Karen J. McGlathery

A stormflow/baseflow comparison of dissolved organic matter concentrations and bioavailability in an Appalachian stream

Patterns of dissolved organic carbon (DOC) and nitrogen (DON) delivery were compared between times of stormflow and baseflow in Paine Run, an Appalachian stream draining a 12.4 km2 forested catchment in the Shenandoah National Park (SNP), Virginia. The potential in-stream ecological impact of altered concentrations and/or chemical composition of DOM during storms also was examined, using standardized bacterial bioassays. DOC and DON concentrations in Paine Run were consistently low during baseflow and did not show a seasonal pattern. During storms however, mean DOC and DON concentrations approximately doubled, with maximum concentrations occurring on the rising limb of storm hydrographs. The rapid response of DOM concentration to changes in flow suggests a near-stream or in-stream source of DOM during storms. Stormflow (4% of the time, 36% of the annual discharge) contributed >50% of DOC, DON and NO3− flux in Paine Run during 1997. In laboratory bacterial bioassays, growth rate constants were higher on Paine Run stormflow water than on baseflow water, but the fraction of total DOM which was bioavailable was not significantly different. The fraction of the total stream DOC pool taken up by water column bacteria was estimated to increase from 0.03 ± 0.02% h−1 during baseflow, to 0.15 ± 0.04% h−1 during storms. This uptake rate would have a minimal effect on bulk DOM concentrations in Paine Run, but storms may still have considerable impact on the bacterial stream communities by mobilizing them into the water column and by supplying a pulse of DOM.

Habitat Cascades: The Conceptual Context and Global Relevance of Facilitation Cascades via Habitat Formation and Modification

The importance of positive interactions is increasingly acknowledged in contemporary ecology. Most research has focused on direct positive effects of one species on another. However, there is recent evidence that indirect positive effects in the form of facilitation cascades can also structure species abundances and biodiversity. Here we conceptualize a specific type of facilitation cascade-the habitat cascade. The habitat cascade is defined as indirect positive effects on focal organisms mediated by successive facilitation in the form of biogenic formation or modification of habitat. Based on a literature review, we demonstrate that habitat cascades are a general phenomenon that enhances species abundance and diversity in forests, salt marshes, seagrass meadows, and seaweed beds. Habitat cascades are characterized by a hierarchy of facilitative interactions in which a basal habitat former (typically a large primary producer, e.g., a tree) creates living space for an intermediate habitat former (e.g., an epiphyte) that in turn creates living space for the focal organisms (e.g., spiders, beetles, and mites). We then present new data on a habitat cascade common to soft-bottom estuaries in which a relatively small invertebrate provides basal habitat for larger intermediate seaweeds that, in turn, generate habitat for focal invertebrates and epiphytes. We propose that indirect positive effects on focal organisms will be strongest when the intermediate habitat former is larger and different in form and function from the basal habitat former. We also discuss how humans create, modify, and destroy habitat cascades via global habitat destruction, climatic change, over-harvesting, pollution, or transfer of invasive species. Finally, we outline future directions for research that will lead to a better understanding of habitat cascades.

CHANGES IN INTRACELLULAR NITROGEN POOLS AND FEEDBACK CONTROLS ON NITROGEN UPTAKE IN CHAETOMORPHA LINUM (CHLOROPHYTA)1

Changes in the size of intracellular nitrogen pools and the potential feedback by these pools on maximum N uptake (NH4+ and NO3−) rates were determined for Chaetomorpha linum (Müller) Kützing grown sequentially under nutrient‐saturating and nutrient‐limiting conditions. The size of individual pools in N‐sufficient algae could be ranked as residual organic N (RON) comprised mainly of amino acids and amino compounds > protein N > NO3− > NH4+ > chlorophyll N. When the external N supply was removed, growth rates remained high and individual N pools were depleted at exponential rates that reflected both dilution of existing pools by the addition of new biomass from growth and movement between the pools. Calculated fluxes between the tissue N pools showed that the protein pool increased throughout the N depletion period and thus did not serve a storage function. RON was the largest storage reserve; nitrate was the second largest, but more temporary, storage pool that was depleted within 10 days. Upon N resupply, the RON pool increased 3 × faster than either the inorganic or protein pools, suggesting that protein synthesis was the rate‐limiting step in N assimilation and caused a buildup of intermediate storage compounds. Maximum uptake rates for both NH4+ and NO3− varied inversely with macroalgal N status and appeared to be controlled by changes in small intracellular N pools. Uptake of NO3− showed an initial lag phase, but the initial uptake of NH4+ was enhanced and was present only when the intracellular NH4+ pool was depleted in the absence of an external N supply. A strong negative correlation between the RON pool size and maximum assimilation uptake rates for both NH4+ and NO3− suggested a feedback control on assimilation uptake by the buildup and depletion of organic compounds. Enhanced uptake and the accumulation of N as simple organic compounds or nitrate both provide a temporary mechanism to buffer against the asynchrony of N supply and demand in C. linum.

Wind-driven Sediment Suspension Controls Light Availability in a Shallow Coastal Lagoon

Light availability is critically important for primary productivity in coastal systems, yet current research approaches may not be adequate in shallow coastal lagoons. Light attenuation in these systems is typically dominated by suspended sediment, while light attenuation in deeper estuaries is often dominated by phytoplankton. This difference in controls on light attenuation suggests that physical processes may exert a greater influence on light availability in coastal lagoons than in deeper estuaries. Light availability in Hog Island Bay, a shallow coastal lagoon on the eastern shore of Virginia, was determined for a summer and late fall time period with different wind conditions. We combined field measurements and a process-based modeling approach that predicts sediment suspension and light availability from waves and currents to examine both the variability and drivers of light attenuation. Total suspended solids was the only significant predictor of light attenuation in Hog Island Bay. Waves and currents in Hog Island Bay responded strongly to wind forcing, with bottom stresses from wind driven waves dominant for 60% of the modeled area for the late fall period and 24% of the modeled area for the summer period. Higher wind speeds in late fall than in summer caused greater sediment suspension (41 and 3 mg l−1 average, respectively) and lower average (spatial and temporal) downwelling light availability (32% and 55%, respectively). Because of the episodic nature of wind events and the spatially variable nature of sediment suspension, conventional methods of examining light availability, such as fair-weather monitoring or single in situ recorders, do not adequately represent light conditions for benthic plants.

Patterns of ammonium uptake within dense mats of the filamentous macroalga Chaetomorpha linum

Abstract The effect of macroalgal uptake on the flux of ammonium across the sediment-water interface was tested in laboratory experiments in which dense mats of Chaetomorpha linum were incubated at high and low surface irradiances and were exposed to a high simulated sediment nutrient flux. Depth profiles of NH + 4 concentrations within the 15-cm deep mats and the timing and magnitude of NH + 4 efflux through the mats to the overlying water reflected differences in macroalgal uptake between the two light treatments. Patterns of algal productivity and NH + 4 uptake with depth in the mats were determined from the accumulation of 13 C and 15 N in the algal tissue. Nitrogen-saturated macroalgae incubated at low irradiance exhibited a strong diel periodicity in NH + 4 uptake that was not present in the N-limited macroalgae incubated at high irradiance. Assimilation by the macroalgal mat at high irradiance was approximately 900 NH + 4 μmol m −2 h −1 , and was sufficient to prevent NH + 4 diffusion from the benthic nutrient source into the overlying water during both the light and dark periods. Uptake of NH + 4 in excess of the N growth demand in the lower half of the high-light mat resulted in a spatial separation of nutrient and light resources; NH + 4 did not diffuse into the upper layers and the most photosynthetically-active macroalgae remained N-deficient. Reduced irradiance decreased the total uptake of the mat by more than 50% (400 NH + 4 μmol m −2 h −1 ), and an efflux of NH + 4 into the overlying water occurred in the dark and early part of the light period. Ammonium diffused through the unproductive bottom layers of the low-light mat and was incorporated primarily in the photic zone in the upper 4 cm of the mat where photosynthesis provided the carbon required for N uptake and assimilation. These results support the hypothesis that actively-growing macroalgal mats efficiently sequester benthic nutrient inputs to the overlying water and reduce nutrient availability to a level that may limit pelagic production. Factors that reduce irradiance within the mat, such as self-shading or decreased insolation, limit macroalgal uptake of benthic flux and result in a release of nutrients into the overlying water.

A meta-analysis of seaweed impacts on seagrasses: Generalities and knowledge gaps

Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than ‘rooted’ seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that ‘rooted’ seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.

A preliminary methods comparison for measurement of dissolved organic nitrogen in seawater

Abstract Routine determination of dissolved organic nitrogen (DON) is performed in numerous laboratories around the world using one of three families of methods: UV oxidation (UV), persulfate oxidation (PO), or high temperature combustion (HTC). Essentially all routine methods measure total dissolved nitrogen (TDN) and calculate DON by subtracting the dissolved inorganic nitrogen (DIN). While there is currently no strong suggestion that any of these methods is inadequate, there are continuing suspicions of slight inaccuracy by UV methods. This is a report of a broad community methods comparison where 29 sets (7 UV, 13 PO, and 9 HTC) of TDN analyses were performed on five samples with varying TDN and DIN concentrations. Analyses were done in a “blind” procedure with results sent to the first author. With editing out one set of extreme outliers (representing 5 out of 145 ampoules analyzed), the community comparability for analyzing the TDN samples was in the 8–28% range (coefficient of variation representing one standard deviation for the five individual samples by 28 analyses). When DIN concentrations were subtracted uniformly (single DIN value for each sample), the comparability was obviously worse (19–46% cv). This comparison represents a larger and more diverse set of analyses, but the overall comparability is only marginally better than that of the Seattle workshop of a decade ago. Grouping methods, little difference was seen other than inconclusive evidence that the UV methods gave TDN values for several of the samples higher than HTC methods. Since there was much scatter for each of the groups of methods and for all analyses when grouped, it is thought that more uniformity in procedures is probably needed. An important unplanned observation is that variability in DIN analyses (used in determining the final analyte in most UV and PO methods) is essentially as large as the variability in the TDN analyses. This exercise should not be viewed as a qualification exercise for the analysts, but should instead be considered a broad preliminary test of the comparison of the families of methods being used in various laboratories around the world. Based on many independent analyses here, none of the routinely used methods appears to be grossly inaccurate, thus, most routine TDN analyses being reported in the literature are apparently accurate. However, it is not reassuring that the ability of the international community to determine DON in deep oceanic waters continues to be poor. It is suggested that as an outgrowth of this paper, analysts using UV and PO methods experiment and look more carefully at the completeness of DIN conversion to the final analyte and also at the accuracy of their analysis of the final analyte. HTC methods appear to be relatively easy and convenient and have potential for routine adoption. Several of the authors of this paper are currently working together on an interlaboratory comparison on HTC methodology.

Seagrass Restoration Enhances “Blue Carbon” Sequestration in Coastal Waters

Seagrass meadows are highly productive habitats that provide important ecosystem services in the coastal zone, including carbon and nutrient sequestration. Organic carbon in seagrass sediment, known as “blue carbon,” accumulates from both in situ production and sedimentation of particulate carbon from the water column. Using a large-scale restoration (>1700 ha) in the Virginia coastal bays as a model system, we evaluated the role of seagrass, Zostera marina , restoration in carbon storage in sediments of shallow coastal ecosystems. Sediments of replicate seagrass meadows representing different age treatments (as time since seeding: 0, 4, and 10 years), were analyzed for % carbon, % nitrogen, bulk density, organic matter content, and 210Pb for dating at 1-cm increments to a depth of 10 cm. Sediment nutrient and organic content, and carbon accumulation rates were higher in 10-year seagrass meadows relative to 4-year and bare sediment. These differences were consistent with higher shoot density in the older meadow. Carbon accumulation rates determined for the 10-year restored seagrass meadows were 36.68 g C m-2 yr-1. Within 12 years of seeding, the restored seagrass meadows are expected to accumulate carbon at a rate that is comparable to measured ranges in natural seagrass meadows. This the first study to provide evidence of the potential of seagrass habitat restoration to enhance carbon sequestration in the coastal zone.

Genetic diversity enhances restoration success by augmenting ecosystem services.

Disturbance and habitat destruction due to human activities is a pervasive problem in near-shore marine ecosystems, and restoration is often used to mitigate losses. A common metric used to evaluate the success of restoration is the return of ecosystem services. Previous research has shown that biodiversity, including genetic diversity, is positively associated with the provision of ecosystem services. We conducted a restoration experiment using sources, techniques, and sites similar to actual large-scale seagrass restoration projects and demonstrated that a small increase in genetic diversity enhanced ecosystem services (invertebrate habitat, increased primary productivity, and nutrient retention). In our experiment, plots with elevated genetic diversity had plants that survived longer, increased in density more quickly, and provided more ecosystem services (invertebrate habitat, increased primary productivity, and nutrient retention). We used the number of alleles per locus as a measure of genetic diversity, which, unlike clonal diversity used in earlier research, can be applied to any organism. Additionally, unlike previous studies where positive impacts of diversity occurred only after a large disturbance, this study assessed the importance of diversity in response to potential environmental stresses (high temperature, low light) along a water–depth gradient. We found a positive impact of diversity along the entire depth gradient. Taken together, these results suggest that ecosystem restoration will significantly benefit from obtaining sources (transplants or seeds) with high genetic diversity and from restoration techniques that can maintain that genetic diversity.

GRACILARIA VERMICULOPHYLLA (RHODOPHYTA, GRACILARIALES) IN HOG ISLAND BAY, VIRGINIA: A CRYPTIC ALIEN AND INVASIVE MACROALGA AND TAXONOMIC CORRECTION1

Gracilaria in Virginia, USA, is abundant and composed of thalli either having relatively flat or cylindrical branches. These two morphologies were referred to previously as G. foliifera (Forsskål) Bøgesen and G. verrucosa (Hudson) Papenfuss. However, G. verrucosa is regarded an invalid name, and the flat specimens are now referred to as G. tikvahiae McLachlan. This has created confusion about the nomenclature of Gracilaria from this region. Here we document that the cylindrical form that dominates Hog Island Bay, Virginia, is G. vermiculophylla (Ohmi) Papenfuss, an alien macroalga from the West Pacific. Most of the ecological studies performed at the Long Term Ecological Research (LTER) site in Hog Island Bay used this cylindrical species. The present study clarifies the taxonomical status of this species, and we identify attributes that make this alien successful in turbid coastal lagoons.