Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

Relative and interactive effects of plant and grazer richness in a benthic marine community

The interactive effects of changing biodiversity of consumers and their prey are poorly understood but are likely to be important under realistic scenarios of biodiversity loss and gain. We performed two factorial manipulations of macroalgal group (greens, reds, and browns) and herbivore species (amphipods, sea urchin, and fish) composition and richness in outdoor mesocosms simulating a subtidal, hard-substratum estuarine community in North Carolina, U.S.A. In the experiment where grazer richness treatments were substitutive, there were no significant effects of algal or herbivore richness on final algal biomass. However, in the experiment in which grazer treatments were additive (i.e., species-specific densities were held constant across richness treatments), we found strong independent and interactive effects of algal and herbivore richness. Herbivore polycultures reduced algal biomass to a greater degree than the sum of the three herbivore monocultures, indicating that the measured grazer richness effects were not due solely to increased herbivore density in the polycultures. Taking grazer density into account also revealed that increasing algal richness dampened grazer richness effects. Additionally, the effect of algal richness on algal biomass accumulation was far stronger when herbivores were absent, suggesting that grazers can utilize the increased productivity and mask the positive effects of plant biodiversity on primary production. Our results highlight the complex independent and interactive effects of biodiversity between adjacent trophic levels and emphasize the importance of performing biodiversity-ecosystem functioning experiments in a realistic multi-trophic context.     

Effects of species diversity on community biomass production change over the course of succession

Over the past decade an increasing number of studies have experimentally manipulated the number of species in a community and examined how this alters the aggregate production of species biomass. Many of these studies have shown that the effects of richness on biomass change through time, but we have limited understanding of the mechanisms that produce these dynamic trends. Here we report the results of an experiment in which we manipulated the richness of freshwater algae in laboratory microcosms. We used two experimental designs (additive and substitutive) that make different assumptions about how patches are initially colonized, and then tracked the development of community biomass from the point of initial colonization through a period of 6-12 generations of the focal species. We found that the effect of initial species richness on biomass production qualitatively shifted twice over the course of the experiment. The first shift occurred as species transitioned from density-independent to dependent phases of population growth. At this time, intraspecific competition caused monocultures to approach their respective carrying capacities more slowly than polycultures. As a consequence, species tended to over-yield for a brief time, generating a positive, but transient effect of diversity on community biomass. The second shift occurred as communities approached carrying capacity. At this time, strong interspecific interactions caused biomass to be dominated by the competitively superior species in polycultures. As this species had the lowest carrying capacity, a negative effect of diversity on biomass resulted in late succession. Although these two shifts produced dynamics that appeared complex, we show that the patterns can be fit to a simple Lotka-Volterra model of competition. Our results suggest that the effects of algal diversity on primary production change in a predictable sequence through successional time.     

Direct and indirect effects of CO2, nitrogen, and community diversity on plant-enemy interactions

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.     

A mathematical model of competition and colonization in a community of marine benthic algae

A mathematical model has been constructed for the algal community on the rocky shores of a Norwegian fjord. We report here on the studies of competition and colonization along a vertical transect from the upper intertidal to the sublittoral habiats. Results on species abundance and distribution (patterns of zonation) and time to reach maturity have been compared to observations both in the fjord area and in other rocky shore areas.Competition coefficients for the algae were inferred from plant morphology and shown to be in agreement with observations of algal abundance and their zone-forming ability. Competition restricts the distribution of the species, especially at the lower elevations, but does not alter their relative position. However, increasing uniform competition prolongs the time in which zone-forming can occur, and it also decreases the overall biomass which an area can sustain. Colonization by a single species may create transient stages in community development of the same order of magnitude as algae longevity, and probably also alters the zonation pattern to some degree.The simulation results indicate that the large-scale algal distribution pattern in the Hardangerfjord area results from global stability of the rocky shore community.     

Forest age influences oak insect herbivore community structure, richness, and density.

Plant succession is one of many factors that may affect the composition and structure of herbivorous insect communities. However, few studies have examined the effect of forest age on the diversity and abundance of insect communities. If forest age influences insect diversity, then the schedule of timber harvest rotation may have consequent effects on biodiversity. The insect herbivore community on Quercus alba (white oak) in the Missouri Ozarks was sampled in a chronoseries, from recently harvested (2 yr) to old-growth (approximately 313 yr) forests. A total of nine sites and 39 stands within those sites were sampled in May and August 2003. Unique communities of plants and insects were found in the oldest forests (122-313 yr). Density and species richness of herbivores were positively correlated with increasing forest age in August but not in May. August insect density was negatively correlated with heat load index; in addition, insect density and richness increased over the chronoseries, but not on the sunniest slopes. Forest structural diversity (number of size classes) was positively correlated with forest age, but woody plant species richness was not. In sum, richness, density, and community structure of white oak insect herbivores are influenced by variation in forest age, forest structure, relative abundance of plant species, and abiotic conditions. These results suggest that time between harvests of large, long-lived, tree species such as white oak should be longer than current practice in order to maintain insect community diversity.     

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.     

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Macro- and mesoherbivores prefer native seaweeds over the invasive brown seaweed <Emphasis Type="Italic">Sargassum muticum</Emphasis>: a potential regulating role on invasions

Herbivory has a strong impact on algal distribution, abundance and community structure and may influence the establishment and spread of introduced seaweed species. In this study, we assess the potential regulating role of herbivory on one of the most invasive brown seaweeds: Sargassum muticum. Multiple choice feeding experiments were conducted with 13 native seaweeds, S. muticum and 5 herbivore species from the Northwest, Southwest and South of Portugal. S. muticum was always the least or among the least preferred seaweeds and attained one of the highest growth rates of the tested seaweeds, with and without herbivores. The addition of herbivores increased the number of cases by 40% in which the invader had higher growth rates. Our results suggest that low grazing pressure on S. muticum by the recipient herbivore community may give the invader a competitive advantage over at least part of the native seaweed community, thereby contributing to the invasiveness of S. muticum along the Portuguese coast.     

Herbivores, tidal elevation, and species richness simultaneously mediate nitrate uptake by seaweed assemblages

In order for research into the consequences of biodiversity changes to be more applicable to real-world ecosystems, experiments must be conducted in the field, where a variety of factors other than diversity can affect the rates of key biogeochemical and physiological processes. Here, we experimentally evaluate the effects of two factors known to affect the diversity and composition of intertidal seaweed assemblages--tidal elevation and herbivory--on nitrate uptake by those assemblages. Based on surveys of community composition at the end of a 1.5-year press experiment, we found that both tide height and herbivores affected seaweed community structure. Not surprisingly, seaweed species richness was greater at lower tidal elevations. Herbivores did not affect richness, but they altered the types of species that were present; seaweed species characterized by higher rates of nitrate uptake were more abundant in herbivore-removal plots. Both tide height and herbivores affected nitrate uptake by seaweed assemblages. Individual seaweed species, as well as entire seaweed assemblages, living higher on the shore had greater rates of biomass-specific nitrate uptake, particularly at high ambient nitrate concentrations. Grazed seaweed assemblages exhibited reduced nitrate uptake, but only at low nitrate concentrations. We evaluated the effect of seaweed richness on nitrate uptake, both alone and after accounting for effects of tidal elevation and herbivores. When only richness was considered, we found no effect on uptake. However, when simultaneous effects of richness, tide height, and herbivores on uptake were evaluated, we found that all three had relatively large and comparable effects on nitrate uptake coefficients and that there was a negative relationship between seaweed richness and nitrate uptake. Particularly because effects of richness on uptake were not apparent unless the effects of tide height and herbivory were also considered, these results highlight the importance of considering the effects of environmental context when evaluating the consequences of biodiversity change in more realistic systems.     

Herbivores on a dominant understory shrub increase local plant diversity in rain forest communities

Indirect effects of trophic interactions on biodiversity can be large and common, even in complex communities. Previous experiments with dominant understory Piper shrubs in a Costa Rican rain forest revealed that increases in herbivore densities on these shrubs caused widespread seedling mortality as a result of herbivores moving from Piper to seedlings of many different plant genera. We tested components of the Janzen-Connell hypothesis by conducting focused studies on the effects of specialist and generalist Piper herbivores on local seedling diversity. Whereas specialist herbivores are predicted to increase mortality to neighboring seedlings that are closely related to the source plant, true generalists moving from source plants may cause density-dependent mortality of many species, and possibly increase richness if new species replace abundant species that have been thinned by herbivores. Therefore, we hypothesized that seedling richness would be greater in understory control plots created in patches of Piper that had normal densities of generalist herbivores compared to plots from which we removed generalist herbivores manually from all Piper shrubs. After 15 months, generalist-herbivore-removal plots had > 40% fewer seedlings, > 40% fewer species, and 40% greater seedling evenness, on average, than control plots with generalist herbivores intact. Using a complementary approach in unmanipulated plots in four forests, we used path analysis to test for a positive association between seedling diversity and herbivore damage on Piper species. In unmanipulated plots, for both generalist and specialist herbivores, our data were significant fits to the causal model that Piper herbivores decrease evenness and increase plant species richness, corroborating the experimental results. Because herbivores changed how individuals were apportioned among the species and families present (lower evenness), one interpretation of these associations between herbivores on Piper shrubs and local seedling richness is that high seedling mortality in dominant families allowed the colonization or survival of less common species. If interspecific or apparent competition allowed for a relative increase in species richness, then the Janzen-Connell hypothesis may extend its predictions to generalist seedling predators. We speculate that apparent competition may explain some of the deviations from neutral model predictions, especially at small scales.     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

Abundance of Fucus spiralis and ephemeral seaweeds in a high eulittoral zone: effects of grazers,canopy and substratum type

According to a widely accepted view of intertidal community organization in the north-west Atlantic Ocean, the abundance of perennial seaweeds is regulated by the activities of herbivores (which control ephemeral algal competitors) and carnivores (which control populations of competing filterfeeders). This concept was examined in 1987–1988 in a eulittoral belt on the Atlantic shores of Nova Scotia, Canada, dominated by a closed canopy of Fucus spiralis. A factorial design was used to test the effects of (a) grazers (there were no carnivores present, (b) adult F. spiralis canopy, and (c) substratum type on the abundance of perennial and ephemeral seaweeds. Grazers had no significant (p>0.05) effect on the density or cover of juvenile F. spiralis, but significantly reduced canopy cover of adults through the winter. The presence of grazers significantly enhanced the cover of ephemeral algae in early spring. Barnacle presence enhanced the cover of juvenile F. spiralis, but reduced the canopy of adult plants through abrasion. The most important regulator of recruitment density in F. spiralis was the presence of a canopy of conspecific adults. Canopy greatly reduced juvenile development. There was no significant relationship between the covers of perennial F. spiralis and ephemeral seaweeds. This study demonstrates that the recruitment of the perennial rockweed population examined is not dependent on the activities consumer animals. The results contrast with work in the midshore intertidal of New England, where the activities of herbivores are thought to regulate the abundances of perennial seaweeds.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

Intertidal community structure and oceanographic patterns around Santa Cruz Island, CA, USA

Recent studies suggest that nearshore oceanographic conditions can have important effects on the structure of benthic communities. On Santa Cruz Island (SCI), CA, USA there is a persistent difference in mean annual sea surface temperature (SST) around the island due to its location at the confluence of opposing cold and warm ocean current systems. Over the course of a 4-year study (1997–2001) seawater nutrient and chl-a concentrations, algal tissue C:N ratios, recruitment and growth of filter-feeders (barnacles and mussels), and intertidal community structure were measured at six intertidal sites around the island. There were strong associations between remotely sensed SST and patterns of community structure. Macrophyte abundance was highest at sites with persistently low SST, while recruitment, abundance, and growth of filter-feeding invertebrates were strongly, positively correlated with SST. The cold-water sites were associated with higher nutrient concentrations and lower algal C:N ratios, particularly in the winter months. Values of chl-a were generally low and variable among sites, and were not correlated with the predominant SST gradient. Recruitment of barnacles and mussels was positively correlated with adult abundance across all sites. While detailed experimental studies are needed to further evaluate the mechanisms underlying community dynamics, these results indicate that the confluence of cold- and warm-water masses around SCI may determine the contrasting patterns of intertidal community structure.     

Hemiparasites generate environmental heterogeneity and enhance species coexistence in salt marshes.

Tidal inundation and salinity are considered to be controlling factors in salt marsh species distributions. Parasitic plants may also influence community organization as parasite-host interactions may play a functional role in stress amelioration due to physiological mechanisms for salinity tolerance and resource acquisition. Endangered root hemiparasites (Cordylanthus maritimus ssp. palustris and Cordylanthus mollis ssp. mollis) occupy unique habitat within fragmented northern California tidal wetlands. My objective was to examine the effects of these root hemiparasites on soil salinity, aeration, and community composition. I compared experimentally established bare patches, shaded and unshaded, and parasite removal patches to controls with hemiparasites across intertidal elevation gradients. Plant community composition, soil salinity, and redox potential were measured as response variables. In this field removal experiment, I demonstrated that parasite-host associations can enhance the amelioration of physical stress conditions in the salt marsh exceeding the passive role of shading by vegetation. Consumer-driven reduction of physical stress resulted in increased plant species richness, and the effect was most pronounced with elevated salinity and hypoxia stress. Although previous studies have demonstrated that removal of dominant plant biomass by herbivores can increase physical stress in salt marshes, this is one of the first examples of a positive indirect effect of a consumer on community diversity through physical stress relief. Greater understanding of biological interactions coupled with abiotic factors may improve rare plant conservation and salt marsh restoration success.