Nutrients, competition and plant zonation in a New England salt marsh

1 We examined the effects of nutrient availability on the competitive interactions of the New England salt marsh perennials that occupy discrete vegetational zones parallel to the shoreline.
2 Fertilized and unfertilized plots of pair-wise mixtures and monocultures of Spartina alterniflora, S. patens and Juncus gerardi were compared in order to assess the effects of nutrient addition on the competitive dynamics of these species in the field. In addition, we examined competition between some of these species and Distichlis spicata , a species common to disturbed marsh habitats.
3 After two growing seasons, changes in above-ground biomass of the species indicated that in fertilized plots, S. alterniflora outcompeted S. patens, S. patens outcompeted J. gerardi, and D. spicata outcompeted both J. gerardi and S. patens. This was the reverse of the interactions seen under ambient marsh conditions, and suggested that, under conditions of nutrient limitation, competitive dominance may result from efficient competition for nutrients.
4 Using a conceptual model of salt marsh zonation as a function of competition, physical stress and nutrient limitation, we hypothesize that a nutrient-induced reversal in the competitive dynamics among salt marsh perennials may result in modification of the pattern of plant zonation in this and similar marshes.     

Alteration of salt marsh plant community composition by grazing snow geese

A notable omission from wetlands ecology has been the study of the influence of herbivores on vegetation. Reported here are the effects of grazing by snow geese Anser caerulescens altantica on the vegetation of salt marshes along the mid-Atlantic coast of the United States, Exclosures were used to compare total and species percent cover between grazed and ungrazed areas in three marshes (Salt Flats, South Pond, and Bodie Island) with differing vegetation communities from 1978 to 1980.
Spartina alterniflora was reduced by ⅔ in grazed versus ungrazed areas of Salt Flats, S. S. patens was reduced by ½ in grazed portions of South Pond but recovered when grazing ceased. In grazed portions of Bodie Island total plant cover was reduced by 16%. Scirpus robustus and S. patens reacted in opposite ways to grazing pressure with S. robustus increasing and S. patens decreasing, Elocharis was found only in grazed areas of Bodie Island. Echinochloa crusgali appeared in grazed portions of this marsh in 1978 but decreased in abundance during subsequent years. Scirpus americanus was unaffected by grazing, maintaining a nearly constant percent cover in grazed and ungrazed areas at Bodie Island.
Differences in responses to grazing are discussed in terms of each species' growth and reproductive strategies as tempered by the physical and biological environment within each marsh.     

Crab herbivory regulates re‐colonization of disturbed patches in a southwestern Atlantic salt marsh

Recent work exploring the effects of physical stress and herbivory on secondary succession in estuarine plant communities agrees with basic stress models and reveal that herbivory is an important force in brackish and oligohaline marshes but negligible in physically stressful salt marshes. In these systems, herbivores are terrestrial, and thus negatively affected by the same stressful factors that affect marsh plants (i.e. frequent flooding or high salinities). We evaluated the effects of a marine herbivore (i.e. the crab Neohelice granulata) on plant secondary succession in a southwestern Atlantic salt marsh. Field surveys revealed that disturbance‐generated bare patches have harsh physical conditions and that their edges suffer higher herbivore pressure compared to the marsh matrix. A factorial experiment demonstrated that asexual expansion of the surrounding plants is the only possible mechanism to re‐colonize disturbed patches and that crab exclusion can increase this colonization rate by more than 30 times. Our results show that even in highly stressful environments, herbivores strongly impact marsh structure by regulating patch recovery. The synergism of physical stress and herbivory may make plant succession an extremely slow process and lead to the prevalence of bare areas.     

Vegetation, substrate and hydrology in floating marshes in the Mississippi river delta plain wetlands, USA

In the 1940s extensive floating marshes (locally called flotant) were reported and mapped in coastal wetlands of the Mississippi River Delta Plain. These floating marshes included large areas of Panicum hemitomon-dominated freshwater marshes, and Spartina patens/Scirpus olneyi brackish marshes. Today these marshes appear to be quite different in extent and type. We describe five floating habitats and one non-floating, quaking habitat based on differences in buoyancy dynamics (timing and degree of floating), substrate characteristics, and dominant vegetation. All floating marshes have low bulk density, organic substrates. Nearly all are fresh marshes. Panicum hemitomon floating marshes presently occur within the general regions that were reported in the 1940's by O'Neil, but are reduced in extent. Some of the former Panicum hemitomon marshes have been replaced by seasonally or variably floating marshes dominated, or co-dominated by Sagittaria lancifolia or Eleocharis baldwinii.     

Emergent interactions influence functional traits and success of dune building ecosystem engineers

Aims Dune building processes are affected by interactions between the growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and sea spray. Research investigating how species interactions influence dune community structure and functional trait responses in high abiotic stress environments is minimal. We investigated how species interactions influence the functional trait responses of three dominant dune grasses to common abiotic stressors.Methods We performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata Fern., Uniola paniculata L. and Spartina patens Muhl.) in different interspecific combinations, using sand burial and sea spray as abiotic stressors. Sand burial was applied once at the beginning of the study. Sea spray was applied three times per week using a calibrated spray bottle. Morphological functional trait measurements (leaf elongation, maximum root length, aboveground biomass and belowground biomass) were collected at the end of the study. The experiment continued from May 2015 to August 2015.Important findings Species interactions between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata, indicating that interactions with U. paniculata could alter dune community structure. Furthermore, A. breviligulata had a negative interaction with S. patens, which decreased S. patens functional trait responses to abiotic stress. When all species occurred together, the interactions among species brought about coexistence of all three species. Our data suggest that species interactions can change traditional functional trait responses of dominant species to abiotic stress.     

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.     

Effects of flooding, salinity and herbivory on coastal plant communities, Louisiana, United States

Flooding and salinity stress are predicted to increase in coastal Louisiana as relative sea level rise (RSLR) continues in the Gulf of Mexico region. Although wetland plant species are adapted to these stressors, questions persist as to how marshes may respond to changed abiotic variables caused by RSLR, and how herbivory by native and non-native mammals may affect this response. The effects of altered flooding and salinity on coastal marsh communities were examined in two field experiments that simultaneously manipulated herbivore pressure. Marsh sods subjected to increased or decreased flooding (by lowering or raising sods, respectively), and increased or decreased salinity (by reciprocally transplanting sods between a brackish and fresh marsh), were monitored inside and outside mammalian herbivore exclosures for three growing seasons. Increased flooding stress reduced species numbers and biomass; alleviating flooding stress did not significantly alter species numbers while community biomass increased. Increased salinity reduced species numbers and biomass, more so if herbivores were present. Decreasing salinity had an unexpected effect: herbivores selectively consumed plants transplanted from the higher-salinity site. In plots protected from herbivory, decreased salinity had little effect on species numbers or biomass, but community composition changed. Overall, herbivore pressure further reduced species richness and biomass under conditions of increased flooding and increased salinity, supporting other findings that coastal marsh species can tolerate increasingly stressful conditions unless another factor, e.g., herbivory, is also present. Also, species dropped out of more stressful treatments much faster than they were added when stresses were alleviated, likely due to restrictions on dispersal. The rate at which plant communities will shift as a result of changed abiotic variables will determine if marshes remain viable when subjected to RSLR. Received: 8 April 1998 / Accepted: 15 June 1998     

Relationship of soil hydrogen sulfide level to net carbon assimilation ofPanicum hemitomon andSpartina patens

Panicum hemitomon Schult andSpartina patens (Ait) Muhl. plants from Louisiana Gulf Coast fresh and brackish marshes were subjected to hydrogen sulfide under controlled sediment redox conditions. Net carbon assimilation responses of both species to the combined sediment anaerobiosis and hydrogen sulfide concentrations was measured.Panicum hemitomon was more sensitive to hydrogen sulfide as compared toSpartina patens. Initiation of reduction in net carbon assimilation inP. hemitomon began when H₂S concentrations of soil solution exceeded 0.22 mgl^-1. Reductions in net carbon assimilation inS. patens were also noted at H₂S concentrations exceeding 0.34 mgl^-1. The reduction in net carbon assimilation of both species measured at elevated H₂S concentrations suggests that extreme anaerobiosis and elevated sulfide could contribute to the growth reduction of these species under certain conditions. However based on H₂S concentration in fresh and brackish marsh soil profiles, levels were too low to cause any adverse effects ofPanicum hemitomon. In brackish marsh soils containing hydrogen sulfide of 3.4 mgl^-1 in soil solution, sulfide could be a major factor limiting growth ofS. patens.     

Response of two oligohaline marsh communities to lethal and nonlethal disturbance

Severity is recognized as an important attribute of disturbance in many plant communities. However, the effects of disturbances of different severity on patterns of regeneration in oligohaline marsh vegetation have not been experimentally examined. In these communities, a critical difference in the effects of disturbance severity may be whether the vegetation dies as a result of the disturbance or is merely damaged and hence capable of resprouting. We described the regeneration of vegetation in two Louisiana marsh community types, one dominated by Sagittaria lancifolia L. and the other by Spartina patens (Ait.) Muhl., following three levels of disturbance: no disturbance, a nonlethal disturbance, and a lethal disturbance. In the nonlethal disturbance, aboveground vegetation was clipped to simulate common disturbances such as fire and herbivory that remove aboveground vegetation but leave rhizomes intact. In the lethal disturbance vegetation was killed using herbicide to simulate disturbances causing plant mortality such as wrack deposition, sedimentation, scouring, and flooding following fire or herbivory. Regeneration was assessed over a 2-year period by measuring plant species richness, relative abundance, relative dominance, cover, and final biomass. To elucidate mechanisms for observed responses of vegetation, the species composition of the seed bank, light penetration, water level, salinity, and soil redox potential were evaluated. Despite differences in the structure of undisturbed vegetation in the two community types, they exhibited the same overall pattern of regeneration. Following nonlethal disturbance, the dominant species resprouted and quickly reestablished the structure of the vegetation. In contrast, recolonization following lethal disturbance occurred primarily via seedling recruitment, which resulted in marked shifts in community structure that persisted throughout the study. While the two communities responded similarly overall to disturbance, the response of individual species was not uniform; abundance, dominance, biomass, or cover increased for some species but decreased for others in response to disturbance. Seed bank species occurred in the vegetation following lethal disturbance in the Spartina community and in both disturbed and undisturbed plots in the Sagittaria community, indicating that the seed bank is a source of propagules for regeneration and maintenance of oligohaline marshes. Of the environmental variables measured, light level was most closely related to the effect of disturbance severity on community structure. Our results suggest that lethal and nonlethal disturbances have differential effects on regeneration of vegetation that can create pattern in oligohaline marshes communities. Received: 29 September 1997 / Accepted: 12 May 1998     

Vegetation change from chronic stress events: Detection of the effects of tide gate removal and long‐term drought on a tidal marsh

Question: Chronic stress events are defined as disturbance events that exceed the lifespan of the dominant plant species, fluctuate in intensity and lack abruptness or physical destruction of biomass. Can the effects of chronic stress events be measured on vegetation communities? Did two chronic stress events, the removal of a tide gate and a four year drought, cause a temporary or permanent shift in the vegetation communities of a tidal marsh? Location: Tidal marsh in southeastern United States. Methods: Change in species composition and dominance and community change on a landscape level salinity gradient were measured between time periods ranging from four months to seven years to construct a statistical baseline reference community at freshwater, oligohaline, and mesohaline sections of a tidal marsh. Statistical shifts in the plant community were defined as changes in the plant community that fell outside of the defined baseline reference community. Results: Plant community changes outside of the reference community occurred in 13 out of 378 community comparisons. Removal of the tide gate had a greater effect on interstitial salinity levels than the drought and was most intense in the oligohaline marsh, where between 20 to 45% of the freshwa‐ter/oligohaline community types permanently converted to oligohaline community types. However, community shifts in the freshwater and oligohaline marsh induced by the drought were temporary, lasting from 1 to 3+ years. Neither chronic stress event permanently altered the mesohaline plant communities. Conclusion: The effects of chronic stress events could be detected; an extended historical record of vegetation change (18 years) was necessary to identify community shifts outside of a reference condition of the community and to determine if those shifts were permanent or temporary.     

Sediments in Marsh Ponds of the Gulf Coast Chenier Plain: Effects of Structural Marsh Management and Salinity

Physical characteristics of sediments in coastal marsh ponds (flooded zones of marsh associated with little vegetation) have important ecological consequences because they determine compositions of benthic invertebrate communities, which in turn influence compositions of waterbird communities. Sediments in marsh ponds of the Gulf Coast Chenier Plain potentially are affected by (1) structural marsh management (levees, water control structures and impoundments; SMM), and (2) variation in salinity. Based on available literature concerning effects of SMM on sediments in emergent plant zones (zones of marsh occasionally flooded and associated with dense vegetation) of coastal marshes, we predicted that SMM would increase sediment carbon content and sediment hardness, and decrease oxygen penetration (O₂ depth) and the silt-clay fraction in marsh pond sediments. Assuming that freshwater marshes are more productive than are saline marshes, we also predicted that sediments of impounded freshwater marsh ponds would contain more carbon than those of impounded oligohaline and mesohaline marsh ponds, whereas C:N ratio, sediment hardness, silt-clay fraction, and O₂ depth would be similar among pond types. Accordingly, we measured sediment variables within ponds of impounded and unimpounded marshes on Rockefeller State Wildlife Refuge, near Grand Chenier, Louisiana. To test the above predictions, we compared sediment variables (1) between ponds of impounded (IM) and unimpounded mesohaline marshes (UM), and (2) among ponds of impounded freshwater (IF), oligohaline (IO), and mesohaline (IM) marshes. An a priori multivariate analysis of variance (MANOVA) contrast indicated that sediments differed between IM and UM marsh ponds. As predicted, the silt-clay fraction and O₂ depth were lower and carbon content, C:N ratio, and sediment hardness were higher in IM than in UM marsh ponds. An a priori MANOVA contrast also indicated that sediments differed among IF, IO, and IM marsh ponds. As predicted, carbon content was higher in IF marsh ponds than in ponds of other impounded marsh types. In contrast to our predictions, C:N ratio and sediment hardness were lowest and silt-clay fraction and O₂ depth were highest in IO and IM marsh ponds. Our results indicated that SMM has affected physical properties of sediments in coastal marsh ponds. Moreover, sediments in IF marsh ponds were affected more so than were those in IO and IM marsh ponds. Our results, in conjunction with those of previous studies, indicated that sediments of marsh ponds and emergent plant zones differed greatly. We predict that changes in pond sediments due to SMM will promote greater epifaunal macroinvertebrate biomass, which in turn should attract larger populations of wintering waterbirds. However, waterbirds that filter or probe soft sediments may be negatively affected by SMM because of the expected decrease in infaunal invertebrate biomass.     

The dynamics of bottom–up and top–down control in a New England salt marsh

Traditionally, salt marsh ecosystems were thought to be controlled exclusively by bottom–up processes. Recently, this paradigm has shifted to include top–down control as an additional primary factor regulating salt‐marsh community structure. The most recent research on consumer impacts in southern US marshes has shown that top–down forces often interact with biotic and abiotic factors, such as secondary fungal infection in grazer‐induced wounds, soil nutrients and climatic variation, to influence ecosystem structure. In a more northern salt marsh, located in New England, we examined the separate and interactive effects of nutrient availability, insect herbivory and secondary fungal infection, on growth of the foundation species, Spartina alterniflora. We used a factorial design with two levels of nutrients (control and addition) insects (control and removal) and fungi (control and removal). Nutrient addition increased plant biomass by 131% in the absence of herbivores. When insect consumers were allowed access to fertilized plots, biomass was reduced by nearly 45% when compared with treatments with nutrients and insecticide. In contrast, insect herbivores did not affect plant biomass in unfertilized control treatments. These differences suggest that consumer effects are triggered under high nutrient levels only. We also found that secondary fungal infections in grazer‐induced wounds, in contrast to lower latitude marshes, did not significantly impact primary production. Our results suggest that while New England salt marshes may typically be under bottom–up control, eutrophication can trigger dual control with inclusion of top–down regulation. However, unlike lower latitude marshes, consumer control of plant growth in northern US salt marshes is not dependent on herbivores facilitating fungal infections that then control grass growth, suggesting that the intensity of disease mediated top–down control by small grazers may be regulated by climate and/or grazer identity that co‐vary with latitude.     

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

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

Interactive effects of elevation and burial with wrack on plant community structure in some Rhode Island salt marshes

1 The effects of disturbances and elevation on marsh plant communities were examined using experimental disturbances along an elevation gradient in marshes with different disturbance histories. In addition, differences in species composition among five marshes were determined at elevations at which the greatest concentration of burial by wrack occurred.
2 Experimental wrack burial generally caused significant mortality of the high-marsh competitive dominants, Juncus gerardi and Spartina patens , and strongly increased the abundance of the fugitive perennial, Distichlis spicata .
3 The effects of experimental wrack burial interacted strongly with abiotic factors associated with elevation to influence the distributions of both competitive dominants and annual fugitive plants.
4 Frequent wrack burial in a marsh appears to lead to a persistent assemblage of plants dominated by competitively subordinate fugitives. This assemblage of fugitives tends to occur at intermediate elevations within the marsh, where wrack gets stranded for long periods of time and where the resistance of Juncus gerardi to wrack burial is lowest.
5 We suggest that wrack-burial disturbances interact strongly with marsh elevation to influence the zonation of plants in New England salt marshes, and discuss some implications of our results.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Experimental warming causes rapid loss of plant diversity in New England salt marshes

Anthropogenic climate change is predicted to cause widespread biodiversity loss due to shifts in species' distributions, but these predictions rarely incorporate ecological associations such as zonation. Here, we predict the decline of a diverse assemblage of mid-latitude salt marsh plants, based on an ecosystem warming experiment. In New England salt marshes, a guild of halophytic forbs occupies stressful, waterlogged pannes. At three sites, experimental warming of < 4 °C led to diversity declines in pannes and rapid takeover by a competitive dominant, Spartina patens . In Rhode Island, near their southern range limit, pannes were more sensitive to warming than farther north, and panne area also declined in control plots over the three-season experiment. These results suggest that warming will rapidly reduce plant diversity in New England salt marshes by eliminating a high diversity zone. Biodiversity in zoned ecosystems may be more affected by climate-driven shifts in zonation than by individual species' distribution shifts.     

Joint control by rodent herbivory and nutrient availability of plant diversity in a salt marsh–salty steppe transition zone

Questions: Do current models that predict shifting effects of herbivores on plant diversity with varying nutrient conditions apply to stressful systems like salt marshes? Do herbivores affect different components of the diversity as nutrient availability varies? Location: Salt marsh–salty steppe transition zone at the SW Atlantic Mar Chiquita coastal lagoon (37°44′52″S, 57°26′6″W), Argentina. Methods: We experimentally evaluated the separate and interactive effect of nutrients and rodent (Cavia aperea) herbivory, using exclosures and applying fertilizer (mostly nitrogen), following a factorial design in 50 cm × 50 cm plots. Results: We found a negative effect of herbivory on diversity in the resource‐poor scenario (due to a reduction in species richness), but a positive effect when nutrients were added, by reducing the abundance of the dominant plant (and hence increasing evenness). Conclusions: Our experimental results contribute to the limited factorial evidence evaluating the role of nutrients and herbivory on the diversity of terrestrial plant communities, even in highly stressful environments like salt marsh–salty steppe transition zones. Our results also support the model that predicts negative effects of herbivores on plant diversity in low‐nutrient conditions and positive effects in nutrient‐enriched scenarios, and also support the mechanism assumed to act in these situations.     

Testing the role of biotic stress in the stress gradient hypothesis. Processes and patterns in arid rangelands

Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.     

Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

Herbaceous Forage and Selection Patterns by Ungulates across Varying Herbivore Assemblages in a South African Savanna

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.