Benthos of the York River by David J
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Benthos of the York River David J. Gillett and Linda C. Schaffner Virginia Institute of Marine Science Gloucester Point, VA 23061 U.S.A. ABSTRACT Benthic organisms and their communities are key components of estuarine systems. We provide an overview of the biology and key ecological features of benthic communities of York River Estuary (YRE), which is the site of the Chesapeake Bay National Es- tuarine Research Reserve in Virginia (CBNERRVA). Major subtidal benthic habitats in YRE include soft mud and sand bottoms, with only limited distribution of submerged aquatic vegetation and oyster shell. Major taxonomic groups of macrofauna dominat- ing muds and sands of YRE include annelids, molluscs and crustaceans; similar to those found in other temperate estuaries of the US Mid-Atlantic. Meiofaunal assemblages of YRE soft bottoms are dominated by nematodes and copepods. Species distribution patterns in YRE are strongly correlated with salinity and bottom type, while other factors such as eutrophication and hypoxia may be growing in importance. Much of the YRE benthos fails to meet the restoration goals set by the Chesapeake Bay Program. The poor condition of the benthos is expressed as low biomass and abundance and may be associated with degraded water quality, hypoxia and sediment disturbance processes. No comprehensive inventory of the benthic biota of the CBNERRS sites is available, which will make it difficult to assess future changes due to human impacts such as climate change or the introduction of exotic species. Given this paucity of data, a systemic cataloging of the benthic resources of the reserve sites and any potential invasive species is a much needed avenue of future research for CBNERRVA. INTRODUCTION TO THE BENTHOS larvae are commonly encountered macrobenthos in estuar- ies. The largest size-based category, the megabenthos, can The soft mud and sand habitats of the York River Estuary, be identified without magnification because individuals are as well as the interspersed patches of aquatic vegetation and typically multiple centimeters in size. This group includes oyster shell, support a wide variety of fauna and flora and are animals such as crabs, bivalves, gastropods, sponges, colonial an important part of this productive coastal ecosystem. These entoprocts and hydrozoans. Benthic organisms may progress bottom habitats and their resident organisms are called the through different categories as they grow. Many animals clas- benthos, derived from the Greek for “bottom of the sea.” The sified as macrobenthos start off as meiobenthic juveniles and 1 animals comprising benthic communities, the zoobenthos , are known as “temporary meiobenthos.” include almost every known phylum and exclusively encom- Beyond size, the mobility of an animal (motile versus ses- pass a number of them. For the purposes of this paper we sile) and how it associates with the sediment or hard substrate have limited ourselves to a discussion of the benthic inverte- (infaunal versus epibenthic) are other common ways benthic brate residents and their communities of the York River Estu- organisms are classified. Epibenthic animals live on or just ary. This is not to slight the countless numbers of bacteria, above the substrate. They may be firmly attached (sessile), Archea, and protozoans that comprise the microbenthos, or relatively sedentary, or fully motile. Animals such as barna- the bottom-dwelling fish and crustaceans of the estuary, all of cles, oysters, sponges, tunicates, entoprocts, gastropods, an- which are discussed in other papers in this issue. thozoans, mud crabs, and certain species of amphipods are Most benthic invertebrates are quite small and can be common representatives of the epibenthos. Animals that live clearly distinguished only with the aid of magnification. They within the substrate are called infauna and include most spe- are classified into three major groups based on adult size. The cies of annelids and bivalves, larval insects, phoronids, as well smallest are the meiobenthos, which pass through a 500-µm as some species of amphipods and anthozoans. mesh, but are retained on a 63-μm screen. Important taxa of meiobenthos include harpactacoid copepods, nematodes, MAJOR TAXONOMIC GROUPS OF BENTHIC FAUNA ostracods and Foraminfera (see HIGGINS and THIEL, 1988). Macrobenthos are retained on a 500-μm mesh screen and IN THE YORK ESTUARY are not readily identifiable without magnification. Annelid A comprehensive checklist of benthic animals in the York worms, bivalves, gastropods, crustaceans, tunicates, and insect River Estuary and the greater Chesapeake Bay was published by Wass (1972). It provides frequency of occurrence and habi- 1The generic terms benthos and benthic, which are used to describe tat preferences of those animals known at the time. There is the bottom realm, have also been variously used to describe any and all of the organisms, from bacteria and microalgae to seagrasses and no complete benthic invertebrate species list exclusively for demersal predators, that are associated with benthic habitats. Use of the York River system; however, most of the benthic fauna the term zoobenthos provides more clarity, but in practice is rarely used found in the York River Estuary are listed in the regularly by benthic ecologists working in the U.S. updated checklist available for the Chesapeake Bay Benthic 77 Monitoring Program (LLANSÓ, 2005). A partial checklist of duction during different benthic organisms in the York River Estuary developed from stages of their life cycle. these and other sources is provided in the Appendix. Asexually, new hydroid zo- oids can be budded off an Poriferans adult in an expansion of the colony, or as separate Sponges are colonial macro- to megabenthic-sized organ- individuals in the non- isms. They filter feed by pumping water through inhalant and colonial forms. Sexual re- exhalent pores called ostia, trapping particles along the body production in hydrozoans, wall, and ingesting them by phagocytosis (BRUSCA and BRUSCA, much like the other types 1990). Most sponges in the York River Estuary are limited to of cnidarians, is somewhat the meso- to polyhaline reaches. Among the most conspicu- more complex. A free- ous are the red beard (Microciona prolifera) and brown (Hali- swimming male or female clona spp.) sponges, both of which grow attached to hard sub- medusa (jellyfish-like) strate (Figure 1). M. prolifera is frequently seen on pier pilings, stage is budded off of the while Haliclona loosanoffi is commonly found on the blades benthic adult form, which of submerged in turn, releases gametes aquatic vegetation into the water column (SAV). The bor- that when fertilized, form Figure 2. Colonial hydroids from the ing sponges, Cliona asexual, benthic individu- lower York River. (Image courtesy of spp., erode galler- als (BRUSCA and BRUSCA, Robert Diaz, VIMS) ies of passageways 1990). through calcareous Though less abundant shell of molluscs, and less diverse in the York River Estuary than hydrozoans, which provides pro- anthozoan (sea anemones) and scyphozoan (jelly fish) cnidar- tection from preda- ians are also found within the benthic communities. Like the tors. These types of hydrozoans, benthic anthozoans are passive filter feeders ca- sponges are consid- pable of both sexual and asexual reproduction. Anthozoans ered nuisance spe- are non-simultaneous hermaphrodites that can bud off new Figure 1. Unidentified red sponge. (Image cies by commercial individuals from the adult form, as well as produce male or courtesy of Southeastern Regional Taxo- shellfish harvesters female gametes. Anthozoans have lost the free-swimming nomic Center/South Carolina Department because the erosion medusa-stage of other cnidarians. The benthic adults directly of Natural Resources) of shell material is release gametes to the water column, where they combine to detrimental to liv- form planular larvae that settle out of the water to from new ing molluscs. All of the sponges found in the York River Es- benthic adults. Common anthozoans include epibenthic spe- tuary are capable of both sexual and asexual reproduction. cies (e.g., Diadumene leucolena) and infaunal species (Cerian- Fragments of a sponge can grow an entire new sponge, given theopsis americanus, Actiniaria sp. or Edwardsia elegans) (SAGASTI an appropriate substrate. Sexual reproduction in sponges is et al., 2001; LLANSÓ 2005). Scyphozoans are only ephemeral through broadcast spawning with most species thought to be benthic organisms, but the benthic stage is an essential part of hermaphroditic, which means that they switch between the their reproductive lifestyle that occurs at various times of the production of male and female gametes during different parts year depending upon the species (see STEINBERG and CONDON of their lives (BRUSCA and BRUSCA, 1990). this S.I.). This benthic stage is referred to as a scyphistoma and is an asexual from that buds off the familiar, pelagic me- Cnidarians dusae seen in the estuary. Representatives of all three classes of cnidarians (Hydro- zoa, Anthozoa, and Scyphozoa) have been observed among Platyhelminthes the macrobenthic fauna of the York River Estuary. All cnidar- Flatworms are a small, relatively obscure component of ians possess nematocysts, responsible for the familiar stinging the benthic community that can be found all along the estua- sensation of jellyfish, which they use for both defensive and rine salinity gradient. Free-living turbellarian flatworms can prey capturing purposes. Hydrozoans, the