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1. Channele

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1. Nutrient R

fish restoratio ries ability to dles et al. 201 oplankton is a ajor consume Management whelk Bus Knobbed whelk nd to associat ch may expla e very adept a ec -1 Ferner an ersole et al. 2 one-month tim l loss from sh alves such as q eproductive po e highly susce al. 2008. system Ser s are processe human life, pr cease to thriv ing feeding, f mn. Reef build ng wave stren s a description lve population ReductionW on, particularl improve wat 13. Asmus an key process w ers of phytopl t Plan, Versio C HAPT sycon carica k Busycon car te with mud a ain why they a at finding pre nd Weissburg 2008 on knob me span show hell chipping, quahogs. otential, limit eptible to ove rvices Pro es through wh roviding esse ve Dame 201 for instance, p ding bivalves, ngth hitting th n of some of t ns. Water Qual ly for reef bui er quality thro nd Asmus 19 within the ma ankton, a loss on II TER 2: E COLOG a rica, illustratio and sand botto are less likely ey through che g 2005. Thei bbed whelk fo wed no shell g which is a co ted movemen r harvesting ovision hich ecosyste ntial life supp 12. Filter feed provide a wate such as muss he shore and t the known im lity Improve ilding species ough filtratio 993 conclude aterial exchan s of filter feed GY ons courtesy of oms, and pref y to enter whe emically med ir slow movin ound that 95 growth, and th ommon occur nt, slow growt Castagna and ems and the sp port services w ding bivalves er quality serv sels and oyste thereby slowin mportant ecosy ement s such as mus n of water by e: “The intera nge between b der concentra N OV f Brandon Full fer live prey o elk traps or po diated cues, ev ng nature may of the indiv hat 47 show rrence in whe th, and large s d Kraeuter 19 pecies that ma without whic s, by the proce vice by remo ers, provide sh ng erosional p ystem service ssels and oyste y filter feeding action between benthic and p ations e.g., oy VEMBER 18, 201 ler, 2014. over carrionb ots. ven at flow sp y assist in che viduals marke wed negative s elk feeding on size at maturi 994; Walker e ake them up ch human esses involve ving particula horeline prote processes. Th es provided by ers, increases g shellfish n mussel beds elagic species yster reefs, m 14 bait peeds emical ed and shell n ity, et al. ed in ates ection he y s an s and s.” mussel N OVEMBER 18, 2014 C HAPTER 2: E COLOGY P AGE 47 OF 373 beds, quahog aggregations in eutrophic systems provides a positive feedback mechanism for hypoxia as phytoplankton are no longer heavily consumed by the bivalves Altieri and Witman 2006. 2. Ecosystem services provided by oyster reefs has been summarized as: 1 production of oysters, 2 water filtration and concentration of pseudofeces, 3 provision of habitat for epibenthic invertebrates, 4 carbon sequestion, 5 augmented fish production, 6 stabilization of adjacent habitats and shoreline, and 7 diversification of the landscape and ecosystem Grabowski and Peterson 2007; Coen et al. 2007. The National Research Council 2005 suggests a similar listing of ecosystem services, but includes enhanced water clarity improvement and alteration of hydrography in shallow water ecosystems as it pertains to shoreline buffering. 3. Bivalves, as filter feeders, remove plankton and particulates from the water column—an adult soft- shell clam can filter 4 liters of water per hour, and an adult Eastern oyster can filter as much as 14 liters of water per hour Bertness 2007; Rice 2001. For the Providence River, Rice et al. 2000 calculated a filtering capacity of 2 x 10 7 m 3 5.3 billion gallons per day in August by the quahog population. Newell 1988 calculated a 3.3 day clearance time 9 for the Chesapeake Bay in the late 1800s, but 325 days in 2007 due to the dramatic decline in oyster reef extent. Dame 2012 estimates that Narragansett Bay has a bivalve clearance time of 25 days. 4. Dense aggregates of bivalves can exert a controlling effect on phytoplankton abundance, resulting in improved water clarity and general eutrophication control Alpine and Cloern 1992; Asmus and Asmus 1993; Dame 2012. As such, filter feeding bivalves serve as a critical link between primary production in the water column and the benthic ecosystem Bertness 2007; Higgins et al. 2011, 2013, particularly since they do so on a consistent, long-term basis as permanent residents of the ecosystem, though impact on water quality may vary seasonally Dame 2012. Loss of this ecosystem service in the Chesapeake Bay has been credited to have resulted in increased suspended sediments and decreased water clarity, and with those, a loss of eelgrass beds Newell 1988; Carmichael et al. 2012. 5. Bivalve aggregations—oyster reefs and mussel beds for instance—are significant elements of nutrient recycling between benthos and water column Asmus and Asmus 1993; Coen and Grizzle 2007; Dame 2012; Grabowski and Peterson 2007; Prins et al. 1998; Sisson et al., 2011; Smaal and Prins 1993. Particulate organic matter is removed during filter feeding and returned to bottom sediments as feces and pseudofeces. Nitrogen is then recycled by sediment microbes where it is nitrified 10 for reuse by phytoplankton, or denitrified 11 and released to the atmosphere Giles and Pilditch 2006; Sisson et al., 2011; Higgins et al. 2013. Sisson et al. 2011 found that 21 of the organic N underwent nitrification, being released back to the environment for biological uptake, and 12 underwent denitrification at an oyster reef in the Chesapeake Bay. The authors do note however, that rate of nitrogen use and conversion were very seasonal September for their work as well as site specific, and some caution should be applied in transferring those rates to other places or times. Ammonia excretion by bivalves is rapidly taken up in the water column by phytoplankton for growth Giles and Pilditch 2006; Newell 2004. In this way bivalve aggregates help control eutrophication while at the same time rapidly recycling nutrients to promote continued phytoplankton growth that sustains the bivalve community. Newell 2004 notes that the biggest impact to benthic-pelagic coupling will come from those species that maintain high clearance rates and produce excess pseudofeces. Smaal and Prins 1993 sum the process of benthic-pelagic coupling in shellfish aggregates as: 1 filtration of large quantities of material from the water column, 2 reduction of phytoplankton with possible 9 Clearance time is the theoretical time needed for the total bivalve-suspension feeder biomass within an ecosystem to filter all particles from the entire volume of water in the aquatic ecosystem in question Dame 2012. 10 Nitrification is a naturally occurring process carried out by nitrifying bacteria in which ammonia, produced by the breakdown of organic materials, is converted to nitrites and nitrates which become available for use by aquatic plants. 11 Denitrification is a naturally occurring process, performed by denitrifying bacteria, that converts nitrates back into nitrogen gas which can then be released back into the atmosphere.