Carpinteria Salt Marsh Research Articles

Marsh Madness: Using Video Games and a Case Study to Explore Food Webs and Ecosystem Services in Carpinteria Salt Marsh Reserve

Marsh Madness: Using Video Games and a Case Study to Explore Food Webs and Ecosystem Services in Carpinteria Salt Marsh Reserve

Shifts in Salt Marsh Vegetation Landcover after Debris Flow Deposition

On 9 January 2018, Carpinteria Salt Marsh Reserve received a large quantity of sediment following debris flows in Montecito, California. Because disturbances potentially impact the ecosystem services and functions that wetlands provide, an understanding of how the ecosystem responded to the debris flows is important for the management of salt marsh systems. However, a lack of field data before and after this disturbance makes this task impossible to complete by field methods alone. To address this gap, we used Sentinel-2 satellite imagery to calculate landcover fractions and spectral indices to produce maps of landcover before, during, and after the debris flow using a random forest classifier. Change detection showed that vegetation extent in November 2020 approached pre-debris flow conditions. While total vegetated area experienced little net change (0.15% decrease), there was a measurable change in the areal extent of vegetation type, with high marsh vegetation transitioning to mid marsh vegetation in regions that initially showed an increase in bare soil cover. These results are uniquely quantifiable using remote sensing techniques and show that disturbance due to debris flows may affect ecosystem function via plant community change. These impacts will need to be taken into consideration when managing wetlands prone to depositional events.

Analyzing ecosystem services as part of ecological networks in three salt marsh ecosystems.

Biodiversity plays important roles in nature's contributions to people (i.e., ecosystem services), but the critical details of how biodiversity contributes are challenging to determine. Efforts to identify the components of an ecosystem that provide services have improved our understanding of which species, functional groups, population, or habitats directly provide services. However, species do not exist in isolation and considerably less is known about how species indirectly influence ecosystem services through interacting with those species directly providing services. This uncertainty is even greater when considering that species interact in complex networks. As such, detailed analyses of species interdependencies are rarely included in ecosystem services assessments or conservation decisions. To date, most studies on food webs and on ecosystem services have developed largely in parallel for many services, but these fields and data are ripe for empirical integration. To further this integration, we compiled data sets that linked three existing ecological networks to seven ecosystem functions and services: wave attenuation, shoreline stabilization, carbon sequestration, water filtration, fisheries, birdwatching, and waterfowl hunting. We leveraged high-resolution ecological interaction network data sets from three coastal salt marsh ecosystems including detailed species information (e.g., consumer strategy, body size, biomass) on several hundred species from Carpinteria Salt Marsh in California, USA, and for Estero de Punta Banda and Bahia Falsa in Baja, Mexico from Hechinger et al. (2011). Through an extensive literature synthesis and use of citizen science data, we identified which species in the Hechinger et al. (2011) data provided each ecosystem services directly. We augmented the Hechinger et al. (2011) data published in Ecology, particularly the link (or edge) list to include species-service links to indicate a species providing a service, in which species are listed as "Resources" and services are listed as "Consumers." Connecting these data to the previously published ecological networks with species interactions (i.e., trophic, parasitism) formed a topological network with species and service nodes. We also provided a protocol for assigning services to ecological networks that can be used in other ecosystems. This data set provides a step toward advancing the knowledge of important supporting species for ecosystem services and to developing new ecological network methods for ecosystem services. There are no copyright restrictions; please cite this data paper when the data are used in publications.

Mapping Coastal Wetland Biomass from High Resolution Unmanned Aerial Vehicle (UAV) Imagery

Salt marsh productivity is an important control of resiliency to sea level rise. However, our understanding of how marsh biomass and productivity vary across fine spatial and temporal scales is limited. Remote sensing provides a means for characterizing spatial and temporal variability in marsh aboveground biomass, but most satellite and airborne sensors have limited spatial and/or temporal resolution. Imagery from unmanned aerial vehicles (UAVs) can be used to address this data gap. We combined seasonal field surveys and multispectral UAV imagery collected using a DJI Matrice 100 and Micasense Rededge sensor from the Carpinteria Salt Marsh Reserve in California, USA to develop a method for high-resolution mapping of aboveground saltmarsh biomass. UAV imagery was used to test a suite of vegetation indices in their ability to predict aboveground biomass (AGB). The normalized difference vegetation index (NDVI) provided the strongest correlation to aboveground biomass for each season and when seasonal data were pooled, though seasonal models (e.g., spring, r2 = 0.67; RMSE = 344 g m−2) were more robust than the annual model (r2 = 0.36; RMSE = 496 g m−2). The NDVI aboveground biomass estimation model (AGB = 2428.2 × NDVI + 120.1) was then used to create maps of biomass for each season. Total site-wide aboveground biomass ranged from 147 Mg to 205 Mg and was highest in the spring, with an average of 1222.9 g m−2. Analysis of spatial patterns in AGB demonstrated that AGB was highest in intermediate elevations that ranged from 1.6–1.8 m NAVD88. This UAV-based approach can be used aid the investigation of biomass dynamics in wetlands across a range of spatial scales.

Nutrient Fluctuations in Marine Systems: Press Versus Pulse Nutrient Subsidies Affect Producer Competition and Diversity in Estuaries and Coral Reefs

Resource subsidy regimes, which range from presses to pulses, are common structuring forces in communities, yet research contrasting their effects is lacking. Many coastal marine ecosystems, including estuaries and coral reefs, have experienced increased nutrient subsidies while concurrently shifting to macroalgal dominance; however, the role of subsidy regime in transitions remains unknown. We created concentration–frequency distributions of nutrients in Cook’s Bay, Moorea, French Polynesia, and Carpinteria Salt Marsh Reserve, CA, USA. Both showed relatively high pulses interspersed by press concentrations. We grew dominant macroalgae alone and together in microcosms approximating these subsidy regimes to quantify individual performance and competitive outcomes. Subsidy regime changed growth and competitive abilities of macroalgae from both ecosystems but with divergent effects. In nutrient-limited reefs, different species were favoured under each enrichment regime, suggesting a fluctuating nutrient environment enhances diversity. In contrast, in eutrophic estuaries, enrichment of both regimes facilitated a single competitive dominant, suppressing diversity. Functional form groups did not predict responses to subsidy regime, likely because classifications ignore temporal variability in resource supplies. Because climate change will alter rainfall patterns globally, further accelerating nutrient subsidies from land to sea, understanding species’ responses to nutrient subsidy regimes is key to predicting the fate of coastal communities.

Host density increases parasite recruitment but decreases host risk in a snail-trematode system.

Most species aggregate in local patches. High host density in patches increases contact rate between hosts and parasites, increasing parasite transmission success. At the same time, for environmentally transmitted parasites, high host density can decrease infection risk to individual hosts, because infective stages are divided among all hosts in a patch, leading to safety in numbers. We tested these predictions using the California horn snail, Cerithideopsis californica (=Cerithidea californica), which is the first intermediate host for at least 19 digenean trematode species in California estuaries. Snails become infected by ingesting trematode eggs or through penetration by free-swimming miracidia that hatch from trematode eggs deposited with final-host (bird or mammal) feces. This complex life cycle decouples infective-stage production from transmission, raising the possibility of an inverse relationship between host density and infection risk at local scales. In a field survey, higher snail density was associated with increased trematode (infected snail) density, but decreased trematode prevalence, consistent with either safety in numbers, parasitic castration, or both. To determine the extent to which safety in numbers drove the negative snail-density-trematode-prevalence association, we manipulated uninfected snail density in 83 cages at eight sites within Carpinteria Salt Marsh (California, USA). At each site, we quantified snail density and used data on final-host (bird and raccoon) distributions to control for between-site variation in infective-stage supply. After three months, overall trematode infections per cage increased with snail biomass density. For egg-transmitted trematodes, per-snail infection risk decreased with snail biomass density in the cage and surrounding area, whereas per-snail infection risk did not decrease for miracidium-transmitted trematodes. Furthermore, both trematode recruitment and infection risk increased with infective-stage input, but this was significant only for miracidium-transmitted species. A model parameterized with our experimental results and snail densities from 524 field transects estimated that safety in numbers, when combined with patchy host density, halved per capita infection risk in this snail population. We conclude that, depending on transmission mode, host density can enhance parasite recruitment and reduce per capita infection risk.

Salt Marsh Reduces Fecal Indicator Bacteria Input to Coastal Waters in Southern California

Abstract We investigated fecal indicator bacteria (FIB) concentrations in water and sediment from Carpinteria Salt Marsh, a medium-sized (93 ha), mostly natural southern California coastal wetland. High FIB concentrations, exceeding recreational water quality standards, were found at inlet sites after winter storm events and during a summer dry weather sampling event. Runoff entering the wetland had the highest concentrations of FIB after large rain events and after rain events following extended periods without rain. The watersheds with the greatest agricultural and urban development draining into the wetland generally contributed the highest loads of FIB, while the largest and least developed watershed contributed the lowest FIB concentrations. Surface water exiting the wetland at the ocean contained relatively low concentrations of FIB and only exceeded recreational water quality standards after the largest rain event of the year. Bacterial concentrations in sediment were only elevated after rain event...

Two Myxozoans from the Urinary Tract of Topsmelt, Atherinops affinis.

Two myxozoan species were observed in the kidney of topsmelt, Atherinops affinis , during a survey of parasites of estuarine fishes in the Carpinteria Salt Marsh Reserve, California. Fish collected on 3 dates in 2012 and 2013 were sectioned and examined histologically. Large extrasporogonic stages occurred in the renal interstitium of several fish from the first 2 collections (5/8, 11/20, respectively) and, in some fish, these replaced over 80% of the kidney. In addition, presporogonic and polysporogonic stages occurred in the lumen of the renal tubules, collecting ducts, and mesonephric ducts. The latter contained subspherical spores with up to 4 polar capsules, consistent with the genus Chloromyxum. For the third collection (15 May 2013, n = 30), we portioned kidneys for examination by histology, wet mount, and DNA extraction for small subunit ribosomal (SSU rDNA) gene sequencing. Histology showed the large extrasporogonic forms in the kidney interstitium of 3 fish and showed 2 other fish with subspherical myxospores in the lumen of the renal tubules with smooth valves and 2 spherical polar capsules consistent with the genus Sphaerospora. Chloromyxum-type myxospores were observed in the renal tubules of 1 fish by wet mount. Sequencing of the kidney tissue from this fish yielded a partial SSU rDNA sequence of 1,769 base pairs (bp). Phylogenetic reconstruction suggested this organism to be a novel species of Chloromyxum, most similar to Chloromyxum careni (84% similarity). In addition, subspherical myxospores with smooth valves and 2 spherical polar capsules consistent with the genus Sphaerospora were observed in wet mounts of 2 fish. Sequencing of the kidney tissue from 1 fish yielded a partial SSU rDNA sequence of 1,937 bp. Phylogenetic reconstruction suggests this organism to be a novel species of Sphaerospora most closely related to Sphaerospora epinepheli (93%). We conclude that these organisms represent novel species of the genera Chloromyxum and Sphaerospora based on host, location, and SSU rDNA sequence. We further conclude that the formation of large, histozoic extrasporogonic stages in the renal interstitium represents developmental stages of Chloromyxum species for the following reasons: (1) Large extrasporogonic stages were only observed in fish with Chloromyxum-type spores developing within the renal tubules, (2) a DNA sequence consistent with the Chloromyxum sp. was only detected in fish with the large extrasporogonic stages, and (3) several Sphaerospora species have extrasporogonic forms, but they are considerably smaller and are composed of far fewer cells.

Food webs including parasites, biomass, body sizes, and life stages for three California/Baja California estuaries

This data set presents food webs for three North American Pacific coast estuaries and a “Metaweb” composed of the species/stages compiled from all three estuaries. The webs have four noteworthy attributes: (1) parasites (infectious agents), (2) body-size information, (3) biomass information, and (4) ontogenetic stages of many animals with complex life cycles. The estuaries are Carpinteria Salt Marsh, California (CSM); Estero de Punta Banda, Baja California (EPB); and Bahia Falsa in Bahia San Quintin, Baja California (BSQ). Most data on species assemblages and parasitism were gathered via consistent sampling that acquired body size and biomass information for plants and animals larger than ∼1 mm, and for many infectious agents (mostly metazoan parasites, but also some microbes). We augmented this with information from additional published sources and by sampling unrepresented groups (e.g., plankton). We estimated free-living consumer–resource links primarily by extending a previously published version of t...

Parasites reduce food web robustness because they are sensitive to secondary extinction as illustrated by an invasive estuarine snail

A robust food web is one in which few secondary extinctions occur after removing species. We investigated how parasites affected the robustness of the Carpinteria Salt Marsh food web by conducting random species removals and a hypothetical, but plausible, species invasion. Parasites were much more likely than free-living species to suffer secondary extinctions following the removal of a free-living species from the food web. For this reason, the food web was less robust with the inclusion of parasites. Removal of the horn snail, Cerithidea californica, resulted in a disproportionate number of secondary parasite extinctions. The exotic Japanese mud snail, Batillaria attramentaria, is the ecological analogue of the native California horn snail and can completely replace it following invasion. Owing to the similarities between the two snail species, the invasion had no effect on predator–prey interactions. However, because the native snail is host for 17 host-specific parasites, and the invader is host to only one, comparison of a food web that includes parasites showed significant effects of invasion on the native community. The hypothetical invasion also significantly reduced the connectance of the web because the loss of 17 native trematode species eliminated many links.

Ecosystem consequences of fish parasites*

In most aquatic ecosystems, fishes are hosts to parasites and, sometimes, these parasites can affect fish biology. Some of the most dramatic cases occur when fishes are intermediate hosts for larval parasites. For example, fishes in southern California estuaries are host to many parasites. The most common of these parasites,Euhaplorchis californiensis, infects the brain of the killifishFundulus parvipinnisand alters its behaviour, making the fish 10–30 times more susceptible to predation by the birds that serve as its definitive host. Parasites likeE. californiensisare embedded in food webs because they require trophic transmission. In the Carpinteria Salt Marsh estuarine food web, parasites dominate the links and comprise substantial amount of biomass. Adding parasites to food webs alters important network statistics such as connectance and nestedness. Furthermore, some free‐living stages of parasites are food items for free‐living species. For instance, fishes feed on trematode cercariae. Being embedded in food webs makes parasites sensitive to changes in the environment. In particular, fishing and environmental disturbance, by reducing fish populations, may reduce parasite populations. Indirect evidence suggests a decrease in parasites in commercially fished species over the past three decades. In addition, environmental degradation can affect fish parasites. For these reasons, parasites in fishes may serve as indicators of environmental impacts.

ENDANGERED LIGHT‐FOOTED CLAPPER RAIL AFFECTS PARASITE COMMUNITY STRUCTURE IN COASTAL WETLANDS

An extinction necessarily affects community members that have obligate relationships with the extinct species. Indirect or cascading effects can lead to even broader changes at the community or ecosystem level. However, it is not clear whether generalist parasites should be affected by the extinction of one of their hosts. We tested the prediction that loss of a host species could affect the structure of a generalist parasite community by investigating the role of endangered Light-footed Clapper Rails (Rallus longirostris levipes) in structuring trematode communities in four tidal wetlands in southern California, U.S.A. (Carpinteria Salt Marsh, Mugu Lagoon) and Mexico (Estero de Punta Banda, Bahia Falsa-San Quintin). We used larval trematode parasites in first intermediate host snails (Cerithidea californica) as windows into the adult trematodes that parasitize Clapper Rails. Within and among wetlands, we found positive associations between Clapper Rails and four trematode species, particularly in the vegetated marsh habitat where Clapper Rails typically occur. This suggests that further loss of Clapper Rails is likely to affect the abundance of several competitively dominant trematode species in wetlands with California horn snails, with possible indirect effects on the trematode community and changes in the impacts of these parasites on fishes and invertebrates.

Characterizing patterns of plant distribution in a southern California salt marsh using remotely sensed topographic and hyperspectral data and local tidal fluctuations

We used LiDAR topographic data, AVIRIS hyperspectral data, and locally measured tidal fluctuations to characterize patterns of plant distribution within a southern California salt marsh (Carpinteria Salt Marsh (CSM)). LiDAR data required ground truthing and correction before they were suitable for use. Twenty to forty percent of the uncertainty associated with LiDAR was due to variance in the elevation of the target surface, the balance was attributed to error inherent in the LiDAR system. The incidence of LiDAR penetration of plant canopy cover (i.e., registration of ground elevation) was only three percent. The depth of LiDAR penetration into the plant canopy varied according to plant species composition; plant species-specific corrections significantly improved LiDAR accuracy (58% reduction in overall uncertainty) and with the use of ground-based surveys, reduced overall RMSE to an average of 6.3 cm in vegetated areas. A supervised classification of AVIRIS data was used to generate a vegetation map with six classification types; overall classification accuracy averaged 59% with a kappa coefficient of 0.40. The vegetation classification map was overlaid with a LiDAR-based digital elevation model (DEM) to compute elevation distributions and frequencies of tidal inundation. The average elevations of the dominant plant classifications found in CSM (e.g., Salicornia virginica, Jaumea carnosa, and salt-grass mix, a mixture of multiple marsh plant species) occurred within a 17 cm range, a vertical change that resulted in a 7% difference in the period of tidal inundation.

USING OPPORTUNISTIC GREEN MACROALGAE AS INDICATORS OF NITROGEN SUPPLY AND SOURCES TO ESTUARIES

Nutrient inputs to estuaries are increasing worldwide, and anthropogenic contributions are increasingly complex and difficult to distinguish. Measurement of integrated effects of salinity and nutrient changes simultaneously can help ascertain whether N sources of similar magnitude and stable isotope (sigma15N) signatures are river dominated. We used Enteromorpha spp., an opportunistic macroalga, to obtain integrated measures of salinity, nutrient supply, and nutrient source in estuaries. We outplanted cultured algae in the field along spatial gradients within three southern California estuaries for 24 hours in wet and dry seasons. Tissue was analyzed for potassium (K+) to measure osmoregulatory changes, total nitrogen to examine changes in nutrient supply, and sigma15N to assess nutrient sources. Discrete measures of water salinity correlated with tissue K+ content; however, there was significant variability in the relationship, suggesting that the algae were subject to considerable variation in salinity over a tidal cycle. Tissue total N was not always related to snapshot measures of water column dissolved inorganic nitrogen (DIN), suggesting that integrated measures may be better at capturing the temporal and spatial complexity of nutrient availability. The combination of tissue K+, total N, and sigma15N measures revealed that inflowing rivers delivered N from watershed sources to Mugu Lagoon and Carpinteria Salt Marsh, while both the inflowing river and a mid estuary source were important sources of high sigma15N N in Upper Newport Bay. These experiments revealed complex patterns of supply and sources of N and demonstrate the usefulness of macroalgal indicators over water sampling to detect these patterns.

Trematodes in Snails near Raccoon Latrines Suggest a Final Host Role for this Mammal in California Salt Marshes

Of the 18 trematode species that use the horn snail, Cerithidea californica, as a first intermediate host, 6 have the potential to use raccoons as a final host. The presence of raccoon latrines in Carpinteria Salt Marsh, California, allowed us to investigate associations between raccoons and trematodes in snails. Two trematode species, Probolocoryphe uca and Stictodora hancocki, occurred at higher prevalences in snails near raccoon latrines than in snails away from latrines, suggesting that raccoons may serve as final hosts for these species. Fecal remains indicated that raccoons fed on shore crabs, the second intermediate host for P. uca, and fish, the second intermediate host for S. hancocki. The increase in raccoon populations in the suburban areas surrounding west coast salt marshes could increase their importance as final hosts for trematodes in this system.

USING LARVAL TREMATODES THAT PARASITIZE SNAILS TO EVALUATE A SALTMARSH RESTORATION PROJECT

We conducted a Before‐After‐Control‐Impact (BACI) study using larval digeneans infecting the California horn snail, Cerithidea californica, to evaluate the success of an ecological restoration project at Carpinteria Salt Marsh in California, USA. Digenean trematodes are parasites with complex life cycles requiring birds and other vertebrates as final hosts. We tested two hypotheses for prevalence and species richness of larval trematodes in C. californica: (1) prior to the restoration, sites to be restored would have lower trematode prevalence and species richness relative to unimpacted control sites, and (2) that these differences would diminish after restoration. The sites to be restored were initially degraded for trematode species. They had a mean trematode prevalence (12%) and species richness (4.5 species) that were lower than control sites (28% trematode prevalence and 7 species). Despite the differences in prevalence, the proportional representation of each trematode species in the total community was similar between sites to be restored and control sites. Over the six years following restoration, trematode prevalence nearly quadrupled at restored sites (43%) while the prevalence at control sites (26%) remained unchanged. In addition, species richness at restored sites doubled (9 species), while species richness at the control sites (7.8 species) did not change. Immediately after restoration, the relative abundance of trematode species using fishes as second intermediate hosts declined while those using molluscs as second intermediate hosts increased. Trematode communities at restored and control sites gradually returned to being similar. We interpret the increase in trematode prevalence and species richness at restored sites to be a direct consequence of changes in bird use of the restored habitat. This study demonstrates a new comparative technique for assessing wetlands, and while it does not supplant biotic surveys, it informs such taxonomic lists. Most importantly, it provides a synthetic quantification of the linkages among species in wetland food webs.

Importance of Vascular Plant and Algal Production to Macro-invertebrate Consumers in a Southern California Salt Marsh

The dietary importance of marsh vascular plants (primarilySalicornia virginica), algae and upland particulate inputs to macro-invertebrate consumers was studied in Carpinteria Salt Marsh, southern California, using stable carbon and nitrogen isotope ratios. This marsh is predominantly a marine or hypersaline system and succulents are the most common vascular plant species. Of invertebrates collected from the vegetated marsh, tidal flats and channels, only detritivores from the vegetated marsh (Traskorchestia traskiana,Melampus olivaceus) had isotope values (δ13C=−20‰) that suggested some use ofSalicornia-derived carbon.T. traskianacultured in the laboratory on decomposingS. virginicaor blue-green micro-algal mat had distinctive isotopic signatures, reflecting the capability of this consumer to assimilate carbon and nitrogen derived from these sources. The δ13C values (generally −16‰ to −15‰) of species from tidal flats and channels (e.g.Cerithidea californica,Protothaca staminea,Mytilus galloprovincialis,Neotrypaea californiensis) were most similar to values for benthic algae and phytoplankton. Specimens ofM. galloprovincialisalong a gradient of presumed increase in marine influence had similar isotope values, suggesting little contribution to diet from upland runoff. The present results differ most noticeably from published values in the13C enrichment of suspension-feeders, suggesting the use of resuspended13C-enriched benthic microalgae in tidal channels by these consumers, and in the13C depletion and15N enrichment of plants and consumers along a portion of the marsh boundary receiving inputs of nutrient-enriched perched groundwater. In general, the isotopic composition of macro-invertebrates indicated the incorporation of algal production rather than ofS. virginicaor upland sources into the marsh food web.

Influence of Watershed Runoff on Nutrient Dynamics in a Southern California Salt Marsh

The effect of freshwater runoff on nutrient loading and dynamics was studied in Carpinteria Salt Marsh, a typically small (93 ha) southern California, U.S.A., estuary adjoining a watershed supporting extensive agricultural and urban development. Concentrations of dissolved nitrate, but not ammonium or phosphate, were elevated in stream flow and perched groundwater in the watershed, relative to marsh surface waters. Loading of nitrate and particulate organic nitrogen (PON) from the watershed, but not ammonium or phosphate, increased as a function of stream discharge. Estimates of net nutrient exchange over selected tidal cycles suggested that the marsh exported nitrate, ammonium and PON to the Santa Barbara Channel. Nitrate and PON were advected from the watershed through the marsh while ammonium was produced within the marsh and exported. The availability of imported nitrogen for primary production may be limited by marsh tidal elevation as most nitrogen inputs are throughput to the Santa Barbara Channel.

Effects of variable precipitation on the structure and diversity of a California salt marsh community

Abstract. We measured the effects of annual variation in climate and experimentally augmented rainfall on patterns of distribution and above‐ground productivity in annual plant communities at Carpinteria Salt Marsh in central California. In the driest year, Hutchinsia procumbens was codominant throughout much of the upper marsh; however, Hutchinsia was very rare or not present in the wetter years. Conversely, Juncus bufonius was common in the wettest year and absent in the driest year. Elevational distributions of other annual species also differed among years with different total precipitation. In 1989–1990, an exceptionally dry season, supplemental water decreased soil salinity, increased above‐ground productivity of annuals, and caused significant changes in spatial patterns and relative density. In the lowest zone, Hutchinsia occurred only in watered plots and supplemental water increased the density of Spergularia marina. At intermediate elevations Lasthenia glabrata occurred only in watered plots and supplemental water increased the density of Spergularia, Hutchinsia, and Parapholis incurva. At upper elevations, Juncus occurred only in watered plots, and Lasthenia was the only species that increased significantly in density with watering. Unlike natural shifts in species abundance, no species declined significantly in cover in any zone in the watered treatment. Although climatic variation has complex affects on annual plant communities, our experiments isolated important affects of total annual rainfall on the structure of annual plant communities that were similar to those that occurred with natural variation in rainfall. We conclude that variation in total annual precipitation promotes dynamic community composition and spatial distributions among years, and thus increases overall species diversity in the salt marsh.

Comparative effects of infaunal bivalves on an epibenthic microalgal community

The effects of three infaunal bivalves on an epibenthic microalgal community were studied in a tidal channel at Carpinteria Salt Marsh, California, USA. Sediment Chl a concentration was patchy, ranging from ≈ 1 to ≈ 5.5 μd·cm −2. Chl a concentrations in 78.5-cm 2 sample plots was negatively correlated with the density of the deposit-feeder Macoma nasuta, but not the suspension-feeders Protothaca staminea and Tagelus californianus within those plots. In field experiments, M. nasuta inhibited the development of microalgal standing stock relative to P. staminea and Control treatments. The effect of M. nasuta on microalgae varied with clam density. M. nasuta also reduced existing microalgal standing stock but without affecting the relative abundance of common diatom species. Our results indicate that the random distribution of the deposit-feeding M. nasuta contributed to small-scale patchiness in microalgal standing stock, locally reducing the availability of microalgae to infaunal as well as epifaunal benthic consumers.