Positive interactions of the smooth cordgrass Spartina alterniflora on the mud snail Heleobia australis, in South Western Atlantic salt marshes

The role of positive interactions is often crucial in communities with intense abiotic stress such as intertidal environments. Grasses acting as ecosystem engineers, for example, may ameliorate intertidal harsh physical conditions and modify the community structure. The mud snails Heleobia australis d'Orbigny frequently inhabit the SW Atlantic marshes, mainly associated to intertidal marsh plants (mainly the smooth cordgrass Spartina alterniflora Loisel) probably due to the plant indirect effects. The purpose of this work was to investigate the magnitude of these association and the processes that generate the pattern. Samples of the snail abundance in six SW Atlantic coastal marshes show that H. australis is associated to coastal areas of low energy and low or none freshwater input. This result is important because this species is being used as bioindicator of coastal estuarine systems during the Holocene. Thus the paleontological interpretation based on this species should be revised. Within the studied areas, snails are associated to intertidal marsh plants. However, stable isotope analysis shows that neither plant nor their epiphytes are their main food sources. Field experiments show that snails actively select areas with plants, although tethering experiments show that plants do not provide shelter from predators. However, plants do buffer physical stress factors such as temperature, which generate important mortality outside plants covered areas. These positive interactions have large effects on H. australis distributions in marsh communities; increasing the habitats available for colonization and affecting their local distribution.     

Abiotic stress mediates top-down and bottom-up control in a Southwestern Atlantic salt marsh

Increasing evidence has shown that nutrients and consumers interact to control primary productivity in natural systems, but how abiotic stress affects this interaction is unclear. Moreover, while herbivores can strongly impact zonation patterns in a variety of systems, there are few examples of this in salt marshes. We evaluated the effect of nutrients and herbivores on the productivity and distribution of the cordgrass Spartina densiflora along an intertidal stress gradient, in a Southwestern Atlantic salt marsh. We characterized abiotic stresses (salinity, ammonium concentration, and anoxia) and manipulated nutrients and the presence of the herbivorous crab Neohelice (Chasmagnathus) granulata, at different tidal heights with a factorial experiment. Abiotic stress increased at both ends of the tidal gradient. Salinity and anoxia were highest at the upper and lower edge of the intertidal, respectively. Nutrients and herbivory interacted to control cordgrass biomass, but their relative importance varied with environmental context. Herbivory increased at lower tidal heights to the point that cordgrass transplants onto bare mud substrate were entirely consumed unless crabs were excluded, while nutrients were most important where abiotic stress was reduced. Our results show how the impact of herbivores and nutrients on plant productivity can be dependent on environmental conditions and that the lower intertidal limits of marsh plants can be controlled by herbivory.     

Predictions of vegetation change following fire in a great salt lake marsh

A Gleasonian model of succession (van der Valk, 1981) was used to predict the species composition of a freshwater marsh following fire. Seed bank samples from 5 vegetation types (Distichlis spicata (L.) Greene, Scirpus lacustris L., S. maritimus L., Typha spp., open water) were used in conjuction with plant life history characteristics to make predictions of post-fire species composition for each vegetation type and the entire study area. In general, predictions for specific vegetation types were not good, but were more accurate for the entire area. Stronge correlations for specific vegetation types may have been prevented by considering marsh vegetation types as discrete units, statistical sampling problems, competitive interactions, or more specific germination requirements of some species.     

土著菌根真菌和混生植物对羊草生长和磷营养的影响

植物种间相互作用直接影响植物生长、根系可塑性及养分吸收,而与植物共生的丛枝菌根真菌可以改变植物个体和种间养分资源的分配,具有协调种间竞争的潜力.以我国北方草甸草原建群种羊草(Leymus chinensis)和混生植物紫花苜蓿(Medicago sativa)及独行菜(Lepidium spetalum)为供试植物,通过模拟盆栽试验,研究了土著菌根真菌和混生植物对羊草生长、根系形态及磷营养的影响.试验结果表明,土著菌根真菌能够与羊草及紫花苜蓿形成良好共生,而独行菜根内基本未形成菌根共生结构.土著菌根真菌显著降低了羊草及独行菜的生物量,但促进了紫花苜蓿的生长;混种紫花苜蓿显著促进了羊草的生长,而混种独行菜则显著抑制了羊草的生长.土著菌根真菌对羊草根系形态的影响表现出与植株生物量类似的趋势,但不同混生植物对羊草根系生长均无显著影响.土著菌根真菌和混生植物对羊草植株磷含量均无显著影响.与混生植物相比,羊草具有较高的比根长和磷吸收能力,这也解释了其负向菌根依赖性.研究证实了菌根真菌和植物种间相互作用均是影响草原优势植物生长和根系发育的重要因素,深入研究其交互作用对于科学管理草地生态系统,维持植物群落的稳定性和生态系统生产力具有重要意义.     

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.     

Plant diversity changes and succession along resource availability and disturbance gradients in Kamchatka

Changes in plant species richness across environmental and temporal gradients have often been explained by the intermediate disturbance hypothesis and a unimodal diversity–productivity relationship. We tested these predictions using two sets of mountain plant communities assembled along postglacial successional and snow depth (disturbance and stress) gradients in maritime Kamchatka. In each community, we counted the number of species in plots of increasing sizes (0.0025–100 m²) and analyzed them using species–area curves fitted by the Arrhenius power function and the Gleason logarithmic function. A comparison of successional communities along a 270-year-old moraine chronosequence behind the receding Koryto Glacier—representing gradients of increasing productivity and resource competition—confirmed the unimodal species richness pattern. The plant diversity peaked in a 60–80-year-old Salix–Alnus stand where light availability was sufficient to sustain a rich understory combining pioneer and late successional herbs. The closed Alnus canopy on older moraines caused a pronounced decrease in species richness for all plot sizes (interactive stage 80–120 years since deglaciation). A slight increase in species richness in the oldest assortative stages (120–270 years), when Alnus stands are mature, was found only at the smaller spatial scales. This reflects (i) the consolidation of clonal understory dominants and (ii) the absence of other woody species such as Betula ermanii whose invasion would eliminate Alnus and increase diversity at larger spatial scales. A comparative study of major mountain plant communities distributed above the Koryto Glacier foreland did not confirm the highest species richness at intermediate levels of disturbance and stress. Contrary to our expectation, the species richness was highest in alpine tundra and snowbed communities, which are subjected to severe winter frost and a short summer season, while less disturbed communities of subalpine meadows, heaths, and Betula ermanii woods were less species-rich. We attribute this pattern to differences in habitat area and species pool size.     

Salt tolerance and stress level affect plant biomass–density relationships and neighbor effects

It has been shown that plant biomass–density relationships are altered under extreme or stressed conditions. We do not know whether variation in biomass–density relationships is a direct result of stress tolerance or occurs via changes in plant–plant interactions. Here, we evaluated biomass–density relationships and neighbor effects in six plant species that differ in salt tolerance in a salt marsh, and conducted a literature review of biomass–density relationship under higher and lower stress levels. Our field study showed that both neighbor effects and the exponent of the biomass–density relationship (α) varied among plant species with different degrees of salt tolerance. There was a positive relationship between neighbor effects (measured as relative interaction index) and α-value among the tested species. The literature review showed that α and its variation increased under higher stress. Our results indicate that plant species with different salinity tolerance differ in the direction and strength of neighbor effects, resulting in variation in biomass–density relationships. Our results support the hypothesis that differences in biomass–density relationships among species are not due to differences in stress tolerance alone, they are mediated by changes in plant–plant interactions.     

基于胁迫梯度假说和互惠理论的海三棱藨草种群恢复技术

滨海湿地生态修复已成为阻止海岸带生态系统退化、保护生物多样性以及提供生态服务的关键措施。以长江口原生盐沼植物海三棱藨草(Scirpus mariqueter)为研究对象,选取崇明东滩新生滩涂湿地为研究区域,通过沿潮滩高程梯度的海三棱藨草植株斑块的移植实验,探究胁迫梯度假说和互惠理论(即种内的正相互作用)对长江口海三棱藨草种群恢复的指导意义。研究结果显示:(1)在一定的胁迫梯度范围内(潮滩高程2.0 m以上),增大种植斑块可以促进海三棱藨草的种内正相互作用,显著提高种植斑块的存活率和植株密度(P0.05);(2)潮滩水文动力沉积条件与潮滩高程梯度密切相关(P0.05),水文动力沉积作用对海三棱藨草定居和生长的胁迫随高程梯度下降而增强。潮滩高程2.0 m以下处强烈的水文动力条件干扰限制了生物-物理因素的正反馈作用。滨海湿地盐沼植被修复工作的成功率可以通过改进种植方式,增强种内的正相互作用得到极大的提高。研究可为开展大规模滨海湿地盐沼植被修复工程和提高生态修复效率提供科学依据和技术支持。     

Patterns in the utilization of the intertidal salt marsh by larvae and juveniles of Fundulus heteroclitus (Linnaeus) and Fundulus luciae (Baird)

Larval and juvenile fishes were collected at low tide from the surface of an intertidal salt marsh on Sapelo Island, Georgia every 6 days from 25 May through 20 December 1982. Larval fishes were present in shallow puddles of tidal water on the marsh from the beginning of the sampling period until the end of October. Juveniles were present throughout the year. Fundulus heteroclitus (Linnaeus) and F. luciae (Baird) accounted for 96.3% (67.0% and 29.3%, respectively) of the 4355 fishes collected. These fishes exhibited synchronous, temporal pulses in larval abundance, suggesting that reproduction was discontinuous and controlled by the same factor(s) in both species. Larval F. heteroclitus inhabited aquatic micro-habitats near the upland edge of the intertidal marsh, but as the larvae grew to juvenile size (≈ 10 mm standard length) they moved to lower elevations near a tidal creek. Large juveniles and adults of F. heteroclitus infrequently occurred in the samples, presumably because they leave the intertidal marsh as the tide ebbs. F. luciae were rarely found in low marsh areas, but all age classes (including adults) occurred at higher elevations, supporting previous suggestions that this species prefers high marsh habitats. The vegetated, intertidal salt marsh appears to be the principal nursery habitat for both of these cyprinodontid species.     

Effects of light and intertidal position on seedling survival and growth in tropical tidal forests

Reciprocal transplant experiments were used to study the effects of tidal inundation and light level on growth and survival of four species of mangroves in Australia: Avicennia marina (Forsk.) Vierh., Bruguiera gymnorrhiza (L.) Lam., Ceriops tagal C.T. White, and Rhizophora stylosa Griff. Seedlings were planted in the high or low intertidal and in light gaps or under the shade of a closed canopy. Survival and growth of the seedlings were monitored for 30–36 months. Significant differences in survival were found among species, between intertidal zones and due to light level. Averaged across intertidal zones and light level, survival was greatest for Rhizophora and decreased in the order Ceriops, Avicennia, and Bruguiera. For all species survival was greater (P ⩽ 0.001) in the high than in the low intertidal treatment, regardless of light level. Within the high intertidal all species survived better in light gaps than under the canopy. Relative growth of Rhizophora stylosa, Avicennia marina, and Ceriops tagal were greater in the high versus low intertidal and in gaps versus under the canopy. For Bruguiera gymnorrhiza growth was not significantly different between gap and canopy or high and low intertidal. Although Avicennia marina, Bruguiera gymnorrhiza, and Rhizophora stylosa survived and/or grew best in the high intertidal they reach maximum abundance in the low intertidal. Ceriops tagal, however, performed best in the region where it is most abundant, yet even there, it was out-performed by the former three species. These results indicate that the species zonation patterns often observed across the intertidal cannot be explained by physiological adaptation alone. Factors such as propagule dispersal, competition and predation on propagules may also be important.     

Inundation,Vegetation, and Sediment Effects on Litter Decomposition in Pacific Coast Tidal Marshes

The cycling and sequestration of carbon are important ecosystem functions of estuarine wetlands that may be affected by climate change. We conducted experiments across a latitudinal and climate gradient of tidal marshes in the northeast Pacific to evaluate the effects of climate- and vegetation-related factors on litter decomposition. We manipulated tidal exposure and litter type in experimental mesocosms at two sites and used variation across marsh landscapes at seven sites to test for relationships between decomposition and marsh elevation, soil temperature, vegetation composition, litter quality, and sediment organic content. A greater than tenfold increase in manipulated tidal inundation resulted in small increases in decomposition of roots and rhizomes of two species, but no significant change in decay rates of shoots of three other species. In contrast, across the latitudinal gradient, decomposition rates of Salicornia pacifica litter were greater in high marsh than in low marsh. Rates were not correlated with sediment temperature or organic content, but were associated with plant assemblage structure including above-ground cover, species composition, and species richness. Decomposition rates also varied by litter type; at two sites in the Pacific Northwest, the grasses Deschampsia cespitosa and Distichlis spicata decomposed more slowly than the forb S. pacifica. Our data suggest that elevation gradients and vegetation structure in tidal marshes both affect rates of litter decay, potentially leading to complex spatial patterns in sediment carbon dynamics. Climate change may thus have direct effects on rates of decomposition through increased inundation from sea-level rise and indirect effects through changing plant community composition.     

Annual plant–shrub interactions along an aridity gradient

Annual plants in semi-arid and arid areas are often closely associated with shrubs. The degree of association largely depends on the balance of negative and positive effects between these contrasting plant life-forms, ranging from interference to facilitation. Since positive interactions are predicted to become less important with increasing rainfall, the interaction balance is expected to shift along aridity gradients. However, this prediction has not been tested on a community level and for different life-history stages across large geographical gradients. Here, we show such changes for annual plant populations and communities in four contrasting sites along a steep climatic gradient, ranging from the arid desert to mesic Mediterranean regions in Israel. Above-ground productivity, richness, seedling density, and seed bank density of the annual plant community, as well as fecundity of annual plant populations, were generally higher under shrubs than in areas between shrubs at the arid end of the gradient, but significantly lower at the humid end. Net effects of shrubs on annuals expressed as relative interaction intensity indicated a steady and consistent shift from net positive or neutral effects in the desert to net negative effects in the mesic part of the gradient. These findings emphasize the usefulness of studies along large-scale gradients encompassing a wide range of environmental conditions for understanding community level interactions among coexisting species.     

Respiratory enzyme activities correlate with anoxia tolerance in salt marsh grasses

Salt marsh communities are known for well-defined species zonation patterns. Lower limits of plant growth are thought to be set by an ability to tolerate anoxic sediments, but the physiological differences between species have not previously been examined. To investigate responses to anoxic sediments, several estuarine species were grown in greenhouse experiments to compare how respiratory processes were affected by flooding. Metabolic characteristics related to respiration and anoxia tolerance were studied in the emergent estuarine species Spartina alterniflora, S. anglica, S. densiflora, S. foliosa, S. alterniflora × S. foliosa hybrids, S. patens, and Distichlis spicata and compared to the inland species maize (Zea mays). All species showed a strong ability to respire anaerobically, indicating flooding tolerance. High intertidal marsh species had significantly higher root aerobic respiration enzyme activities compared to low intertidal species that may suggest lower aerobic demand in low marsh species. Some higher marsh species showed an apparent high sensitivity to sulfide that may be related to high cytochrome c oxidase activities. In contrast, the low marsh species S. alterniflora and S. anglica had lower aerobic respiration enzyme activities and a lower sensitivity to sulfide. Thus differences in aerobic demand and sulfide sensitivity may influence estuarine species zonation.     

Changes in marsh nekton communities along the salinity gradient of the Schelde river, Belgium and The Netherlands

Nekton was sampled in five marshes along the salinity gradient of the Schelde River. The utilisation of three different habitats (large and small creek, marsh pond) by fish and macrocrustacean species was compared among the five sampling sites. In the larger channels fyke nets were deployed to capture fish and macrocrustaceans leaving the marsh at ebb while block nets were set in smaller intertidal creeks. Fish traps passively sampled fish and shrimp in the marsh ponds. The tidal freshwater marsh had a species poor fauna and only a low number of fish was caught. Besides some freshwater species (Alburnoides bipunctatus, Carassius carassius) the European eel, Anguilla anguilla was still present. The four other marshes had a similar community structure although Platichthys flesus was absent from the euhaline area. Among fish species, dominance of Dicentrarchus labrax, Platichthys flesus and Pomatoschistus microps was observed. Carcinus maenas and Palaemonetes varians were the most abundant macrocrustacean species in every marsh. Between the large and small intertidal creeks there was no difference in nekton species composition. The main species used both habitats. Marsh ponds were utilized intensively only by two species, Pomatoschistus microps and Palaemonetes varians in every marsh.     

Role of biotic interactions and physical factors in determining the distribution of marsh species along an estuarine salinity gradient

Understanding the mechanisms that shape plant distribution patterns is a major goal in ecology. We investigated the role of biotic interactions (competition and facilitation) and abiotic factors in creating horizontal plant zonation along salinity gradients in the Elbe estuary. We conducted reciprocal transplant experiments with four dominant species from salt and tidal freshwater marshes at two tidal elevations. Ten individuals of each species were transplanted as sods to the opposing marsh type and within their native marsh (two sites each). Transplants were placed at the centre of 9‐m² plots along a line parallel to the river bank. In order to disentangle abiotic and biotic influences, we set up plots with and without neighbouring vegetation, resulting in five replicates per site. Freshwater species (Bolboschoenus maritimus and Phragmites australis) transplanted to salt marshes performed poorly regardless of whether neighbouring vegetation was present or not, although 50–70% of the transplants did survive. Growth of Phragmites transplants was impaired also by competition in freshwater marshes. Salt marsh species (Spartina anglica and Puccinellia maritima) had extremely low biomass when transplanted to freshwater marshes and 80–100% died in the presence of neighbours. Without neighbours, biomass of salt marsh species in freshwater marshes was similar to or higher than that in salt marshes. Our results indicate that salt marsh species are precluded from freshwater marshes by competition, whereas freshwater species are excluded from salt marshes by physical stress. Thus, our study provides the first experimental evidence from a European estuary for the general theory that species boundaries along environmental gradients are determined by physical factors towards the harsh end and by competitive ability towards the benign end of the gradient. We generally found no significant impact of competition in salt marshes, indicating a shift in the importance of competition along the estuarine gradient.     

Nurse plant effects on plant species richness in drylands: The role of grazing,rainfall and species specificity

The outcome of plant–plant interactions depends on environmental (e.g. grazing and climatic conditions) and species-specific attributes (e.g. life strategy and dispersal mode of the species involved). However, the joint effects of such factors on pairwise plant–plant interactions, and how they modulate the role of these interactions at the community level, have not been addressed before. We assessed how these species-specific (life strategy and dispersal) and environmental (grazing and rainfall) factors affected the co-occurrence of 681 plant species pairs on open woodlands in south-eastern Australia. Species-specific attributes affected the co-occurrence of most species pairs, with higher co-occurrence levels dominating for drought-intolerant species. The dispersal mechanism only affected drought-tolerant beneficiaries, with more positive co-occurrences for vertebrate-dispersed species. Conversely, the percentage of facilitated species at the community scale declined under higher rainfall availabilities. A significant grazing × rainfall interaction on the percentage of facilitated species suggests that grazing-mediated protection was important under low to moderate, but not high, rainfall availabilities. This study improves our ability to predict changes in plant–plant interactions along environmental gradients, and their effect on community species richness, by highlighting that: (1) species-specific factors were more important than environmental conditions as drivers of a large amount (～37%) of the pairwise co-occurrences evaluated; (2) grazing and rainfall interaction drive the co-occurrence among different species in the studied communities, and (3) the effect of nurse plants on plant species richness will depend on the relative dominance of particular dispersal mechanisms or life strategies prone to be facilitated.     

Does the intensive grazing and aridity change the relations between the dominant shrub Artemisia kopetdaghensis and plants under its canopies?

The interspecific plant interactions along grazing and aridity stress gradients represent a major research issue in plant ecology. However, the combined effects of these two factors on plant–plant interactions have been poorly studied in the northeast of Iran. To fill this knowledge gap, 144 plots were established in 12 study sites with different grazing intensities (high vs. low) and climatic characteristics (arid vs. semiarid) in northeastern Iran. A dominant shrub, Artemisia kopetdaghensis, was selected as the model species. Further, we studied changes in plant life strategies along the combined grazing and aridity stress gradients. In this study, we used relative interaction indices calculated for species richness, Shannon diversity, and species cover to determine plant–plant interactions using linear mixed‐effect models (LMM). The indicator species analysis was used to identify the indicator species for the undercanopy of shrub and for the adjacent open areas. The combined effects of grazing and aridity affected the plant–plant interactions and plant life strategies (CSR) of indicator species. A. kopetdaghensis showed the highest facilitation effect under high stress conditions (high grazing, high aridity), which turned into competition under the low stress conditions (low grazing, low aridity). In the arid region, the canopy of the shrub protected ruderals, annual forbs, and grasses in both high and low grazing intensities. In the semiarid region and high grazing intensity (low aridity/high grazing), the shrubs protected mostly perennial forbs with C‐strategy. Our findings highlight the importance of context‐dependent shrub management to restore the vegetation damaged by the intensive grazing.     

The effects of crude oil on the growth of Spartinaalterniflora Loisel. and Spartina cynosuroides (L.) Roth

Greenhouse studies were conducted to examine the effects of crude oil on the growth of Spartinaalterniflora Loisel. and S. cynosuroides (L.) Roth from North Carolina. The way in which crude oil came into contact with the plant tissue and/or substratum was an important factor in determining the responses of both species to oil pollution. Plants recovered from a single application of oil to aerial tissue with relatively little impact on productivity. The presence of an oil layer on the surface of an overlying layer of water had little impact on existing aerial portions of S. alterniflora plants; however, regrowth following harvest was completely inhibited. Incorporation of oil into the substratum significantly reduced aerial productivity and regrowth of S. alternflora and S. cynosuroides. Observations suggest that decreased productivity and regrowth may have been caused by decreased root and rhizome growth. Regrowth potential of S. alterniflora grown in oiled substratum was greater in fine-textured marsh substratum than in sand substratum.     

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.