Reciprocal facilitation between ants and small mammals in tidal marshes.

The role of facilitation in shaping natural communities has primarily been studied in the context of plant assemblages, while its relevance for mobile animals remains less understood. Our study investigates whether reciprocal interspecific facilitation may exist between fire ants (Solenopsis richteri) and cavies (Cavia aperea), two mobile animals, in the SW Atlantic coast brackish marshes. Field samples showed a spatial association between ant mounds and cavies, and that ants prefer to use cavy runways for movement within the marsh. Through experiments involving transplanting the dominant plant, cordgrass (Spartina densiflora), and manipulating cavy presence in areas with and without ant mounds, we observed that cavies forage extensively (and defecate more) near ant mounds. The ants actively remove cavy droppings in their mound vicinity. These ant activities and interactions with cavy droppings led to reduced moisture and organic content while increasing nitrate and phosphate levels in marsh sediment. Consequently, this enhanced plant growth, indirectly facilitating the cavies, which preferred consuming vegetation near ant mounds. These cascading indirect effects persisted over time; even four months after cavies left the marshes, transplanted plants near ant mounds remained larger and exhibited more leaf senescence when exposed to cavy herbivory. Therefore, the networks of positive interactions appear to generate simultaneous selection among species (populations), promoting coexistence within the community. Although complex, these reciprocal facilitative effects among mobile animals may be more common than currently believed and should be further studied to gain a better understanding of the underlying mechanisms driving species coexistence in natural communities.

Herbivores and nutrients control grassland plant diversity via light limitation.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation.

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change - fertilisation and herbivore loss - are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.

Nitrogen enrichment suppresses other environmental drivers and homogenizes salt marsh leaf microbiome.

Microbial community assembly is affected by a combination of forces that act simultaneously, but the mechanisms underpinning their relative influences remain elusive. This gap strongly limits our ability to predict human impacts on microbial communities and the processes they regulate. Here, we experimentally demonstrate that increased salinity stress, food web alteration and nutrient loading interact to drive outcomes in salt marsh fungal leaf communities. Both salinity stress and food web alterations drove communities to deterministically diverge, resulting in distinct fungal communities. Increased nutrient loads, nevertheless, partially suppressed the influence of other factors as determinants of fungal assembly. Using a null model approach, we found that increased nutrient loads enhanced the relative importance of stochastic over deterministic divergent processes; without increased nutrient loads, samples from different treatments showed a relatively (deterministic) divergent community assembly whereas increased nutrient loads drove the system to more stochastic assemblies, suppressing the effect of other treatments. These results demonstrate that common anthropogenic modifications can interact to control fungal community assembly. Furthermore, our results suggest that when the environmental conditions are spatially heterogeneous (as in our case, caused by specific combinations of experimental treatments), increased stochasticity caused by greater nutrient inputs can reduce the importance of deterministic filters that otherwise caused divergence, thus driving to microbial community homogenization.

Increased grassland arthropod production with mammalian herbivory and eutrophication: a test of mediation pathways.

Increases in nutrient availability and alterations to mammalian herbivore communities are a hallmark of the Anthropocene, with consequences for the primary producer communities in many ecosystems. While progress has advanced understanding of plant community responses to these perturbations, the consequences for energy flow to higher trophic levels in the form of secondary production are less well understood. We quantified arthropod biomass after manipulating soil nutrient availability and wild mammalian herbivory, using identical methods across 13 temperate grasslands. Of experimental increases in nitrogen, phosphorus, and potassium, only treatments including nitrogen resulted in significantly increased arthropod biomass. Wild mammalian herbivore removal had a marginal, negative effect on arthropod biomass, with no interaction with nutrient availability. Path analysis including all sites implicated nutrient content of the primary producers as a driver of increased arthropod mean size, which we confirmed using 10 sites for which we had foliar nutrient data. Plant biomass and physical structure mediated the increase in arthropod abundance, while the nitrogen treatments accounted for additional variation not explained by our measured plant variables. The mean size of arthropod individuals was 2.5 times more influential on the plot-level total arthropod biomass than was the number of individuals. The eutrophication of grasslands through human activity, especially nitrogen deposition, thus may contribute to higher production of arthropod consumers through increases in nutrient availability across trophic levels.

Physical stress modifies top-down and bottom-up forcing on plant growth and reproduction in a coastal ecosystem.

Bottom-up and top-down effects act together to exert strong control over plant growth and reproduction, but how physical stress modifies those interactive forces remains unclear. Even though empirical evidence is scarce, theory predicts that the importance of both top-down- and bottom-up forces may decrease as physical stress increases. Here, we experimentally evaluate in the field the separate and interactive effect of salinity, nutrient availability, and crab herbivory on plant above- and belowground biomass, as well as on sexual and clonal reproduction in the salt marsh plant Spartina densiflora. Results show that the outcome of the interaction between nutrient availability and herbivory is highly context dependent, not only varying with the abiotic context (i.e., with or without increased salinity stress), but also with the dependent variable considered. Contrary to theoretical predictions, our results show that, consistently across different measured variables, salinity stress did not cancel bottom-up (i.e., nutrients) or top-down (i.e., consumers) control, but has additive effects. Our results support emerging theory by highlighting that, under many conditions, physical stress can act additively with, or even stimulate, consumer control, especially in cases where the physical stress is only experienced by basal levels of the trophic chain. Abiotic stress, as well as bottom-up and top-down factors, can affect salt marsh structure and function not only by affecting biomass production but also by having other indirect effects, such as changing patterns in plant biomass allocation and reproduction.

Herbivory affects salt marsh succession dynamics by suppressing the recovery of dominant species.

Disturbance can generate heterogeneous environments and profoundly influence plant diversity by creating patches at different successional stages. Herbivores, in turn, can govern plant succession dynamics by determining the rate of species replacement, ultimately affecting plant community structure. In a south-western Atlantic salt marsh, we experimentally evaluated the role of herbivory in the recovery following disturbance of the plant community and assessed whether herbivory affects the relative importance of sexual and clonal reproduction on these dynamics. Our results show that herbivory strongly affects salt marsh secondary succession by suppressing seedlings and limiting clonal colonization of the dominant marsh grass, allowing subordinate species to dominate disturbed patches. These results demonstrate that herbivores can have an important role in salt marsh community structure and function, and can be a key force during succession dynamics.

Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

The blue carbon of southern southwest Atlantic salt marshes and their biotic and abiotic drivers.

Coastal vegetated ecosystems are acknowledged for their capacity to sequester organic carbon (OC), known as blue C. Yet, blue C global accounting is incomplete, with major gaps in southern hemisphere data. It also shows a large variability suggesting that the interaction between environmental and biological drivers is important at the local scale. In southwest Atlantic salt marshes, to account for the space occupied by crab burrows, it is key to avoid overestimates. Here we found that southern southwest Atlantic salt marshes store on average 42.43 (SE = 27.56) Mg OC·ha-1 (40.74 (SE = 2.7) in belowground) and bury in average 47.62 g OC·m-2·yr-1 (ranging from 7.38 to 204.21). Accretion rates, granulometry, plant species and burrowing crabs were identified as the main factors in determining belowground OC stocks. These data lead to an updated global estimation for stocks in salt marshes of 185.89 Mg OC·ha-1 (n = 743; SE = 4.92) and a C burial rate of 199.61 g OC·m-2·yr-1 (n = 193; SE = 16.04), which are lower than previous estimates.

Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

Areal changes of lentic water bodies within an agricultural basin of the Argentinean pampas. Disentangling land management from climatic causes.

Wetland loss is a frequent concern for the environmental management of rural landscapes, but poor disentanglement between climatic and land management causes frequently constrains both proper diagnoses and planning. The aim of this study is to address areal changes induced by non-climatic factors on lentic water bodies (LWB) within an agricultural basin of the Argentinean Pampas, and the human activities that might be involved. The LWB of the Mar Chiquita basin (Buenos Aires province, Argentina) were mapped using Landsat images from 1998-2008 and then corrected for precipitation variability by considering the regional hydrological status on each date. LWB areal changes were statistically and spatially analyzed in relation to land use changes, channelization of streams, and drainage of small SWB in the catchment areas. We found that 12 % of the total LWB in the basin had changed (P < 0.05) due to non-climatic causes. During the evaluated decade, 30 % of the LWB that changed size had decreased while 70 % showed steady increases in area. The number of altered LWB within watersheds lineally increased or decreased according to the proportion of grasslands replaced by sown pastures, or the proportion of sown pastures replaced by crop fields, respectively. Drainage and channelization do not appear to be related to the alteration of LWB; however some of these hydrologic modifications may predate 1998, and thus earlier effects cannot be discarded. This study shows that large-scale changes in land cover (e.g., grasslands reduction) can cause a noticeable loss of hydrologic regulation at the catchment scale within a decade.

Network analysis suggests changes in food web stability produced by bottom trawl fishery in Patagonia.

Demersal fisheries are one of the top anthropic stressors in marine environments. In the long term, some species are more vulnerable to fishery impacts than others, which can lead to permanent changes on the food web. The trophic relationships between predator and prey constitute the food web and it represents a network of the energy channels in an ecosystem. In turn, the network structure influences ecosystem diversity and stability. The first aim of this study was to describe for the first time the food web of the San Jorge Gulf (Patagonia Argentina) with high resolution, i.e. to the species level when information is available. The San Jorge Gulf was subject to intense fisheries thus our second aim is to analyse the food web structure with and without fishery to evaluate if the bottom-trawl industrial fishery altered the network structure and stability. We used several network metrics like: mean trophic level, omnivory, modularity and quasi-sign stability. We included these metrics because they are related to stability and can be evaluated using predator diets that can weight the links between predators and prey. The network presented 165 species organized in almost five trophic levels. The inclusion of a fishery node adds 69 new trophic links. All weighted and unweighted metrics showed differences between the two networks, reflecting a decrease in stability when fishery was included in the system. Thus, our results suggested a probable change of state of the system. The observed changes in species abundances since the fishery was established, could represent the state change predicted by network analysis. Our results suggests that changes in the stability of food webs can be used to evaluate the impacts of human activity on ecosystems.

Between-habitat comparison of digestive enzymes activities and energy reserves in the SW Atlantic euryhaline burrowing crab Neohelice granulata.

The digestive and metabolic characteristics at the biochemical level underlying between-habitat dietary shift of the SW Atlantic euryhaline burrowing crab Neohelice granulata under natural conditions are unknown. We made studies on adult males of N. granulata from the open mudflat and the vegetated saltmarsh in a SW Atlantic costal lagoon (Mar Chiquita, 37°32'-37°45'S; 57°19'-57°26'W, Argentina). We determined and compared amylase, maltase, sucrase, proteolytic, lipase and alkaline phosphatases activities in the hepatopancreas; glycemia, and glycogen, free glucose, triglycerides and protein concentrations in hepatopancreas, chela muscle, and anterior and posterior gills. The results show that N. granulata exhibits characteristics and between-habitat differences at the biochemical level (i.e. high amylase and disaccharidase activities, differences in total proteolytic, lipase and levamisole-insensitive AP activities in the hepatopancreas, and in the concentrations of glycogen in the gills, triglycerides in the hepatopancreas and of protein in the chela muscle) which could represent adaptive digestive and metabolic strategies to face the differences in environmental conditions (i.e. food availability). The possible relationship between digestive and metabolic characteristics and feeding patterns, type of food available and environmental conditions in each habitat is discussed.

Density affects mating mode and large male mating advantage in a fiddler crab.

Fiddler crabs show two different mating modes: either females search and crabs mate underground in male burrows, or males search and crabs mate on the surface near female burrows. We explored the relationship between crab density, body size, the searching behavior of both sexes, and the occurrence of both mating modes in the fiddler crab Uca uruguayensis. We found that crabs change their mating mode depending on their size and crab density. Crabs mated mostly on the surface at low densities, and underground at high densities. The proportion of wandering receptive females but not courting males accounted for the variation in mating modes. This suggests that whether crabs mate underground (or on the surface) is determined by the presence (or absence) of searching females. We found that the change in the mating mode affected the level of assortative mating; males mating underground were bigger than those mating on the surface, suggesting active female choice. Given that fiddler crabs experience multiple reproductive cycles, they are prone to showing behavioral plasticity in their mating strategy whenever the payoffs of using different mating modes differ between reproductive events. Our results suggest that the incorporation of different levels of environmental variability may be important in theoretical models aimed at improving our understanding of the evolution of alternative mating tactics and strategies.

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

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

Linking the scientific knowledge on marine frontal systems with ecosystem services.

Primary production hotspots in the marine environment occur where the combination of light, turbulence, temperature and nutrients makes the proliferation of phytoplankton possible. Satellite-derived surface chlorophyll-a distributions indicate that these conditions are frequently associated with sharp water mass transitions named "marine fronts". Given the link between primary production, consumers and ecosystem functions, marine fronts could play a key role in the production of ecosystem services (ES). Using the shelf break front in the Argentine Sea as a study case, we show that the high primary production found in the front is the main ecological feature that supports the production of tangible (fisheries) and intangible (recreation, regulation of atmospheric gases) marine ES and the reason why the provision of ES in the Argentine Sea concentrates there. This information provides support to satellite chlorophyll as a good indicator of multiple marine ES. We suggest that marine fronts could be considered as marine ES hot spots.

Beyond competition: the stress-gradient hypothesis tested in plant-herbivore interactions.

The stress-gradient hypothesis predicts that interactions among plants are context dependent, shifting from facilitation to competition as environmental stress decreases. Although restricted to facilitation/competition, the mechanistic model behind the hypothesis is easily modified to include other negative interactions that are as important as competition in structuring natural communities, e.g., herbivory. To evaluate this hypothesis we experimentally tested if the balance between the facilitative and trophic effect of an intertidal, burrowing, herbivorous crab in marsh plants is context dependent and shifts from positive to negative as stress decreases. By sampling salt marshes differing in sediment size characteristics, we show that sites with larger sediment particle size had less stressful oxygen levels than sites with fine sediment particles, and that the level of stress was reduced by the presence of crab burrows. We then conducted a factorial experiment manipulating sediment size and crab presence. Results show that, by decreasing soil anoxic stress, crabs increase plant growth in stressful zones, but their ecological importance as herbivores increases in more benign zones. Our findings suggest that the balance between positive and negative interactions along stress gradients is more important than previously perceived and also applies to facilitation and herbivory between animals and plants.

Eutrophication in a semi-desert coastal ecosystem promotes increases in N and C isotopic signatures and changes in primary sources.

Using C and N isotopic signatures of food web components, we evaluated the land-marine coupling through nutrient flows and the likely changes in the food web structure in tidal channels with contrasting anthropogenic nutrient inputs at a semi desert-macrotidal coastal system (northern Argentine Patagonia). The results showed an increase in the δ13C signatures of primary producers and in the δ15N signatures in all levels of the benthic food web, from primary producers to predators, with possible changes in the relative contribution of primary food sources for consumer in the tidal channel with high anthropogenic N input. This is an example on the extent of the distribution of anthropogenic N into natural systems, flowing through the food web from terrestrial origin to coastal marine components.

Crab herbivory regulates plant facilitative and competitive processes in Argentinean marshes.

Interactions among plants have been hypothesized to be context dependent, shifting between facilitative and competitive in response to variation in physical and biological stresses. This hypothesis has been supported by studies of the importance of positive and negative interactions along abiotic stress gradients (e.g., salinity, desiccation), but few studies have tested how variation in biotic stresses can mediate the nature and strength of plant interactions. We examined the hypothesis that herbivory regulates the strength of competitive and facilitative interactions during succession in Argentinean marshes dominated by Spartina densiflora and Sarcocornia perennis. Spartina densiflora is preferred by the dominant herbivore in the system, the crab Chasmagnathus granulatus. We experimentally manipulated crab herbivory, plant structure, and shade, and we found that, when herbivory was low in the spring and summer, competitive interactions between plants were dominant, but in the fall, when herbivory was highest, facilitative interactions dominated, and Spartina densiflora survival was completely dependent upon association with Sarcocornia perennis. Moreover, experimental removal of Sarcocornia perennis across recently disturbed tidal flats revealed that, while Sarcocornia perennis positively affected small Spartina densiflora patches by decreasing herbivory, as patch size increases and they can withstand the impact of herbivory, competitive interactions predominated and Spartina densiflora ultimately outcompeted Sarcocornia perennis. These results show that herbivory can mediate the balance between facilitative and competitive processes in vascular plant communities and that the strength of consumer regulation of interactions can vary seasonally and with patch size.

Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands-those with both species-rich local communities (α-diversity) and large compositional differences among localities (ß-diversity)-had higher levels of multifunctionality. Moreover, α- and ß-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.