Quantifying Vegetation and Nekton Response to Tidal Restoration of a New England Salt Marsh

Tidal flow to salt marshes throughout the northeastern United States is often restricted by roads, dikes, impoundments, and inadequately sized culverts or bridge openings, resulting in altered ecological structure and function. In this study we evaluated the response of vegetation and nekton (fishes and decapod crustaceans) to restoration of full tidal flow to a portion of the Sachuest Point salt marsh, Middletown, Rhode Island. A before, after, control, impact study design was used, including evaluations of the tide‐restricted marsh, the same marsh after reintroduction of tidal flow (i.e., tide‐restored marsh), and an unrestricted control marsh. Before tidal restoration vegetation of the 3.7‐ha tide‐restricted marsh was dominated by Phragmites australis and was significantly different from the adjacent 6.3‐ha Spartina‐dominated unrestricted control marsh (analysis of similarities randomization test, p < 0.001). After one growing season vegetation of the tide‐restored marsh had changed from its pre‐restoration condition (analysis of similarities randomization test, p < 0.005). Although not similar to the unrestricted control marsh, Spartina patens and S. alterniflora abundance increased and abundance and height of Phragmites significantly declined, suggesting a convergence toward typical New England salt marsh vegetation. Before restoration shallow water habitat (creeks and pools) of the unrestricted control marsh supported a greater density of nekton compared with the tide‐restricted marsh (analysis of variance, p < 0.001), but after one season of restored tidal flow nekton density was equivalent. A similar trend was documented for nekton species richness. Nekton density and species richness from marsh surface samples were similar between the tide‐restored marsh and unrestricted control marsh. Fundulus heteroclitus and Palaemonetes pugio were the numerically dominant fish and decapod species in all sampled habitats. This study provides an example of a quantitative approach for assessing the response of vegetation and nekton to tidal restoration.     

Early Responses of Fishes and Crustaceans to Restoration of a Tidally Restricted New England Salt Marsh

Nekton (fishes and decapod crustaceans) is an abundant and productive faunal component of salt marshes, yet nekton responses to tidal manipulations of New England salt marshes remain unclear. This study examined nekton use of a tidally restricted salt marsh in Narragansett, Rhode Island relative to an unrestricted marsh during summer. In addition, a before‐after‐control‐impact design was used to examine early responses of nekton to the reintroduction of natural tidal flushing. Species richness and densities of Cyprinodon variegatus, Lucania parva, Menidia beryllina, and Palaemonetes pugio were higher in the restricted marsh compared with the unrestricted marsh. The unrestricted marsh supported higher densities of Menidia menidia and Fundulus majalis. Mean lengths of Carcinus maenas and P. pugio were greater in the restricted marsh. Tidal restoration resulted in increased tidal flushing, salinity, and water depth in the restricted marsh. Densities of Fundulus heteroclitus, F. majalis, and Callinectes sapidus were higher after 2 years of restoration. Density of L. parva decreased after restoration, probably in response to a loss of macroalgal habitat. Species richness also decreased after 2 years, from 20.9 species when the marsh was restricted to 13.0 species. Total nekton density did not change with restoration, but shifts in community composition were evident. In this study restoration induced rapid changes in the composition, density, size, and distribution of nekton species, but additional monitoring is necessary to quantify longer‐term effects of salt marsh restoration on nekton.     

Isolation of a New Polysaccharide-Digesting Bacterium from a Salt Marsh

A new marine bacterium that digested a variety of storage and structural polysaccharides, including agar, was isolated. Strain 2-40 is a nonfermentative gram-negative, polarly flagellated rod that sometimes grew as a filamentous helix and secreted a melaninlike pigment. Its characteristics conform to those of no previously described species.     

Survivorship and Spatial Development of Spartina alterniflora Loisel. (Gramineae) Seedlings in a New England Salt Marsh

Patterns of survival and spatial arrangement of tillers of Spartinaalterniflora were examined in natural and in artificially producedbare areas, and in pure stands of adult S. alterniflora in aNew England salt marsh. Seedling growth and survival were highin naturally occurring bare patches and in artificial patches,whether created by continual clipping of adult plants to groundlevel throughout the growing season or by providing bare substrateafter removal of adult plants. Seedling growth and survivorshipincreased with increasing size of bare area. S. alternifloraseedlings were also common in areas dominated by adult plants,but no seedlings survived a whole first growing season underthe mature canopy, probably because of competition from adultplants. In large bare areas, S. alterniflora seedlings grew non-directionally,reaching heights of 0.5 m, and producing as many as 36 tillersin one growing season. Examination of leaf area ratios suggestedthat the production of tillers increased photosynthetic capacityin seedlings with several tillers in contrast to seedlings withouttillers. Thus, seedlings can apparently tiller and colonizefree space radially without a loss of photosynthetic capacity. These results suggest that while seedling success is generallylimited by adult plants in monocultures of S. alterniflora,in disturbed spaces seedling success is high and results ina rapid non-directional proliferation of sexually generatedclones that ultimately preclude the successful invasion of futureseedlings. Clones, directional growth, leaf area ratio, salt marsh cordgrass, seedling survivorship, Spartina alterniflora Loisel     

Molluscan Community Recovery in a New England Back‐Barrier Salt Marsh Lagoon 10 Years after Partial Restoration

Like many Eastern U.S. salt marshes, East Harbor salt marsh lagoon on Cape Cod was isolated from tidal flow in the 1800s, resulting in near‐freshwater conditions and loss of native salt marsh species. After its partial restoration in 2002, a variety of marine and estuarine fauna recolonized East Harbor, and soft shell clam (Mya arenaria) recolonization was particularly prolific. The goal of our study was to evaluate molluscan community composition, density, and distribution at regular intervals for 10 years following restoration, and to relate molluscan community recovery to various physical properties at the site. In 2007, 2008, and 2011, we sampled mollusks at several points across East Harbor, and we also recorded water salinity and temperature, particle size distribution, and submerged aquatic vegetation density. In 2007 and 2008, we encountered 12 and 11 mollusk species, respectively; M. arenaria was the most abundant species in 2007 and the second most abundant species in 2008. In 2011, we encountered eight mollusk species and M. arenaria was the most abundant species. Mollusk species richness declined from 12 to 8 species between 2008 and 2011. Our results show that mollusk species richness and density have declined significantly since the first few years following restoration; related studies attribute this to high summer water temperatures in the Main Lagoon and severe macroalgal blooms during 2005–2006. This suggests that East Harbor is still equilibrating to baseline conditions and that full tidal restoration may be necessary to sustain a diverse mollusk community at East Harbor.     

New England Salt Marsh Recovery: Opportunistic Colonization of an Invasive Species and Its Non-Consumptive Effects

Predator depletion on Cape Cod (USA) has released the herbivorous crab Sesarma reticulatum from predator control leading to the loss of cordgrass from salt marsh creek banks. After more than three decades of die-off, cordgrass is recovering at heavily damaged sites coincident with the invasion of green crabs ( Carcinusmaenas ) into intertidal Sesarma burrows. We hypothesized that Carcinus is dependent on Sesarma burrows for refuge from physical and biotic stress in the salt marsh intertidal and reduces Sesarma functional density and herbivory through consumptive and non-consumptive effects, mediated by both visual and olfactory cues. Our results reveal that in the intertidal zone of New England salt marshes, Carcinus are burrow dependent, Carcinus reduce Sesarma functional density and herbivory in die-off areas and Sesarma exhibit a generic avoidance response to large, predatory crustaceans. These results support recent suggestions that invasive Carcinus are playing a role in the recovery of New England salt marshes and assertions that invasive species can play positive roles outside of their native ranges.     

Physiological Diversity of the Rhizosphere Diazotroph Assemblages of Selected Salt Marsh Grasses

Rhizosphere diazotroph assemblages of salt marsh grasses are thought to be influenced by host plant species and by a number of porewater geochemical parameters. Several geochemical variables can adversely affect plant productivity and spatial distributions, resulting in strong zonation of plant species and growth forms. This geochemically induced stress may also influence the species compositions and distributions of rhizosphere diazotroph assemblages, but little is currently known about these organisms. The diversity and key physiological features of culturable, O₂-tolerant rhizosphere diazotrophs associated with the tall and short growth forms of Spartina alterniflora and with Juncus roemerianus were examined. A total of 339 gram-negative strains were isolated by a root stab culture approach and morphologically and physiologically characterized by using API and BIOLOG tests. Eighty-six distinct groups composed of physiologically similar strains were identified. Of these groups, 72% were shown to be capable of N₂ fixation through molecular analyses, and a representative strain was chosen from each diazotroph group for further characterization. Cluster and principal-components analysis of BIOLOG data allowed the designation of physiologically distinct strain groupings. Most of these groups were dominated by strains that were not identifiable to species on the basis of API or BIOLOG testing. Representatives of several families including the Enterobacteriaceae, Vibrionaceae, Azotobacteraceae, Spirillaceae, Pseudomonadaceae, and Rhizobiaceae were recovered, as well as strains with no clear taxonomic affiliations. This study identifies numerous potentially important physiological groups of the salt marsh diazotroph assemblage.     

Populations of Methanogenic Bacteria in a Georgia Salt Marsh

Methanogens represented about 0.5% of the total bacteria in sediments from a Georgia salt marsh in which Spartina alterniflora is the predominant vegetation. The population of methanogens was composed of at least two groups of nearly equal size. One group was represented by cocci which were able to utilize trimethylamine and were unable to use H₂ or acetate. The second group was composed of two subgroups which were able to utilize H₂ but were unable to use trimethylamine or acetate. The more common subgroup included rod- or plate-shaped methanogens which could utilize isopropanol in addition to H₂ and formate. The second subgroup included Methanococcus maripaludis, which utilized only H₂ and formate. Other groups of methanogens were also present, including Methanosarcina sp. which utilized acetate, H₂, and methylamines. In addition to the overall variability in the types of methanogens, the numbers of methanogens in sediments also exhibited significant spatial variability both within and between tall- and short-Spartina zones.     

Physiological Integration Ameliorates Negative Effects of Drought Stress in the Clonal Herb Fragaria orientalis

Clonal growth allows plants to spread horizontally and to establish ramets in sites of contrasting resource status. If ramets remain physiologically integrated, clones in heterogeneous environments can act as cooperative systems - effects of stress on one ramet can be ameliorated by another connected ramet inhabiting benign conditions. But little is known about the effects of patch contrast on physiological integration of clonal plants and no study has addressed its effects on physiological traits like osmolytes, reactive oxygen intermediates and antioxidant enzymes. We examined the effect of physiological integration on survival, growth and stress indicators such as osmolytes, reactive oxygen intermediates (ROIs) and antioxidant enzymes in a clonal plant, Fragaria orientalis, growing in homogenous and heterogeneous environments differing in patch contrast of water availability (1 homogeneous (no contrast) group; 2 low contrast group; 3 high contrast group). Drought stress markedly reduced the survival and growth of the severed ramets of F. orientalis, especially in high contrast treatments. Support from a ramet growing in benign patch considerably reduced drought stress and enhanced growth of ramets in dry patches. The larger the contrast between water availability, the larger the amount of support the depending ramet received from the supporting one. This support strongly affected the growth of the supporting ramet, but not to an extent to cause increase in stress indicators. We also found indication of costs related to maintenance of physiological connection between ramets. Thus, the net benefit of physiological integration depends on the environment and integration between ramets of F. orientalis could be advantageous only in heterogeneous conditions with a high contrast.     

Responses of Plant Communities to Incremental Hydrologic Restoration of a Tide-Restricted Salt Marsh in Southern New England (Massachusetts, U.S.A.)

Hydrologic restoration of Hatches Harbor, a tide-restricted marsh on Cape Cod (Massachusetts), has resulted in significant plant community changes 7 years following the reintroduction of seawater. Since 1999, incremental increases in flow through a tide-restricting dike have facilitated the rapid decline of salt-intolerant vegetation, while encouraging the expansion of native salt marsh taxa. These changes show strong spatial gradients and are correlated with marsh surface elevation, distance from the point of seawater entry, and porewater salinity. Common reed ( Phragmites australis ) has not decreased in abundance but has migrated a considerable distance upslope. In the wake of this retreat native halophytes have proliferated. Now that maximum flow through the existing dike structure has been reached, continued recovery may be limited less by changing physicochemical conditions and more by rates of growth, seed dispersal, and seed germination of salt marsh taxa.     

间隔子长度对美丽箬竹克隆分株水分生理整合效应的影响

克隆植物分株间隔子长度(即相邻分株间距离)可能对生理整合产生显著的影响,其抗氧化系统与光合色素的变化可能影响植物对水分异质性的响应能力。该研究以克隆植物美丽箬竹(Indocalamus decorus)为材料,设高[H,(90±5)%]、中[M,(60±5)%]和低[L,(30±5)%]3个基质相对含水率和2个水势梯度(H-L和M-L),测定了3种间隔子长度(10、25和40cm)根状茎相连克隆分株叶片抗氧化酶活性、相对电导率以及丙二醛(MDA)、可溶性蛋白质和光合色素含量,分析美丽箬竹水分生理整合的水分梯度效应、间隔子长度效应与时间效应。结果显示:(1)随处理时间的延长,分株叶片CAT活性、相对电导率及MDA含量均先升高后降低,而叶片SOD活性则与之相反;叶片POD活性和光合色素含量总体上呈"N"型变化,而可溶性蛋白质含量的变化趋势与POD活性呈倒"N"型。(2)水分梯度和间隔子长度均会对美丽箬竹分株间水分生理整合产生影响,随分株间隔子长度增大和水势梯度差减小,低水势受体分株叶片抗氧化酶活性、可溶性蛋白质含量和光合色素含量明显降低,而其相对电导率和丙二醛(MDA)含量明显升高;供体分株水势越高,分株间间隔子越长,供体分株叶片抗氧化酶活性、可溶性蛋白质和光合色素含量越高,而其相对电导率和MDA含量明显降低。研究表明,在异质水分环境下美丽箬竹克隆系统发生了从高水势向低水势分株的水分转移,分株间水势梯度差越大、间隔子长度越短,水分传输的整合强度越高,低水势受体分株获益越明显;随着处理时间的延长,分株间水分生理整合强度在处理前期增强,而在处理后期减弱,反映出供体分株与受体分株间损耗-受益在时间序列上是有变化的,且处理前期损耗-受益表现更为明显。     

The Roles of Competition and Disturbance in a Marine Invasion

Two hypotheses for the decline of native species are the superior exploitation of disturbance by exotic species and the competitive displacement of native species by their exotic counterparts. Theory predicts that functional similarity will increase the intensity of competition between native and invasive species. Ecologically important “foundation” species, Zostera marina and other seagrasses have globally declined during the past century. This study used transplant and vegetation removal experiments to test the hypotheses that disturbance and competitive interactions with an invasive congener (Z. japonica) are contributing to the decline of native Z. marina in the northeastern Pacific. Interspecific competition reduced Z. marina and Z. japonica above-ground biomass by 44 and 96%, respectively, relative to intraspecific competition. Disturbance substantially enhanced Z. japonica productivity and fitness, and concomitantly decreased Z. marina performance, effects that persisted two years following substratum disturbance. These results demonstrate that disturbance and competitive interactions with Z. japonica reduce Z. marina performance, and suggest that Z. japonica’s success as an invasive species stems dually from its ability to persist in competition with Z. marina and its positive response to disturbance. These results highlight the importance of understanding the interconnected roles of species interactions and disturbance in the decline of seagrass habitats, and provide a rationale for amending conservation policy in Washington State. In the interest of conserving native eelgrass populations, the current policy of protecting both native and invasive Zostera spp. should be refined to differentiate between native and invader, and to rescind the protection of invasive eelgrass.     

Populations of Anaerobic Phototrophic Bacteria in a Spartina alterniflora Salt Marsh

Habitat-simulating media were used with the Hungate anaerobic roll tube technique to enumerate culturable anaerobic photosynthetic bacteria in sediment, tidal waters, and Spartina alterniflora plant samples collected from the salt marsh at Sapelo Island, Ga. No phototrophs were detected in samples of creekside (low marsh) sediment or in tidal waters in creekside regions. In the high marsh region, 90% of anaerobic phototrophic bacteria occurred in the top 5 mm of sediment and none were detected below 6 mm. There was a seasonal variation, with maximal populations occurring in summer and fall (mean, 4.4 × 10⁵ phototrophs g of dry sediment⁻¹) and minimal numbers occurring in winter (mean, 3.9 × 10³ phototrophs g of dry sediment⁻¹). During winter and late spring, phototrophs had a patchy distribution over the high marsh sediment surface. In contrast, during late summer they had a random uniform distribution. Tidal water collected over high marsh sediment contained an average of 8.7 × 10² phototrophs ml⁻¹, with no significant seasonal variation. Anaerobic phototrophic bacteria were also cultured from the lower stem tissue of S. alterniflora growing in both the high (4.3 × 10⁴ phototrophs g of dry tissue⁻¹) and creekside (4.9 × 10⁴ phototrophs g of dry tissue⁻¹) marsh regions. Chromatium buderi, Chromatium vinosum, Thiospirillum sanguineum, Rhodospirillum molischianum, and Chlorobium phaeobacteroides were the predominant anaerobic phototrophic species cultured from high marsh sediment. The two Chromatium species were dominant.     

Accelerating the Restoration of Vegetation in a Southern California Salt Marsh

Re-establishing plant cover is essential for restoring ecosystem functions, but revegetation can be difficult in severe sites, such as salt marshes that experience hypersalinity and sedimentation. We tested three treatments (adding tidal creeks, planting seedlings in tight clusters, and rototilling kelp compost into the soil) in a site that was excavated to reinstate tidal flows and restore salt marsh. The magnitude of responses was the reverse of expectations, with tidal creeks having the least effect and kelp compost the most. On the marsh plain, kelp compost significantly increased soil organic matter (by 17% at 0–5 cm; p = 0.026 and 11.5% at 5–20 cm; p = 0.083), total Kjeldahl nitrogen (45% at 5–8 cm; p < 0.001) and inorganic nitrogen (35% at 5–8 cm; p < 0.006), and decreased bulk density (16% at 0–5 cm; p < 0.001 and 21% at 5–8 cm depth; p < 0.001) compared to control plots. Survivorship of kelp compost treated plantings increased, along with growth (> 50% increase in a growth index at 20 months after planting; p < 0.0001). In Spartina foliosa plots, kelp compost did not affect soil organic matter, but plants were taller (by ~11 cm; p = 0.003) and denser (47% more stems; p = 0.003). Planting seedlings 10-cm apart in tight clusters on the marsh plain increased survivorship by 18% (compared to 90-cm apart in loose clusters; p = 0.053), but not growth. Tidal creek networks increased survivorship of Batis maritima and Jaumea carnosa by ≥20% (p = 0.060 and 0.077, respectively). Kelp compost had a strong, positive influence on vegetation establishment by ameliorating some of the abiotic stress.     

Tracer Analysis of Methanogenesis in Salt Marsh Soils

Differences in paths of carbon flow have been found in soils of the tall (TS) and short (SS) Spartina alterniflora marshes of Sapelo Island, Ga. Gaseous end products of [U-¹⁴C]glucose metabolism were ¹⁴CO₂ and ¹⁴CH₄ in the SS region and primarily ¹⁴CO₂ in the TS region. Sulfate concentration did not demonstrably affect glucose catabolism or the distribution of end products in either zone. [U-¹⁴C]acetate was converted to ¹⁴CO₂ and ¹⁴CH₄ in the SS soils and almost exclusively to ¹⁴CO₂ in the TS soils. Sulfate concentration did not affect acetate metabolism in the SS soils; however, a noticeable effect of sulfate dilution was seen in TS soils. Sulfate dilution in TS samples resulted in increased methane formation. Total glucose and acetate metabolism were similar in TS and SS soils despite differences in end products. A microbial community characterized by fermentative/sulfate-reducing processes has developed in TS soils as opposed to the fermentative/methanogenic/sulfate-reducing community found in SS soils.     

Diversity of Ascomycete Laccase Gene Sequences in a Southeastern US Salt Marsh

The diversity of ascomycete laccase sequences was surveyed in a southeastern US salt marsh using a degenerate primer set designed around copper binding sites conserved in fungal laccases. This gene was targeted for diversity analysis because of its potential function in lignin degradation in the salt marsh ecosystem and because few studies have assessed functional gene diversity in natural fungal communities. Laccase sequences were amplified from genomic DNA extracted from 24 isolates (representing 10 ascomycete species) cultured from decaying blades of Spartina alterniflora, and from DNA extracted directly from the decaying blades. Among the ascomycete isolates, 21 yielded a PCR product of expected size (˜900 bp) that was tentatively identified as laccase based on sequence similarities to previously published laccase sequences from related organisms. Overall, 13 distinct sequence types, containing 39 distinct sequences, were identified among the isolates, with several species yielding multiple distinct laccase types. PCR amplifications from early and late decay blades of S. alterniflora yielded seven laccase types. Of these, five were composed of sequences >96% similar at the amino acid level to sequences from three cultured ascomycetes previously found to be dominant members of the fungal communities on decaying S. alterniflora blades. Two of the laccase types from the natural-decay clone library were novel and did not match any of the sequences obtained from the cultured ascomycetes. The 39 distinct sequences and 15 distinct laccase sequence types retrieved from the S. alterniflora decay system demonstrate high sequence diversity of this functional gene in a natural fungal community.     

Seasonal Variability and Transport of Suspended Microfungi in a Southeastern Salt Marsh

Tidally induced fluctuations and transports of microfungi were investigated. Samples were collected at three depths from three stations positioned at a transect in a large salt marsh creek. Samples were taken every 1.5 h for 50 consecutive h during neap tides and 50 consecutive h during the corresponding spring tides. In each season, microfungi concentrations fluctuated out of phase with the tides during both neap and spring tides. Mean concentrations of suspended microfungi did not vary appreciably throughout the year. Fungi were exported from the marsh during the majority of the tidal cycles studied. The results suggest that microfungi may serve as indicators of water mass movements.     

Clonal Integration Enhances the Performance of a Clonal Plant Species under Soil Alkalinity Stress

Clonal plants have been shown to successfully survive in stressful environments, including salinity stress, drought and depleted nutrients through clonal integration between original and subsequent ramets. However, relatively little is known about whether clonal integration can enhance the performance of clonal plants under alkalinity stress. We investigated the effect of clonal integration on the performance of a typical rhizomatous clonal plant, Leymus chinensis, using a factorial experimental design with four levels of alkalinity and two levels of rhizome connection treatments, connected (allowing integration) and severed (preventing integration). Clonal integration was estimated by comparing physiological and biomass features between the rhizome-connected and rhizome-severed treatments. We found that rhizome-connected treatment increased the biomass, height and leaf water potential of subsequent ramets at highly alkalinity treatments but did not affect them at low alkalinity treatments. However, rhizome-connected treatment decreased the root biomass of subsequent ramets and did not influence the photosynthetic rates of subsequent ramets. The biomass of original ramets was reduced by rhizome-connected treatment at the highest alkalinity level. These results suggest that clonal integration can increase the performance of clonal plants under alkalinity stress. Rhizome-connected plants showed dramatically increased survival of buds with negative effects on root weight, indicating that clonal integration influenced the resource allocation pattern of clonal plants. A cost-benefit analysis based on biomass measures showed that original and subsequent ramets significantly benefited from clonal integration in highly alkalinity stress, indicating that clonal integration is an important adaptive strategy by which clonal plants could survive in local alkalinity soil.