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Studies of competition in mangroves are mostly limited to seedlings and artificial settings like forestry projects. We conducted the first experimental study of intraspecific competition among adult mangroves in a natural mangrove forest to examine how important competition is in determining tree size compared with abiotic conditions. We conducted a study near Port Aransas, TX, USA, which is near the geographical limit of mangroves and dominated by monospecific stands of ‘scrub’ form black mangroves, Avicennia germinans. We thinned 10 plots to create a gradient of mangrove cover, and quantified the effects of mangrove cover on the growth of tagged mangroves from 2013 to 2019, and the mangrove canopy height in 2019. The relative growth rate of tagged mangroves declined as mangrove cover increased, and plants in the plot with 100% mangrove cover did not grow, indicating that they had attained their maximum size. In plots with reduced mangrove cover, plant height increased sharply, with plants in the plot with 11% mangrove cover growing ~52% taller over 6 years. Canopy height was ~30% taller in the plot fringe than in the interior, and canopy height in both fringe and interior declined as mangrove cover increased. Measures of leaf chlorophyll concentration and light interception suggested that plants were primarily limited by nitrogen. Our results showed that scrub mangroves compete strongly despite being limited by abiotic conditions, and that the importance of competition was greater in magnitude than that of abiotic differences between the fringe and interior.

1. Global variation in litter decomposition rates is driven by climate, decomposer taxa and litter quality. Most large-scale studies of litter quality have made comparisons across species, rather than within a species sourced from different latitudes. Here, we aim to explore latitudinal variation in leaf litter decomposition rate and litter quality within a species. 2. We investigated whether the typical interspecific pattern of increased litter lability at higher latitudes also holds within plant species, by comparing leaf litter decomposition rates and litter traits from stands of native Phragmites australis and exotic Spartina alterniflora in Chinese coastal wetlands spanning 20 degrees of latitude using parallel greenhouse and field transplant experiments. 3. We found that leaf litter from high-latitude plants decomposed 46.6 +/- 5.1% faster than that from low-latitude conspecifics, with similar patterns in both plant species in both decomposition experiments. Higher lability of leaves from high latitudes was associated with lower levels of plant defensive compounds (phenolics and tannins) and higher levels of nutrients (Ca, Mg and Na). Litter from the exotic S. alterniflora decomposed faster than litter from the native P. australis, being consistent with lower defences and higher nutrient concentrations in S. alterniflora litter. 4. Our results show that individual species follow the same pattern of increasing litter lability at higher latitudes previously reported in cross-species geographical comparisons. Moreover, this pattern can develop rapidly (

Ecologists use multiple methods for studying interspecific competition, but different approaches may give different answers. We compared four methods to quantify the competitive interactions between Avicennia germinans (black mangroves) and salt marsh vegetation in Texas, USA: two different methods of sampling a large (24 x 42 m) mangrove removal experiment, a transplant experiment conducted within the large experiment, and an observational study comparing sites naturally dominated by marsh or mangrove vegetation. As expected, mangroves strongly suppressed the cover and biomass of salt marsh plants. But our understanding of the strength of these interactions varied depending on the study method used, the plant species studied, and the spatial scale considered; different processes influencing community assembly. The transplant experiment isolated the effects of competition driven only by the presence or absence of mangroves in the immediate (3x3 m plot) vicinity of the transplanted plants. In contrast, the observational study examined the combined effects of dispersal, abiotic suitability and competition as a function of the cover of mangroves at the larger plot scale. Combining the approaches in areas with levels of mangrove cover varying from 45% to 97% provided insight into how results from the local scale could be extrapolated to the landscape. Although mangroves may compete strongly with neighboring marsh plants growing in their immediate vicinity, marsh plants may not be totally eradicated from sites colonized by mangroves, but instead may persist on the landscape at low densities in stressful habitats that offer a refuge from competition.

Herbivores are important drivers in both above‐ and below‐ground ecosystems. Despite their importance, studies of herbivores often focus on the effects of individual species, potentially missing the effects of herbivore diversity. Here we examine how orthopteran herbivore species and functional (i.e. diet) diversity interact with nutrient availability to shape above‐ and below‐ground ecosystems. We stocked six species of grasshoppers (Orthoptera) with two different feeding modes (i.e. grass‐only or grass‐ and forb‐feeders) at varying taxonomic compositions (0, 1 or 6 species) in field enclosures and measured their effects on above‐ground plant biomass, chemistry and richness, as well as below‐ground microbial community composition and function. We treated half the cages with nitrogen fertilizer to examine how herbivore effects may be mitigated by soil nutrient availability. Above‐ground, we predicted that grasshoppers would decrease plant biomass and richness, and increase the %N of the plant community. We also predicted that high diversity cages would have the largest decreases in plant biomass and richness, but these effects would be offset by fertilizer additions. Below‐ground, we predicted that herbivore taxonomic composition and diet would differentially shift soil microbial community composition and function. We also predicted that fertilization would decrease the soil pH and increase nutrient availability creating an interaction with herbivores that leads to shifts below‐ground community composition. We found that grasshoppers decreased plant biomass and richness, and this effect was not dependent on nutrient additions. Contrary to our predictions, we did not see the largest above‐ground changes in high diversity cages. Below‐ground, herbivore diet and fertilization, but not their interaction, shifted bacterial communities. Grasshopper taxonomic composition did not influence bacterial communities. Grasshopper taxonomic composition and diet interacted with fertilizer to increase below‐ground levels of bioavailable C and microbial biomass. Fungal communities did not respond to any treatments. The differing effects of herbivore diet versus taxonomic composition highlight the need for studies that examine multiple diversity metrics when exploring herbivore‐meditated effects on above‐ and below‐ground ecosystems. Combined, our results suggest that bottom‐up and top‐down controls are important factors to consider when studying the composition and function of grassland ecosystems. A free Plain Language Summary can be found within the Supporting Information of this article.

AIM: Introduced species may display or foster novel latitudinal clines because they are not well adapted to their new habitats. We tested the hypothesis that the latitudinal cline in nematode diversity in salt marshes would differ between the native (United States) and introduced (China) ranges of Spartina alterniflora. LOCATION: East Coasts of the United States (30.32–43.33°N) and China (30.36–39.14°N). METHODS: We extracted nematodes from soil samples collected at 32 sites along the United States East Coast and 41 sites along the Chinese coast. We compared latitudinal patterns in nematode diversity and composition between the native and introduced ranges. RESULTS: In the native range of S. alterniflora, nematode richness at lower latitudes was almost twice as high as that at higher latitudes. In contrast, we found no latitudinal pattern in nematode richness or diversity in the introduced range of S. alterniflora. Nematode genus richness at all sites in China was about half that at lower latitudes in the United States. Beta diversity of nematodes increased with geographic distance in the United States, but not China. MAIN CONCLUSIONS: Nematode diversity did not show latitudinal clines in salt marshes dominated by introduced S. alterniflora in China. A likely explanation is that the recently introduced populations are still relatively genetically homogenous, whereas in the native range, genetic variation in plant populations across latitude drives different nematode communities. We suggest that future studies of introduced species will gain additional insights by taking an explicitly geographic perspective.

As global change alters the composition and productivity of ecosystems, the importance of subsidies from one habitat to another may change. We experimentally manipulated black mangrove (Avicennia germinans) cover in 10 large plots and over a 5-year period (2014-2019) quantifying the effects of mangrove cover on subsidies of floating organic material (wrack) into coastal wetlands. As mangrove cover increased from 0% to 100%, wrack cover and thickness decreased by ~60%, the distance that wrack penetrated into the plots decreased by ~70%, and the percentage of the wrack trapped in the first 6 m of the plot tripled. These patterns observed during 4 "normal" years disappeared in a fifth year following Hurricane Harvey (2017), when large quantities of wrack were pushed far into the interior of all the plots, regardless of mangrove cover. Prior to the storm, the abundance of animals collected in grab samples increased with wrack biomass. Wrack composition did not affect animal abundance or composition. Experimental outplants of two types of wrack (red algae and seagrass) revealed that animal abundance and species composition varied between the fringe and interior of the plots, and between microhabitats dominated by salt marsh versus mangrove vegetation. The importance of wrack to overall carbon stocks varied as a function of autochthonous productivity: wrack inputs (per m

Biological invasions offer model systems of contemporary evolution. We examined trait differences and evolution across geographic clines among continents of the intertidal grass Spartina alterniflora within its invasive and native ranges. We sampled vegetative and reproductive traits in the field at 20 sites over 20° latitude in China (invasive range) and 28 sites over 17° in the US (native range). We grew both Chinese and US plants in a glasshouse common garden for 3yr. Chinese plants were c.15% taller, c. 10% denser, and set up to four times more seed than US plants in both the field and common garden. The common garden experiments showed a striking genetic cline of seven-fold greater seed set at higher latitudes in the introduced but not the native range. By contrast, there was a slight genetic cline in some vegetative traits in the native but not the introduced range. Our results are consistent with others showing that introduced plants can evolve rapidly in the new range. S.alterniflora has evolved different trait clines in the native and introduced ranges, showing the importance of phenotypic plasticity and genetic control of change during the invasion process.

Author(s): Liu, W; Chen, X; Strong, DR; Pennings, SC; Kirwan, ML; Chen, X; Zhang, Y | Abstract: Introduced plants provide a unique opportunity to examine how plants respond through plasticity and adaptation to changing climates. We compared plants of Spartina alterniflora from the native (United States, 27–43°N) and introduced (China, 19–40°N) ranges. In the field and greenhouse, aboveground productivity of Chinese plants was greater than that of North American plants. Aboveground biomass in the field declined with increasing latitude in the native range, a pattern that persisted in the greenhouse, indicating a genetic basis. Aboveground biomass in the field displayed hump-shaped relationships with latitude in China, but this pattern disappeared in field and greenhouse common gardens, indicating phenotypic plasticity. Relationships in both geographic regions were explained by temperature, which is probably the underlying environmental factor affecting aboveground biomass. S. alterniflora has evolved greater biomass in China, but in the four decades since it was introduced, it has not yet evolved the genetic cline in biomass seen in its native range. By working at lower latitudes in the introduced range than have been sampled in the native range, we identified an optimum temperature in the introduced range above which aboveground productivity decreases.

Phenology studies mostly focus on variation across time or landscapes. However, phenology can vary at fine spatial scales, and these differences may be as important as long-term change from climate warming. We used high-frequency “PhenoCam” data to examine phenology of Spartina alterniflora, a foundation species native to salt marshes on the US East and Gulf coasts, and a common colonizer elsewhere. We examined phenology across three microhabitats from 2013 to 2017 and used this information to create the first spring green-up model for S. alterniflora. We then compared modern spatial variation to that exhibited over a 60-year climate record. Marsh interior plants initiated spring growth 17 days earlier than channel edge plants and spent 35 days more in the green-up phenophase and 25 days less in the maturity phenophase. The start of green-up varied by 17 days among 3 years. The best spring green-up model was based on winter soil total growing degree days. Across microhabitats, spring green-up differences were caused by small elevation changes (15 cm) that drove soil temperature variation of 0.8°C. Preliminary evidence indicated that high winter belowground biomass depletion triggered early green-up. Long-term change was similar: winter soil temperatures warmed 1.7 ± 0.3°C since 1958, and green-up advanced 11 ± 6 days, whereas contemporary microhabitat differences were 17 ± 4 days. Incorporating local spatial variation into plant phenology models may provide an early warning of climate vulnerability and improve understanding of ecosystem-scale productivity. Microscale phenology variation likely exists in other systems and has been unappreciated.

Plants adjust their size and reproductive effort in response to numerous selection pressures and constraints. The self‐thinning law describes a well‐known trade‐off between size and density. Plants also trade‐off investment into growth vs. sexual reproduction, as described by life‐history theory. We build on past work on plant allometry and life history by examining both self‐thinning and size‐dependent reproduction in a single plant species, the saltmarsh grass Spartina alterniflora, across a wide range of settings: three landscape positions, two habitats and eight sites, across sixteen years. Plants in different landscape positions and years varied tremendously in size and shoot density. However, all this variation could be explained by a single allometric relationship consistent with the self‐thinning law, but with a lower slope. Flowering was size‐dependent, and the size at which plants had a 50% probability of flowering varied among habitat, sites and years. Plants that were stressed reproduced at a smaller size than plants that were growing under good conditions, and this pattern was consistent among habitat, sites and years. Finally, reproductive biomass and the proportion of shoots flowering increased with increasing vegetative size (plant height or shoot biomass). Combining these two patterns, S. alterniflora plants growing high density are small and reproduce at a smaller size than large plants growing at low density. Although there is tremendous spatial and temporal variation in S. alterniflora growth and reproductive patterns, all this variation can be understood as resulting from two simple allometric trade‐offs. Because saltmarsh plants often occur in monospecific stands, they may serve as simple, model systems for studies of plant life history. A free Plain Language Summary can be found within the Supporting Information of this article.

Consumers often deplete local resources and aggregate along edges of remaining resources, forming "consumer fronts." We examined the factors that promote Sesarma reticulatum crab aggregations at saltmarsh creek heads to explain the directional but slow movement of these fronts. We also created artificial creek heads to test the hypothesis that hydrological conditions at creek heads create superior habitat for crabs. Soil temperatures were ˜11-12% cooler, hydrogen sulfide concentrations lower (0.0 vs. ˜0.58 mg/L), and dissolved oxygen concentrations twofold higher at the creek head versus the marsh platform. In the artificial creek-head experiment, altering hydrological conditions led to lower dissolved sulfide levels, higher dissolved oxygen levels, and increased densities of crab burrows and Sesarma crabs. Moreover, the elevation of the soil surface declined rapidly at artificial creek heads versus controls, suggesting that crabs were increasing erosion. Our results suggest that abiotic conditions for crabs are better at the leading edge of the creek head than the trailing edge, explaining the directional movement of the front. Moreover, the speed at which the front propagates appears to be limited by the rate at which the creekhead erodes, rather than by crab mobility. The directional and slow movement of Sesarma fronts compared to consumer fronts of other invertebrates appears to result from the inextricable link between Sesarma and marsh geomorphology, whereas other consumer fronts are associated mostly with food resources.

We tested the hypothesis that mangroves provide better coastal protection than salt marsh vegetation using ten 1,008 m2 plots in which we manipulated mangrove cover from 0 to 100 percent. Hurricane Harvey passed over the plots in 2017. Data from erosion stakes indicated up to 26 cm of vertical and 970 cm of horizontal erosion over 70 months in the plot with 0 percent mangrove cover, but relatively little erosion in other plots. The hurricane did not increase erosion, and erosion decreased after the hurricane passed. Data from drone images indicated 196 m2 of erosion in the 0 % mangrove plot, relatively little erosion in other plots, and little ongoing erosion after the hurricane. Transects through the plots indicated that the levee (near the front of the plot) and the bank (the front edge of the plot) retreated up to 9 m as a continuous function of decreasing mangrove cover. Soil strength was greater in areas vegetated with mangroves than in areas vegetated by marsh plants, or nonvegetated areas, and increased as a function of plot-level mangrove cover. Mangroves prevented erosion better than marsh plants did, but this service was non-linear, with low mangrove cover providing most of the benefits.

Nitrogen and phosphorus are thought to be the most important limiting nutrients in most terrestrial ecosystems, but little is known about how other elements may limit the abundance of arthropods. We utilized a fully factorial fertilization experiment that manipulated macronutrients (N and P, together) and micronutrients (calcium, sodium, potassium, separately), in large 30 × 30 m plots and sampled litter arthropods via pitfall trapping to determine the nutrients that limit this group. An invasive ant, Nylanderia fulva, numerically dominated the community and increased in abundance 13% in plots fertilized by Ca. Detritivores were not limited by any nutrient combination, but macronutrients increased predator abundance 43%. We also found that some combinations of macronutrients and micronutrients had toxic or stressful effects on the arthropod community: detritivores decreased in abundance 23% with the combination of macronutrients, Ca, and K, and 22% with macronutrients and K; and N. fulva decreased in abundance 24% in plots fertilized by K and 45% in plots fertilized by the combination of Na and K. Our work supports growing evidence that micronutrients, especially Ca and K, may be important in structuring grassland arthropod communities, and suggests that micronutrients may affect whether or not invasive ants reach numerical dominance.

Spatially synchronous population dynamics are important to ecosystem functioning and have several potential causes. By looking at synchrony in plant productivity over 18 years across two elevations in three types of coastal marsh habitat dominated by different clonal plant species in Georgia, USA, we were able to explore the importance of plant species and different habitat conditions to synchrony. Synchrony was highest when comparing within a plant species and within a marsh zone, and decreased across species, with increasing distance, and with increasing elevational differences. Abiotic conditions that were measured at individual sites (water column temperature and salinity) also showed high synchrony among sites, and in one case (salinity) decreased with increasing distance among sites. The Moran Effect (synchronous abiotic conditions among sites) is the most plausible explanation for our findings. Decreased synchrony between creekbank and mid-marsh zones, and among habitat types (tidal fresh, brackish and salt marsh) was likely due in part to different exposure to abiotic conditions and in part to variation in sensitivity of dominant plant species to these abiotic conditions. We found no evidence for asynchrony among species, sites or zones, indicating that one habitat type or zone will not compensate for poor production in another during years with low productivity; however, tidal fresh, brackish and salt marsh sites were also not highly synchronous with each other, which will moderate productivity variation among years at the landscape level due to the portfolio effect. We identified the creekbank zone as more sensitive than the mid-marsh to abiotic variation and therefore as a priority for monitoring and management.

Decomposition of vegetal detritus is one of the most fundamental ecosystem processes. In complex landscapes, the fate of litter of terrestrial plants may depend on whether it ends up decomposing in terrestrial or aquatic conditions. However, (1) to what extent decomposition rates are controlled by environmental conditions or by detritus type, and (2) how important the composition of the detritivorous fauna is in mediating decomposition in different habitats, remain as unanswered questions. We incubated two contrasting detritus types in three distinct habitat types in Coastal Georgia, USA, to test the hypotheses that (1) the litter fauna composition depends on the habitat and the litter type available, and (2) litter mass loss (as a proxy for decomposition) depends on environmental conditions (habitat) and the litter type. We found that the abundance of most taxa of the litter fauna depends primarily on habitat. Litter type became a stronger driver for some taxa over time, but the overall faunal composition was only weakly affected by litter type. Decomposition also depends strongly on habitat, with up to ca. 80% of the initial detrital mass lost over 25 months in the marsh and forest habitats, but less than 50% lost in the creek bank habitat. Mass loss rates of oak versus pine litter differed initially but converged within habitat types within 12 months. We conclude that, although the habitat type is the principle driver of the community composition of the litter fauna, litter type is a significant driver of litter mass loss in the early stages of the decomposition process. With time, however, litter types become more and more similar, and habitat becomes the dominating factor in determining decomposition of older litter. Thus, the major driver of litter mass loss changes over time from being the litter type in the early stages to the habitat (environmental conditions) in later stages., Decomposition of vegetal detritus is one of the most fundamental ecosystem processes. In complex landscapes, the fate of litter of terrestrial plants may depend on whether it ends up in decomposition of terrestrial or aquatic conditions. While litter type is a significant driver of litter mass loss in the early stages of decomposition, litter types become more and more similar over time, and habitat becomes the dominating factor in determining the decomposition of older litter.

The outcome of species interactions may manifest differently at different spatial scales; therefore, our interpretation of observed interactions will depend on the scale at which observations are made. For example, in ladybeetle–aphid systems, the results from small‐scale cage experiments usually cannot be extrapolated to landscape‐scale field observations. To understand how ladybeetle–aphid interactions change across spatial scales, we evaluated predator–prey interactions in an experimental system. The experimental habitat consisted of 81 potted plants and was manipulated to facilitate analysis across four spatial scales. We also simulated a spatially explicit metacommunity model parallel to the experiment. In the experiment, we found that the negative effect of ladybeetles on aphids decreased with increasing spatial scales. This pattern can be explained by ladybeetles strongly suppressing aphids at small scales, but not colonizing distant patches fast enough to suppress aphids at larger scales. In the experiment, the positive effects of aphids on ladybeetles were strongest at three‐plant scale. In a model scenario where predators did not have demographic dynamics, we found, consistent with the experiment, that both the effects of ladybeetles on aphids and the effects of aphids on ladybeetles decreased with increasing spatial scales. These patterns suggest that dispersal was the primary cause of ladybeetle population dynamics in our experiment: aphids increased ladybeetle numbers at smaller scales because ladybeetles stayed in a patch longer and performed area‐restricted searches after encountering aphids; these behaviors did not affect ladybeetle numbers at larger spatial scales. The parallel experimental and model results illustrate how predator–prey interactions can change across spatial scales, suggesting that our interpretation of observed predator–prey dynamics would differ if observations were made at different scales. This study demonstrates how studying ecological interactions at a range of scales can help link the results of small‐scale ecological experiments to landscape‐scale ecological problems.

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.

Small-scale armoring placed near the marsh-upland interface to protect single-family homes is widespread but understudied. Using a nested, spatially blocked sampling design on the coast of Georgia, USA, we compared the biota and environmental characteristics of 60 marshes adjacent to either a bulkhead, a residential backyard with no armoring, or an intact forest. We found that marshes adjacent to bulkheads were at lower tidal elevations and had features typical of lower elevation marsh habitats: high coverage of the marsh grass Spartina alterniflora, high density of crab burrows, and muddy sediments. Marshes adjacent to unarmored residential sites had higher soil water content and lower porewater salinities than the armored or forested sites, suggesting that there may be increased freshwater input to the marsh at these sites. Deposition of Spartina wrack on the marsh-upland ecotone was negatively related to elevation at armored sites and positively related at unarmored residential and forested sites. Armored and unarmored residential sites had reduced densities of the high marsh crab Armases cinereum, a species that moves readily across the ecotone at forested sites, using both upland and high marsh habitats. Distance from the upland to the nearest creek was longest at forested sites. The effects observed here were subtle, perhaps because of the small-scale, scattered nature of development. Continued installation of bulkheads in the southeast could lead to greater impacts such as those reported in more densely armored areas like the northeastern USA. Moreover, bulkheads provide a barrier to inland marsh migration in the face of sea level rise. Retaining some forest vegetation at the marsh-upland interface and discouraging armoring except in cases of demonstrated need could minimize these impacts.

The Deepwater Horizon spill (2010) was the largest marine oil spill in US waters to date and one of the largest worldwide. To examine effects of the oil spill on an important salt marsh species over time, we conducted a meta-analysis on marsh periwinkles Littoraria irrorata using published and unpublished sources spanning more than 5 yr (2010−2015), including newly available Natural Resources Damage Assessment (NRDA) and Gulf of Mexico Research Initiative (GoMRI) data sets. We tested the hypotheses that the spill decreased mean periwinkle density, reduced mean snail shell length, and changed periwinkle size distribution. Averaged across multiple studies, sites, marsh zones (edge versus interior), and years, our synthesis revealed a negative effect of heavy oiling on periwinkles. Snail densities were reduced by 73% in heavily oiled sites across all study-zone-by-year combinations, including adverse effects for both the oiled marsh edge and oiled marsh interior, with impacts observed over more than 5 yr. Mean periwinkle shell length was somewhat reduced at the oiled marsh edge in a few cases; however, periwinkle size distributions displayed greater relative proportions of smaller adults and sub-adults, and fewer large adults, across all years. Given the spatial and temporal extent of data analyzed, this synthesis provides evidence that the Deepwater Horizon spill suppressed populations of marsh periwinkles in heavily oiled marshes for over 5 yr, and that impacts were ongoing and recovery was incomplete, likely affecting other ecosystem components, including marsh productivity, organic matter and nutrient cycling, marsh−estuarine food webs, and associated predators.

© 2017 by the Ecological Society of America Ecological invasions are facilitated by pre-adaptation and phenotypic plasticity, upon which evolution can act. The rapid invasion of the intertidal grass Spartina alterniflora in China during the last 36 yr is a test case for the roles of these mechanisms. A previous study of S. alterniflora in China found strong latitudinal clines in vegetative and sexual traits and concluded that most of this variation was due to phenotypic plasticity. Recent observations suggested provenance-by-environment interactions, and we employed common gardens at multiple latitudes as a test of this idea. Phenotypically, field plant height, which correlates strongly with biomass and other indices of vegetative performance in this species, showed a hump-shaped relationship across 10 sites, covering 19° of latitude; field seed set increased linearly with latitude. To assess the role of plasticity vs. genetic differentiation in these patterns, we grew plants from the ten field sites in three common gardens at low (20.9° N), mid (28.3° N), and high (38.0° N) latitudes to maturity, at 18 months. Plant height varied among common gardens, with the tallest plants at mid latitude, mirroring the field pattern, consistent with the previous study. Within the gardens, latitude of origin also affected plant height. Seed set varied among the gardens, with the greatest values at high latitudes, again mirroring the field pattern and indicating substantial plasticity. Evidence of evolution was found as increasing seed set with latitude among provenances within common gardens. However, the effect differed among common gardens, with the greatest slope in the high-latitude garden, lower slope in the mid-latitude garden, and no relationship in the low-latitude garden, indicating a provenance-by-environment interaction. The number of surviving plants also suggested a provenance-by-environment interaction; no relationship with latitude among provenances in the two southern gardens and increasing survival with latitude in the northern garden. Field seedling density was ~200-fold greater at high than at low latitude sites. The profuse seed germination and recruitment in the north could have created high selection intensity resulting in evolution of reproductive traits at high latitudes with the result that the mechanisms of the invasion differ with latitude.

Despite overall global declines, mangroves are expanding into and within many subtropical wetlands, leading to heterogeneous cover of marsh-mangrove coastal vegetation communities near the poleward edge of mangroves' ranges. Coastal wetlands are globally important carbon sinks, yet the effects of shifts in mangrove cover on organic-carbon (OC) storage remains uncertain. We experimentally maintained black mangrove (Avicennia germinans) or marsh vegetation in patches (n = 1,120, 3 × 3 m) along a gradient in mangrove cover (0-100%) within coastal wetland plots (n = 10, 24 × 42 m) and measured changes in OC stocks and fluxes. Within patches, above and belowground biomass (OC) was 1,630% and 61% greater for mangroves than for recolonized marshes, and soil OC was 30% greater beneath mangrove than marsh vegetation. At the plot scale, above and belowground biomass increased linearly with mangrove cover but soil OC was highly variable and unrelated to mangrove cover. Root ingrowth was not different in mangrove or marsh patches, nor did it change with mangrove cover. After 11 months, surface OC accretion was negatively related to plot-scale mangrove cover following a high-wrack deposition period. However, after 22 months, accretion was 54% higher in mangrove patches, and there was no relationship to plot-scale mangrove cover. Marsh (Batis maritima) leaf and root litter had 1,000% and 35% faster breakdown rates (k) than mangrove (A. germinans) leaf and root litter. Soil temperatures beneath mangroves were 1.4°C lower, decreasing aboveground k of fast- (cellulose) and slow-decomposing (wood) standard substrates. Wood k in shallow soil (0-15 cm) was higher in mangrove than marsh patches, but vegetation identity did not impact k in deeper soil (15-30 cm). We found that mangrove cover enhanced OC storage by increasing biomass, creating more recalcitrant organic matter and reducing k on the soil surface by altering microclimate, despite increasing wood k belowground and decreasing allochthonous OC subsidies. Our results illustrate the importance of mangroves in maintaining coastal OC storage, but also indicate that the impacts of vegetation change on OC storage may vary based on ecosystem conditions, organic-matter sources, and the relative spatiotemporal scales of mangrove vegetation change.

We compared coastal restoration projects in a developing country, China, and a developed country, the United States of America, both of which are facing loss and degradation of coastal habitats at similar latitudes, for the period of 1992–2014. To document the scale of coastal habitat restoration projects in the two countries, we identified 914 coastal restoration projects with an accumulated area of 300,521 acres in China, with most of our information coming from scientific papers, and 1,620 coastal restoration projects with an accumulated area of 243,064 acres in the USA, with most of our information coming from public databases. In both countries, about half the projects were in wetland habitats, but China had a greater proportion of projects in submerged habitats (43% versus 28% in the USA) and the USA a greater proportion in coastal upland habitats (21% versus 9% in China). The number of new projects steadily increased over time in China, but dropped after 2006 in the USA, although the total cost of new projects continued to increase. The number of projects in China and the total cost of projects in the USA were correlated with national GDP. Restoration projects in China used fewer techniques, had fewer partners, and took longer to complete than projects in the USA. Information about projects was incomplete, especially in China, and both countries could do more to make information publically available. We know more about project construction than project outcomes, and it is unclear whether projects are achieving their goals or whether the techniques used are optimal.

Fil: Komatsu, Kimberly J. Smithsonian Environmental Research Center, Edgewater. United States. Fil: Avolio, Meghan L. Johns Hopkins University. Department of Earth and Planetary Sciences. Baltimore, United States. Fil: Lemoine, Nathan P. Marquette University. Department of Biological Sciences. Milwaukee, United States. Fil: Chaneton, Enrique José. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Chaneton, Enrique José. CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Tognetti, Pedro Maximiliano.Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Tognetti, Pedro Maximiliano.CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Yahdjian, María Laura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Yahdjian, María Laura. CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Isbell, Forest. University of Minnesota. Department of Ecology, Evolution and Behavior. Saint Paul, United States. Fil: Grman, Emily. Eastern Michigan University. Department of Biology. Ypsilanti, United States. Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of CDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term ( minor to 10 y). In contrast, long-term (major or equal to 10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously. tbls., grafs.

Sea level rise is expected to increase inundation and saltwater intrusion into many tidal freshwater marshes and forests. Saltwater intrusion may be long-term, as with rising seas, or episodic, as with low river flow or storm surge. We applied continuous (press) and episodic (pulse) treatments of dilute seawater to replicate 2.5 × 2.5 m field plots for three years and measured soil attributes, including soil porewater, oxidation-reduction potential, soil carbon (C), and nitrogen (N) to investigate the effects of continuous and episodic saltwater intrusion and increased inundation on tidal freshwater marsh elemental cycling and soil processes. Continuous additions of dilute seawater resulted in increased porewater chloride, sulfate, sulfide, ammonium, and nitrate concentrations. Plots that received press additions also had lower soil oxidation-reduction potentials beginning in the second year. Episodic additions of dilute seawater during typical low flow conditions (Sept.-Oct.) resulted in transient increases in porewater chloride and sulfate that returned to baseline conditions once dosing ceased. Freshwater additions did not affect porewater inorganic N or soil C or N. Persistent saltwater intrusion in freshwater marshes alters the N cycle by releasing ammonium-N from sorption sites, increasing nitrification and severely reducing N storage in macrophyte biomass. Chronic saltwater intrusion, as is expected with rising seas, is likely to shift tidal freshwater marshes from a sink to a source of N whereas intermittent intrusion from drought may have no long term effect on N cycling.

Global changes are causing broad-scale shifts in vegetation communities worldwide, including coastal habitats where the borders between mangroves and salt marsh are in flux. Coastal habitats provide numerous ecosystem services of high economic value, but the consequences of variation in mangrove cover are poorly known. We experimentally manipulated mangrove cover in large plots to test a set of linked hypotheses regarding the effects of changes in mangrove cover. We found that changes in mangrove cover had strong effects on microclimate, plant community, sediment accretion, soil organic content, and bird abundance within 2 yr. At higher mangrove cover, wind speed declined and light interception by vegetation increased. Air and soil temperatures had hump-shaped relationships with mangrove cover. The cover of salt marsh plants decreased at higher mangrove cover. Wrack cover, the distance that wrack was distributed from the water's edge, and sediment accretion decreased at higher mangrove cover. Soil organic content increased with mangrove cover. Wading bird abundance decreased at higher mangrove cover. Many of these relationships were non-linear, with the greatest effects when mangrove cover varied from zero to intermediate values, and lesser effects when mangrove cover varied from intermediate to high values. Temporal and spatial variation in measured variables often peaked at intermediate mangrove cover, with ecological consequences that are largely unexplored. Because different processes varied in different ways with mangrove cover, the "optimum" cover of mangroves from a societal point of view will depend on which ecosystem services are most desired.

Introduced species may homogenize biotic communities. Whether this homogenization can erase latitudinal patterns of species diversity and composition has not been well studied. We examined this by comparing nematode and microbial communities in stands of native Phragmites australis and exotic Spartina alterniflora in coastal wetlands across 18° of latitude in China. We found clear latitudinal clines in nematode diversity and functional composition, and in microbial composition, for soils collected from native P. australis. These latitudinal patterns were weak or absent for soils collected from nearby stands of the exotic S. alterniflora. Climatic and edaphic variables varied across latitude in similar ways in both community types. In P. australis there were strong correlations between community structure and environmental variables, whereas in S. alterniflora these correlations were weak. These results suggest that the invasion of S. alterniflora into the Chinese coastal wetlands has caused profound biotic homogenization of soil communities across latitude. We speculate that the variation in P. australis nematode and microbial communities across latitude is primarily driven by geographic variation in plant traits, but that such variation in plant traits is largely lacking for the recently introduced exotic S. alterniflora. These results indicate that widespread exotic species can homogenize nematode communities at large spatial scales.

The canonical factors typically thought to determine herbivore community structure often explain only a small fraction of the variation in herbivore abundance and diversity. We tested how macronutrients and relatively understudied micronutrients interacted to influence the structure of insect herbivore (orthopteran) communities. We conducted a factorial fertilisation experiment manipulating macronutrients (N and P, added together) and micronutrients (Ca, Na and K) in large plots (30 × 30 m2 ) in a Texas coastal prairie. Although no single or combination of micronutrients affected herbivore communities in the absence of additional macronutrients, macronutrients and sodium added together increased herbivore abundance by 60%, richness by 15% and diversity by 20%. These results represent the first large-scale manipulation of single micronutrients and macronutrients in concert, and revealed an herbivore community co-limited by macronutrients and Na. Our work supports an emerging paradigm that Na may be important in limiting herbivore communities.

Direct interactions among plant species may be highly modified by indirect or diffuse effects within a multispecies community. We investigated the direct and diffuse effects of two salt marsh perennials, Monanthechloe littoralis and Arthrocnemum subterminale, on winter annuals and the perennial herb Limonium californicum in a salt marsh in central California. In permanent plots, Monanthechloe had expanded substantially in the upper marsh over the past 13 yr, while Arthrocnemum and all annual species had decreased. These dynamics suggest that Monanthechloe may directly outcompete most other species in the upper marsh. In contrast, Arthrocnemum is known to facilitate some annual species. In our field experiments, Monanthechloe strongly suppressed Arthrocnemum, all four common annual species, and Limonium in direct interactions. In contrast, Arthrocnemum directly facilitated the winter annuals Parapholis incurva and Lasthenia glabrata, competed with Spergularia marina, and did not have a significant effect on Limonium. However, when the combined effects of Monanthechloe and Arthrocnemum were tested, Arthrocnemum ameliorated the negative effect of Monanthechloe on all four species. Although isolated Arthrocnemum competed with Spergularia and had no direct effects on Limonium, Arthrocnemum in the presence of Monanthechloe facilitated both species. We hypothesize that Arthrocnemum buffered the strong competitive effects of Monanthechloe on Lasthenia and Parapholis via direct positive effects and benefited Spergularia and Limonium through its competitive effect on Monanthechloe. These findings add to the growing body of literature emphasizing the importance of diversity and interdependence in the functioning of plant communities.

Understanding how biodiversity affects ecosystem processes is a key question in ecology. Previous research has found that increasing plant diversity often enhances many ecosystem processes, but less is known about the role of consumer diversity to ecosystem processes, especially in terrestrial ecosystems. Furthermore, we do not know how general biodiversity responses are among ecosystem types. We examined the role of insect herbivore (Orthoptera) diversity on plant production using parallel field experiments in three grassland ecosystems (mixed grass prairie, tallgrass prairie and coastal tallgrass prairie) to determine whether the effects of grasshopper diversity were consistent among sites. Using mesocosms, we manipulated orthopteran species richness (0, 1, 2, 3 or 4 species), functional richness (number of functional feeding groups present; 0, 1 or 2 functional groups) and functional composition (composition of functional groups present; mixed-feeders only, grass-feeders only, both mixed-feeders and grass-feeders). Diversity treatments were maintained throughout the experiment by replacing dead individuals. Plant biomass was destructively sampled at the end of the experiment. We found no effect of species richness or functional richness on plant biomass. However, herbivore functional composition was important, and effects were qualitatively similar across sites: The presence of only grass-feeding species reduced plant biomass more than either mixed-feeding species alone or both groups together. Orthopterans had consistent effects across a range of abiotic conditions, as well as different plant community and orthopteran community compositions. Our results suggest that functional composition of insect herbivores affects plant communities in grasslands more than herbivore species richness or functional richness, and this pattern was robust among grassland types.

Coastal wetlands are important to human well-being and vulnerable to oil spills. Research on biotic effects of oil has been focused on microbes, plants, and benthic invertebrates, neglecting terrestrial arthropods. We review how oil affects terrestrial arthropods in coastal marshes and suggest future research topics. Terrestrial arthropods play important ecological roles in coastal marshes, affecting primary production and decomposition and providing food to terrestrial and marine vertebrates. Some species are pests and disease vectors for humans and livestock. Terrestrial arthropods are sensitive to oil exposure and are suppressed even in lightly oiled sites where plants are not affected. Some arthropods later reinhabit oiled marshes, which demonstrates that portions of the arthropod community may be resilient to oil exposure. However, oil that remains in sediments may affect terrestrial populations for years after the spill. The sensitivity of arthropods to oil exposure makes them useful indicators of marsh health following environmental impacts.

Interactions between plants and herbivores often vary on a geographic scale. Although theory about plant defenses and tolerance is predicated on temporal or spatial variation in herbivore damage, no single study has compared the pattern of herbivory, plant defenses and tolerance to herbivory of a single species across a latitudinal gradient. In 2002–2005 we surveyed replicate salt marshes along the Atlantic coast of the United States from Florida to Maine. At each field site we scored leaves of Iva frutescens for herbivore damage. In laboratory experiments we measured constitutive resistance and induced resistance in I. frutescens from high and low latitude sites along the Atlantic Coast. In another common garden experiment we studied tolerance to herbivory of I. frutescens from various sites. Theory predicts that constitutive resistance should matter more when damage is high, and induced resistance when herbivory is high but variable. In the field, average levels of herbivore damage, and spatial and temporal variation in herbivore damage were all greater at low versus high latitudes, indicating that constitutive as well as induced resistance should be stronger at low latitudes. Consistent with this prediction, constitutive resistance to herbivory was stronger at low latitudes. Induced resistance to herbivores was also stronger at low latitudes: it was deployed faster and lasted longer. Theory also predicts that tolerance to herbivory should be greater where average herbivory damage is greater; however, tolerance to herbivory in Iva did not depend on geographic origin. Our results emphasize the value of considering multiple ways in which plants respond to herbivores when examining geographic variation in plant–herbivore interactions.

Tidal wetlands are threatened by global changes related not only to sea level rise but also to altered weather patterns. To predict consequences of these changes on coastal communities, it is necessary to understand how temporally varying abiotic conditions drive wetland production. In 2000–2011, we conducted annual surveys of Spartina alterniflora biomass in tidal marshes at nine sites in and around the Altamaha river estuary on the coast of Georgia, USA. End of the year live biomass was assessed in the creekbank and midmarsh zones to estimate annual net primary production (ANPP). River discharge was the most important driver of S. alterniflora ANPP, especially in creekbank vegetation. Increased river discharge reduces water column salinity, and this was most likely the proximate driver of increased production. In the midmarsh zone, the patterns were less distinct, although river discharge was again the best predictor, but maximum temperature had similar predictive ability. In contrast to results from terrestrial grasslands, we found no consistent evidence for a sharply delimited critical period for any climate driver in the tidal marsh, which indicates that plant growth was responsive to abiotic drivers at any time during the growing season. Results were broadly consistent across multiple sites within a geographic region. Our results differ from previous analyses of production in S. alterniflora marshes, which either identified oceanic drivers of S. alterniflora production or were unable to identify any drivers, likely because the low-latitude sites we studied were hotter and more affected by river discharge than those in previous studies.

Salt marshes often contain remarkable spatial heterogeneity at multiple scales across the landscape. A combination of advanced remote-sensing approaches (hyperspectral imagery and lidar) and conventional field survey methods was used to produce detailed quantifications and maps of marsh platform geomorphology, vegetation composition and biomass, and invertebrate patterns in a Georgia (USA) salt marsh. Community structure was largely related to hydrology, elevation, and soil properties. Both abiotic drivers and community patterns varied among subwatersheds and across the landscape at larger spatial scales. The authors conclude that measurements of marsh ecosystem structure and processes are spatially contextual and not scalable without detailed geospatial analysis. Efforts to protect and restore coastal marshes must strive to document, understand, and conserve this inherent spatial complexity.

Summary 1. An important recent advance in food web ecology has been the application of theory regarding spatial gradients to studies of the factors that affect animal population dynamics. Building on extensive studies of the Spartina alterniflora food web at the local scale, we hypothesized that geographic variation in S. alterniflora quality is an important bottom-up control on food web structure and that geographic variation in S. alterniflora quality would interact with the presence of predators and top omnivores to mediate herbivore densities. 2. We employed a four-factor fully crossed experiment in which we (i) collected plants from highand low-latitude locations and grew them in a common garden and varied (ii) plant fertilization status (mimicking the plant quality differences due to marsh elevation), (iii) mesopredator density and (iv) omnivore density. 3. Our results suggest that the single most important factor mediating insect herbivore densities is local variation in plant quality – induced in our experiment by fertilization and demonstrated repeatedly as a consequence of marsh elevation. 4. Top-down effects were generally weak and in those cases where predators did exert a significant suppressing effect on herbivores, that impact was itself mediated by host-plant characteristics. 5. Finally, despite observed variation in plant quality with latitude, and the separately measurable effects of this variation on herbivores, geographic-scale variation in plant quality was overwhelmed by local conditions in our experiments. 6. Synthesis. We suggest that a first-order understanding of variation across large latitudinal ranges in the Spartina alterniflora arthropod food web must begin with local variation in plant quality, which provides strong bottom-up forcing to herbivore populations. A second-order understanding of the arthropod food web should consider the role of predation in controlling herbivores feeding on lowquality plants. Finally, while latitudinal variation in plant quality probably explains some variation in herbivore densities, it is probably more of a response to herbivore pressure than a driver of the herbivore dynamics. Although extrapolating from local to geographic scales presents multiple challenges, it is an essential task in order for us to develop an understanding that is general rather than site-specific.

In salt marshes, disturbance by wrack (floating mats of dead vegetation) is common and affects plant productivity and species composition, but little is known about how the timing of disturbance mediates these effects, nor how it interacts with herbivory. Using a field experiment on the Georgia coast, we simulated the effects of wrack disturbance at different times of the year on the marsh grass Spartina alterniflora and its stem-boring herbivorous insects. The timing of disturbance throughout the growing season strongly affected fall biomass, stem height, the proportion of stems flowering, and the proportion of stems colonized by stem-boring herbivorous insects. End-of-season biomass in plots disturbed in March did not differ from undisturbed controls, but biomass was reduced by 50% in plots disturbed in May, and by over 90% in plots disturbed in September. Disturbance in March and May stimulated flowering, but disturbance later in the growing season suppressed it. Plots disturbed late in the growing season had a low frequency of stem-boring herbivores. Stems containing stem borers rarely flowered. These results indicate that the timing of disturbance matters in coastal salt marshes. Late-season disturbances had the strongest effects on S. alterniflora and its herbivores. Disturbances early in the growing season did not affect end-of-season biomass, and stimulated flowering, suggesting parallels between fire disturbance in grasslands and wrack disturbance in salt marshes. Late-season disturbance did reduce herbivory by stem-boring insects, but not enough to compensate for the direct effects of disturbance on the plants. Future studies of disturbance in salt marshes should consider how the timing of experimental disturbance treatments relates to the timing of natural disturbances.

We documented the frequency and effect on live biomass of five different types of disturbance over 14 years in creekbank and mid-marsh zones of eight salt marshes dominated by Spartina alterniflora in Georgia, USA. Wrack (floating debris) and creekbank slumping were the most common disturbances at the creekbank, and snails were the most common disturbance agent in the mid-marsh. Disturbance frequency varied among sites due to differences in plot elevation and landscape position. Wrack disturbance at the creekbank was positively correlated with plot elevation, and both initial slumping and terminal slumping of creekbank plots were negatively correlated with plot elevation. Wrack disturbance at the creekbank and snail disturbance in the mid-marsh were also most common at barrier island vs. interior marshes. Disturbance varied up to 14-fold among years. Wrack disturbance at the creekbank was negatively correlated with river discharge and sea level, and initial slumping of creekbank plots was also negatively correlated with sea level. The different disturbance types varied in their effects on end-of-year standing plant biomass. At the creekbank, wrack disturbance reduced biomass in affected plots by ~46%, but slumping did not affect biomass until the plot was totally lost. In the mid-marsh, slumping and wrack were not important disturbances, but snail disturbance reduced biomass in affected plots by ~70%. In addition, abiotic conditions (river discharge, maximum monthly temperature, sea level, and precipitation) strongly affected year-to-year variation in biomass. Across the entire landscape, fewer than a quarter of the plots on average were disturbed, and disturbance reduced overall standing biomass by ~18% in the creekbank zone and ~3% in the mid-marsh zone. Our results indicate that wrack has fairly strong effects on end-of-year biomass at the creekbank. Overall, however, variation in abiotic conditions among years had stronger effects on end-of-year standing biomass in both marsh zones than did disturbance.