Composition of soil seed banks in southern California coastal sage scrub and adjacent exotic grassland

Soil seed banks are important to many plant communities and are recognized as an important component of management plans. Understanding seed bank composition and density is especially important when communities have been invaded by exotic species and must be managed to promote desirable species. We examined germinable soil seed banks in southern California coastal sage scrub (CSS) that is heavily invaded by exotic grasses and in adjacent exotic grassland. Soils from both communities had similar seed banks, dominated by high densities of exotic grass and forb species. Up to 4,000 exotic grass seeds and at least 400 exotic forb seeds/m² were found in most soils, regardless of aboveground vegetation type. Native forbs averaged 400 seeds/m² in grass-dominated areas and about 800 in shrub-dominated soils. Shrub seed density was <1 and <10 seeds/m² in grass- and shrub-dominated areas, respectively, indicating that the shrub seed bank is not persistent compared to annuals. We also compared pre- and post-burn soil seed banks from one location that burned in October 2003. Late-season burning in both grass- and CSS-dominated areas disproportionately reduced exotic grass seed densities relative to native seed densities. The similarity of the seed banks in adjacent grass and shrub communities suggests that without intervention, areas currently dominated by CSS may become more similar to grass-dominated areas in terms of aboveground vegetation. In such areas, the first growing season following a wildfire is a window of opportunity for increasing native diversity at a time when density of exotic grass seeds is low. At time of research, Robert D. Cox was graduate student.     

Seed dispersal in the dung of large herbivores: implications for restoration of Renosterveld shrubland old fields

Species-rich, winter-rainfall, microphyllous Renosterveld vegetation in the Western Cape Province of South Africa has largely been transformed for production of wheat and wine. Remaining fragments thus have high conservation value. Abandoned old fields adjacent to natural vegetation fragments could potentially be restored as corridors and habitat for indigenous flora and fauna. We hypothesised that indigenous antelope maintained in a matrix of natural vegetation and abandoned field could play a role in restoration of Renoserveld via seed dispersal.We collected dung of indigenous ungulates in an abandoned field at various distances from natural Renosterveld vegetation, in order to assess the potential of large herbivores to contribute to restoration of plant diversity through seed dispersal. Emerged seedlings from the collected dung represented 29 forb, 13 grass, four sedge, four geophyte and one shrub species. The most abundant emerging seedlings were lawn grass Cynodon dactylon (38%), alien pasture grasses (31%) and indigenous geophyte Romulea rosea (12%). Whereas seeds of annual forbs and grasses were dispersed, only one shrub species was dispersed at very low density. We concluded that large herbivores could retard the rate of recovery of Renosterveld vegetation because viable seeds of herbaceous plants, particularly alien annual grasses and lawn-grasses were more abundant in the dung than the shrub, geophyte or perennial tussock grass species that characterise this vegetation type.     

Community dynamics of evergreen broadleaf forests in southwestern Japan. II. Species composition and density of seeds buried in the soil of a climax evergreen oak forest

The species composition and density of seeds in the soil of a climax evergeen oak forest in the Kasugayama Forest Reserve, Nara, Japan were studied by directly extracting seeds from soil samples and using soil samples in planting boxes as the basis of germination tests. A total of 33 species were identified from all seeds collected in 6 stands: 11 evergreen broadleaf, 15 deciduous broadleaf, 2 coniferous, 2 liana, 1 herb, and 2 grass species. Species producing sap fruits and dry fruits accounted for 60% and 40% of the total number of species, respectively. The species composition of all the seeds was independent of the species composition of the forest vegetation. The mean density of the seeds was 22,134 seeds/m²·5 cm.Eurya japonica, of which seeds were found in all soil samples, was the most abundant species, followed byCryptomeria japonica, Ilex micrococca, andBoehmeria longispica. Pioneer species such asMallotus japonicus, Zanthoxylum ailanthoides, andAralia elata were found in all soil samples in spite of the paucity of adult trees in the forest. Seeds of evergreen oaks were relatively aboundant but no viable seeds were found. ViableE. japonica, I. micrococca, Symplocos prunifolia, andB. longispica seeds were abundant.     

Germination,survival, and growth of grass and forb seedlings: Effects of soil moisture variability

Seed germination and seedling growth, survivorship, and final biomass and their responses to watering interval were studied in two grass and six forb species to assess germination and seedling growth responses to increased soil moisture variability as might occur with future increases in precipitation variability. Seeds were planted in prairie soil and watered at 1, 2, 4, or 7 d intervals (I). Seed germination peaked at I = 4 d whereas leaf growth in grasses and forbs, and final biomass in grasses peaked at I = 7 d, suggesting that growth and biomass were favored at greater soil moisture variability than seed germination. Biomass responses to I were stronger than the germination responses, suggesting that soil moisture variability more strongly influenced post germination growth. Individual species responses to I fell into three groups; those with responses to I for: (1) seed germination and seedling survival, (2) biomass, or (3) both germination and biomass production. These species groups may be more useful than life form (i.e., grass/forb) for understanding seed germination and seedling dynamics in grasslands during periods of soil moisture variability. Seed germination and early growth may assume more importance in grassland plant community dynamics under more variable precipitation patterns.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

Spatial patterns of the germinable soil seed bank of coexisting perennial-grass species in grazed shrublands of the Patagonian Monte

We compared the size and spatial pattern of the germinable soil seed bank (GSB) of the three dominant perennial-grass species in the arid Patagonian Monte of Argentina. These species differ in plant functional traits, seed morphology, and in their preference by grazers. During three consecutive years, we sampled the soil seed bank at 1 m intervals and the intercepted diameter of perennial-grass species and shrub patches along 50-m linear transects at three sites with different grazing pressure. Soil samples were incubated at field capacity during 12 weeks and emerged seedlings counted. The largest-seeded and most xerophytic Stipa speciosa did not form a GSB. The other two grass species with more mesophytic traits and smaller seeds than S. speciosa formed different GSB. Poa ligularis formed a smaller and more heterogeneous GSB than Stipa tenuis. Grazing had not effect on GSB size of P. ligularis but increased the proportion of seeds under dense shrub canopies. In contrast, grazing reduced the size and also increased the proportion of the GSB of S. tenuis under shrub canopies. We conclude that the size and the spatial pattern of the GSB of the dominant perennial-grass species in the arid Patagonian Monte were related not only to seed and plant traits but also to the spatial distribution of grass plants and their exposure to grazers.     

Disturbance regimes as determinants of seed banks in coastal dune vegetation of the southeastern Cape

Soil-stored seed banks of grassland, fynbos and thicket, all growing on calcareous dunes and each subject to different disturbance regimes, were examined. Seed banks were determined from counts of germinants from 50 soil cores from each type. Aboveground estimates of plant species cover in 10 1-m² plots were used in determining vegetation/seed bank similarities. There was no evidence for seed bank densities to be markedly higher in the most frequently disturbed community (grassland -4273 seeds/m²) than the least disturbed community (thicket - 3417 seeds/m²). Highest similarity between seed bank and above-ground vegetation composition in terms of species and growth form/life-span classes was recorded for grassland (CC = 50%). Lowest similarity (CC = 13%) was found in the less frequently disturbed thicket where no seeds of climax trees were recorded in the seed bank. A fynbos community on a north-facing (warm, dry) slope had intermediate-sized seed banks (1683 seeds/m²) with intermediate vegetation/seed bank similarity (CC = 46%). However, on the south-facing slope, which has a large post-fire ephemeral herb component, seed banks were larger (4518 seeds/m²) but less similar to above-ground vegetation (CC = 39%o). Ordination (DCA) of vegetation data from the four communities was different from an ordination of their seed bank data. Fynbos shrub species were absent from seed banks of both grassland and thicket, even though secondary succession proceeds from grassland, through fynbos to thicket. Their seed banks appear less persistent than those of European heath or Californian chaparral shrubs.     

Seed dispersal and vegetation development on a debris avalanche on the Ontake volcano,Central Japan

Seed dispersal and early revegetation processes were studied in an area devastated by a debris avalanche that occurred on volcano Ontake, Central Japan in 1984. The avalanche was initiated by a volcanic earthquake, and a 3.4 x 10⁷ m³ land mass destroyed the vegetation over 700 ha. The revegetation of the first five years was slow in the area of higher elevation (subalpine area). Both the speed of revegetation and the species richness of trapped seeds varied with the elevation. The post-disturbance vegetation on the deposit with surface soil or plant fragments was richer in species than that without any surface soil. Almost all of the seeds caught by sticky traps and most of the species with frequent occurrence in the vegetation after disturbance were wind dispersed. Wind dispersal was especially important on the deposit without surface soil or plant fragments.     

Composition,spatial and temporal variability of the soil seed pool in a Eucalyptus populnea shrub woodland in central New South Wales

Three sites in a Eucalyptus populnea woodland in central New South Wales were sampled for the number and composition of seed in the surface cm of soil. There was a total of 3200 seeds/m² beneath eucalypt canopy, 13800 seeds/m² in grassy inter-tree areas and 4000 seeds/m² in bare inter-tree areas in April, 1975. These numbers were reduced by 23, 73 and 66% respectively during the subsequent 10 months, when there was no addition of seed. This decline was thought to be mainly due to predation, with germination as a secondary factor. Species composition of the seed pool differed between the three sites with seed of some species being only found beneath the eucalypt canopy or in the inter-tree areas. There were no seeds that were not present as plants in the community. Any attempts to restore this woodland to its former grassy nature should take into account the low seed population of many of the formerly common grass species and the high number of shrub seeds (>200/m²). Species distribution of seed was highly heterogeneous making adequate sampling very difficult. Extracting and identifying the seed took 8 to 12 man-h per 100 g sample.     

Seed dynamics during forest succession in Costa Rica

Soil seed banks and current seed inputs each play a role in tropical succession. We compared the abundance and floristic composition of seeds from these two sources at a Costa Rican site by germinating seeds from the soil, measuring seed inputs for 3 yr, and monitoring the earliest colonists in a forest clearing.There were an estimated 6800 viable seeds/m² in the soil of 3.3-yr-old vegetation, 9500 seeds/m² in 11-yr-old vegetation, and 7000 seeds/m² in a 75-yr-old forest. An estimated 10100 seeds/m² fell on the soil surface of the young successional vegetation during 3 yr and 3700 seeds/m² fell during that same time in the forest.Locally produced seeds accounted for about 75% of the seed input to the soil surface early in succession. Seeds dispersed out of young successional vegetation increased the quantity and species richness of the seed input and storage in an adjacent forest. Much of the species richness of the young successional vegetation resulted from seeds dispersed there from other communities by animals.Deforestation stimulated germination of most seeds in the surface soil of the old forest, including seeds of the dominant canopy tree. The recruitment of seedlings from the soil seed bank numerically overwhelmed that from post-disturbance seed rain and sprouts.We evaluated patterns of soil seed storage during succession and predicted the ability of vegetation of differing ages to respond to disturbance. Immediately after disturbance the number of seeds in the soil plummeted due to mortality, low inputs, and germination. As the vegetation regrew, the soil seed bank increased to a peak after 4 to 7 yr, then gradually decreased to its pre-disturbance size. High-frequency pulses of disturbance should result in reduced species richness, dominance by species with long-lived seeds, and fast recovery by seedling recruitment from the soil seed bank.Journal series number 6459 from the Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611, USA.Reprint requests to J. J. E. at Florida.     

Nitrogen fertilization and water supply affect germination and plant establishment of the soil seed bank present in a semi-arid Mediterranean scrubland

Anthropogenic nitrogen (N) inputs in terrestrial ecosystems are higher than those that occur naturally and have been related to global biodiversity loss and altered ecosystem functioning. However, its effects on Mediterranean-type ecosystems, where production is water-limited and N regulated, remain unclear. We conducted a green-house experiment where we evaluated the effects of four simulated scenarios of N pollution (0, 10, 20 and 50 kg N ha⁻¹ year⁻¹) and two differential water supply regimes on the germination (experiment 1) and early plant establishment (experiment 2) of a seed bank from a semi-arid Mediterranean ecosystem of central Spain. Seed bank density was estimated as 62,374 ± 3,279 seeds m⁻². Approximately 99.5% of emerged seeds corresponded to only 14 species of a total of 52, the majority of which were the annual forb Sagina apetala. The responses for N treatments were species-specific, mainly positive or unimodal, with watering treatments having some interactive effects. N and water supply also affected total and specific productivity; the responses found for N treatments were mainly humpback-shaped and an increased water supply had additive effects on community establishment in terms of total plant biomass. This response was linked to forb responsiveness. Contrary to predictions, grass biomass did not change with N supply; however, grass to forb ratio was affected because of changes in the latter. Overall, these experiments suggest a critical load for plant biomass production and conclude that N and water availability and supply can modify germination and plant establishment. This should be taken into account when analysing the effects of global change on the dynamics of plant communities where annuals are dominant or vegetation must establish from seed following a natural or anthropogenic disturbance regime.     

Spatial distribution and population dynamics of Striga hermonthica seeds in naturally infested farm soils

Densities and spatial distribution in soil of seeds of Striga hermonthica were analysed for four naturally infested farm fields in Western Kenya. A revised method for extraction of Striga seeds from soil was used, combining centrifugation with existing techniques based on flotation. Tests showed that 85% of Striga seeds were retrieved from soil samples. In all fields the majority of seeds were found in the plough layer (0 – 20 cm). New seeds entering the soil from the surface after seed shedding created a strong gradient with depth. Downward penetration from the soil surface was larger in sandy soil than in clay soil. In tilled soils no significant vertical density gradient was found within the plough layer. At a fine scale (0.2 m) seed densities showed little horizontal variation, but significant differences in seed densities in the horizontal plane were found at larger scale distances (several m) between locations in all fields. At 125 days after sowing the estimated average number of seeds produced per emerged Striga shoot was 4,827, excluding an approximately similar amount of seeds present in maturing capsules. The estimated average number of seeds produced per mature Striga seed capsule was 1188. Large seasonal fluctuations in the Striga seedbank were measured. An average net increase of 88,825 Striga seeds m^-2 (equivalent to 340%) was calculated from seedbank analyses in 16 sorghum plots. The level of Striga infestation in one field had decreased by 62% from 34,250 seeds m^-2 to 13,125 seeds m^-2 after keeping it fallow for a year. A sharp decline in Striga seed density was found in samples taken at increasing distances from highly infested fields, irrespective of wind direction or slope, suggesting very limited dispersal of Striga seeds by wind or water. Parasite emergence was non-linearly related to initial Striga seed densities in the soil, but this relationship was only observable at the scale of individual plant holes. Seed production was also non-linearly related to numbers of emerged parasites, when measured at plot scale (25 m²), but not at the scale of individual plant holes. In the fields we studied, seed densities below levels of 13,000 Striga seeds m^-2 could be considered to suppress the number of emerging parasites. However, if two or three emerged Striga plants per m^-2 were left to seed, enough seeds would be produced to keep the seedbank in balance.     

Reproductive effort and phenology of seed production of savanna grasses with different growth form and life history

The phenology of seed production in natural savanna grasslands was studied in the grass speciesAristida congesta, Cymbopogon plurinodis, Cynodon dactylon, Digitaria eriantha ssp.pentzii, Eragrostis rigidior, Eragrostis superba, Panicum coloratum, Schmidtia pappophoroides, Tragus berteronianus andUrochloa panicoides. Maximum seed production varied according to life history strategy and growth form from 0.03 mg seed g^-1 shoot dry weight in the perennialD. eriantha ssp.pentzii which produces long stolons and 14.8 mg seed g^-1 shoot inE rigidior, which produces short geniculate stolons, to 169.1 mg g^-1 in the annualT. berteronianus. Seed production was in most species divided over several peaks during the season. Peaks of seed production were observed 3 to 7 months after the onset of the growth season depending on the start of the rains and the life history strategy and growth form of the species. Seed production varied from maxima of 180 seeds m^-2 inD. eriantha ssp.pentzii to 47000 seeds m^-2 in annual stands ofT. berteronianus. Except for annual grasslands withU. panicoides, seedling emergence data reported are smaller by at least a factor of 10 than the observed seed production. Among other factors, a low quality of produced seeds, predation by birds and insects and previous grazing by livestock may have contributed to this difference.     

Seed and microsite limitation in Rheum nobile,a rare and endemic plant from the subnival zone of Sino-Himalaya

Background: Species persistence, particularly in monocarpic species, depends on the successful recruitment of individuals. An understanding of the factors that limit the recruitment of rare monocarpic plant species is therefore vital for their conservation.

Aims: To identify the factors limiting the recruitment of Rheum nobile, a rare and highly specialised monocarpic giant herb endemic to the high eastern Himalayas.

Methods: Seed sowing (seeds added or not added) and seedling transplanting experiments were conducted in disturbed (vegetation removed) and undisturbed plots in the vicinity of established populations of R. nobile to explore the mechanisms of recruitment limitation. Four levels of photosynthetically active radiation (0, 15, 30 and 50 μmol m^?2 s^?1) and two sowing positions (beneath and above grass litter or moss layer) were manipulated in the laboratory to determine how ground cover limited seedling emergence.

Results: Seed addition increased seedling recruitment. Disturbance significantly increased seedling emergence and establishment. Seed germination significantly decreased with the reduction of light availability, but 31.7% of all seeds germinated in complete darkness. Seedling emergence was close to zero when seeds were sown on top of a layer of grass litter or moss, but rose to 34.5% when the seeds were sown beneath such layers.

Conclusions: Our results indicate that the recruitment of R. nobile is limited by a combination of seed and microsite availability. Therefore, in order to conserve this species, we suggest adding seeds to suitable sites and implementing soil disturbances in existing populations to create suitable microsites.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

El Niño effects on soil seed bank dynamics in north-central Chile

The soil seed bank was monitored in four 75×75 m plots over 6 years (1990–1995) in an arid thorn scrub community in north-central Chile. Sixty-six species were identified. Total seed densities ranged from 2,000 to 42,000/m². Average mass of shrub seeds was significantly greater than that of other growth-forms. Between 70 and 90% of the seeds were less than 1 mg, with those in the 0.51–1.00 mg size class being most numerous. Seed densities were highly variable between years as well as within years, but were also closely associated with plant cover patterns and rainfall regime. Higher seed densities were found in wet years, and in samples taken in early summer and early autumn (i.e., after seed set); the lowest seed densities were in late winter (i.e., after annual plant germination). The annual plant species with the highest cover were also the most abundant in the soil seed bank and exhibited the largest seed density fluctuations. In general, seed densities were 5- to 10-fold higher during the 1991–1992 El Niño/southern oscillation (ENSO) years than non-ENSO years, showing the importance of this phenomenon for seed bank replenishment in the arid region of Chile.     

Seed and microsite limitations of recruitment and the impacts of post-dispersal seed predation at the within population level

Seed predation may cause important seed losses in plant populations, but its impact on the dynamics of populations will depend on the degree of seed or microsite limitations for recruitment. Seed losses will only affect recruitment if it is seed limited. The nature of recruitment limitation (seeds or microsites) is usually ascribed to whole plant populations but it may vary within populations among microhabitats and habitats. Thus, the potential impact of seed predation will also vary within the population, being highest where recruitment is seed limited. The impact to the whole population will depend on the spatial concordance between the intensity of seed predation and that of seed limitation. Recruitment limitations (with seed addition experiments), seed predation (with seed removal experiments), and the dynamics of seed availability in the soil (with soil samples taken both after seed dispersal and before the following dispersal event) of the shrub Corema album (Empetraceae) were investigated in dunes in NW Spain, at microhabitats ‘open ground’, ‘underneath C. album ♀’, and ‘underneath C. album ♂’ at two habitats, sparse and dense scrub. The nature of recruitment limitation (seeds vs. microsites) varied within the population. It was seed limited in the microhabitat ‘open ground’ and microsite limited under shrub cover. The spatial patterns of seedling recruitment were unrelated to seed availability but strongly affected by germination requirements. The spatial discordance between seed availability and recruitment implies a crucial constraint for processes affecting seed availability (seed predation but also e.g., dispersal) to impact recruitment. They will not affect its spatial pattern but only its quantity as long as they act in those sites selected by seeds to germinate. Seed predation was highest underneath mother plants and lowest in open ground. Thus, its potential impact is low, as it is centred where recruitment is not seed limited. This study shows that the analysis of seed predation in relation to recruitment limitations at smaller spatial scales within the population provides more insight to understand its impact.     

The impact of flooding and drought on seeds of Cnidium dubium, Gratiola officinalis, and Juncus atratus, three endangered perennial river corridor plants of Central European lowlands

The effects of flooding and drought on the post-flooding survival and germinability of seeds in the soil were investigated for three endangered river corridor angiosperms Cnidium dubium (Schkuhr) Thell., Gratiola officinalis L. and Juncus atratus Krocker. Freshly matured seeds were placed in small nylon bags and were buried in pots filled with soil collected from a wetland along the Lower Havel River. The pots were subjected to different winter/spring flooding treatments (30, 60, 120, and 180 days) and to drought for up to 3 years. Every year in spring, after drawdown, the seeds were exhumed, counted and tested for germination. Seed mortality of flooded seeds was low, except for C. dubium. Flooding did not affect germination of seeds of J. atratus. Flooding of seeds of C. dubium and G. officinalis had inconsistent, but no beneficial effects on germination of either. The year of harvest and the interannual variability of weather are found to be the main factors affecting germinability of C. dubium and G. officinalis in the course of the three consecutive years.The experiment showed that the capacity of the flooded seed bank to maintain germination ability and viability for extended periods of time could be directly related to the flooding duration of natural habitats of the three species. The three species are resilient to variable flooding. Seeds appeared desiccation tolerant, and single flooding events do not deplete their seed bank.     

Seed viability and seed dormancy of non-native Phragmites australis in suburbanized and forested watersheds of the Chesapeake Bay, USA

The non-native, invasive haplotype of Phragmites australis is rapidly invading tidal and non-tidal wetlands across North America. Phragmites has the potential to spread by seeds and rhizomes. Seed viability and dormancy differences were quantified among 18 patches of non-native Phragmites in subestuarine wetlands in developed (i.e., suburbanized) vs. forested watersheds of the Chesapeake Bay. We used tetrazolium and germination assays to assess seed viability and compared germination percentages and rate of germination among fresh seeds, cold–moist treated seeds, and warm–dry treated seeds to evaluate seed dormancy. Seed viability was <1% in most patches but a few patches produced abundant viable seeds (5–21%). Seed viability, however, did not differ significantly between wetlands in forested vs. developed watersheds. Contrary to studies of Phragmites seed dormancy in European populations, some Phragmites seeds were dormant at maturity; cold–moist treated seeds germinated faster and to higher percentages than fresh seeds or warm–dry treated seeds.     

Endozoochory of Mediterranean aquatic plant seeds by teal after a period of desiccation: Determinants of seed survival and influence of retention time on germinability and viability

The capacity of eight wetland seeds to be internally dispersed by common teal (Anas crecca L.) after a dry period typical of natural Mediterranean wetlands was experimentally quantified. These plant taxa (Chara spp., Echinochloacrus-galli (L.) P. Beauv., Eleocharis palustris (L.) Roem. & Schult, Polygonum lapathifolium L., Potamogeton nodosus Poir., Potamogeton pusillus L., Schoenoplectus mucronatus (L.) Palla, Scirpus maritimus L.) were frequently ingested by teal in the field. Captive teal were fed with known quantities of dry seeds and faecal samples were collected at intervals of 1-2 h for 48 h. The proportion of seeds retrieved after gut passage that germinated varied between taxa from 3% to 83%. From 2% to 63% of the seeds ingested were recovered intact, and 18-95% of these were still viable. Therefore, an overall 32% of the seeds ingested were evacuated in a viable condition, ranging from 0.2% for Chara spp. to 54% for S. mucronatus. Mean seed retention time was 3-5 h. Overall, 94% of seeds were evacuated within 12 h, but 2% were recovered after more than 24 h. Seed viability was reduced at longer retention times, although viable seeds of all taxa except Chara spp. were recovered 44-48 h after the experiment began. Germinability was increased by gut passage, although for some species this effect was only significant after stratification, indicating that they exhibit both mechanical and physical dormancy. Seed germination was accelerated by gut passage, especially at longer retention times. Gut passage increased the proportion of viable seeds, suggesting selective digestion of non-viable ones. Endozoochorous transport by teal appears to be a widespread dispersal mechanism among wetland plants.