Katharyn E. Boyer

Effects of macroalgal species identity and richness on primary production in benthic marine communities

Plant biodiversity can enhance primary production in terrestrial ecosystems, but biodiversity effects are largely unstudied in the ocean. We conducted a series of field and mesocosm experiments to measure the relative effects of macroalgal identity and richness on primary productivity (net photosynthetic rate) and biomass accumulation in hard substratum subtidal communities in North Carolina, USA. Algal identity consistently and strongly affected production; species richness effects, although often significent, were subtle. Partitioning of the net biodiversity effect indicated that complementarity effects were always positive and species were usually more productive in mixtures than in monoculture. Surprisingly, slow growing species performed relatively better in the most diverse treatments than the most productive species, thus selection effects were consistently negative. Our results suggest that several basic mechanisms underlying terrestrial plant biodiversity effects also operate in algal-based marine ecosystems, and thus may be general.

Elevated nutrient content of tropical macroalgae increases rates of herbivory in coral, seagrass, and mangrove habitats

We explored the role of food quality in herbivore preference for macroalgae by comparing consumption of Acanthophora spicifera with and without elevated tissue nitrogen and phosphorus concentrations. Algal enrichment effects on herbivory were examined in coral, seagrass, and mangrove habitats along a sparsely populated Honduran island protected from fishing. Nutrient enrichment led to significantly increased grazing by herbivores across habitats. Consumption of enriched algae increased by 91% compared to controls among the mangrove roots, where herbivory rates were generally lowest. In the heavily grazed seagrass and coral habitats, nutrient enrichment increased consumption by 30 and 20%, respectively, with the effect more spatially variable than among the mangrove roots. We suggest that, at least on the local scale, intact herbivore populations may be able to compensate for effects of increased nutrient supply by locating and consuming nutrient-enriched algae, but that the importance of this mechanism varies both among and within habitats.

Developing an indicator of nutrient enrichment in coastal estuaries and lagoons using tissue nitrogen content of the opportunistic alga, Enteromorpha intestinalis (L. Link)

Abstract We explored the use of an opportunistic green alga, Enteromorpha intestinalis (L. Link), as an indicator of N enrichment in a southern California salt marsh. In conjunction with N additions to cordgrass ( Spartina foliosa , Trin) in April, June and August 1995, mesh bags containing N-starved algal tissue were placed within cordgrass patches, at their edges along islands, and in adjacent channels. After 1 week in the field, recovered algal tissue was used to test detection of two levels of total N supply (one twice as high as the other), as well as no added N (control). Tissue N concentration, calculated as the percentage change in N, was the best of several algal measures at discerning differences in N availability in any month. In both April and June, tissue N declined from the marsh plain to the channels, reflecting declining N supply. Tissue N concentration also reflected differences in the total quantity of N added. Within the channels adjacent to fertilized areas, algal tissue N was similar to control areas, suggesting that N additions to cordgrass are not resulting in eutrophication of open waters. In August, the algae detected N additions on the marsh plain, but survivorship was poor; other algal species may be better indicators of enrichment in late-summer. With further investigation, the technique presented in this paper has the potential to be developed into a useful bioassay for detecting eutrophication of coastal salt marshes and lagoons.

EFFECTS OF NITROGEN ADDITIONS ON THE VERTICAL STRUCTURE OF A CONSTRUCTED CORDGRASS MARSH

Because tall vegetation can enhance habitat quality in intertidal wetlands, we examined the effects of N fertilization on the height growth of a constructed cordgrass (Spartina foliosa) marsh in San Diego Bay, where plants are short and soil N low. We varied the duration (therefore also the quantity; 30 g N/M2 every month for 1, 2, 4, or 6 mo) and timing (30 g N/M2 in March, April, June, or August) of N additions during 1993. By fertilizing a second set of replicates for the 6-mo treatment during 1994, we were able to examine the persistence of effects from 1993 vs. the effects of amendments in both years. Traditional canopy response measures (total stem length, maximum height, foliar N) increased most with the greatest duration/quantity of additions in 1993. All duration and timing treatments produced canopies with >100 stems/M2 and ?30 stems/M2 taller than 90 cm, considered suitable for nesting by the endangered Light-footed Clapper Rail (Rallus longirostris levipes). However, without additional fertilizer in 1994, the previous years canopy responses were not retained, suggesting insufficient storage of N in belowground tissues. Only plots fertilized in both years continued to produce ?30 tall stems/M2. The coarse dredge-spoil sediments of the constructed marsh had only 1/4 to 1/3 the total N found in the fine-textured soils of a nearby natural marsh, where cordgrass grows tall and clapper rails nest. We recommend that future marsh restoration projects begin with fine soils that promote N retention and accumulation to sustain tall canopies, rather than sug- gesting long-term fertilization.

Salinity stress, nitrogen competition, and facilitation: what controls seasonal succession of two opportunistic green macroalgae?

Differential tolerance of low salinity, competition for nitrogen (N), and facilitation by altering N supply all may act to determine the pattern of seasonal succession of Enteromorpha intestinalis (L.) Link and Ulva expansa (Setch) S. and G. in estuaries and lagoons of southern California. Low salinity negatively affected both of these algae. However, when N was in sufficient supply, salinities of 15 ppt favored E. intestinalis while oceanic salinity (35 ppt) favored U. expansa; neither alga had a clear advantage at 25 ppt. When starved of N, E. intestinalis and U. expansa competed directly for nutrients. When grown alone, they had similar N uptake and growth rates; when grown together, E. intestinalis was the superior competitor, negatively affecting growth of U. expansa. In addition, U. expansa facilitated the growth of E. intestinalis when N was in short supply; when grown together, there was a positive effect of U. expansa on E. intestinalis. The mechanism of this effect may have been the release or ‘leaking’ of DON when U. expansa no longer had sufficient tissue N to grow. Thus, E. intestinalis would be favored immediately after a rain, but would be replaced by U. expansa when N is available and tidal action reestablishes oceanic salinity. However, at the end of the rainy season when N becomes scarce, E. intestinalis would outcompete U. expansa. We hypothesize that U. expansa may facilitate the dominance of E. intestinalis by leaking N that can be assimilated by E. intestinalis.

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.

SALICORNIA VIRGINICA IN A SOUTHERN CALIFORNIA SALT MARSH: SEASONAL PATTERNS AND A NUTRIENT-ENRICHMENT EXPERIMENT

Salicornia virginica (common pickleweed) is the dominant vascular plant of many saline marshes of the US west coast, yet little is known about seasonal patterns or abiotic factors controlling it. In a southern California salt marsh, quarterly sampling revealed strong seasonal trends, with 2x greater S. virginica biomass in summer than in winter. Tissue nitrogen (N) and phosphorus (P) concentrations were highest in winter and lower in spring and summer, suggesting a dilution of nutrients as plants accumulated biomass during the growing season. Despite high sediment nutrient levels in this marsh, an experiment examining N and P effects still found strong S. virginica responses to N applied biweekly for > 1 year. Increases in succulent tissue biomass after N addition were first seen in April 1998 (after fertilization for 11 months); two-fold increases in biomass and the number of branches resulted by the end of the experiment in August 1998. Addition of N increased N concentration in the woody tissues when sampled in August. The N:P ratio increased with N addition beginning in winter (7 months after fertilization began) and continuing through the remainder of the experiment. Effects of P addition were less marked, as adding P did not result in biomass responses; however, it did influence tissue nutrient levels. These amendments increased P concentrations in the woody tissue in August 1998. In contrast to N amendments, which did not affect root nutrient concentrations, P addition led to increases in P content of root tissues in the latter portion of the growing season. These data suggest that increases in nutrients (especially N, but also P) can lead to large changes in S. virginica characteristics even in estuaries with high sediment nutrient levels.

Influence of physical processes on the design, functioning and evolution of restored tidal wetlands in California (USA)

The performance of two intertidal wetland mitigation projects constructed by the California Department of Transportation (Caltrans) in the Sweetwater Marsh National Wildlife Refuge (SMNWR) in San Diego Bay was evaluated over 5 years. Most of the Sweetwater wetland complex has been altered this century, including diking (with subsequent subsidence), filling, modification of the tidal regime, freshwater inflow and sediment fluxes. The mitigation project goals included a range of functional criteria intended to support two endangered bird species (light-footed clapper rail and California least tern) and one endangered plant (salt marsh birds-beak). While the mitigation projects have achieved some of the performance criteria established in the regulatory permits (particularly, those related to fish), vegetation criteria for one of the bird species have not been met. The initial grading (in relation to local tidal datums) should support the target plant species, but growth has been less than required. Shortcomings of the habitat include elevated soil and groundwater salinity, low nutrient levels (especially nitrogen, which is readily leached from the coarse substrate), and eroding topography (where a single oversized and overly sinous channel and the lower-than-natural marshpalin result in high velocity surface water flow and erosion). The failure to achieve a large plain at low-marsh elevations highlights the importance of a more complete understanding of the relationship between the site physical processes (topography, hydrology, climate, geomorphology), substrate conditions, and biotic responses.

WETLAND RESTORATION THRESHOLDS: CAN A DEGRADATION TRANSITION BE REVERSED WITH INCREASED EFFORT?

Previous attempts to reverse the degradation of a coastal wetland and restore nesting habitat for an endangered bird showed that adding nitrogen could temporarily increase the height of Spartina foliosa, but not produce self-sustaining tall canopies. We asked if increased effort (up to five years of N fertilization) would shift canopy attributes across the hypothesized threshold. Thirty plots were treated with 0–5 yr of urea addition, and all were followed for 5 yr. Canopies were robust while urea was being added, but Spartina reverted to short stature soon after fertilization ended, supporting R. J. Hobbs and D. A. Nortons concept of an irreversible transition. However, specific outcomes depended on the choice of response variable (six comparisons), the choice of reference data (initial conditions, same-year data, and pooled data), and the choice of statistical design (repeated measures vs. complete design), indicating the need to assess experiments thoroughly before making strong recommendations for ma...

Damage to Cordgrass by Scale Insects in a Constructed Salt Marsh: Effects of Nitrogen Additions

Because tall cordgrass (Spartina foliosa) is needed for nesting by the endangered light-footed, clapper rail, managers of constructed salt marshes in southern California are proposing large-scale nitrogen fertilization to improve cordgrass growth. How this might affect an existing infestation of scale insects (Haliaspis spartina) and the degree of damage these insects cause to their cordgrass hosts was unknown. We explored the effects of timing and duration of fertilization onHaliaspis damage to cordgrass, as well as the timing ofHaliaspis dispersal, in a constructed marsh at Sweetwater Marsh National Wildlife Refuge in San Diego Bay, California. Fertilization did not result in increasedHaliaspis abundance. After a large dispersal pulse in late May,Haliaspis establishment in the long-term fertilized plots was greater than in the controls; however, this trend reversed in August, when many more stems in the control plots were infested with large numbers ofHaliaspis. Since adultHaliaspis cannot leave a feeding site, losses of individuals in the fertilized plots were apparently due to mortality, perhaps resulting from mechanical or chemical changes in the fertilized plants or increased predation. Late in the growing season, plots fertilized with 10 applications of urea over 20 wk had the lowest meanHaliaspis abundance. Plots fertilized only in March, April, June, or August did not differ from controls in meanHaliaspis abundance.Haliaspis was never abundant in the fertilized or control plots in the adjacent natural marsh. This study suggests that fertilization, of constructed salt marshes in San Diego Bay may proceed without concern that furtherHaliaspis outbreaks will be facilitated.