Crab: snail size-structured interactions and salt marsh predation gradients

We studied size-structured predator-prey interactions between blue crabs (Callinectes sapidus) and marsh periwinkles (Littoraria irrorata) with a combination of field studies, laboratory experiments and individual-based modeling. Size distributions of Littoraria differed among years at the same sites in a salt marsh and could largely be explained by dominance of strong cohorts in the population. At a given site, abundance increased with elevation above tidal datum. Size-selective predation by blue crabs does not appear to be an important regulator of snail size distributions but may have a major effect on local abundance. Laboratory studies indicated that predator-prey interactions between Callinectes and Littoraria are strongly size-dependent. Crabs were generally effective at feeding on periwinkles at size ratios greater than approximately 6 (crab width: snail length). At lower size ratios crabs were far less effective at manipulating the snails, which often survived but with damaged shells. An individual-based model which incorporated information about incidence of snail shell scarring (resulting from non-lethal interactions) and snail density, predicted reduced predation rates and smaller average crab size with distance from the low tide refugium for crabs.     

Prescribed fire and cutting as tools for reducing woody plant succession in a created salt marsh

This paper reports on efforts to reduce woody successional growth by the native shrub Iva frutescens L. in a created salt marsh by using prescribed fire and cutting. Experimental treatments included a winter burn, cutting plants at ground level, and a combination burn-and-cut treatment, with replicate plots of each. Iva frutescens proved to be extremely hardy, with zero mortality following the cutting, burning, or combination treatment; similar levels of regrowth were observed for all treatments. Individual shrub response, however, was found to be related to initial plant size, ground water level and salinity, and two fire characteristics (total heating >60°C and total heat index >60°C). Fire severity, sediment nutrient concentrations, and other abiotic factors had no observable effects.     

Lacunal allocation and gas transport capacity in the salt marsh grass Spartina alterniflora

Summary Lacunal allocation as the fraction of the total cross sectional area of leaves, stem bases, rhizomes, and roots was determined in both tall and short growth forms of Spartina alterniflora collected from natural monospecific stands. The results indicate that in both growth forms lacunal allocation is greater in stem bases and rhizomes than in leaves and roots and that tall form plants allocate more of their stem and rhizome to lacunae than short form plants.Measurements made in natural stands of Spartina alterniflora suggest that total lacunal area of the stem base increases with increasing stem diameter and that stem diameter increases with increasing plant height and above-ground biomass. However, the fraction of cross section allocated to lacunae was relatively constant and increased only with the formation of a central lacuna.Experimental manipulations of surface and subsurface water exchange were carried out to test the influence of flooding regime on aerenchyma formation. No significant differences in lacunal allocation were detected between plants grown in flooded (reduced) and drained (oxidized) sediments in either laboratory or field experiments. While aerenchyma formation in Spartina alterniflora may be an adaptation to soil waterlogging/anoxia, our results suggest that lacunal formation is maximized as a normal part of development with allocation constrained structurally by the size of plants in highly organic New England and Mid-Atlantic marshes.The cross sectional area of aerenchyma for gas transport was found to be related to the growth of Spartina alterniflora with stands of short form Spartina alterniflora exhibiting a lower specific gas transport capacity (lacunal area per unit below ground biomass) than tall form plants despite having a similar below-ground biomass supported by a 10 fold higher culm density. The increased specific gas transport capacity in tall vs. short plants may provide a new mechanism to explain the better aeration, higher nutrient uptake rates and lower frequency of anaerobic respiration in roots of tall vs. short Spartina alterniflora.     

Oxygen loss from Spartina alterniflora and its relationship to salt marsh oxygen balance

Spartina alterniflora has been reported to lose significant amounts of oxygen to its rhizosphere with potentially important effects on salt-marsh biogeochemical cycling and plant productivity. The potential significance of this oxidative pathway was evaluated using laboratory split-chamber experiments to quantify oxygen loss from intact root systems under a wide variety of pre-treatment and incubation conditions including antibiotics to inhibit microbial respiration. The aerenchyma system of S. alterniflora was found to transport O₂, N₂, Ar, and CH₄ from above-ground sources to its below-ground roots and rhizomes. While non-respiratory gases were observed to move from the lacunae to water bathing the root systems, net O₂ loss did not occur; instead oxygen present outside of the roots/rhizomes was consumed. Net oxygen loss was found when resistance to gas transport was reduced in the lacunae-rhizosphere pathway by placing the root systems in a gas phase and when plant respiration was significantly reduced. Root system respiration appeared to be the major variable in the plant oxygen balance. When root and rhizome respiration was inhibited using poisons or lowered by cooling, the oxygen deficit was greatly reduced and oxygen loss was indicated. The effect of seasonal temperature changes on root system oxygen deficit presents a possible explanation as to why Spartina produces root systems with respiration rates that cannot be supported by gas transport. Overall, while oxygen loss from individual plant roots is likely, integrating measured root system oxygen loss with geochemical data indicates that the mass amount of oxygen lost from S. alterniflora root systems is small compared to the total oxygen balance of vegetated salt marsh sediments.     

Spatial and temporal changes in salt marsh distribution in the Dee estuary,NW England,determined from aerial photographs

Vegetation changes in salt marsh communities of the Dee estuary, northwest England, were analysed with a combination of remote sensing techniques using data dating back to the 1950s. The distribution of communities in 1997 was classified using Airborne Thematic Mapper data and used to develop a methodology for the analysis of black and white photographs of the marsh. These methods were then applied retrogressively to a time sequence of monochrome photographs running from 1955 to 1975. At the apex of the salt marshes on the English shore of the Dee estuary, the marsh expanded dramatically to 1975, and consisted predominantly of pioneer and low marsh vegetation types. Between 1975 and 1997, however, there was only a slight increase in salt marsh area, but with an increase in mid and high marsh vegetation, replacing pioneer marsh. In a second area of the salt marsh on the English shore, a different pattern of salt marsh expansion was observed. The area occupied by marsh continued to increase right up to 1997, with extensive pioneer vegetation suggesting a process of continuing expansion. However, the pattern of marsh colonisation appeared to be different in 1997 compared to 1975. The significance of the changes in salt marsh distribution within the Dee estuary are discussed in relation to the historical pattern of salt marsh colonisation, the importance of Spartina anglica in the process and the implications for strategic management of the estuarine resources.     

Nitrogen resorption from senescing leaves of three salt marsh plant species

Seasonalvariation in leaf nitrogen of mature green and senescent leaves and nitrogenresorption efficiency in three plants (Spartina maritima, Halimioneportulacoides and Arthrocnemum perenne) of aTagus estuary salt marsh are reported. Total nitrogen concentrations in greenand senescent leaves were higher during winter (December and March). Soilinorganic nitrogen availability showed an opposite pattern with higherconcentrations during summer (June and September) when total leaf biomass washigher. Nitrogen resorption efficiency ranged between 31 and 76% andH. portulacoides was the plant that better minimizednitrogen loss by this process. Nitrogen resorption occurred mainly from thesoluble protein pool, although other fractions must have been broken down duringthe resorption process. No significant seasonal variation in nitrogen resorptionefficiency and no relation to leaf total nitrogen or soil nitrogen availabilitywere found. This suggests that the efficiency of the resorption process is notdetermined by the plant nitrogen status nor by the availability of the nutrientin the soil. Nevertheless, resorption from senescing leaves may play animportant role in the nitrogen dynamics of salt marsh plants and reduce thenitrogen requirements for plant growth.     

Variation with depth and season in metal sulfides in salt marsh soils

Seasonal monitoring of metal sulfides was carried out in four soils ofthe Ría de Ortigueira salt marshes. Soils from the high salt marsh (withsuboxic redox conditions at the surface), had low concentrations of ironsulfides (AVS and pyrite fraction) and thus a low degree of trace metalpyritization (DTMP) in surface layers (0–10 cm), butconcentrations of metals associated with the pyrite fraction increasedconsiderably at depth (27.5 cm). In the low salt marsh soils (withanoxic conditions at the surface) maximum concentrations of metal sulfides werefound in the surface layers of soils colonized by Spartina maritima. These results are explained by the double effectexerted by roots in strongly reduced soils. On the one hand, they stimulate theactivity of sulfate-reducing bacteria and on the other, they favour the partialoxidation of the soil, thus generating polysulfides with which Fe²⁺immediately precipitates as pyrite, whereas in the deepest, permanently anoxiclayers, pyrite must be formed in a reaction in which FeS is an intermediate, asfollows: FeS + H₂S FeS₂ + H₂.Concentrations of metal sulfides also varied greatly with the season, with twopatterns being distinguished. In soils colonized by S. maritima in both high and low salt marshes, the lowestconcentrations were found in summer. At this time of the year there is a netloss of metal sulfides throughout the profile, presumably due to physiologicalactivity of plants (evapotranspiration and release of oxygen from roots). Incontrast, maximum concentrations of AVS and pyritic metals were found in thesummer in the low salt marsh soils not colonized by vascular plants (creekbottom). In this case, the higher temperatures increased the activity ofsulfur-reducing bacteria leading to synthesis and accumulation of metalsulfidesin the soil.     

Arbuscular mycorrhizal fungi enhance root cadmium and copper accumulation in the roots of the salt marsh plant Aster tripolium L.

It is known that vegetation plays an important role in the retention of heavy metals in salt marshes by taking up and accumulating the metals. In this study, we investigated whether arbuscular mycorrhizal fungi (AMF) increase Cd and Cu uptake and accumulation in the root system of the salt marsh species Aster tripolium L., and whether indigenous AMF isolated from polluted salt marshes have higher capacity to resist and alleviate metal stress in A. tripolium than isolates of the same species originated from non-polluted sites. Plants inoculated with Glomus geosporum, either isolated from a polluted salt marsh site (PL isolate) or from a non-polluted site (NP isolate), and non-mycorrhizal (NM) plants were compared in a pot experiment at four different Cd and Cu concentrations. Cd had no effect in root colonization, whereas high concentrations of Cu decreased colonization level in plants inoculated with the NP isolate. AM colonization did not increase plant dry weight or P concentration but influenced root Cd and Cu concentrations. Inoculation with PL and NP isolates enhanced root Cd and Cu concentrations, especially at highest metal addition levels, as compared to NM plants, without increasing shoot Cd and Cu concentrations. There was no evidence of intraspecific variation in the effects between AMF isolated from polluted and non-polluted sites, since there were no differences between plants inoculated with PL or NP isolate in any of the tested plant variables. The results of this study showed that AMF enhance metal accumulation in the root system of A. tripolium, suggesting a contribution of AMF to the sink of metals within vegetation in the salt marshes.     

Effects of grazing and inundation on pasture quality and seed production in a salt marsh

During a six-year period, changes in the composition of dominant plant species of importance to foraging birds in a salt marsh on the Swedish west coast were followed inside and outside exclosures to document effects of grazing on herbage quality and seed production. Since marshes provide an important habitat for foraging geese and ducks, it was of interest to determine how cattle grazing would affect herbage production inAgrostis stolonifera andPuccinellia maritima and seed and root-tuber production inScirpus maritimus. Measurements of cover and height in permanent plots revealed that a wetter weather type favouredAgrostis, probably through reduced salinity, at the expense ofPuccinellia, which was the most favoured food of both cattle and birds.Agrostis out-competedPuccinellia when grazing pressure was low. Seed production inScirpus maritimus was reduced by cattle grazing, particularly whenPhragmites australis formed part of the vegetation. In the absence of cattle grazing, both herbage- and seed producing plants were gradually reduced, andPhragmites increased. Since high herbage consumption and high seed production are mutually exclusive, grazing rotation in combination with mowing is suggested as a management strategy.     

Root and rhizome distribution as an indicator of upper salt marsh wetland limits

Four wetland plant species, Deschampsia cespitosa, Distichlis spicata, Grindelia integrifolia, and Salicornia virginica, were tested to determine the effect of soil moisture on the vertical distribution of roots and rhizomes. In an 8-month greenhouse experiment involving plants grown from seeds and rhizomes the occurrence of more than 65% of the root and rhizome mass in the upper 10 cm of soil was indicative of saturated conditions. Roots and rhizomes were more evenly distributed under field capacity and low moisture conditions. The percent of total root and rhizome biomass in the upper 10 cm of a 35 cm core may be useful in determining whether or not a plant is experiencing saturated soil conditions, a criterion often used for defining wetlands. Limited field studies support the conclusion of the greenhouse study.     

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.     

Environmental gradients and herbivore feeding preferences in coastal salt marshes

Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.     

Population dynamics of annual Salicornia species in the tidal salt marshes of the Oosterschelde,The Netherlands

1. The population dynamics of two Salicornia species from the Bergen op Zoom salt marsh (south-west Netherlands) was examined. Based on the results of several field studies three preliminary life tables were constructed, two for S. procumbens agg. populations growing respectively on the mud flats and in the salt marsh, and one for S. europaea agg. living in the upper marsh.
2. The life cycles are described and quantified in terms of eight phases and the transition probabilities between them, starting from a notional individual representative of each population.
3. The models depicting the life cycle of S. procumbens show a mean offspring number of 4.26 individuals per parent for the mud-flat population and 0.18 for the salt-marsh population. The S. europaea model gives an output of 0.44 individuals per parent. These results reflect the fluctuations in population size observed in sample plots over the years 1976–78.
4. Comparison of the transition probabilities reveals that on the mud flats most S. procumbens individuals die during pollination and seed germination, while the population in the salt marsh proper is thinned especially during the seed phase in winter time and during the growth from established seedlings to maturation. S. europaea behaves in a similar but less pronounced way to S. procumbens in the salt marsh.
5. Probabilities for one flower or one seed to produce a mature flowering plant were calculated, and were compared with those found in the literature. They are roughly of the same order of magnitude as the probabilities for other annual species, but much higher than those reported for biennial species.

     

The fish assemblage of the intertidal salt marsh creeks in North Bull Island, Dublin Bay: seasonal and tidal changes in composition, distribution and abundance

An intertidal salt marsh fish assemblage inhabiting two creeks on North Bull Island, Dublin Bay was sampled monthly from June 2000 until May 2002. Water temperature and salinity were recorded in situ and samples were also taken for Suspended Particulate Matter (SPM) and Chlorophyll a. All fish caught were weighed and measured and classified into functional guilds. A total of 6,549 individuals comprising 10 fish species from 10 families were recorded within the two creeks. The community was dominated by a few species, a feature common to other estuarine fish populations. Of the 10 species found, the common goby, Pomatoschistus microps, the 3-spined stickleback, Gasterosteus aculeatus, the thick-lipped grey mullet, Chelon labrosus and the flounder, Platichthys flesus contributed 98.4% of all fish sampled. The fish population of the channels at Bull Island, Dublin, was dominated by the resident gobies (true estuarine resident species), but also hosted juveniles of species such as the bass, Dicentrarchus labrax (marine juvenile migrant species). In turn, the nekton populations were dominated by the brown shrimp, Crangon crangon and the fairy shrimp, Palaemonetes varians especially in winter when fewer fish (numbers of species and abundance) were found. Multivariate analysis of fish diversity and abundance revealed a strong seasonal pattern but there was little evidence of difference between creeks, nor of tidal (spring/neap) effects. The estuarine fish using the intertidal marsh creeks have been little studied in Europe yet they play a major role with the decapods in these habitats. This role needs to be quantified for a proper understanding of the system’s function.     

Decomposition of Spartina anglica,Elytrigia pungens and Halimione portulacoides in a Dutch salt marsh in association with faunal and habitat influences

Decomposition of Spartina anglica, Elytrigia pungens and Halimione portulacoides was studied for 20.5 months in situ in two habitats on a salt marsh in The SW Netherlands. Litter bags of three different mesh sizes were used to exclude meio- and/or macrofauna. The middle-marsh habitat was flooded more frequently than the plant-debris habitat in the highest marsh zone. Decomposition of the three species followed an exponential pattern of decay: instantaneous decay rates varied from 0.0026 to 0.0054 per day. Decay rates were significantly influenced by habitat factors and fauna, while there was a significant interaction between plant species and habitat. In case of a significant meio- and/or macrofauna effect, this became noticeable 12–16 weeks after the start of decomposition and resulted in a difference of 5–10% ash-free dry weight remaining after 20.5 months. Nematodes were the dominant microfaunal group in the plant litter. Densities were influenced by habitat conditions but not by resource quality, season and meio- and/or macrofauna. Only initial C/N and C/P ratios were correlated with differences in decomposition rates between the plant species. During the later stages of decomposition N and P concentrations of the plant litter were higher in the plant-debris habitat than in the middle-marsh habitat, probably as a result of fluctuating detritivores densities. The course of the decomposition process differed per plant species and per habitat. The results of this study underline the importance of knowledge of long-term decomposition rates.     

Consumer pressure and seed set in a salt marsh perennial plant community

Summary Seed predation can be an important determinant of plant success, but has received little attention in wetland plant communities. Here, we examine the role of flower and seed predators in limiting the seed production of the dominant perennial plants in a salt marsh plant community. Of the four perennial investigated, direct ovule loss to consumers ranged from 51 to 80%, resulting in seed set reductions ranging from 50% to over 20-fold. Most losses were due to generalist grazing by the grasshopper, Conocephalus spartinae. More species-specific losses were inflicted by planthoppers, and microlepidopteran and dipteran larval seed parasites.Insect abundance and consumer pressure on flowers and seeds increased over the early summer, peaked in the middle of July, and declined through August, and this temporal pattern was reflected in the natural consumer damage incurred by each of the marsh perennials. Juncus gerardi flowers earlier than other marsh perennials and largely escapes heavy consumer losses. Spartina patens and Distichlis spicata flower in the middle of the summer during the peak consumer activity and incur extremely heavy seed losses. Spartina alterniflora flowers late in the summer as consumer pressure is subsiding, which appears to minimize its seed loss. In addition to destroying seeds directly, consumers also markedly reduce the frequency and affect the timing of sexual expression in these plants. In particular, predation drastically reduces the frequency of male flowers, which could lead to pollen limitation of seed set.Intense flower and seed predation on these marsh perennials may be an important determinant of the success of marsh plant populations as well as a potent selective force on their flowering phenologies and reproductive effort.     

Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

The influence of tidal channels on the distribution of salt marsh plant species in Petaluma Marsh,CA, USA

Tidal channels influence the distribution and composition of salt marsh vegetation in a San Francisco Bay salt marsh. Two channel networks in the Petaluma Marsh, Sonoma County, CA, were mapped and characterized using global positioning and geographic information systems. Plant species abundance was sampled on transects placed perpendicular to and extending away from the channel banks. The vegetation showed significant increases in species richness along channel banks and larger areas of effect which increased approximately linearly with channel size. Composition of species assemblages varies with distance from the channel bank and channel size. These results demonstrate that salt marsh plant assemblages, composed of both major and minor species, are distributed with respect to the channel network in Petaluma Marsh.     

Mobility of metals in salt marsh sediments colonised by <Emphasis Type="Italic">Spartina maritima</Emphasis> (Tagus estuary,Portugal)

Chemical associations of Zn, Pb, Cu, Co and Cd were determined using a sequential extraction procedure in sediments colonised by S. maritima in three salt marshes within the Tagus estuary: Rosário, Corroios and Pancas. Concentrations of these metals were also analysed in above- and belowground parts of Spartina maritima, as well as in sediments colonised by the plant. The highest metal concentrations in sediments were found in the marshes near the industrial and urban areas, whereas metal concentrations in plants were not significantly different among sites. This was thought to be a consequence of differences observed in metal bioavailability: Metals in Pancas, the least polluted location, were largely associated to easily accessible fractions for plant uptake, probably as a result of low organic matter content and high sandy fraction in sediments. S. maritima was able to induce the concentration of metals between its roots in the three salt marshes. The results obtained in this study indicate that S. maritima could be useful to induce phytostabilisation of metals in sediments, although the effectiveness to modify chemical associations is highly dependent on existing sediment parameters, and thus different results could be obtained depending on site characteristics. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

Short-term sedimentation in Tagus estuary,Portugal: the influence of salt marsh plants

Short-term sediment deposition was studied at four salt marsh areas in the Tagus estuary. In areas covered with Sarcocornia perennis, Sarcocornia fruticosa, Halimione portulacoides and Spartina maritima and also in the non-vegetated areas, sedimentation was measured as the monthly accumulation of sediments on nylon filters anchored on the soil surface, from August 2000 to May 2001. Our experiments were used also to determine the influence of the different plant species in vertical accretion rates. Short-term sedimentation rates (from 2.8 to 272.3 g m⁻² d⁻¹) did show significant differences when the four salt marshes studied in the Tagus estuary were compared to each others. Salt marshes closer to the sediment sources had higher sedimentation rates. Our results suggest that the salt marsh type and surface cover may provide small-scale variations in sedimentation and also that sediment deposition values do change according to the position of the different plant species within the salt marsh. Sedimentation is an essential factor in salt marsh vertical accretion studies and our investigation may provide support to help forecast the adaptative response of the Tagus estuary wetlands to future sea level rise.