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Community Structure and Abundance of Benthic Infaunal Invertebrates in Maine Fringing Marsh Ecosystems

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Community Structure and Abundance of Benthic Infaunal Invertebrates in Maine Fringing Marsh Ecosystems Estuaries and Coasts DOI 10.1007/s12237-015-9977-8 Community Structure and Abundance of Benthic Infaunal Invertebrates in Maine Fringing Marsh Ecosystems Richard A. MacKenzie1 & Michele Dionne2 & Jeremy Miller2 & Michael Haas2 & Pamela A. Morgan3 Received: 1 October 2013 /Revised: 21 April 2015 /Accepted: 22 April 2015 # Coastal and Estuarine Research Federation (outside the USA) 2015 Abstract Fringing marshes are abundant ecosystems that biomass, but the less abundant and larger invertebrates such as dominate the New England coastline. Despite their abun­ green crabs, tanaids, and bivalves were also large contributors dance, very little baseline data is available from them and to biomass in the low marsh zone. Low marsh invertebrate few studies have documented the ecosystems services that communities were characterized by significantly higher den­ they provide. This information is important for conservation sities of nematodes, Nereis virens,anunidentifiedoligochaete, efforts as well as for an increased understanding of how fring­ the bivalves Gemma gemma and Mya arenaria,and ing marshes function compared to larger marsh meadow sys­ Leptochelia rapax. High marsh invertebrate communities tems. Benthic infaunal invertebrates were sampled from cores were characterized by higher densities of insects, specifically collected from Spartina alterniflora-dominated low marsh, Culicoides sp. ceratopogonid larvae and Anurida maritima,as Spartina patens-dominated high marsh, and Phragmites well as an unidentified species of mite. Our results revealed a australis-invaded high marsh zones of nine fringing marsh diverse and abundant infaunal invertebrate community that ecosystems in Casco Bay, Maine, USA. Infaunal densities likely supports similar ecosystem services in fringing marshes and biomass were generally higher in low marsh than high as invertebrates in larger marsh meadows. marsh or P. australis cores. Invertebrate community structure was significantly different between low marsh and high marsh Keywords Fringing marsh . Spartina patens . Spartina and P. australis cores, which was attributed to significantly alterniflora .Phragmitesaustralis .Saltmarsh .Benthiccore . higher pore water salinity, lower organic matter, total plant Infaunal invertebrates . Sediment percent cover, and S. patens cover in low marsh zones. There were no differences in invertebrate densities, biomass, or community structure when high marsh and P. australis Introduction cores were compared. Invertebrate densities and community structure were dominated by oligochaetes in all zones. Fringing salt marshes are a common component of New Oligochaetes were also an important component of infaunal England coastlines (Jacobsen et al. 1987; Morgan et al. 2009; Roman et al. 2000). Compared to larger Spartina patens-dominated marsh meadows, fringing marshes are Communicated by Wayne S. Gardner much smaller in size but have greater edge to area ratios (Morgan et al. 2009). Despite their small size, fringing * Richard A. MacKenzie marshes provide similar ecosystem services to coastal popu­ [email protected] lations of humans as their larger marsh meadow counterparts. Narrow fringing salt marshes that line edges of bays and rivers 1 USDA Forest Service, Pacific Southwest Research Station, Institute regulate water quality through the removal of human-induced of Pacific Islands Forestry, 60 Nowelo St, Hilo, HI 96720, USA nitrogen (Tobias et al. 2001; Wigand et al. 2004) and sediment 2 Wells National Estuarine Research Reserve, 90 Laudholm Farm Rd, loads (Morgan et al. 2009). These ecosystems can also reduce Wells, ME 04090, USA impacts to coastlines from storm events. S. patens dominated 3 University of New England, Biddeford, ME 04005, USA fringe marshes reduced wave height by more than 60 % Estuaries and Coasts (Morgan et al. 2009), while Spartina alterniflora-dominated that while the majority of these marshes are relatively intact, systems reduced mean flow velocities by 40–60 % (Bruno and their integrity is threatened from invasive plant species and Kennedy 2000). The latter often resulted in the establishment various human activities (e.g., shoreline development, oil of diverse beach strand plant communities (Bruno and spills, sea level rise) (Hayes et al. 2008). Shifts in hydrological Kennedy 2000). Fringing marshes also support coastal biodi­ regimes from increased development in this region have al­ versity by providing habitat for many species of plants tered soil salinity and resulted in the increased spread of both (Morgan et al. 2009), resident and transient nekton (Morgan exotic Phragmites australis (common reed) (Silliman and et al. 2015; Rountree and Able 1992), and marsh birds Bertness 2004)and Typha angustifolia (narrow-leaved (Shriver et al. 2004). Fringe marshes likely support cattail) (Burdick et al. 1997), both of which could potentially abundant and diverse assemblages of benthic infaunal alter certain habitat values of these ecosystems. For ex­ invertebrates that are valuable food resources for higher ample, significantly lower densities of resident mummi­ organisms. However few, if any studies, have document­ chogs (Fundulus heteroclitus) have been reported from ed invertebrate communities in New England fringing or the surface of P. australis-invaded marsh meadows vs. larger marsh meadow ecosystems and what factors in­ native S. alterniflora-dominated ones (Able and Hagan 2000; fluence these communities. Able and Hagan 2003;Hunteretal.200 6). Lower densities of Benthic infaunal invertebrate communities are largely larval mummichogs in P. australis-invaded marshes were at­ comprised of small invertebrates (i.e., annelids, insects, am­ tributed to higher evapotranspiration (ET) rates and litter in­ phipods) found within or on the sediment of the vegetated puts of P. australis compared to S. alterniflora. Higher ET marsh surface (Kneib 1984;Rader 1984). Salt marshes are rates and increased litter inputs result in fewer water-filled often perceived as supporting nuisance insect species such depressions on the marsh surface that provide valuable as mosquitoes, green head flies, and biting no-see-ums. subtidal refuge for resident fish during low tide (Able and However, salt marshes also support diverse assemblages of Hagan 2003;Ableetal. 2003; Weinstein and Balletto 1999). insect species that include chironomid midge flies, a salt Invertebrate densities and species richness can also be marsh dragonfly, and a salt marsh caddisfly (MacKenzie lower in P. australis or Typha spp. compared to S. alterniflora 2003;MacKenzie200 5). These diverse invertebrate assem­ marshes (Angradi et al. 2001;GrattonandDenno2006; Talley blages support many ecosystems services that marsh ecosys­ and Levin 2001; Warren et al. 2002). This was also attributed tems provide. Marsh invertebrates support provisional ser­ to reduced standing water and a resulting decrease in vices as they are an important food resource for many species microalgal food resources (Angradi et al. 2001). Other of transient and migratory birds and nekton that utilize these studies have reported no changes in biomass, species habitats (Allen et al. 1994; Deegan and Garritt 1997; richness, or overall diversity in invertebrate communities MacKenzie and Dionne 2008; Shriver et al. 2004). Many of in P. australis-invaded marshes (Fell et al. 1998; Warren et al. these fish are of high commercial importance (e.g., winter 2001; Yuhas 2001). flounder, American eel, blue fish) (Deegan and Garritt The main objectives of this project were to 1) document the 1997). Benthic macroinvertebrates also indirectly provide benthic infaunal invertebrate community structure, biomass, food resources by breaking down plant material into more and densities in Casco Bay fringe marshes and 2) to determine readily available particulate and dissolved food resources if these invertebrate parameters differed between high and low for higher trophic groups (Graça et al. 2000). Marsh fringing marsh areas or between exotic and native plant as­ invertebrates can also support the regulating services semblages. This information also provided a much needed provided by plants (e.g., increased water quality, storm baseline data set that could be used to assess how infaunal protection) by supporting plant growth and community communities may change after natural (e.g., hurricane) or composition. For example, marsh plant productivity can human-induced disturbances (e.g., oil spill). Impacts from be significantly influenced by top–down control of her­ oil spills are of special concern as Casco Bay is the largest bivorous snails and amphipods by predaceous crabs and oil port in Northern New England (Hayes et al. 2008). fish (Pennings and Bertness 2001; Silliman and Bertness Invertebrate assemblages were compared between high and 2002), increased oxygen in sediments by burrowing crabs low marsh zones of nine different fringing marshes. Two of (Andersen and Kristensen 1988), and increased nutrient avail­ these sites included high marsh areas with established popu­ ability by filter feeders (Bertness 1984; Bertness 1985; lations of exotic P. australis.Wehypothesizedthatinverte­ Pennings and Bertness 2001). brate community structures, biomass, and densities would dif­ The nearly 41 ha of fringing salt marsh that make up over fer 1) between high and low marsh areas due to differences in 10 % of the Casco Bay shoreline (Morgan et al. 2009) and the plant communities and hydrological influences on sediment majority of marsh habitat found there are expected
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