traits that increase fitness in stressful environments.

     

Parental effects in Lychnis flos-cuculi. I: Seed size,germination and seedling performance in a controlled environment

Selection responses in natural plant populations depend on how the phenotypic variation of traits is composed. The contributions of nuclear genetic, maternal, paternal, environmental and inbreeding effects to variation in time to germination, germination percentage, and seed- and seedling size were studied in a population of Lychnis flos-cuculi. It was found that: (1) Maternal effects predominated in the determination of progeny seed size and germination characteristics; (2) Maternal environment during seed development was less important than maternal genotype; (3) Small but significant variation within maternal families could be observed among individuals sired by different fathers; (4) Additive genetic variance was significant for seedling size 4 weeks after germination. In conclusion, selection shortly after emergence will mainly favour particular maternal genotypes, while selection later in the life cycle may act upon zygotic genotypes. Inbreeding depression was significant, especially for vegetative growth. Consistent differences were found among maternal genotypes in the degree of variation in the time to germination, suggesting that selection could operate to favour polymorphic or uniform germination behaviour.     

Parental effects in Lychnis flos-cuculi. II: Selection on time of emergence and seedling performance in the field

Selection on the timing of seedling emergence was investigated in an experimental population of Lychnis flos-cuculi, a perennial hay-meadow species. Seeds obtained from a full diallel cross of 8 genotypes from a field population were sown along an environment gradient that included the parental site. Significant directional selection for early emergence was found and the intensity of selection varied among sites. Emergence time varied significantly among progeny families of different maternal and paternal genotypes. These differences could be attributed to parental effects whereas narrow-sense heritabilities were close to zero. Survivorship until autumn differed among progeny of paternal families. Survivorship of maternal progeny varied among sites. Whereas differences in survival and plant size among individuals from different emergence cohorts persisted over the winter, the significance of these differences among progeny from different parental genotypes disappeared. It is suggested that a response to selection on emergence time might be low since (1) the narrow sense heritability was low, (2) parental genotypes differed in their effect on offspring emergence time when used as female parent or as pollen donor and (3) there was a family x site interaction for survival. Families with relatively early emerging seedlings also had a significantly higher seed weight, emergence percentage, and plant weight although the strength of these among-family correlations varied among sites. It is therefore not likely that simultaneous selection on emergence time and either of these traits would retard a response to selection on emergence time.     

Seed germination and seedling emergence of Euryodendron excelsum H. T. Chang: implications for species conservation and restoration

Euryodendron excelsum H. T. Chang is a critically endangered plant from the family Theaceae that is endemic to China. It is now present in only one remnant population in a very narrow range and with a highly isolated and fragmented distribution pattern. The species is distributed close to the local villages and faces extinction because of the high level of anthropogenic disturbance. Thus, conservation and restoration of this species is urgent. The species reproduces by seed, but its germination requirements and seedling emergence are rarely understood. In this study, the germination requirements, desiccation tolerance and seedling emergence of the species were studied under controlled laboratory conditions. The results indicated that seeds of E. excelsum were non‐dormant. Optimal temperatures for germination were 15°C to 25°C; the germination percentage decreased and the mean germination time increased at high temperature. Seed germination was inhibited in the dark, suggesting that the seeds were positive photoblastic. Water stress also significantly inhibited germination percentage; no seeds germinated at 15% polyethylene glycol 6000. The fresh seeds had moderate moisture content of 28.6% and showed strong tolerance of dehydration. Thus, the seeds of E. excelsum tended to be orthodox. Seeds on the soil surface had the highest emergence percentage, which declined with depth of burial. Soil types significantly affected seedling emergence; seeds had higher emergence percentage in sandy soil than in mixed soil and clayed soil. Our study demonstrated that ex situ conservation and reintroduction of E. excelsum using seedling propagation from seeds is feasible.     

GYNODIOECY IN SILENE VULGARIS (CARYOPHYLLACEAE): PROGENY SUCCESS,EXPERIMENTAL DESIGN,AND MATERNAL EFFECTS

This study documents the comparative success of seeds and seedlings of the perennial gynodioecious-gynomonoecious weed, Silene vulgaris, in the greenhouse. The importance of experimental design is stressed by comparing two different statistical analyses of the data. Seeds were obtained from artificial pollinations in the field: self-fertilization of hermaphrodites, and cross-fertilizations of both hermaphrodites and females. One-way analysis of variance using progeny type (selfed hermaphrodites, outcrossed hermaphrodites, and outcrossed females) as the treatment effect for each seed and seedling variable showed statistically discernable differences among progeny from different cross types. The significance of this type of ANOVA resulted from a reduced error term and positively-biased F ratios. A factorial design showed no significant differences due to type of progeny in seed mass, days to germination, leaf number, area, or biomass at six weeks of age. There were, however, significant differences among seeds attributable to maternal parent for all seed and seedling variables. A higher proportion of seeds from outcrossed individuals germinated compared to that from self-fertilization. When the data were analyzed to include and partition all sources of variation, differences among offspring appeared during germination, rather than during later development. Seed mass, cross type, and sex of maternal parent all significantly affected the likelihood of germination; however, they had decreasing predictive power, respectively. Inbreeding depression in Silene vulgaris may help maintain gynodioecy; however, the pleiotropic effects of both nuclear and cytoplasmic genes for sex expression also may affect fitness and the maintenance of females.     

Recruitment in a Mediterranean annual plant community: seed bank, emergence, litter, and intra- and inter-specific interactions

A small‐scale field study was conducted to explain seedling emergence and recruitment of reproductive individuals in a four‐year‐old Mediterranean annual plant community. The analyzed levels were populations, functional types, and total number of individuals in the community. We hypothesized that the number of germinable seeds positively affects the number of emerged seedlings, which positively affects the number of reproductive individuals. We hypothesized as well that litter mass, biomass of established adults, and number of non‐conspecific emerged seedlings negatively affect the numbers of emerged seedlings and reproductive individuals. The results supported our multivariate causal explanation of plant recruitment, although concordances with the initial hypotheses were more frequent at the community level. The effect of the number of germinable seeds on the number of emerged seedlings was found to be robust only at the community level. At the population level, this relationship did not have a consistent tendency and depended on species identity and range of seed density experienced. Litter had negative effects on total number of individuals in the community, and usually non‐significant effects at the population level. Litter effects were found to be inversely related to seed mass, and sharper cotyledons did not improve seedling emergence likely by favoring litter mat penetration. Interactions among plant species appeared not to have any effect on seedling emergence and survival, and had positive effects on certain populations. Biomass of established adult plants exerted no influence on seedling emergence and survival, probably due to their low abundance in this community during the germination period. Results permitted the inference of the main stages in the recruitment process and causal factors. They provided evidence for the greater importance of germination and emergence in comparison with seedling survival to insure reproductive recruitment in this community.     

THE EFFECT OF SEED SIZE AND MATERNAL SOURCE ON INDIVIDUAL SIZE IN A POPULATION OF LUDWIGIA LEPTOCARPA (ONAGRACEAE)

Seed size is normally distributed for many annual species, while mature plant size is frequently positively skewed. A study was conducted to determine the influence of seed size and the role of genetic differences in determining relative seedling size for Ludwigia leptocarpa. Seed size had a significant effect on percentage germination and time of seed germination but no effect on dry weight or leaf area of seedlings. Seed size and spacing had a significant effect on seedling dry weight for plants grown under competition, while relative day of emergence had no effect. Familial (genetic) differences were found in average seed weight between maternal plants, but not in average number of days to germination, average weight of seeds which germinated, or shoot dry weight. It is concluded that neither seed size alone nor genetic differences between plants are directly responsible for the development of size hierarchies in Ludwigia leptocarpa populations. Large seed size does convey an advantage in growth when plants from seeds of differing initial size interact.     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

EXPERIMENTAL STUDIES OF THE EVOLUTIONARY SIGNIFICANCE OF SEXUAL REPRODUCTION. III. MATERNAL AND PATERNAL EFFECTS DURING SEEDLING ESTABLISHMENT

To determine whether genetic differences in fitness components exist among seeds and seedlings in a natural population, weighed propagules of six parents of Anthoxanthum odoratum from a reciprocal diallel cross were planted into the parental source population, a mown field. Seed families of maternal genotypes differed in germination success, while paternal families showed no detectable differences. Differential germination success could not be attributed to propagule weight. Seed families ranked differently in germination percentage in different blocks. No survivorship differences among parental seed families could be detected. There were significant cross × block × germination and cross × block × survivorship interactions; different crosses performed better or worse in different blocks. In some cases, crosses sired by different fathers within a maternal seed family differed in germination or survivorship, suggesting that natural selection may be capable of discriminating among juvenile genotypes within a maternal family despite the presence of large overall maternal effects. These results indicate that seedling establishment may differ according to genotype and that microsite heterogeneity may maintain genetic variation in juvenile traits in natural plant populations.     

Polyandry restores female fertility and paternal effects diminished by inbreeding in Hippodamia convergens

1. The fertility restoration hypothesis posits that polyandry can evolve when female fertility is reduced by matings with related males, but restored by matings with unrelated ones. 2. Using a promiscuous ladybird, this hypothesis was tested by mating mature, virgin females twice with sib and non‐sib males in all four permutations and observing female fertility. The development of progeny from the first and 10th clutches was also followed to test for differences in paternal effects. 3. Mating treatment did not affect fecundity, but egg viability was reduced by sib matings, and restored by non‐sib matings, regardless of mating sequence. In addition, negative paternal effects of sib matings on progeny (lower survival) were compensated by non‐sib matings. 4. The survival of offspring in first clutches was sensitive to the paternal mating sequence and was higher if a sibling male was preceded by a non‐sib male, compared with the reverse, consistent with a time lag in response to male epigenetic signals. Offspring survival did not differ between these two treatments in the 10th clutch, suggesting a blending of paternal effects over time. 5. The results are indicative of interactions between the paternal effects contributed by different males, and between paternal and maternal effects, the latter causing faster development in later clutches. 6. Thus, the reproductive benefits of polyandry for H. convergens females are potentially both genetic and epigenetic, as both egg fertility and beneficial paternal effects were diminished by sib matings, but restored by matings with unrelated males.     

Seed germination and seedling growth of two Pseudobombax species (Malvaceae) with contrasting habitats from Brazilian Cerrado

Pseudobombax tomentosum and P. longiflorum are common trees in the Cerrado region, but the former species is more common in forest edges while the later is present in open cerrado areas. This work aimed to investigate differences in seed germination and seedling growth in these species, from seed collected from Cerrado areas in Central Brazil. For this, a seed germination experiment was designed and included four replicates with 25 seeds per species; seeds were randomly distributed in the germination chamber. To evaluate initial seedling growth, seedlings height was measured up to 67 days after seedling emergence; besides, some of these seedlings were grown for biomass evaluation during nine months. Results showed that seeds of the two species had the same germinability (near 100%) and mean germination time (ca. 12 days). However, P. longiflorum showed a more spread seed germination through time, with higher values of coefficient of variation in germination time and uncertainty index; and lower values of synchronization than P. tomentosum. The two species showed basically the same growth pattern, but lower values for height of apical meristem, diameter of underground structures (mostly roots), dry mass of shoots, underground structure and total mass of seedlings in P. tomentosum were obtained, compared to P. longiflorum. Both species allocated more dry mass to underground structures in detriment of shoot. This probably allows resprouting behavior which prevents hydric stress and detrimental fire action typical of the open Cerrado areas.     

Do pollen donors with fastest-growing pollen tubes sire the best offspring in an anemophilous tree, Betula pendula (Betulaceae)?

The relationship between pollen and progeny performance has been a subject of many studies but the evidence for pollen-tube growth rate as an indicator of progeny fitness is equivocal. We used an anemophilous tree, Betula pendula, to examine the relationship between pollen-tube growth rate and seed and seedling performance. We crossed nine maternal plants with pollen from six pollen donors in a clonal B. pendula seed orchard, measured the pollen-tube growth rates for every cross, and analyzed the performance of the resulting seeds and seedlings. The only significant positive correlation was found between pollen-tube growth rate and seed mass when we controlled for seed number per inflorescence. Using seed mass as a covariate, we found that only maternal parent had a significant effect on the number of seeds per inflorescence, the percentage of germinable and embryonic seeds, and early seedling growth. Both maternal and paternal parents had significant effects on seedling height after 85 d of growth. These results are in concordance with the general view that maternal effects are usually most apparent in seed characters and during early plant growth. This study does not provide strong evidence for the theory of pollen-tube growth rate as an indicator of progeny quality.     

Experimental determination of seed emergence and carry‐over in the soil seed bank of the herbaceous perennial,Trachymene incisa (Apiaceae)

Abstract Many populations of herbaceous perennial plants contain seeds stored in a soil seed bank. The contribution of seeds to population persistence is an important parameter in population models but germination rates of known‐age seeds are difficult to obtain because individual seeds cannot easily be followed. Although Trachymene incisa Rudge plants produce copious seeds that are dispersed into the soil, the existence of a seed bank has not been confirmed. To quantify the potential for a seed bank fresh seeds of T. incisa were sown into experimental seed banks in the eucalypt‐dominated Agnes Banks Woodland in western Sydney, NSW. A recent fire provided the opportunity to compare germination in the burnt and unburnt vegetation. Density of seed sowing and time of maturation/dispersal of seeds were manipulated in 75 seed cages. Emergence of seeds after 5 months was significantly higher for the earliest planting date but after 1 year, germination of seeds planted in the later weeks increased, and the final germination for all weeks was 28%. Density of sowing and the recent fire did not affect emergence. A second experiment planted over a broader time span (9 weeks instead of 3 weeks) confirmed the effect of planting date but also found significant spatial variation on a scale of tens of metres. Laboratory germination rates of over 70% confirmed that the seeds were viable and non‐dormant when sown in the field cages. The carry‐over of non‐germinated seed in the soil seed bank is estimated to be about 70% after 2 years, implying that a cohort of seeds would not be depleted through germination alone for up to 40 years. The potential for a long‐lived seed bank in this species is interesting because the plants are also capable of resprouting from their rootstock after fire, giving them characteristics of both resprouters and seeders.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

ENVIRONMENTAL AND GENETIC MATERNAL EFFECTS ON SEED CHARACTERS IN NEMOPHILA MENZIESII

Nuclear genetic, maternal genetic and maternal environmental effects on seed characters were estimated in the California native annual plant Nemophila menziesii using two greenhouse crosses. In one cross, according to a nested mating design, the narrow sense heritability of seed weight was small (3.9%). A subset of full-sib progenies produced in this cross was grown singly and in competition with the introduced grass Bromus diandrus. In a second cross, these plants were used as mothers (dams) and were each mated to the same three sires. Seeds produced by mothers competing with B. diandrus showed a significant reduction in weight, increase in time to germination, and increase in the incidence of dormancy, when compared to seeds from mothers grown singly. Significant sire components were found for time to germination and incidence of dormancy. Maternal genetic variation for seed weight was largely expressed as maternal genotype by maternal environment interaction, and showed no significant maternal genetic main effect. Time to germination and dormant fraction showed a relatively large maternal genetic effect. Evolution of seed characters in N. menziesii is more likely to occur via indirect response to selection among maternal plants than among the seeds themselves. Maternal genotype by maternal environment interaction could potentially contribute to the maintenance of maternal genetic variation in seed weight, but this does not appear likely for dormancy.     

Individual variation in body color in the field cricket Teleogryllus occipitalis (Orthoptera: Gryllidae)

Body color of animals may affect individual fitness through direct effects on various processes, such as predatory avoidance, thermoregulation, UV resistance, and mating behavior. Body color variation of the field cricket Teleogryllus occipitalis (Audinet‐Serville) (Orthoptera: Gryllidae) is often observed. We quantified the individual variation in body color of this species by measuring the luminance of hind leg femur. Thereafter, we examined whether the body color was a heritable trait or not using parent–offspring analysis. In addition, the effect of body color of both parents on body size, sex ratio, and survival rate of progeny was investigated. The results showed that maternal body color was not heritable, whereas paternal body color was correlated with the body color of the progeny. Body size and sex ratio were not correlated with body color of parents. Although paternal body color did not affect the survival rate of the progeny beyond the stage of final instar nymph, progeny survival rate significantly increased with darkness of maternal body.     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.     

Seasonal cycle of seed dormancy controls the recruitment of Butia odorata (ARECACEAE) seedlings in savanna‐like palm tree formations in southern Brazil

Butia odorata (Barb. Rodr.) Noblick is a palm tree that grows in savanna‐like formations in subtropical regions of South America, and whose regeneration is threatened by agricultural management. Its diaspores are dormant after dispersal which takes place during the summer and early autumn. The aim of this study was to investigate seasonal and microhabitat effects on the germination and seedling recruitment of this palm species. Diaspores were sown in the field, in both open lands and forest patches. During 2 years, we measured seed germination, viability and moisture, seedling emergence and germination response to warm stratification of those seeds that failed to germinate in the field. Germination was concentrated during the summer, when soil temperatures were highest, whilst seedling emergence peaked in the autumn and early winter, when temperature and humidity conditions became less extreme. In open lands, there were two pulses of germination (first and second summer), whilst in forest patches, a single pulse (second summer) was detected. Although overall germination did not differ between microhabitats, the percentage of seedling emergence from seeds that remained buried until the end of the experiment was almost twice as large in the forest patches compared with open areas. The viability of seeds declined over time, particularly in open areas. Laboratory‐induced warm stratification was found to act on seed dormancy release in a cyclic way, being far more effective on seeds retrieved from the field in spring–summer months than in those retrieved in the winter. This cyclic pattern of dormancy in B. odorata seeds results in major seedling recruitment after the summer, under wetter and cooler conditions, thus reducing mortality risk. This process can be enhanced by the presence of surrounding vegetation, which both increases seedling emergence and/or prolongs seed viability.     

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.     

THE EFFECT OF SEED DIMORPHISM ON THE GERMINATION AND SURVIVAL OF SALICORNIA EUROPAEA L. POPULATIONS

Seed dimorphism influenced the germination behavior of Salicornia europaea L., with small seeds being more dormant and less salt tolerant than large seeds. All of the large less dormant seeds of S. europaea germinated prior to May, and all seedlings produced after this time were from small seeds. A persistent seed bank was maintained by the small dormant seeds. Survivorship was relatively constant during the normal germination season, but increasing salinity stress at any time during the spring reduces the chance of seedling survival. Little germination occurred from July through September because of the high surface soil salinities during this period.