海滨猫尾木种子萌发和实生苗抗逆性研究

本文研究不同播种时间对海滨猫尾木种子萌发的影响,以及淹水、盐度、低温条件对其实生苗生长的影响。结果表明,海滨猫尾木种子千粒重为52g,种子密度0.021g·cm^-3,种子首次出芽时间在9~16d不等,发芽率和发芽势随着播种时间推移逐渐降低。水淹至根颈对幼苗生长影响不显著,完全水淹22d后幼苗开始死亡。幼苗耐受盐分浓度为10g·kg^-1,盐分浓度超过10g·kg^-1不利于幼苗正常生长,盐分浓度超过18g·kg^-1幼苗叶片开始卷曲、脱落。3℃低温胁迫对幼苗正常生长有不利影响,0℃会导致幼苗呈现半致死状态。海滨猫尾木种子适宜随采随播,幼苗具有较强的耐水淹特性,适生盐度范围广,不耐低温。     

Seed bank of seasonally flooded grassland: experimental simulation of flood and post-flood

Wetland seed banks comprise the propagules of plant species that have species-specific germination requirements for germination in either flooded or dry conditions. At the community level, wetland structure and succession during and after a seasonal flooding event depends upon the early life-history requirements of species, including germination under flooded and dry conditions. We examined the effects of simulated flood and post-flood scenarios on seedling emergence from a seed bank of seasonally flooded grassland in the Pantanal, Brazil. Field samplings were conducted in both wet and dry seasons, both of which were subject to flood and post-flood conditions. A total of 70 species emerged from the seed bank, dominated by Poaceae and Cyperaceae. Sixteen species were exclusive to the wet and one exclusive to the dry season. The richness of perennial species was higher under flood conditions, while the richness of annuals was greater post-flood. In general, the aquatic and amphibious species exhibited a significant germination response to flooding. Terrestrial species only germinated in post-flood conditions, with higher richness in the dry season. Four species had high seedling abundance in both treatments. The capacity of regeneration by seeds is high in these grasslands and can be increased by seasonal flooding and drawdown. In these seasonally flooded grasslands, we observed three main germination strategies: under flooded conditions, aquatic and amphibious species; post-flood conditions, an explosion of annual amphibious and terrestrial species; and in moist soil, perennial terrestrial species. The differential responses to flooding versus post-flood conditions help to maintain the structure and species richness in the community over time.     

The effect of extended exposure to hypersaline conditions on the germination of five inland halophyte species

In order to determine how salinity and exposure time affect seed viability and germination, seeds of five halophytes, Atriplex prostrata, Hordeum jubatum, Salicornia europaea, Spergularia marina, and Suaeda calceoliformis were exposed to 3.0, 5.0, and 10.0% NaCl solutions for 30, 60, 90, 365, and 730 d. Recovery experiments in distilled water indicated significantly different species responses to salinity over time. Percentage germination and rate of germination in H. jubatum were dramatically reduced following extended exposure and all seeds exposed to 10% NaCl for > 1 yr failed to germinate. Spergularia marina seeds were stimulated following short-term exposure to 3% NaCl; however, germination was delayed and overall germination was significantly reduced with exposure time in the two higher salinity levels. Percentage germination in A. prostrata decreased over time, but salinity level was not related to this reduction. Germination of S. europaea and S. calceoliformis, the most salt-tolerant species being tested, was stimulated by exposure to high salinity. Both species had a significant increase in percentage germination and in the germination rate when compared to seeds germinated in distilled water. Baseline germination data from seeds placed in 0, 1, 2, and 3% NaCl solutions indicated that S. europaea and S. calceoliformis were the only species to germinate in the 3% NaCl solution. Spergularia marina failed to germinate in the 2% NaCl treatment, and germination of A. prostrata and H. jubatum was significantly reduced at this salinity level. It is concluded that prolonged exposure to saline solutions can inhibit or stimulate germination in certain species, and the resulting germination and recovery responses are related to the duration and intensity of their exposure to salt in their natural habitats.     

Linking seed germination and plant height: a case study of a wetland community on the eastern Tibet Plateau

• Seed germination is the earliest trait expressed in a plant's life history, and it can directly affect the expression of post‐germination traits. Plant height is central to plant ecological strategies, because it is a major determinant of the ability of a species to compete for light. Thus, linking seed germination and plant height at the community level is very important to understanding plant fitness and community structure.
• Here, we tested storage condition and temperature requirements for germination of 31 species from a wetland plant community on the eastern Tibet Plateau and analysed correlation of germination traits with plant height in relation to storage condition.
• Germination percentage was positively related to plant height, and this relationship disappeared when seeds were incubated at a low temperature (i.e. 5 °C) or after they were stored under wet‐cold conditions. The response of seeds to dry+wet–cold storage was negatively related to plant height. Based on the scores of each species on the first two principal components derived from PCA, species were classified into two categories by hierarchical clustering, and there was a significant difference between germination and plant height of species in these two categories.
• These results suggest that the requirements for seed germination together with seasonal change in environmental conditions determine the window for germination and, in turn, plant growth season and resource utilisation and ultimately plant height.

     

The influence of vegetation,salinity, and inundation on seed banks of oligohaline coastal marshes

Sea level rise may alter salinity and inundation regimes and create patches of open water in oligohaline coastal marshes, potentially affecting the composition and germination of seed bank species. We conducted seedling emergence experiments to: (1) examine the effects of standing vegetation on the seed banks of three oligohaline marsh communities in coastal Louisiana (dominated by Paspalum vaginatum Sw., Sagittaria lancifolia L., or Spartina patens (Ait.) Muhl., respectively); and (2) investigate the effects of salinity and inundation regime on germination of seed bank species. We also studied the effect of a temporary increase in salinity (to simulate a salt water intrusion event) on the viability of buried seeds. We found that the presence or absence of vegetation within a community affected the abundance of some species in the seed bank but had little effect on species composition. Also, the seed banks of the three communities exhibited considerable overlap in species composition and had similar species richness (10–11) and diversity (antilog Shannon-Weaver diversity index = 6.5–7.1), despite differences in vegetation type. Higher salinities and flooding reduced seedling emergence for most species; few species emerged at salinities above four parts per thousand (ppt), and only Sagittaria lancifolia and Eleocharis parvula germinated well under flooded conditions. A temporary increase in salinity did not affect species richness or seedling emergence of most species. Our results suggest that differences in vegetation may have little effect on the composition of seed banks of oligohaline marshes. However, higher salinities and greater depth and duration of inundation (anticipated as global sea level continues to rise) may decrease recruitment of seed bank species, reducing their abundance in oligohaline marsh communities.     

Can elevated CO<Subscript>2</Subscript> modify regeneration from seed banks of floating freshwater marshes subjected to rising sea-level?

Higher atmospheric concentrations of CO₂ can offset the negative effects of flooding or salinity on plant species, but previous studies have focused on mature, rather than regenerating vegetation. This study examined how interacting environments of CO₂, water regime, and salinity affect seed germination and seedling biomass of floating freshwater marshes in the Mississippi River Delta, which are dominated by C₃ grasses, sedges, and forbs. Germination density and seedling growth of the dominant species depended on multifactor interactions of CO₂ (385 and 720 μl l⁻¹) with flooding (drained, +8-cm depth, +8-cm depth-gradual) and salinity (0, 6% seawater) levels. Of the three factors tested, salinity was the most important determinant of seedling response patterns. Species richness (total = 19) was insensitive to CO₂. Our findings suggest that for freshwater marsh communities, seedling response to CO₂ is species-specific and secondary to salinity and flooding effects. Elevated CO₂ did not ameliorate flooding or salinity stress. Consequently, climate-related changes in sea level or human-caused alterations in hydrology may override atmospheric CO₂ concentrations in driving shifts in this plant community. The results of this study suggest caution in making extrapolations from species-specific responses to community-level predictions without detailed attention to the nuances of multifactor responses.     

南亚热带森林24种乔木的种子萌发和幼苗生长

以膨胀珍珠岩为基质,在光和暗的条件下,对２４种南亚热带森林乔木的种子萌发和幼苗生长进行了研究。种子的形态和重量与种的演替阶段有关,种子的重量也与不同的种有关。不同种的萌发率差异较大,种子较大的种,萌发率较高。肉质果实的种子,开始萌发的时间较长。光和暗条件对萌发率和萌发速度无明显影响。幼苗高度和种子重量呈正相关,这一相关在光条件下比暗条件下更明显。暗条件明显地引起群落演替早期的树种的茎徒长,限制根的生长,而对演替后期种无显著的影响。幼苗地上部分和根系生物量的分配与种的演替阶段有关,也与不同的科有关。幼苗地上部分和根系生物量的分配也受光暗条件的影响,而不受种子重量的影响。根和叶的生物量分配和种在其群落演替阶段有关。幼苗的生物量与种子重量有显著的正相关。幼苗的相对生长率相差较大,在没有营养供应的生长基质中,以演替中间阶段的种的相对生长率较大。幼苗从种子的物质利用效率与种子重量呈负相关,而与种的演替阶段无明显的相关。     

Gap disturbance triggers the recolonization of the clonal plant Ambrosia tenuifolia in a flooding grassland of Argentina

Abstract Flooding provokes the death of many dicotyledonous species in grazed grasslands of the Flooding Pampa in Argentina, including the clonal plant Ambrosia tenuifolia, which produce the opening of numerous gaps. The objective of this study was to investigate the recolonization of grassland by A. tenuifolia after this species disappeared due to the occurrence of prolonged flooding events. To this end, responses of seed germination to environmental factors associated with gaps, such as light quality and temperature regime, conditions related to seedling survival, and clonal growth of ramets outside the gaps were studied in two different experiments in the field. Environmental factors related to gaps promoted the recruitment of new genets. The combined effect of alternating temperatures and the high red : far‐red ratio set off germination from the soil seed bank; germination also was enhanced when signals were generated artificially under the intact canopy in the field. Higher resource availabilities and maximum seedling survival were recorded in canopy gaps, which were the focus of invasion. Grassland recolonization outside the gaps continued rapidly by clonal growth, from small gaps and large ones, even within the dense surrounding canopy. This provoked an intense competition with the other species. Gap opening by disturbances, seed germination in gaps and clonal growth were decisive for the recolonization of A. tenuifolia populations. This sequence of events triggered the recolonization of the plant community by this species, in sites where it had been eliminated by prolonged flooding. This process represents one of the most significant fluctuations in the vegetation dynamics of the Flooding Pampa Grasslands.     

Seed banks and seed population dynamics of halophytes

In this review I will describe the importance of seed banks and thepopulation dynamics of seeds on the distribution of species in salinehabitats. The main questions being examined in this review include: 1.Does the seed bank represent the flora of the entire salinity gradient or isit restricted to the species in each zonal community? 2. Is the size andspecies composition of the persistent seed bank regulated by the degree ofsalt stress in habitats along an environmental gradient? 3. Does thepopulation dynamics of seeds influence the temporal and spatial distributionof plant species in saline habitats? Seed banks may be transient orpersistent depending upon the physiological responses of species and thesoil environment in which the seeds are found. The formation of zonalcommunities in salt marsh environments is affected by changes in soilsalinity and flooding along an elevational gradient. Population dynamics ofseeds have been found to determine the spatial and temporal distributionof species along salinity gradients. The flora and relative density of speciesof zonal communities are significantly dependent upon the stress toleranceof species at different stages of development and the presence of transientor persistent seed banks. The occurrence of a seed bank is related to thesalinity tolerance of species at the germination stage of development, aseeds ability to tolerate hypersaline conditions and flooding, and whetheror not species are able to maintain a persistent seed bank until hypersalineconditions are alleviated.

     

Effects of waterlogging on seed germination of three Mediterranean oak species: Ecological implications

Soil water saturation during prolonged periods of time generates a negative impact on nearly all terrestrial plants. In Mediterranean woodlands, precipitation can be very abundant during the wet season, inducing temporary soil waterlogging, coinciding with the seed dispersal and germination time of many species. We investigated the effects of waterlogging on seed germination and early root growth of three coexisting oak species (Quercus canariensis, Q. suber and Q. pyrenaica), by completely flooding of seeds for various periods of time. The three oak species showed a certain level of tolerance to waterlogging, only being affected those seeds subjected for long periods of submersion (over 30 days). Waterlogging during prolonged periods of time decreased the probability of seed germination in the three oak species, lengthened the time to germination, and hampered root development in two of the studied species. The main differences between oak species occurred in terms of root growth (Q. canariensis being the less affected, and Q. suber the most); these differential responses could be related to a species rank of waterlogging tolerance. Thus inter-specific differences in germination responses to waterlogging could contribute to explain, at least partially, species habitat and distribution patterns across landscapes. Seed mass also played an important role on different aspects of germination, though its relative importance varied as function of species and waterlogging treatment. The tolerance to stress induced by waterlogging increased with seed mass, but only in the case of Q. canariensis.     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Seed flotation and germination of salt marsh plants: The effects of stratification, salinity, and/or inundation regime

We examined the effects of cold stratification and salinity on seed flotation of eight salt marsh species. Four of the eight species were tested for germination success under different stratification, salinity, and flooding conditions. Species were separated into two groups, four species received wet stratification and four dry stratification and fresh seeds of all species were tested for flotation and germination. Fresh seeds of seven out of eight species had flotation times independent of salinity, six of which had average flotation times of at least 50 d. Seeds of Spartina alterniflora and Spartina patens had the shortest flotation times, averaging 24 and 26 d, respectively. Following wet stratification, the flotation time of S. alterniflora seeds in higher salinity water (15 and 36 ppt) was reduced by over 75% and germination declined by more than 90%. Wet stratification reduced the flotation time of Distichlis spicata seeds in fresh water but increased seed germination from 2 to 16% in a fluctuating inundation regime. Fresh seeds of Iva frutescens and S. alternflora were capable of germination and therefore are non-dormant during dispersal. Fresh seeds of I. frutescens had similar germination to dry stratified seeds ranging 25-30%. Salinity reduced seed germination for all species except for S. alterniflora. A fluctuating inundation regime was important for seed germination of the low marsh species and for germination following cold stratification. The conditions that resulted in seeds sinking faster were similar to the conditions that resulted in higher germination for two of four species.     

Seed germination and salinity tolerance in plant species growing on saline wastelands

Seven plant species including three chenopods:Suaeda fruticosa, Kochia indica, Atriplex crassifolia and four grasses:Sporobolus arabicus, Cynodon dactylon, Polypogon monspeliensis, Desmostachya bipinnata, varied greatly in their seed germination and growth responses to soil moisture or salinity. The germination percentage of each species was significantly lower at soil moisture level of 25 % of water holding capacity than at the levels ranging from 50 to 125 %. Increase in salinity resulted in gradual decrease in seed germination of each species. Growth responses of species to salinity varied widely from significant decrease with slight salinity to stimulation up to salinity levels of 20 dS m^-2. Higher K⁺Na⁺ratios in plant shoots of all species compared to that in the root medium indicated selective K⁺uptake. Higher tolerance in chenopod species seems to be attendant on their ability for internal ion regulation. We are thankful to Mr. Noor Ahmad for his assistance in experimental work.     

High salinity impacts germination of the halophyte Cakile maritima but primes seeds for rapid germination upon stress release

Seed germination recovery aptitude is an adaptive trait of overriding significance for the successful establishment and dispersal of extremophile plants in their native ecosystems. Cakile maritima is an annual halophyte frequent on Mediterranean coasts, which produces transiently dormant seeds under high salinity, that germinate fast when soil salinity is lowered by rainfall. Here, we report ecophysiological and proteomic data about (1) the effect of high salt (200 mM NaCl) on the early developmental stages (germination and seedling) and (2) the seed germination recovery capacity of this species. Upon salt exposure, seed germination was severely inhibited and delayed and seedling length was restricted. Interestingly, non‐germinated seeds remained viable, showing high germination percentage and faster germination than the control seeds after their transfer onto distilled water. The plant phenotypic plasticity during germination was better highlighted by the proteomic data. Salt exposure triggered (1) a marked slower degradation of seed storage reserves and (2) a significant lower abundance of proteins involved in several biological processes (primary metabolism, energy, stress‐response, folding and stability). Yet, these proteins showed strong increased abundance early after stress release, thereby sustaining the faster seed storage proteins mobilization under recovery conditions compared to the control. Overall, as part of the plant survival strategy, C. maritima seems to avoid germination and establishment under high salinity. However, this harsh condition may have a priming‐like effect, boosting seed germination and vigor under post‐stress conditions, sustained by active metabolic machinery.     

Seed dimorphism and germination behaviour in Salicornia patula

One of the annual species of the genus Salicornia from the Mediterranean coast, Salicornia patula presents a seed dimorphism. This morphological seed dimorphism is linked with a physiological ‘dimorphism’ with regard to some requirements for germination. Central seeds, in the inflorescence, are larger, are dispersed attached to the perianth, have no light requirement for germination and seem to have a low germination response to salinity. The lateral seeds, which are free from perianth envelopes, require light and cold pretreatment and seem to have a high sensitivity to salinity. Salicornia patula colonizes unstable transition zones between permanently flooded muds and perennial vegetation. These zones have a winter flooding period. The beginning of the flooding period and of the dry period may vary from year to year, depending on the erratic distribution of autumn and winter rains on the Mediterranean coast. So, in some years, a seedling population may be destroyed before reaching the stage of seed production. Seed dimorphism might cause germination to the more distributed in time, thus reducing the chances of extinction of a complete generation. In the present paper salinity and temperature response are described more precisely, are investigated, some aspects of the longevity of the two types of seed, and the adaptative significance of the corresponding dispersal and germination syndromes are discussed in more detail.     

Effects of storage,mucilage presence,photoperiod, thermoperiod and salinity on germination of Farsetia aegyptia Turra (Brassicaceae) seeds: implications for restoration and seed banks in Arabian Desert

Seed viability and germination are key factors in the success of restoration efforts, especially when stored seeds are used. However, the effect of seed storage on germination of most of the native Arabian species is not well documented. We investigated the effect of storage time and role of the seed mucilage in regulating germination, dormancy, salinity tolerance and consequential survival strategy of F. aegyptia in an unpredictable arid desert setting. Effect of light and temperature during germination was studied under two photoperiods and two thermoperiods using intact and de-mucilaged seeds. Presence of mucilage and thermoperiod did not affect the germination. However, seed collection year and photoperiod had a highly significant effect on the germination. Increasing salinity levels decreased the germination of F. aegyptia but ungerminated seeds were able to germinate when salinity stress was alleviated. Seed storage at room temperature enhances the germination percentage, indicating that F. aegyptia seeds have physiological dormancy and it can be alleviated by after-ripening at dry storage. In addition, F. aegyptia seeds show ability to germinate at lower salinity concentration and remain viable even at higher saline conditions, indicating their adaptability to cope with such harsh environmental conditions.     

Differential effects of four abiotic factors on the germination of salt marsh annuals

Interspecific differences in responsiveness to temperature, photoperiod, soil salinity, and soil moisture confirm the hypothesis that abiotic factors differentially affect the germination of salt marsh plants. In growth chamber experiments, four of eight annual species responded to small differences in temperature or photoperiod. Increasing soil salinity decreased the final proportion of seeds germinating and slowed germination for each of the seven species tested. Higher soil moisture increased the proportion germinating of five species and germination speed of all seven species. Salinity and moisture interacted to affect the proportion germinating of five species and germination speed of all seven species. Although the abiotic factor with the largest effect on germination varied among species, more species responded to, and the magnitudes of the responses were larger for, soil salinity than for the other abiotic factors. These germination tests partially explained interspecific differences in the timing of germination in the field. Patterns of Hutchinsia procumbens, Lythrum hyssopifolium, Parapholis incurva, and possibly Lasthenia glabrata ssp. coulteri germination in response to a nonseasonal rainfall could be explained by their response to salinity, temperature, or photoperiod. Fine-scale differences in the timing of establishment within the typical germination window and spatial distributions along salinity and moisture gradients were not explained.     

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

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

Flooding and grazing promote germination and seedling establishment in the perennial grass Paspalum dilatatum

Seed germination and seedling emergence are key processes for population recruitment. Flooding and grazing are disturbances forming gaps that may strongly influence recruitment patterns in space and time, but their combined effects and action mechanisms have rarely been addressed. In this study we analysed the effects of microhabitat conditions associated with winter flooding and spring‐summer defoliation on seed germination and seedling establishment of Paspalum dilatatum, a dominant perennial C₄ grass in native grasslands of the Flooding Pampa, Argentina. The dynamics of seedling emergence from natural seed banks and buried seeds was studied in a factorial experiment with flooding and defoliation treatments applied to soil monoliths (mesocosms) collected from natural grassland. Additional laboratory experiments were applied to investigate seed germination under different combinations of temperature, light quality and simulated flooding. Seed germination and seedling emergence of P. dilatatum were promoted by flooding and high intensity defoliation. Gaps generated by flooding were maintained by high intensity defoliation exercising a synergistic effect on survival seedlings. Flooding resulted in the breaking of seed dormancy and higher germination rates associated with alternating temperature and the activation of the phytochrome system. Our results indicate that microhabitat conditions associated with the disturbances forming gaps, such as flooding and heavy grazing, synergistically promote the recruitment process of this dominant grass species.     

Establishment of four Salicaceae species on river bars in interior Alaska

Patterns of seed and vegetative reproduction were investigated for three floodplain species, balsam poplar Populus balsamifera L., feltleaf willow Salix alaxensis (Anderss.) Cov. and sandbar willow Salix interior Rowlee, and one upland species, trembling aspen Populus tremuloides Michx., in interior Alaska. All four species have similar patterns of seed germination in response to moisture stress and high salt concentrations when tested under laboratory conditions. In field experiments, percent germination of all four species was also very similar, ranging from 0% on dry sandy sites, to greater than 60% on mesic silty sites. Germination on salt crusts ranged from 0–40% for all species, depending on the physical characteristics of the soil surface. Colonization of mesic silty sites was almost exclusively by seed, whereas colonization of dry sandy sites was limited to those species which were able to root sucker under floodplain conditions. Root suckering was also an important means of establishment on frequently inundated sites where establishment by seed was limited by flooding. Differences between the species in their distribution on the floodplain were related to differences in patterns of vegetative reproduction, but not seed germination.