Salt Marsh Plant Zonation Research Articles

Germination patterns along a salinity gradient of closely-related halophytes in sympatry

Sympatric closely-related species co-occur within heterogeneous environments by occupying separate ecological niches as result of adaptive differentiation. Soil salinity is one of the most relevant drivers leading plant zonation in salt marshes, where germination is a critical stage in the life cycle of many halophytes. Salinity influence on germination patterns was evaluated for six sympatric Limonium species (L. admirable, L. caesium, L. cossonianum, L. delicatulum, L. supinum and L. tobarrense), which grow under arid environmental conditions in an inland salt marsh from the southeastern Spain. Experiments were conducted to determine the effects of different salinities (0–6% NaCl) on seed germination under 12 h in light at 25 °C and for 12 h in dark at 20 °C. Best seed germination was obtained under non-saline conditions, and germination was delayed and significantly reduced by increasing the salinity levels for all the studied taxa. The germination variables related to final germination percentage, mean time to germination and recovery were directly influenced by salinity. Seeds recovered their germination capacity after transferring the seeds from saline solutions to distilled water, especially those seeds previously treated under high salinity conditions. Our results suggest that the successful establishment of Limonium communities depends on the germination strategy which implies a rapid germination velocity when the soil salinity decreases. Two different germination syndromes based on germination percentage and germination recovery have been found. Hence, these germination strategies would favour the adaptation of different Limonium species under favorable conditions, shaping the plant asssembly within the saline habitat.

Leaf fatty acid remodeling in the salt-excreting halophytic grass Spartina patens along a salinity gradient

Spartina patens is a highly dispersed halophytic grass invader in Mediterranean marshes. It is also characterized by having a high degree of resistance to salinity, one of the main drivers of plant zonation in salt marshes. Nevertheless, the physiological basis behind the extreme resistance of S. patens requires more detailed studies. In the present work, we aimed to study how membrane fatty acid remodeling could contribute to the resistance of this plant to salt. Spartina patens individuals exposed to increasing levels of salinity and its leaf fatty acid profile under lipid peroxidation products evaluated under all tested concentrations. A significant increase in the relative amounts of the saturated fatty acids (SFA) was observed, namely palmitic acid (C16:0), essential for PS II functioning, and stearic (C18:0) acid. The chloroplastidial trans-hexadecenoic acid (C16:1t) as well as the polyunsaturated linoleic (C18:2) and linolenic (C18:3) acids showed significant decreases in all the salt treatments. These changes led to a reduction in the double bond index in salt-treated plants which reflects reduction of the fluidity of the chloroplast membranes, which could contribute to maintain the membrane impermeable to the toxic exogenous Na. Despite the decrease observed in the total fatty acid contents in plants exposed to high salt concentrations the amounts of lipid peroxidation products decreased highlighting the resistance of this species towards toxic exogenous salt concentrations. Membrane fatty acid remodeling could represent an efficient mechanism to maintain the photosynthetic machinery of S. patens highly efficient under salt stress.

Biogeography of salt marsh plant zonation on the Pacific coast of South America

AbstractAimThe aim of this study was to investigate the biogeography of plant zonation in salt marshes on the Pacific coast of South America; to examine whether salt marsh plant zonation varies with latitude; and to explore the relative importance of climatic, tidal, edaphic and disturbance factors in explaining large‐scale variation in salt marsh plant community structure.LocationA 2,000‐km latitudinal gradient on the Pacific coast in Chile, with a climate shift from hyper‐arid at low to hyper‐humid at high latitudes.MethodsPlant zonation was quantified in field surveys of ten marshes. Climate, tidal regimes, edaphic factors and disturbances (tsunami and rainfall floods) were determined. We used multivariate analyses to explore their relative importance in explaining large‐scale variation in salt marsh plant community structure.ResultsAcross latitude, marshes were dominated by distinct plant communities in different climate regions, especially at the extreme dry and wet latitudes. Intertidal plant species zonation was present in hyper‐arid and semi‐arid climates, but not in arid, humid and hyper‐humid climates. Latitudinal variation in low‐marsh plant communities (regularly flooded at high tide) was largely a function of precipitation, while at high marshes (never flooded at high tide) latitudinal variation was explained with precipitation, temperature, tidal cycles, soil salinity and disturbances.Main conclusionsSalt marshes on the Pacific coast of South America belong to Dry Coast and Temperate biogeographic types. Salt marsh plant zonation varies across latitude, and is explained by climatic, tidal, edaphic and disturbance factors. These patterns appear to be mechanistically explained by extrapolating experimentally generated community assembly models and have implications for predicting responses to climate change.

Microbial Community Composition and Extracellular Enzyme Activities Associated with Juncus roemerianus and Spartina alterniflora Vegetated Sediments in Louisiana Saltmarshes.

Saltmarshes are typically dominated by perennial grasses with large underground rhizome systems that can change local sediment conditions and be important in shaping the sediment microbial community. Factors such as salinity that control plant zonation in saltmarshes are also likely to influence the microbial community, but little is known as to whether microbial communities share distribution patterns with plants in these systems. To determine the extent to which microbial assemblages are influenced by saltmarsh plant communities, as well as to examine patterns in microbial community structure at local and regional scales, we sampled sediments at three saltmarshes in Louisiana, USA. All three systems exhibit a patchy distribution of Juncus roemerianus stands within a Spartina alterniflora marsh. Sediment samples were collected from the interior of several J. roemerianus stands as well as from the S. alterniflora matrix. Samples were assayed for extracellular enzyme activity and DNA extracted to determine microbial community composition. Denaturing gradient gel electrophoresis of rRNA gene fragments was used to determine regional patterns in bacterial, archaeal, and fungal assemblages, while Illumina sequencing was used to examine local, vegetation-driven, patterns in community structure at one site. Both enzyme activity and microbial community structure were primarily influenced by regional site. Within individual saltmarshes, bacterial and archaeal communities differed between J. roemerianus and S. alterniflora vegetated sediments, while fungal communities did not. These results highlight the importance of the plant community in shaping the sediment microbial community in saltmarshes but also demonstrate that regional scale factors are at least as important.

Herbivory drives zonation of stress-tolerant marsh plants.

Ecological studies of plant distributions along environmental gradients, such as plant zonation in salt marshes, have primarily focused on abiotic stress and plant interactions (competition and facilitation). A decades-old paradigm is that the stressful and benign boundaries of salt marsh plants are determined by abiotic stress and competition, respectively. Although consumers have long been recognized as mediating algal and sessile animal zonation in the rocky intertidal, their role in generating plant zonation in salt marshes remains largely unexplored. We examined the zonation of two annual succulents, Salicornia europaea and Suaeda salsa, along an elevation gradient in a northern Chinese salt marsh, with and without manipulating the common herbivorous crab Helice tientsinensis. Salicornia occupies waterlogged, low-salinity habitats, whereas Suaeda dominates non-waterlogged, hypersaline habitats at higher elevations. We first conducted a pot experiment crossing salinity, waterlogging, and competition, followed by a field experiment with removal of competitors, and found that neither waterlogging nor salinity stress explained the absence of either species from the other's zone, while Suaeda competitively excluded Salicornia from the upper non-waterlogged zone. We then conducted field and lab herbivory experiments, which showed that Helice preferentially grazed Suaeda at waterlogged low elevations and that Helice grazing on Suaeda increased with waterlogging. These results reveal that while competition plays a role in the zonation by excluding Salicornia from the upper Suaeda zone, crab grazing limits the success of Suaeda in the lower Salicornia zone. These findings challenge the idea that plant interactions and abiotic stress are sufficient to explain marsh zonation in all cases, and highlight an overlooked role of consumers, a role potentially general across diverse intertidal ecosystems. Future models of plant distributions should consider how consumer pressure interacts with plant interactions and abiotic stress across environmental gradients.

Community Structure and Abiotic Determinants of Salt Marsh Plant Zonation Vary Across Topographic Gradients

We investigated compositions of plant communities and their abiotic determinants in the Yellow River estuary, China. Along a topographic gradient, we quantified plant compositions and abiotic factors in different vegetation zones, and examined the relationships between plant communities and abiotic factors using canonical correspondence analysis and the effects of vegetation shading using a removal experiment. The relationships between plant communities and abiotic factors differed between high elevations and low elevations. Salinity and flooding oppositely related with distributions of plant communities at low elevations, but they appeared to operate synergically at high elevations. The effects of vegetation shading were found to vary across the topographic gradient, indicating spatial variations in potential positive interactions among plants. These results suggest that spatial variations in determinants of community structure should be addressed in future studies in estuarine and coastal systems, as well as in other natural habitats.

Effects of environmental gradients on the performances of four dominant plants in a Chinese saltmarsh: implications for plant zonation

AbstractPhysical conditions and biotic interactions are believed to be the determinants of plant zonation in saltmarshes. However, in rapidly developing estuarine marshes, succession is regarded as the primary process responsible for plant zonation and it is controlled mainly by environmental factors. Salinity and inundation are two important factors responsible for the distribution pattern of dominant plants in coastal saltmarshes. Here we conducted a common garden experiment as well as a field transplanting to examine the responses of four dominant saltmarsh plants (native Scirpus mariqueter, Scirpus triqueter and Phragmites australis, and exotic Spartina alterniflora) in the Yangtze River estuary to environmental gradients, which may help us understand their current and potential zonation. The results showed that Scirpus adapted to freshwater and less inundated habitats, Phragmites performed well in brackish or freshwater environments with less inundation, and Spartina tolerated the highest salinity and deepest inundation. In the harshest environments (the highest salinity and water level), only Spartina performed well. In the mild environments, however, there were only minor differences in the performances among the four species. The potential ranges of Phragmites and Spartina were predicted to be larger than their current ones, and their lower boundaries might be set by tidal scour rather than edaphic factors. With the saltmarsh succession, invasive Spartina in the Yangtze River estuary might ultimately replace Scirpus, and alter the zonal patterns of native saltmarsh plants, which will lead to severe ecosystem consequences. Thus, proper management measures (e.g., repeated mowing) need to be implemented to control this invasive exotic plant, and restore the vulnerable ecosystems invaded by Spartina in the Yangtze River estuary.

Tidal regime, salinity and salt marsh plant zonation

Salt marsh morphology is known to be strongly correlated to vegetation patterns through a complex interplay of biological and physical processes. This paper presents the results of field surveys at several study salt marshes within the Venice Lagoon (Italy), which indicate that salt-marsh macrophyte species may indeed be associated with narrow ranges of soil topographic elevation. Statistical analyses show that several properties of the frequency distributions of halophytes presence are sensitive not only to variations in soil elevation, but also to the specific marsh considered. Through direct in situ sampling and by use of a finite-element hydrodynamic model the role of plant submersion duration and frequency in determining the observed variability of vegetation species is then studied. Measurements of soil salinity have also been performed at selected salt marshes to address its influence on vegetation occurrence. With implications for tidal marshes in general, the distribution of halophytes in the salt marshes considered is found not to be responding to simple rules dictated by the tidal cycle or to salinity, and that such factors, when singularly considered, cannot explain the observed spatial distribution of halophytes. On the basis of observations and modelling results it is thus concluded that a combination of multiple factors, likely dominated by saturated/unsaturated flow in the soil, may be responsible for the observed macrophyte distribution.

Plant zonation in irregularly flooded salt marshes: relative importance of stress tolerance and biological interactions

Summary 1 Most studies of salt marsh plant zonation have been at middle to high latitudes of the northern hemisphere, in euhaline or periodically hypersaline marshes with regular tides. In this study, we examined plant zonation in an irregularly flooded marsh in southern Brazil. Pore water characteristics were compared in four vegetation zones across a marsh elevation gradient. Reciprocal transplants between vegetation zones and removal experiments were performed to examine species interactions in low and mid marshes. 2 There was no distinctive gradient of physical stress across the elevation of irregularly flooded low and mid marshes. Moreover, the three dominant plants, Spartina alterniflora, Spartina densiflora and Scirpus maritimus, were able to grow across the entire elevation gradient, i.e. within zones normally occupied by the other species. The only exception was Spartina alterniflora, which was strongly limited by selective herbivory by the crab Chasmagnathus granulata in the Scirpus maritimus zone. 3 Although intra- and interspecific competition reduced growth of all three species, no competitive hierarchy was found in any vegetation zone. 4 These results suggest that, as in tidal marshes of the north hemisphere, competition is important in structuring salt marsh plant communities. In contrast, however, plant zonation in irregularly flooded marshes cannot be explained by displacement of competitive subordinates to physically stressful habitats. The roles of founder effects and selective herbivory in such marshes therefore merits further investigation.

Nitrogen addition does not reduce belowground competition in a salt marsh clonal plant community in Mississippi (USA)

Previous studies have suggested that belowground competition for nutrients influences plant zonation in salt marshes. In this study, I tested the hypothesis that competition for nitrogen structured a clonal plant community in a nitrogen-limited salt marsh in coastal Mississippi, USA. In contrast to most previous field studies that have investigated mechanisms of competition, I examined clonal growth responses of established genets of a nitrogen-demanding low-intertidal species (Spartina alterniflora) to nitrogen addition and the removal of a nitrogen-conserving high-intertidal species (Juncus roemerianus). Nitrogen addition stimulated clonal invasion of the Juncus zone by Spartina but did not reduce the significant competitive effects of Juncus on Spartina. Simulated Juncus shade did not reduce invasion of the Juncus zone by Spartina, indicating that belowground competition reduced clonal invasion. In the last year of the study, the border shifted unexpectedly towards the Spartina zone, resulting in competitive displacement of Spartina by Juncus. Nitrogen addition did not prevent or slow this displacement, further contradicting the nitrogen competition hypothesis. Although growth rates were much more strongly limited by nitrogen in Spartina than in Juncus, nitrogen addition did not cause the displacement of Juncus by Spartina after three growing seasons. I conclude that zonation of Spartina and Juncus is maintained by preemption of space and greater tolerance of low nitrogen supplies by Juncus in the high marsh. These results contrast sharply with findings of reduced belowground competition with nutrient addition in previous studies and highlight the important role of nutrient-mediated competition for space between clonal plants.

Competition and Salt-Marsh Plant Zonation: Stress Tolerators May Be Dominant Competitors

Competition and Salt-Marsh Plant Zonation: Stress Tolerators May Be Dominant Competitors

Edaphic characterization and soil ionic composition influencing plant zonation in a semiarid Mediterranean salt marsh

Soil characteristics and plant zonation were studied in a semiarid Mediterranean salt marsh in SE Spain. According to topographic sequences and plants distribution, two transects were established from the border of La Mata lagoon to the upland vegetation limit and soils were described and analysed. Regularly spaced plots were established in these transects in accordance with the stands of vegetation and surface soil samples were taken every 2 months for 2 years. The following edaphic factors were determined: soil moisture content, pH and concentration of Cl −, SO 4 2−, Ca 2+, Mg 2+, Na + and K + in the saturation extract. In addition, the groundwater level was measured and the duration of the flooding periods established in each plot. Soil–plant relationships were studied by means of canonical correspondence analysis. Based on rainfall data for the study period, dry and wet seasons were separated and the habitats of the plant communities were compared for salt quantity and quality independently for each season. Soils were classified, according to FAO (1998), as Hypercalcic, Sodic and Mollic Solonchaks and Hypercalcic Sodic Calcisols. The most important variables which explained plant zonation were: the flooding period, total salinity, minimum Ca 2+/Na + ratio and the mean sodium adsorption ratio. When the habitats of the main plant communities were compared, differences in salt quantity, quality and seasonal variations were found. Two chenopod shrubs, Arthrocnemum macrostachyum and Sarcocornia fruticosa, predominated in the most saline areas. More pronounced seasonal variations in soil salinity were found in the A. macrostachyum zone than in the Sarcocornia fruticosa zone. The highest value for K +/Na + and Ca 2+/Na + ratios were measured in the Suaeda vera stand. The Lygeum spartum zone was distinguished by the high Ca 2+/Na + and Ca 2+/Mg 2+ ratios in the wet period. Among the rushes, Schoenus nigricans predominated in the less saline areas, where the K +/Na + ratio was higher and the Ca 2+/Na + ratio lower than where Juncus maritimus predominated. Limonium cossonianum communities occupied an intermediate position with respect to soil salinity, between the chenopod shrubs and the other communities. Our results suggest that salt marsh plant zonation is influenced by temporal and spatial edaphic gradients which must be jointly considered if soil–plant relationships in saline soils are to be fully understood.

Salt Marsh Plant Zonation: The Relative Importance of Competition and Physical Factors

In Carpinteria Salt Marsh, Salicornia virginica (pickleweed) grows at lower marsh elevations than does Arthrocnemum subterminalis (Parish's glasswort). Standing biomass of both species was greatest immediately adjacent to their abrupt border, suggesting that conditions for plant growth were best here. We utilized field experiments, in which growth rates of naturally occurring and transplanted individuals of both species were measured in four marsh zones, to investigate the role of edaphic factors and competition in maintaining this zonation pattern. The frequency of flooding, and hence soil waterlogging, was greatest at lower marsh elevations, whereas salinity was highest at higher marsh elevations. Consequently, it was not clear, a priori, which part of the marsh had the most severe physical conditions. In our field experiments, both Salicornia and Arthrocnemum grew better in the two middle marsh zones (high Salicornia zone and Arthrocnemum zone) than in either the low marsh (low Salicornia zone), where flooding was frequent and soils were waterlogged, or the high marsh (transition zone), where soil salinity was extremely high during much of the year and plant water potentials very low. However, Salicornia appeared better able to tolerate flooding, and so persisted in the low Salicornia zone, whereas Arthrocnemum appeared better able to tolerate high salinities, and so persisted in the transition zone. Interspecific competition was most important in the relatively benign middle marsh zones, where each species excluded the other from a portion of this prime habitat. In this marsh, flooding, soil salinity, and competition all interacted to determine plant zonation patterns, but the relative importance of these factors varied at different elevations.

Determinants of Pattern in a New England Salt Marsh Plant Community

In New England salt marshes, Spartina alterniflora dominates the low–marsh habitat, which is covered daily by tides. The high–marsh habitat, which is not flooded daily, is dominated on its seaward border by Spartina patens, and on its terrestrial border by Juncus gerardi. Each of these vegetation zones has a characteristic suite of physical factors associated with differences in tidal inundation. In particular, substrate redox increases and salinity decreases with decreasing marsh elevation. Although correlations between physical factors and the occurrence of specific marsh plants have been suggested to be causal, a 5–mo transplant experiment suggested that the distribution of perennials across the marsh does not correspond to their potential performance across the marsh in the absence of surrounding vegetation. While the high–marsh perennials appear to be restricted to the high–marsh habitat by harsh physical conditions in the low–marsh habitat, the low–marsh dominant, S. alterniflora, is capable of vigorous growth across the entire marsh and appears to be excluded from the high–marsh habitat by the high–marsh perennials. Throughout the high marsh, two other plant species, Distichlis spicata and Salicornia europaea, are found associated with areas that have been disturbed recently. Physical disturbance, in the form of mats of dead plant material (wrack) rafted by tides onto the marsh, is most severe in the spring and early summer, and decreases with increasing marsh elevation. Differential plant mortality results from short–term disturbance events. D. spicata and S. alterniflora are more tolerant of wrack burial than are the other marsh plants, and short–term disturbance increases the relative abundance of these species in the community. Longer lasting disturbance events kill all the underlying vegetation, leaving discrete bare patches throughout the high marsh. D. spicata rapidly colonizes these patches with vegetative runners, while S. alterniflora and Sa. europaea recruit to these patches by seed. The relative abundance of these plants in recently created bare patches exceeds greatly their relative abundance in the surrounding vegetation. Over time, however, these early colonizers are overgrown and displaced in high–marsh patches by S. patens and J. gerardi, which grow slowly, as dense turfs of roots, rhizomes, and tillers. Physical disturbance and interspecific competition appear to be major determinants of the spatial pattern of marsh plant communities. These processes will need to be considered in relation to edaphic factors in elucidating the underlying mechanisms of salt marsh plant zonation.