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Microphytobenthos and Benthic Macroalgae Determine Sediment EGU Journal Logos (RGB) Open Access Open Access Open Access Advances in Annales Nonlinear Processes Geosciences Geophysicae in Geophysics Open Access Open Access Natural Hazards Natural Hazards and Earth System and Earth System Sciences Sciences Discussions Open Access Open Access Atmospheric Atmospheric Chemistry Chemistry and Physics and Physics Discussions Open Access Open Access Atmospheric Atmospheric Measurement Measurement Techniques Techniques Discussions Open Access Biogeosciences, 10, 5571–5588, 2013 Open Access www.biogeosciences.net/10/5571/2013/ Biogeosciences doi:10.5194/bg-10-5571-2013 Biogeosciences Discussions © Author(s) 2013. CC Attribution 3.0 License. Open Access Open Access Climate Climate of the Past of the Past Discussions Microphytobenthos and benthic macroalgae determine sediment Open Access Open Access organic matter composition in shallow photic sedimentsEarth System Earth System Dynamics 1,* 1 1 2 3 Dynamics4 A. K. Hardison , E. A. Canuel , I. C. Anderson , C. R. Tobias , B. Veuger , and M. N. Waters Discussions 1Virginia Institute of Marine Science, College of William & Mary, P.O. Box 1346, Gloucester Point, VA 23062, USA 2Department of Marine Sciences, University of Connecticut, Groton, CT 06340, USA Open Access Open Access 3The Royal Netherlands Institute for Sea Research (NIOZ), Yerseke, the Netherlands Geoscientific Geoscientific 4Valdosta State University, Valdosta, GA 31698, USA Instrumentation Instrumentation *current address: University of Texas Marine Science Institute, 750 Channel View Dr., Port Aransas,Methods TX 78373,and USA Methods and Correspondence to: A. K. Hardison ([email protected]) Data Systems Data Systems Discussions Open Access Received: 7 January 2013 – Published in Biogeosciences Discuss.: 15 February 2013 Open Access Geoscientific Revised: 30 May 2013 – Accepted: 19 June 2013 – Published: 20 August 2013 Geoscientific Model Development Model Development Discussions Abstract. Microphytobenthos and benthic macroalgae play to half of the light treatments, decreased SOM accumula- an important role in system metabolism within shallow tion compared to light treatments without macroalgae, with Open Access coastal bays. However, their independent and interactive in- TOC and TN increasing by only 130 ± 32 andOpen Access 94 ± 24 %, fluences on sediment organic matter (SOM) are not well respectively. This decreaseHydrology likely resulted and from shading by Hydrology and understood. We investigated the influence of macroalgae macroalgae, which reducedEarth production System of microphytoben- Earth System and microphytobenthos on SOM quantity and quality in an thos. The presence of macroalgae decreased SOM lability experimental mesocosm system using bulk and molecular as well, which resulted in diminishedSciences buildup of bacterial Sciences level (total hydrolyzable amino acids, THAA; phospholipid biomass. By the final day of the experiment, principal com- Discussions Open Access linked fatty acids, PLFA; pigment) analyses. Our experiment ponent analysis revealed that sediment compositionOpen Access in treat- used an incomplete factorial design made up of two fac- ments with macroalgae was more similar to dark treatments Ocean Science tors, each with two levels: (1) light (ambient vs. dark) and and less similar to lightOcean treatments Science without macroalgae. Discussions (2) macroalgae (presence vs. absence of live macroalgae). Overall, microphytobenthos and benthic macroalgae funda- Over the course of the 42-day experiment, total organic car- mentally altered SOM quality and quantity, which may have bon (TOC) and total nitrogen (TN) increased under ambi- notable ecological consequences for shallow-water systems Open Access Open Access ent light by 173 ± 14 and 141 ± 7 %, respectively, compared such as increased hypoxia/anoxia, sulfide accumulation, en- to in the dark (78 ± 29 and 39 ± 22 %). THAA comprised a hanced mineralization and/or stimulated denitrification. Solid Earth ∼ Solid Earth substantial fraction of SOM ( 16 % of TOC, 35 % of TN) Discussions and followed TOC and TN accumulation patterns. Mole per- cent composition of the THAA pool indicated that SOM was 1 Introduction composed of more labile organic material (e.g., L-glutamic Open Access Open Access acid, phenylalanine) under ambient light conditions while SOM in dark treatments was more degraded, with higher Shallow coastal bays make up approximately 13 % of the The Cryosphere world’s coastline, areThe among Cryosphere the most highly productive proportions of glycine and D-alanine. PLFA content, which Discussions represents viable biomass, made up ∼ 1 % of TOC and con- ecosystems on earth, and are distinctly vulnerable to effects tained high levels of algal fatty acids in the light, particu- from the growing problem of nutrient over-enrichment due larly PLFA derived from diatoms. In the presence of micro- to increased human activities (Nixon, 1995; NRC, 2000; phytobenthos (i.e., light and macroalgae treatments), SOM Pedersen et al., 2004). One consequence of nutrient load- lability increased, resulting in the observed increases in bac- ing to many of these systems has been a shift in primary terial PLFA concentrations. Macroalgae, which were added producer community structure. Because much of the sedi- ment surface resides within the euphotic zone in these bays, Published by Copernicus Publications on behalf of the European Geosciences Union. 5572 A. K. Hardison et al.: SOM composition in shallow photic sediments benthic autotrophs often are the dominant primary produc- ganic matter (SOM) consists of material from a variety of ers. Observations from a number of systems have shown living and non-living sources and performs important ecosys- that as nutrient loading increases, ephemeral macroalgae, tem functions such as providing food for infauna and epi- phytoplankton, and epiphytes increase, while slow-growing fauna, aiding in sediment stability (e.g., extracellular poly- perennial macrophytes such as seagrasses decrease (Valiela meric substances produced by benthic microalgae; Wolfstein et al., 1992; Hauxwell et al., 2001; Duarte, 1995; Sand- et al., 2002), and providing temporary or permanent storage Jensen and Borum, 1991). In Waquoit Bay, MA, for exam- for carbon (C) and nutrients (Hardison et al., 2011b; Middel- ple, ephemeral populations of green (Cladophora) and red burg et al., 2000). While sources contributing to SOM vary (Gracilaria) macroalgae replaced Zostera marina seagrass by system, microbial biomass often contributes a significant when nutrient loadings increased six-fold (Hauxwell et al., fraction of SOM in shallow systems (Volkman et al., 2008; 2003). The mechanisms underlying this shift in community Canuel and Martens, 1993; Bouillon and Boschker, 2006). structure relate to differences among plant types in nutrient Specifically, microphytobenthos may be a particularly good uptake and growth strategies (Sand-Jensen and Borum, 1991; source of labile organic matter, which may support bacte- Nielsen et al., 1996). Microphytobenthos, including ben- rial production (Volkman et al., 2008; Hardison et al., 2011b; thic microalgae (e.g., diatoms) and cyanobacteria, often con- Middelburg et al., 2000). tribute significantly to primary production within these shal- The objectives of this study were to examine the influ- low systems; however, their response as the autotrophic com- ence of microphytobenthos on SOM quality and quantity, munity structure shifts in the face of nutrient over-enrichment and to investigate how the microphytobenthic contribution is not well understood. to SOM changes in the presence of a macroalgal bloom. Be- The dominant plants in a community greatly affect both cause gross measurements of SOM (e.g., total organic C, to- the physical and biological conditions of a system, includ- tal nitrogen (N)) do not provide information on source and/or ing overall community structure (Orth et al., 1984; Heck et lability, we combined bulk and molecular-level analyses to al., 2003; Norkko, 1998), ecosystem processes such as nu- more accurately characterize the SOM of a shallow coastal trient cycling (Tyler et al., 2001; Risgaard-Petersen, 2003), bay. Specific organic compounds (biomarkers) were used to and hydrodynamic conditions (Fonseca and Calahan, 1992; attribute organic matter to different sources. Jumars et al., 2001; Paterson and Black, 1999). For exam- ple, the presence of ephemeral macroalgae often leads to episodic anoxia and increased sulfide concentrations (Sfriso 2 Methods et al., 1992; Krause-Jensen et al., 1999), which negatively 2.1 Site description affect fish and benthic fauna (Norkko et al., 2000; Raf- faelli et al., 1998; Gray et al., 2002) as well as other au- Sediments and macroalgae were collected from Hog Island totrophs (Hauxwell et al., 2003; Sundback and McGlath- Bay, Virginia (HIB), which is located along the Delmarva ery, 2005). Macroalgae also affect other primary producers Peninsula and part of the Virginia Coast Reserve, a Long- directly through shading and/or competition for nutrients. Term Ecological Research (LTER) site. HIB is a shallow Because of their location at the sediment surface or float- coastal lagoon (< 2 m deep at mean low water), typical of ing just above the sediments, macroalgae may reduce the temperate lagoons along the US east coast, and is dominated amount of light available for microphytobenthos, thereby de- by benthic autotrophs (McGlathery et al., 2001; Thomsen creasing or inhibiting microphytobenthic production (Sund- et al., 2006).
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