Reef Habitats in the Middle Atlantic Bight: Abundance, Distribution, Associated Biological Communities, and Fishery Resource Use

Item Type article

Authors Steimle, Frank W.; Zetlin, Christine

Download date 25/09/2021 09:28:45

Link to Item http://hdl.handle.net/1834/26392 Reef Habitats in the Middle Atlantic Bight: Abundance, Distribution, Associated Biological Communities, and Fishery Resource Use

FRANK W. STEIMLE and CHRISTINE ZETLIN

Introduction soft sediments, mostly sands, but grad- line jetties and groins, submerged pipe­ Spatial distribution and perhaps the ing to silt-clay in deeper areas (Stumf lines, cables, artificial reefs, and similar abundance of fishery resources are in- and Biggs, 1988; Poppe et al., 1994). objects or material placed in the marine fluenced by physical and other habitat Except for relic sand and gravel ridges, environment by the human population. factors. The identification of significant exposed Holocene to Pleistocene clay Some of these human additions are marine habitats and strong or critical or sandstone in some areas (Allen et considered objectionable “litter” (Gal­ associations between living marine re- al., 1969; Wigley and Theroux, 1981; gani et al., 2000), but larger objects can sources (LMR’s) and these habitats can Stumf and Biggs, 1988; Poppe et al., function as seabed structures that de­ lead to a better understanding of how 1994; NOAA National Data Center­ velop and support diverse and special environmental influences affect LMR’s NGDC, 1999), and glacially exposed biological communities, even if they and fisheries and support their manage- rock along the southern New England can be patchy in distribution. These ment (NMFS, 1999a). coast, this habitat characterization of communities differ significantly from The Middle Atlantic Bight (the area the Bight was basically true until Euro­ those of the surrounding well surveyed, of the U.S. east coast and continental pean colonization. soft sediment seabed of the Bight. shelf between Cape Cod, Mass., and Within the last two centuries there The expansion of this habitat type in Cape Hatteras, N.C.) hereafter referred has been an increase in hard bottom the Bight by man’s addition of solid to as the Bight, is characterized as or reef (“reef” is used hereafter to material has probably had an effect on being a homogeneous habitat of rel- refer to this multi-dimensional, hard­ LMR distributions and fisheries (such atively flat topography, composed of substrate, structural habitat) habitats in as American lobster, Homarus ameri­ the Bight, which is not commonly rec- canus; cod, Gadus morhua; red hake, ognized by marine geologists and re­ Urophycis chuss; ocean pout, Macro­ The authors are with the Sandy Hook Labora- tory, Northeast Fisheries Science Center, National source managers, e.g. shipwrecks, lost zoarces americanus; scup, Stenotomus Marine Fisheries Service, NOAA, 74 Magruder cargos, disposed solid materials, shore- chrysops; black sea bass, Centropristis Road, Highlands, NJ 07732. striata; and tautog, Tautoga onitis) and possible effect on other resources, but these effects are not well known nor well understood. In fact, reef habitats ABSTRACT—One particular habitat type tribution, abundance, use by living marine in the Middle Atlantic Bight is not well rec- resources and associated biological commu­ in general seem underappreciated by ognized among fishery scientists and man- nities (except on estuarine oyster reefs), and northeastern U.S. habitat managers or agers, although it is well known and used fishery value or management. This poorly researchers. For example, no type of by recreational and commercial fisheries. studied and surveyed habitat can provide fish reef habitat is even listed as a fishery This habitat consists of a variety of hard-sur- refuge from trawls and can be a factor in face, elevated relief “reef” or reef-like envi- studies of the distribution and abundance of habitat in recent reviews of northeast ronments that are widely distributed across a variety of reef-associated fishery resources. fish habitat, except for boulders (Lang­ the predominantly flat or undulating, sandy This review provides a preliminary summary ton et al., 1995; Auster and Langton, areas of the Bight and include both natural of information found on relative distribution 1999), and they are not considered as rocky areas and man-made structures, e.g. and abundance of reef habitat in the Bight, demersal fish nursery habitat in the shipwrecks and artificial reefs. Although the living marine resources and biological there are natural rock and shellfish reefs in communities that commonly use it, threats Bight (Steves et al., 2000). In the waters southern New England coastal waters and to this habitat and its biological resources, south of Cape Hatteras, reef habitats estuaries throughout the Bight, most reef and the value or potential value of artificial are recognized as important to fisher­ habitats in the region appear to be man- reefs to fishery or habitat managers. The pur­ ies and some of the species found in made, mostly wrecks and “obstructions,” and pose of the review is to initiate an awareness man-made reef habitat modification/creation among resource managers about this habi­ the Middle Atlantic Bight are also part may be increasing. Very little effort has been tat, its role in resource management, and the of those fisheries (Miller and Richards, devoted to the study of this habitat’s dis- need for research. 1979; Parker and Mays, 1998).

24 Marine Fisheries Review Although hard bottom habitats off these varied roles involving fishery and cludes references to species, such as southern New England were explored habitat management, there is need to aquatic birds, that interact with reef­ briefly by naturalist dredge in the latter begin to better understand the commu­ associated fishes. half of the nineteenth century (Verrill, nity dynamics and fishery value of this 1872), submarine canyon and tilefish habitat type in the Bight. Characteristics of Reef habitats were examined by submarine This paper summarizes available in­ Habitats in the Bight (Valentine et al., 1980), and some at­ formation in four parts: A reef habitat can be composed of tention was given to biological fouling natural materials (such as rocks used (Redfield and Ketchum, 1952), little 1) A characterization and preliminary for shoreline rip rap jetties), sometimes other work has been done to examine assessment of the abundance and placed by man, or composed of manu­ this habitat type in the Bight. Reef distribution of reef habitats in the factured materials (such as sunken ves­ habitats in the Bight, although not Bight; sels). Some biogenic micro-structures, as wide in occurrence and coverage 2) Known or probable fishery resource e.g. coralline algae; anemone, poly­ as the glacially scoured, rocky areas associations with reef habitats, sev­ chaete, or amphipod tubes; or cobble or of the Gulf of Maine (Oldale et al., eral species of which are currently dead or fossil molluscan shell patches, 1973), or the fossil coral rock and live considered “overfished” (NMFS, have some reduced characteristics of coral patch reefs south of Cape Hat­ 1999b), including some endangered a reef habitat. These can support small­ teras (Menzies et al., 1966), may have marine species; er organisms or life stages; Auster et become common enough to warrant 3) What is known of the biological al. (1995) discusses this use of micro­ consideration of their role in fishery communities that are associated with habitat. After a period of submersion management in the Bight. Although various estuarine, coastal, and conti­ and epifaunal colonization, most reef not as common or as spectacular as in nental shelf reef habitats; and habitats have a similar appearance and the tropics, reef-like habitats (especial­ 4) Discusses status and trends in reef function, but there can be subtle differ­ ly shipwrecks) also support recreation­ habitats, threats to these habitats, ences between natural and man-made al diving in the Bight, and many divers some uses of man-made reefs to reef habitats; the characteristics of each harvest reef fish resources by spear manage marine resources, and infor­ type are reviewed separately. The basic (fish) or hand (lobsters). This recre­ mational or research needs. source of information on the distribu­ ational diving generates economic ben­ tion of reef habitats is NOAA’s NOS efits to nearby businesses through sales This summary is intended to create Hydrographic Surveys Division Auto­ and services. an awareness of this habitat in the Bight mated Wreck and Obstruction Informa­ The introduction of manufactured and serve the information needs of hab­ tion System (AWOIS, 1997), although materials as reef habitat, by both ac­ itat and fishery resource managers. this database only includes structures cidental and intentional depositions, is or reefs that might be of concern to expanding in the Bight. These habitats The Middle Atlantic Bight Reefs navigation, and many small “reefs” or and their associated biological commu­ The Bight generally is defined to in­ “snags” are not included in the data­ nities need the ecosystem/community­ clude estuarine and continental shelf base. This database is augmented by a level attention given similar reef habi­ waters and seabed between Cape Cod, summary of artificial reef construction tats in the adjacent Gulf of Maine and Mass., and Cape Hatteras, N.C. It is from sources involved in various state South Atlantic Bight areas (e.g. McCar­ a broad indentation in the coast line artificial reef programs, and a survey thy et al., 1979; Hulbert et al., 1982; between these boundaries with inflec­ of relevant literature. The very abun­ Wenner et al., 1983; Chester et al., tions at the mouth of Chesapeake Bay dant and widely distributed shellfish 1984; Sedberry and Van Dolah, 1984; and New York harbor (Fig. 1). Its reefs, submerged pipelines, and inter­ Witman, 1985; Witman and Sebens, nonreef benthic environment and as­ tidal man-made structures, e.g. jetties 1988; Kirby-Smith, 1989). Man-made sociated communities have been re­ and bulkheads, are only generally con­ or artificial reef habitats are also often ported by Wigley and Theroux (1981) sidered. No effort is made in this review suggested as replacement habitats for and Theroux and Wigley (1998), and to estimate the total spatial seabed cov­ losses of other habitats, especially in es­ were reviewed by Pacheco (1988), and erage of all types of reef habitats in the tuaries where in-kind mitigation oppor­ are characterized as being composed Bight. Some of the targets listed on the tunities are often absent (Sheehy and of sediments that range from clay to maps that seem to be inland are actu­ Vic, 1992; Foster et al., 1994). They gravel, with sand being the dominant ally within rivers. are also used to create new habitat, sediment (Fig. 1). But within this soft e.g. artificial reefs to enhance fishing sediment matrix, unnoted natural and Natural reefs or surrounding artificial islands (Wil­ man-made reef habitats occur in estu­ Natural reef habitats in the Bight are liams and Duane, 1975; Seaman and aries, along the coast, across the con­ found in some areas consisting of bio­ Sprague, 1991), or are segregated as tinental shelf, and in deeper waters. A genic or rock material. Biogenic reefs special fishery management areas or review of these reef habitats and bio­ are created by living stone coral, Astran­ reserves (GMFMC, 1999). Because of logical associations follows, which in­ gia poculata, certain shellfishes (east­

62(2), 2000 25 ern oyster, Crassostrea virginica, and garis. These reef-building organisms cause they add structural complexity (or blue mussel, Mytilus edulis), and poly- have been called “physical ecosystem micro habitats) to environments, which chaete worms, such as Sabellaria vul- engineers” by Jones et al. (1997) be- in turn attracts and supports other organ­ isms. Nonbiogenic natural reef habitats are exposed rock outcrops or random boulders left by retreating glaciers or rafted from icebergs, about 12,000 years before present (YBP), or erosion of sed­ iment-covered rock or deltaic deposits of rock, cobble, and gravel along former river channels across a retreating shore­ line since the last glacial period. There are reports of submerged ridges of ara­ gonitic sandstones, thought to be relict beach deposits, mid-shelf off Delaware (Allen et al. 1969). Some natural “reefs” can be ephemeral, such as tree trunks that are washed down rivers, become water logged, and sink to the bottom to provide temporary habitat until wood­ borers gradually degrade them. Collec­ tions of dead molluscan shells, such as surf clam, Spisula solidissima; and whelks, Busycon sp.; and exposed semi­ fossil oyster shell can serve as micro­ reef habitats. Shellfish reef habitats (primarily oyster and blue mussels) are primarily known to occur in polyhaline estuaries and coastal areas in the Bight, but mus­ sels occur offshore, too. Oyster beds and reefs are found in Chesapeake Bay and its tributaries, Delaware Bay, the Hudson-Raritan Estuary, in coastal areas of Long Island Sound and South­ ern New England, including bays on Martha’s Vineyard (Ford, 1997; MacK­ enzie, 1997a, b). Oyster beds were more extensive in distribution and abundance in the nineteenth century than current­ ly, and were enhanced in many areas in the 19th and early 20th century by transplanting cultch and spat. Blue mussel beds are attached to hard surfaces in more marine and cooler coastal waters, (e.g. Steimle and Stone, 1973; Langton et al., 1995; MacKen­ zie, 1997a). They can also be found adjacent to larger reef structures after being sloughed off by strong currents or storm surges and there they continue to grow and serve as satellite, low-re­ lief reef habitats. Life spans of a decade or less, predation, and harvesting make the presence and size of these beds Figure 1.—The underlying dominant sediments types and distributions in the Middle somewhat dynamic, and they are not Atlantic Bight (from Wigley and Theroux, 1981). usually mapped as reef structures.

26 Marine Fisheries Review Rocky reefs and associated fauna are mostly found in New York Harbor, Long Island Sound, and along the Southern New England coast, and out­ crops of glauconitic marl (a soft sedi­ mentary rock) occur off northern New Jersey (the Shrewsbury Rocks and El­ beron Grounds). The AWOIS (1997) database represents a rough estimate of reef habitats, including natural rock. For example, there is evidence of ara­ gonitic fossilized sandstone ridges re­ ported off Delaware at about 80 m depth (Allen et al., 1969) that do seem to be in the AWOIS database, and there are “rocky” areas defined by AWOIS at the mouths of Delaware and Ches­ apeake Bays (Fig. 2) that are not nat­ ural, but are protective rip rap. Some natural reef habitat is also known from areas of the outer continental shelf and within submarine canyons where other outcrops of sandstone and clay occur; the exposed clay in this area is further enhanced as fishery habitat by the bur­ rowing activities of tilefish, Lopholati­ lus chamaeleonticeps, and crustaceans, such as American lobster (Cooper et al., 1987; Steimle et al., 1999f), and by soft coral colonization. These deeper reef-like habitats are not included in the AWOIS database nor on Figure 2. There are anecdotal reports by com­ mercial fishermen of cobbles and loose rock patches associated with gravelly Figure 2.—Distribution of submerged rocky reef areas (●) in the Middle Atlantic areas in coastal areas, these could rep­ Bight, based on NOAA NOS’s AWOIS (1997). The targets that appear inland are in the AWOIS database for rivers. resent river deltaic deposits during peri­ ods of lower sea levels; but some could be ballast stones from old wooden ship­ line, they added objects or structures is also created by epifauna colonizing wrecks. Off coastal Delaware and south that functioned as reefs to estuaries, the system of coastal navigation aids, these rocky patch are also associated coasts, and the shelf by building piers, such as buoys and their anchor chains with “live bottom,” i.e. the rocks are docks, bulkheads, and leaving wooden and the submerged supports of light colonized by sea whips, stone coral, shipwrecks. These structures had hab­ towers; there are presently no oil or and other biogenic structural enhancers itat value similar to submerged trees gas rigs in the Bight which would add (see below). washed down river into estuarine waters to this type of habitat. Human activi­ by storms, and in some ways they miti­ ties have introduced rocks, as protec­ Man-made reefs gated the loss of other structured, veg­ tive structures (jetties, groins, breakwa­ Although native Americans used etated habitats (coastal marshes and eel ters, and ice blockers) along or within brush and stone weirs to trap fish in grass beds) that were covered or de­ most coastal areas or bays, or as former estuaries and rivers, which might be graded by shoreline development, al­ waste material as off the mouth of New considered a form of artificial habitat though the hard structures did not fully York Harbor, i.e. the Subway Rocks, which sheltered some fish as well as mitigate lost primary productivity. In which are material removed while con­ aggregating them for collection, most approximately the last century and a structing the New York City subway man-made contributions to Bight reef half, metal vessels have gradually re­ system. habitat have occurred since European placed large wooden vessels and cre­ Shipwrecks constitute one of the most colonization in the seventeenth cen­ ated more enduring sunken structures. abundant types of man-made reef hab­ tury. As settlers developed the shore- Mid-depth to surface “reef-like” habitat itat in the Bight. A summary plot of

62(2), 2000 27 wrecks from the AWOIS (1997) data­ base suggest that they are most common near the mouths of major estuaries or ports (Fig. 3), as might be expected from the volume of shipping traffic in and out of these ports. However, many smaller or older, partially degraded or buried wrecks are not in the database or shown on these charts, and their dis­ tribution is likely to be similar to the plotted shipwrecks. Many of the ship­ wrecks on the outer continental shelf are products of WW II submarine attacks. As shipwrecks degrade, their structural complexity changes and this affects their habitat value for fishery resources. Another type of “reef” habitat noted on NOS charts and in theAWOIS (1997) database, is “Obstructions.” These are objects on the seabed of unknown com­ position, and many are probably small or degraded shipwrecks, lost anchors or deck cargo, a few airplane wrecks, and similar objects. The most notable of these are summarized on Figure 4, and, like shipwrecks, the plot focuses on larger targets of potential concern to navigation. In the last half century or so, fish­ ermen and fishery managers have rec­ ognized the value of “reef” and wreck habitats to fisheries, and they have been constructing artificial reefs in coastal waters. The use of artificial reefs for fishery enhancement has continued to Figure 3.—Distribution of Middle Atlantic Bight shipwrecks (●) in the NOAA expand, although they are a relatively NOS’s AWOIS (1997) considered to be a concern for navigation. The targets that small part of the plotted man-made or appear inland are in the AWOIS database for rivers. overall reef habitat available in the Bight (Fig. 5), in comparison with wrecks and militarized combat vehicles and a va­ abundance of this habitat in the Bight, obstructions (Fig. 3, 4). Areas where ar­ riety of ships as available at a reason­ because of the reasons previously men­ tificial reefs have been constructed are able cost (Steimle, 1982; Joint Artificial tioned. The term, “non-biogenic” is used noted by an older term, “fish havens,” Reef Technical Committee, 1998). In on this figure to note that live coral, on NOS navigation charts but are not in­ the last decade, prefabricated artificial mussels, and oyster beds/reefs are not cluded in the AWOIS (1997) database. reef units that are specifically designed included. Artificial reefs placed on the seabed spe­ and constructed as habitat for fishery re­ The concept of a reef habitat is cifically for fishery enhancement tend to sources are increasingly being deployed more complex than being either nat­ be found in coastal and estuarine waters (Sheehy and Vic, 1992). The contribu­ ural or man-made and involves a vari­ (Fig. 5) and serve primarily the rec­ tion of artificial reefs to reef-fish pro­ ety of conditions under which a reef or reational fishery. They have been built ductivity has been controversial and the reef-like conditions exists in the Bight. and developed in the Bight since the debate continues and fish or fishery pro­ Some characteristics of different reef 1920’s or 1930’s, and especially since ductivity may vary among reef locality or reef-like habitats found in the Bight the 1960’s. Initially they were devel­ and other factors (American Fisheries are summarized in Table 1. oped with a variety of recycled materi­ Society, 1997). als, ranging from Christmas trees stuck A summary of all reef or reef-like Reef-associated Fishery in concrete bases, wooden beer cases habitats, i.e. Fig. 2Ð5, is shown on Fig. 6, Resources in the Bight half filled with concrete, rubber auto­ which should be considered a minimum Reef habitats of all types within the mobile tires, to the recent use of de- estimate of the distribution and relative Bight are used by a wide variety of fish­

28 Marine Fisheries Review ery resources (Hildebrand and Schro­ eder, 1928; Bigelow and Schroeder, 1953) and a few threatened or endan­ gered species (Lutz and Musick, 1996). Table 2 summarizes most of these spe­ cies and notes their known or suspect­ ed reef habitat associations; however, it should be noted that very little in­ formation is available on the inverte­ brate or fish fauna on reef habitats in the Bight, especially in relatively deep (>30 m) waters. Although many fishery species are closely associated with reef habitats, the reef may not adequately supply all of their needs, especially food, and the availability of non-reef resources can be important to these species. For example, black sea bass, which is a common reef habitat-associated fish mostly found during the warmer months in the Bight, may obtain much of its food from the sandy bottom or water column around a coastal artificial reef habitat (Steimle and Figley, 1996). Thus, the near-reef, open bottom/water habitat and its bio­ logical resources are linked to the fish­ ery resource production function of a reef habitat. The open sandy bottom fauna near reef habitat, conversely, can be affected by the presence of a reef and its predatory fauna. The habitat needs of reef-associated fishery resources often shift during on­ togenetic development. Many reef spe­ Figure 4.—Distribution of “obstructions” (●) in the NOAA NOS navigational chart cies use the marine water column as database. The targets that appear inland are in the AWOIS database for rivers. larvae, estuarine structures as juveniles, Table 1.—Summary of physical and biological characteristics of natural and man-made reef or reef-like habitats in and gradually return to deeper and more the Middle Atlantic Bight. marine habitats at the end of their first Natural: These are submerged rocks and other hard materials or solid structures made by living organisms (biogenic materials). season. In the Bight there can also be Estuarine: Reefs in this environment consist of oysters, mussel, sponge, and tube worm beds: exposed stiff clay, peat, a seasonal shift in habitat use among or rocky outcrops; waterlogged trees; or boulder or cobble fields. Oyster and mussels are fishery resource species in their own right, and these estuarine biogenic and non-biogenic habitats provide shelter and food for a variety of juvenile subadults and adults of certain species, to adult fish (see below). i.e. winter and summer habitat use can Coastal (< 12 miles): Mussel and stone coral beds form biogenic reefs here, and other reefs consist of rocky outcrops, such as soft marl off northern N.J. and harder rock from N.Y. Harbor east along the southern New England coast also differ significantly (Steimle et al., (Fig. 2); glacial erratic boulders or cobble accumulations from eastern Long Island to Cape Cod with other submerged 1999 b-f; Steimle and Shaheen, 1999; cobble/ gravel banks reported off New Jersey, Delaware, and Maryland1; relict shell fields; exposed stiff clay or peat deposits; and kelp beds are found along southern New England. NMFS, 1999a). Shelf: Reef habitats are scarcer in deeper water but glacial erratic boulders; exposed rock or stiff clay at or near the edge of the continental shelf or at the shelf edge heads of submarine canyons; relict clay or peat deposits; and shell Other Potential or Transient fields provide patches or bands of reef-like habitat. Reef-associated LMR’s in the Bight Man-made: These types of structured habitats have been available since the 17th century, and a good part of this habitat is from shoreline construction, such as piers and jetties, but also include the remains of shipwrecks and various materials Some reef associated taxa or species, deposited to provide an artificial reef-type habitat, as per Fig. 3-5. Estuarine: This type of habitat is often formed by shoreline development, including functional and decaying bulkheads, that are not included in Table 2 and are bridge abutements, piers and docks, protective rip rap, groins and jetties; navigational aids such as lighthouses and buoys; clay or rock exposed by dredging; submerged natural gas, storm water, processed sewage effluent pipelines; not presently considered a “living re­ exposed communication cables; sunk or abandoned vessels (including ballast rock piles); and other small to medium source” in a fishery management sense, sized materials ranging from beverage containers to vehicles; waterlogged timber; and artificial reefs build of various might be of greater value in the future, reused and some specifically designed materials to support shell- and finfisheries. Coastal (< 12 miles) and Shelf: Man-made structured habitats in this broad marine zone consist of basically many of e.g. for biomedical-pharmaceutical re­ the same materials or structures as are found in estuaries, although some structures are larger in size, e.g. shipwrecks search and industry (Faulkner, 1984). and artificial reefs, but can include lost ship cargos, and exposed exploratory oil and gas pipe heads in deeper waters. Reef-associated taxa known to have 1 Monty Hawkins, partyboat captain, Ocean City Md, personal commun., 2000.

62(2), 2000 29 niidae). Below is an overview of the communities known to be commonly found on subtidal reef structures in the Bight. The type of reef surface, i.e. rock, wood, metal, or other, has a vari­ able effect on the community formed on that surface, but this effect is only briefly discussed here because of the limited available information. Intertid­ al, semi-hard surface communities (e.g. peat banks) and submerged aquatic veg­ etation also provide some of the hab­ itat characteristics of reefs, and these functions are becoming better known (Able et al., 1988) and are also not discussed here. Ducks and other verte­ brates are included in the habitat use synopsis below, when appropriate, as a reminder of their possible predatory or competitive interactions with reef fish and other reef-associated LMR’s. Sci­ entific names are included only for spe­ cies not found in Table 2.

Estuarine Reef Communities Epibenthic and Epibiotic (Organisms Attached to Reef or Shellfish Surfaces) Several types of polyhaline estua­ rine reef epibenthic communities exist in the Bight. These communities in­ clude oyster beds; blue mussel beds; communities attached to nonbiogenic Figure 5.—Distribution of artificial reefs, or fish havens, (●) in the Middle Atlantic hard surfaces such as rock, wood, and Bight, based on individual state artificial reef program data. metal; and those using semi-hard sur­ faces such as stiff clay and peat. such potential qualities include: algae many reefs inhibit the use of towed Oyster reefs The shells of oysters species, sponges, coelenterates, nudi­ fishing gear, or the reefs provide shelter support a diverse epibiotic community branch mollusks, and tunicates (Lustig­ when this gear passes over a low pro­ that can include barnacles, Balanus man et al., 1992). file structure; however, lobster and fish sp.; ribbed mussels, Geukensia demis­ Other fishery resources commonly traps and gill nets are often effective sa; and blue mussels depending on sa­ caught or observed above or near high on or near reefs. Most reefs are heav­ linity, algae, sponge, tube worms (Spi­ profile reef habitats are bluefish, Poma­ ily used by recreational fishermen and robis sp., Polydora sp., and other spe­ tomus saltatrix; mackerels and tunas, divers. cies), anemones, hydroids (Obelia sp. Scombridae; jacks, Carangidae; and and other species), bryozoa ( Membra­ some benthic species such as summer Reef Communities in the Bight nipora sp. and other species), and other flounder, Paralichthys dentata. These Reef habitats support biological com­ taxa (Watling and Maurer, 1972; Maurer fish predators take advantage of aggre­ munities (used here to mean organ­ and Watling, 1973; Kinner and Maurer, gations of prey on the reef or may be isms that are commonly found togeth­ 1977; Larsen, 1985; Zimmerman et al., behaviorally attracted to the reef struc­ er with some degree of interaction) that 1989, Coen et al., 1999). Silt accumu­ ture or the flow refuge effects of struc­ are dependent upon or which signifi­ lated between the oyster shells can sup­ ture (Westman, 1958; Figley, 1996). cantly benefit from this habitat type. port benthic invertebrates that are also The fish that frequent reef habitats These communities range from micro­ found on the soft bottoms of the area. are often less subject to commercial scopic algae and large kelp (in cooler Blue mussel beds Many organisms fishing pressure, because the relatively waters) growing on reef surfaces to are found epibiotically on mussel shells; high relief and complex structure of fishes and possibly sea turtles (Chelo­ these can include many of the same epi­

30 Marine Fisheries Review biotic organisms found on oyster beds: barnacles, algae, sponge, the same types of tube worms, hydroids, anemones, bryozoa, and slipper shells Crepidula sp.(Kinner and Maurer, 1977; Newell, 1989). Additional, nonepibiotic macro­ fauna (mainly a diversity of polychaetes and amphipods) also benefit from the mussels and live within the interstitial spaces among the mussel shells and byssus threads. Other hard surfaces (including a diversity of natural and man-made submerged materials) These surfac­ es, like that of oysters and mussels, can support algae where light is sufficient, barnacles, sponge, tube worms (includ­ ing Sabellaria vulgaris and others), hy­ droids, anemones, encrusting bryozo­ ans, oysters, blue mussels, the jingle shell Anomia sp., northern stone coral, Astrangia poculata (in more marine waters), sea whips Leptogorgia sp. (in Chesapeake Bay), tunicates Molgula sp., and caprellid amphipods (Malo­ ney, 1958; Westman, 1958; Watling and Maurer, 1972; Dean, 1977; Otsuka and Dauer, 1982). Wooden structures within estuaries can also be infested with destructive borers such as Teredo navalis, and gribbles, Limnoria ligno­ rum (Nigrelli and Ricciuti, 1970) and weathered creosoted or other antiborer­ treated pilings can become substrates for some epifaunal colonization, even Figure 6.—Summary of all reef habitats (except biogenic, such as mussel or oyster though borers and others may be tem­ beds) (●), Figs. 2-5, in the Middle Atlantic Bight. The targets that appear inland are porarily inhibited by the chemical treat­ in the AWOIS database for rivers. ment (Stewart, 1983). Semi-hard clay and Spartina peat Motile Epibenthic Invertebrates clude adults and juveniles, and species “reefs” These softer surfaces can sup­ that can be prey (e.g. gobies, Gobiso­ These mostly include decapod crus­ port burrowing mollusks (piddocks ma sp.), predators (e.g. toadfish, Opsa­ taceans, such as mud (xanthid) crabs; such as Pholus sp.,Cyrtopleura costata, nus tau), or competitors (e.g. cunner) blue crabs, Callinectes sapidus; rock Barnea truncata, Zirfaea crispata) and with resource species (Breitburg, 1999). crabs; spider crabs, Libinia emargina­ epibenthic algae (Able et al., 1988); There is a gradual shift in the fish spe­ ta; and juvenile American lobsters (al­ motile organisms, such as juvenile cies assemblages that are commonly though this species is scarce south of American lobsters and American eels, associated with reef habitats from the Delaware Bay, except in deeper waters), also occur in this habitat and are dis­ warmer waters off Virginia to the cooler and sea stars, Asterias sp. and Henri­ cussed below. waters off southern Massachusetts, and cia sp. (Jeffries, 1966; Briggs, 1975; Of note and potential interest to they are thus discussed by subregions. Leathem and Maurer, 1980; Able et al., fish recruitment success is that the Chesapeake Bight (Delaware–North 1988; Barshaw et al., 1994; Wilk et al., larvae of certain epibenthic (reef) or­ Carolina) Gobies, spot, striped bass, 1998). ganisms can be very abundant at times black sea bass, white perch, Morone in the meroplankton, e.g. barnacle cy­ americanus; toadfish, scup, drum, prids and mussel larvae, and be a sig­ Fish croaker, spot, sheepshead porgy, pinfish, nificant source of planktonic food for A number of fishes are commonly tautog, and northern puffer, Sphaer­ some larval fishery resources (Rich­ or seasonally found on estuarine reef oides maculatus; have been reported ards, 1963). or reef-like estuarine habitats. They in­ common on reef habitats in this area

62(2), 2000 31 Table 2.—List of fishery species that are commonly found on reef or reef-like habitats in the Middle Atlantic Bight.

Species Life stage/reef habitat use Notes

Algae All stage grow attached to estaurine/ Grows on inter/subtidal surfaces along southern New England coast as deep as light penetration (kelp, Laminaria sp., dulse, etc.) marine hard surfaces. allow and provides shelter; some are harvested.

Invertebrates Mollusks Blue mussel All stages grow attached to hard surfaces Colonizes intertidal/subtidal surfaces but becomes scarcer towards N.C.; important prey for many reef Mytilus edilis in polyhaline-marine waters. fishery resources; harvested as adults; increases habitat structural complexity and biodiversity.

Eastern oyster All stages grow attached to hard surfaces Colonizes hard surfaces and/or creates low profile reefs; harvested as juveniles (spat for transplanting) Crassostrea virginica in polyhaline-estuarine waters. and adults; increases habitat structural complexity and biodiversity.

Longfin squid Eggs are attached to hard objects in Hard surfaces of all sizes seem important for egg mass attachment. Eggs and larvae can be prey. Loligo pealei marine waters.

Crustaceans American lobster All post-larval stages use shelter in Lobsters are common reef habitat dwellers but are less common south of Delaware Bay; maintain Homarus americanus polyhaline-marine waters. reef habitat structural complexity by clearing burrows.

Jonah crab All post-larval stages use shelter in This larger crab is common to reef habitats; claws are harvested. Cancer borealis polyhaline-marine waters.

Rock crab All post-larval stages use shelter in Common on reef habitats as well as on most other habitats; juveniles or smaller sizes important prey Cancer irroratus polyhaline-marine waters. for fish and lobsters; claws are harvested.

Fish American eel Adults found in estuarine to coastal marine This eel is found seasonally in estuarine areas, including holes in peat banks; harvested by trap and Anguilla rostrata reefs as well as elsewhere. by recreational fishery.

Conger eel Juveniles and adults common in This larger eel preys on smaller reef fish; hard to catch but desirable. Conger oceanicus polyhaline-marine structures.

Atlantic cod Juveniles and adults common on This species feeds on reef organisms; uses structure for shelter; but only found during cooler seasons Gadus morhua polyhaline-marine reefs. south of Long Island, N.Y. to about Delaware.

Pollack Juveniles and adults common on Uses structure for shelter or for feeding; but only found during cooler seasons south of Long Island, Pollachius virens polyhaline-marine reefs. N.Y. to about Delaware.

Red hake Juveniles and adults common on Common reef habitat dweller; preys on small crabs and other organisms found on or near reefs; Urophycis chuss polyhaline-marine reefs. commercially and recreationally harvested.

Striped bass Juveniles and adults common on Juveniles use estuarine structures for shelter; adults find prey near estuarine and coastal structures. Morone saxitilus estuarine and coastal reefs.

Black sea bass Juveniles and adults common on Juveniles use estuarine and coastal structures, and adults mostly use coastal and midshelf structures Centropristis striata estuarine and coastal reefs. during warmer seasons.

Gag grouper Juveniles and adults common on southern Important but variably available fishery species off Virginia and North Carolina. Mycteroperca microlepis Bight reef habitats.

Scup (porgy) Juveniles and adults common on Small schools of this species visit coastal reefs for prey and shelter during warmer seasons; found Stenotomus chrysops estuarine and coastal reefs. offshore and to the south in the winter.

Spot Juveniles and adults common on A warm season user of reef habitats north on Chesapeake Bay. Leiostomus xanthurus estuarine and coastal reefs.

Sheepshead (porgy) Juveniles and adults common on southern Common on estuarine (including oyster beds) and coastal reefs, mostly south of Delaware Bay. Archosargus probatocephalus Bight reef habitats.

Atlantic croaker Juveniles and adults common on Common on estuarine (including oyster beds) and coastal reefs, mostly south of Delaware Bay. Micropogonias undulatus estuarine and coastal reefs.

Black drum Juveniles and adults common on Common on estuarine (including oyster beds) and coastal reefs, mostly south of Delaware Bay. Pogonias cromis estuarine and coastal reefs.

Tilefish Juveniles /adults use rocky areas or holes This species contributes to the creation and persistence of the rough bottom habitat and associated Lopholatilus chamaeleonticeps in stiff clay at the edge of continental shelf biological community found in certain areas of the outer shelf and upper slope. and upper slope.

Cunner All post-larval stages are associated with A very common small reef fish, especially in the northern Bight; prey for other fish found on or visiting Tautogolabrus adspersus marine-polyhaline reef habitats. reefs. Hibernates on reefs in cold winters.

Tautog All post-larval stages are associated with A common larger reef fish that prey heavily upon mussels; youngest juvenile found in estuaries; may Tautoga onitis marine-polyhaline reef habitats. hibernate during cold winters off New England.

Gray triggerfish Juveniles/adults are warm-season reef Found on marine reefs and preys on reef dwellers; growing in popularity as food fish. Balistes capriscus dwellers.

Ocean pout All life stages found on reef habitat, Adults make and possibly guard egg nests within reef structures during the winter. Macrozoarces americanus including eggs which are nested.

Reptilia Sea turtles Juveniles and adults of several species Sea turtles are common summer visitors to the Bight and are known to use reef structures as Eucheloniodea are associated with reefs. sheltered resting areas and can prey on reef crabs.

Mammalia Harbor seal Juveniles and adults use the above water Harbor seals are winter visitors to the northern Bight and are commonly observed on dry parts of Phoca vitulina parts of reefs as resting areas. submerged structures and may prey on associated reef fish.

32 Marine Fisheries Review An assemblage of resident Tau­ tog, Tautoga onitis, and cunner, Tautogolabrus adspersus, on some concrete pipe artificial reef material off New York. Photographer: Christopher J. LaPorta.

Tautog are common at the inter­ face between the predominant open sandy bottom of the Middle Atlantic Bight and reef-like struc­ tures. Photographer: Christopher J. LaPorta.

A tautog, Tautoga onitis, that sometimes seeks shellfish prey on nearby sandy habitats, always returns to a reef structure.

(Arve, 1960; Richards and Castagna, can eel, ocean pout, red hake, white histrionicus; other species can feed on 1970; Feigenbaum et al., 1985; Breit­ hake, cod, juvenile pollack, and various small fish they find on shallow reefs, burg, 1999; Coen et al., 1999). nonfishery species have been reported e.g. wintering loons,Gavia sp.; and New York Bight (New Jersey–south­ on mostly rocky estuarine reefs in this mergansers, Mergus sp.; and cormo­ ern Long Island, N.Y.) Cunner, toad­ area (Nichols and Breder, 1927; Able et rants, Phalacrocorax sp.; most of the fish, spot, gobies, striped bass, sculpins, al., 1988). year (Martin et al., 1951). In the winter, juvenile Atlantic cod, juvenile tautog, harbor seals, Phoca vitulina, visit the black sea bass, scup, rock gunnel, Other Vertebrates (Diving birds, seals) Bight, to at least New Jersey, and may Pholis gunnellus; conger eel, American Several species of diving ducks and find prey around reef habitats near their eel, red hake, and northern puffer have geese (Anatidae) feed seasonally upon haul out, resting places. been reported on reef habitats in estau­ submerged algae, mussels, and other ries of this area (Briggs, 1975; Auster, organisms growing upon shallow reef Coastal (to depths of ~25 m) 1989; Able et al., 1998). habitats, e.g. brant, Branta branta; Southern New England (Long Island scaup, Aythya sp.; goldeneye, Bucepha- Epibenthic Sound–Cape Cod) Cunner, toadfish, la sp., scoters, Melanitta sp., old squaw, Exposed rock–soft marl (e.g. Shrews- striped bass, scup, tautog, black sea Clangula hyemalis; eiders, Somateria bury Rocks off northern N.J.) Certain bass, rock gunnel, conger eel, Ameri­ sp.; and Harlequin ducks, Histrionicus boring mollusks (piddocks such as Cyr­

62(2), 2000 33 Sea anemones, Metridium senile, and other epifauna that are typi­ cal of a well-established hard sur­ face epifauna community in the Middle Atlantic Bight. Photogra­ pher: Christopher J. LaPorta.

White frilly patches of northern stone coral, Astrangia poculanta; anemones, Metridium senile; and various hydroids often dominate reef habitat epifaunal communities, when blue mussels, Mytilus edulis, are absent. Pho­ tographer: Christopher J. LaPorta.

Dense epifaunal growth on ex­ posed hard structure or rocks pro­ vides abundant opportunities for juvenile or large fishery resources to find shelter.

Small American lobsters, Ho marus americanus, are common within the shelter provided by reef habitats in the Middle Atlantic Bight, and come out at night to feed on or near this habitat. Photographer: Christopher J. LaPorta.

34 Marine Fisheries Review In the southern Middle Atlantic Bight large sheepshead porgy, Archosargus probatocephalus, are common residents of reef struc­ tures and feed on epifauna.

Juvenile and adult cunner, Tauto­ golabrus adspersus, are perma­ nent residents of reef habitats in the Middle Atlantic Bight. Photog­ rapher: Christopher J. LaPorta.

Blue mussels, Mytilus edulis, commonly dominate the epifaunal community on reef structures in the Middle Atlantic Bight and are readily eaten by a variety of fish and crustacean shellfish, including the cunner, Tautogolabrus adspersus, that is present in this photo. topleura costata and Zirfaea crispata) anemones (Metridium senile, Tealia sp., modiolus in deeper and cooler waters add complexity as they gradually de­ or Stomphia careola); various hydroids off southern New England; the jingle grade this substrate. The epifauna on (Tubularia sp., Obelia sp., Campan- shell Anomia simplex; bryozoans, in­ this substrate also includes that noted ularis sp.); northern stone coral, soft cluding Bugula sp.; skeleton (caprel­ below for harder substrate (Westman, coral (Alcyonaria sp.) off New England, lid) and tubiculous amphipods, such 1958). and sea whips (Leptogorgia sp.) south as Jassa falcata; and tubiculous poly­ Harder rock When available, this of New Jersey where it becomes part of chaetes, such as Sabellaria vulgaris and rock can be colonized by red algae a “live bottom” community that is most Hydroides dianthus. Much hard rock (Phyllophora sp.); sponges such as Ha­ common south of Virginia; barnacles; reef structure has been added to the lichondria sp. and Polymastia sp.; large blue mussels, horse mussels Modiolus shoreline of the Bight in the form of

62(2), 2000 35 jetties, groins, rip rap, and groins that genbaum et al., 1985; Lavalli and Bar­ most of the species noted in the above abut the shoreline as beach and other shaw, 1986; Karnofsky et al., 1989a, b; Coastal section. The epifaunal coloni­ protection; these can support less sta­ Figley and Dixon, 1994; Mercaldo-Al­ zation of the exposed outer shelf sand­ bile communities because of the great­ len and Kuropat, 1994; Chee1). Mass stone ridges, reported by Allen et al. er environmental extremes and stresses aggregations of Jonah and rock crabs (1969), is unreported. that occur in this littoral zone, although were reported on Southern New England Wrecks and artificial reefs Size, fishery resources frequent them when rocky ledges (Auster and DeGoursey, composition, location, and age affect the rocks are covered by water. 1983). Loligo squid usually attach their the structure and habitat value of these Wrecks and artificial reefs (of various egg clusters to a hard object, such as a reefs. These factors also affect the hab­ compositions) Wood: When exposed, reef surface, or shells, gravel, or other itat value of artificial reefs that were this provides a substrate similar to soft hard surfaces on the seabed (Griswold specifically constructed to support fish­ rock, but can include wood borers (e.g. and Prezioso, 1981; Roper et al., 1984). ing. Few scientific studies are known of Teredo sp. and Xylophaga atlantica) the epibenthic fauna of wrecks and arti­ that also degrade this type of reef mate­ Fish ficial reefs on the Bight shelf, and those rial, which is why they do not usually Chesapeake Bight Black sea bass, that are known are mostly within the persist as significant three-dimensional pinfish, Lagodon rhomboides; scup, coastal zone (Figley, 1989; Figley and structures or habitat for more that a cunner, red hake, gray triggerfish, black Dixon, 1994); or are not quantitative or few decades, unless periodically cov­ grouper, Mycteroperca bonaci; smooth are anecdotal (Bulloch, 1965). ered and protected by sediment. dogfish, Mustelus canis; summer floun­ Other solid substrates Exposed sub­ Metal: As per hard rock, but the ten­ der, scads, Decapterus sp.; bluefish, and marine communication cables can be dency of sheets of rust to slough off amberjack, Seriola dumerili, have been colonized by borers and epifauna (Snoke, the wreck surface creates a less stabile reported as common over these reefs 1957), and can serve as limited reef hab­ community; these surfaces are often (Feigenbaum et al., 1985; Chee1). itat for organisms of suitable size. colonized by hydroids, like Tubularia New York Bight Atlantic cod, gray crocea; anemones, mostly Metridium trig ger fish, scup, black sea bass, tau­ Motile Invertebrates senile; northern stone coral; blue mus­ tog, ocean pout, red hake, conger eel, Most of the motile organisms noted sels; barnacles; sea stars; and related cunner, sea raven, Hemitripterus amer­ above for the coastal areas are also com­ fauna (Bulloch, 1965; Feigenbaum et icanus; and rock gunnel have been re­ monly found on the shelf, although at the al., 1985: Chee1). ported on reefs in this area (Westman, shelf edge, some deepwater taxa occur Kelp Laminaria sp. beds occur in 1958; Briggs, 1975; Steimle and Ogren, (Cooper and Uzmann, 1971; Wigley and Long Island Sound and north and are 1982; Woodhead et al., 1985; Figley Theroux, 1981; Theroux and Wigley, noted separately from the seaweeds and Dixon, 1994). 1998). listed elsewhere. Kelp grows only on Southern New England Scup, black hard reef-like surfaces and adds verti­ sea bass, tautog, Atlantic cod, ocean pout, Fish cal relief and more complexity to reef red hake, conger eel, cunner, sea raven, Chesapeake Bight Reef fish include habitats, as well as contributing to pri­ and radiated shanny, Ulvaria subbifurca­ resident species (black sea bass, scup, mary and detrital production (Alfieri, ta, have been reported on these reefs (Al­ tautog, and cunner), seasonal residents 1975). fieri, 1975; Carr and Amaral, 1981). (gag, sheepshead porgy, round herring, Other materials (such as various and sardines), or transients (amber­ plastics and synthetic materials, such as Other Vertebrates (Diving birds, seals) jack, spadefish, gray triggerfish, vari­ rubber and concrete) These epiben­ These use shallow coastal or shore­ ous mackerels and small tunas, and the thic communities can be similar to that line reefs, e.g. jetties, as per estuarine spot-tailed pinfish, Diplodus holbroo­ found on rock and metal reefs (Alfieri, reefs, above. ki) (Eklund and Targett, 1991; Adams, 1975; McCullough, 1975; Woodhead 1993). and Jacobson, 1985; Figley, 1989). Shelf (generally depths >25 m) New York Bight Gray triggerfish, scup, black sea bass, tautog, Atlantic Motile invertebrates Epibenthic cod, ocean pout, red hake, conger eel, American lobsters, rock crabs, Jonah Rocks and boulders Few rocky cunner, sea raven, rock gunnel, pollack, crabs, spider crabs, sea stars, and ur­ ledges are found on the Bight’s conti­ and white hake have been commonly chins, Arbacia punctulata, are found nental shelf (these occur off southern reported on these deeper reefs (Bulloch, on these reefs (Westman, 1958; Cobb, New England), and most rocky habitat 1965; Woodhead et al., 1985). 1971; Briggs and Zawacki, 1974; Briggs, consists of boulder and cobble residue Southern New England Scup, black 1975; Alfieri, 1975; Sheehy, 1976; Fei­ from periods when glaciers covered sea bass, tautog, Atlantic cod, ocean New England and icebergs drifted in pout, red hake, conger eel, and cunner the Bight, the last glacial period being occur here (Auster, 1984), as well as 1 Chee, P. K. 1976. The ichthyofauna of the Ches­ apeake Light Tower. Old Dominion Univ., Nor­ about 12,000 YBP. Reports of the epi­ other species that are found in the New folk, Va., Unpubl. Rep., 26 p. fauna in this Bight substrate include York Bight.

36 Marine Fisheries Review Outer shelf reefs and clay burrows Island Sound, and along the Southern Wooden shipwrecks and other sub­ (and the “pueblo village community” New England coast, and may be fairly merged wooden structures can be res­ along southern Georges Bank) Tile­ static, although some rock reefs have ervoirs and sources of the nuisance fish, white hake, and conger eel are been removed or reduced because of borers, noted above, which can attack reported using this habitat, as well as their hazard to deep-draft ship naviga­ wooden (and some concrete and plas­ smaller species (Valentine et al., 1980; tion, or been covered by shoreline de­ tic-coated) pilings within harbors (Ni­ Cooper et al., 1987). velopment or silt. The reports of sand­ grelli and Riccuiti, 1970). All man­ stone outcrops on the continental shelf made artificial reefs are subject to deg­ Other Vertebrates off New Jersey and Maryland need fur­ radation processes to variable degrees This habitat is too deep for those bird ther investigation as to their significance (depending on the material used and or seal species that could use estuarine as fishery habitat. Oyster reef habitats the severity on environmental condi­ and coastal reefs. in many estuaries are greatly reduced, tions they are exposed to) and some especially since the 1950’s (MacKen­ types were formerly subject to move­ Significance, Status, and Trends zie, 1997a, b). Made-man “reefs” of all ment of certain reef material out of per­ of Bight Reef Habitats and LMR’s sizes are continuing to add this type of mitted reef sites, such as automobile The reef habitats in the Bight are sig­ habitat to the environment in all areas tires, which interfered with other uses nificant to fisheries because they expand via accidental, careless, and intention­ of the coastal zone, e.g. trawling and the range of some reef-associated spe­ al means. South of Long Island Sound, bathing. This problem mostly occurred cies and perhaps their population abun­ man-made reef habitats probably con­ in the early formative and experimental dance, possibly beyond the apparent tribute significantly to the wider distri­ phases of artificial reef development, extent of seabed area the reefs cover, bution of reef-associated species. although some old artificial reefs are as per Figure 6. The growing array or Although there have been studies still the source of material lost from the network of reef habitats (all types), that of epibenthic fouling (Maloney, 1958) reef sites. occur or have become established or and fishery use of reef habitats in the Another aspect of the creation of specifically placed along the coast and Bight (Figley, 1996), reef habitats in reef habitat (accidental or intentional) across the shelf, can also provide corri­ this area have not been well studied or on open bottom is that the reef and its dors of supporting habitat for some reef­ examined holistically, with all biologi­ associated community displaces previ­ associated species, such as black sea cal components and their interactions ous open bottom communities (Shipp, bass, to use during seasonal migrations defined. Questions remain about how 1999). It is normally assumed that open (Fig. 6). As more fine-scale, side-scan man-made reef habitats have compen­ bottom habitat is not limited, at least sonar seabed mapping becomes avail­ sated for functional losses in natural in the Bight, and that open habitat loss able, more low profile hard bottom and reef habitats or other sheltering habi­ caused by the development of new reef reefs will undoubtedly become known tats, such as eel grass beds, caused by habitats will not significantly effect the or identified, including many of the man-made alterations, such as exces­ function of this habitat to support desired objects classified as “obstructions” by sive nutrient and silt inputs, and toxic population levels of other managed de­ NOS or known as “snags” by fisher­ pollution. mersal species. It is also a value judge­ men. Table 2 shows that a variety of Not all reef-associated organisms are ment of whether the reef habitat fishery important commercial and recreational benign or beneficial. Some “reef” spe­ resource community is of equal or great­ fishery resources are known to be as­ cies can be nuisances or cause human­ er value as a harvestable or economic re­ sociated or depend on reef habitats for interaction problems, e.g. wood-de­ source, than to that of the soft-bottom some or most of their life history, and stroying Teredo “shipworms,” gribbles, community it replaces. These issues are reef structures can serve as refuges from and other borers that attack wooden not easy to resolve, and little informa­ trawling for some species. Certain com­ pilings and vessels; fouling organisms tion is available for such comparisons. mercial fisheries, such as American lob­ that inhibit vessel use efficiency, weigh Several reef-associated fishery spe­ ster or fish trapping and gill netting, down navigational aids, and fill pipe­ cies are presently considered over- or favor reef habitats, and recreational fish­ lines that cycle estuarine or marine fully exploited; these include Atlantic ermen have long valued wrecks and waters. Other species can be a source cod, haddock, pollack, scup, black sea reefs as fishing grounds. It is also be­ of larval resource species predators, bass, ocean pout, tilefish, striped bass, coming evident that several forms of es­ e.g. carnivorous coelenterates such as and American lobster (Clark, 1998; tuarine and coastal reef habitats serve as anemones, or support the sessile stages NMFS, 1999b). Oysters (as a fishery juvenile fish nurseries in the Bight, e.g. of “jellyfish,” such as Chrysaora quin­ resource in this region) persist in de­ oyster and mussel beds, and artificial quecirrha and Cyanea capillata. clining natural beds and under cul­ reefs can increase this function (Heise Besides these nuisance organisms, ture; but the profitability of labor-in­ and Bortone, 1999). the obvious hazards to vessel naviga­ tensive culture is declining because of Natural reef habitats occur primarily tion and safety, and causing the loss increasing costs and stagnant prices in the Bight from within and just out­ of towed fishing gear and anchors, per bushel, and continued investment side New York Harbor, through Long reefs have other negative side effects. in maintaining these beds is in ques­

62(2), 2000 37 tion (C. MacKenzie, Jr.2). Although the 5) Dischar ges or accidental coastal habitat fragmentation for reef-as­ role of habitat in the conservation of spills of toxic materials. sociated species and perhaps sup­ these species is still poorly understood, 6) Removal of docks and piling fields porting their population expansions, reef habitat losses, at least in estuarine in urbanized estuaries; these fields especially on the continental shelf nursery grounds, can be suspected as may replace lost shoreline trees, south of Long Island where reef hab­ being contributory (Rothschild et al., and SAV or oyster beds and other itat is naturally limited (Fig. 2). The 1994). submerged or partially submerged system of wrecks and man-made natural habitat formerly available reef-like structures across and along Human Threats to Bight as LMR habitat. the continental shelf and coastal Reef Habitats and LMR’s 7) Nonpoint source pollution in estu­ areas has created an array of reef Threats to reef habitats, in general, arine and coastal waters, some of habitats (only partially evident on have been discussed by Bohnsack which can be toxic to reef dwellers. Figure 6 owing to the limitations of (1992), with emphasis on coral reefs. 8) Loss of biogenic reef habitat, such the NOS database) that can act as He noted inadequate knowledge of reef as oyster and mussel beds, because corridors or a network that better functions, a variety of habitat effects dis­ of disease, overharvesting, other supports the migrations or other cussed below, and overfishing as being factors in this listing or unknown movements of shelter-using species, responsible for causing alterations in factors (Rothschild et al., 1994). such as black sea bass and Ameri­ reef community structure and fishery 9) A strong preference or dependence can lobster, than occurred a century productivity. There are the “normal” on reef habitats can cause reef or two ago. Although, in the absence or natural threats to reef communities, fish and lobster mortalities during of better shelter, fish and megafauna as well, including damage from severe occasional estuarine-coastal epi­ often create or use previously creat­ storms or other climatic events, and sodes of anoxia/hypoxia (Ogren ed shallow depressions in soft sedi­ geophysical processes, such as changes and Chess, 1969; Steimle and Sin­ ments or low-profile biogenic struc­ in sea level and temperatures, but the dermann, 1978). ture (e.g. amphipod, polychaete, or scope of these natural threats is usually 10) Power plant water use can kill the anemone tubes) or molluscan shell beyond human management. eggs and larvae of reef-associated accumulations for shelter (Auster et A listing of most other threats to species that are generally available al., 1995). temperate reef habitat in the Bight, in waters drawn into the plants, 2) Maintain biodiversity: Hard-bottom besides those named by Wilbur and but especially from the spawning or reef habitats maintained or in­ Pentony (1999), includes, in no partic­ of species that use the rip rap or troduced among other habitat types, ular order: other hard surface linings of water such as flat sand or mud bottoms, can intake canals or conduits as reef increase habitat structural and bio­ 1) Removal of reef habitats deemed habitat. diversity for algae, invertebrates, and navigational hazards or for channel 11) Coastal or estuarine sand mining fish (Sebens, 1991). deepening. and redistribution activities. 3) Provide refuge from excessive or 2) Siltation of reef habitats owing to damaging fishing: Introduction of ineffective land or coastal soil ero­ Man-made Reefs as LMR and solid high profile and complex-sur­ sion control, or increased resuspen­ Habitat Management Tools faced reef structures into an area sion and distribution by vessel traf­ The planned placement of reef struc­ can restrict the use of certain towed fic, dredging, etc., which also causes tures is a habitat management action, fishing gear in that area and pro­ the loss of low-structure, submerged and this management action can be mote a refuge function (Bombace, aquatic vegetation (SAV) beds. applied for various purposes, besides 1997). Artificial reefs that are under 3) Damage to older wrecks and to algae that of traditional fishery enhancement the control of state or Federal fish­ and coral beds by towed fishing gear or one-to-one reef habitat replacement. ery resource agencies can be estab­ and large anchors (and in the 19th Below is a list of habitat or fishery lished as special management zones, century by dynamiting wrecks to get issues that can be addressed by reef reserves, or refuges from all or most valuable metals and to harvest fish). habitat conservation, expansion, or ma­ harvesting (Bohnsack, 1992). 4) Burial by dredged material disposal, nipulation, in no particular order: 4) Expand limiting habitat: In some including beach sand nourishment areas of the Bight, reef-associated loss of shallowwater wrecks, and by 1) Habitat fragmentation: Artificial fauna can be habitat limited because shoreline jetties and groins. reefs can mediate loss of structured of inter- and intra-species competi­ habitat (reefs and SAV), especially tion for shelter (Richards and Cobb, in estuaries (Rothschild et al., 1994; 1986). As the density of all types Eggleston et al., 1998). The contin­ of reef structures increase in the 2 C. L. MacKenzie, Jr., U.S. Dep. Commer., ued input of man-made reef mate­ Bight and are more widely distrib­ NOAA, NMFS, J. J. Howard Marine Science Laboratory, Highlands N.J. Personal commun., rial (intentionally or otherwise) into uted across the open, sandy bottom, 2000. the environment is reducing natural there will be more opportunity for

38 Marine Fisheries Review epifaunal species with relatively short less is known about these habitats and 6) Ecological stability and dynamics lived larvae to find suitable habitat their associated biological and fishery of epifaunal communities should for colonization and thus gradually communities than is known for open be better studied because many expand the distribution of hard sur­ seabed habitats and communities, and “fouling” species are relatively un­ face, reef dwellers. often only anecdotal information or ob­ stable and unpredictable in time 5) Maintain access for land-based fish­ servations are available. However, ship­ and space because of their unpre­ eries: Some reef-like structures in wrecks and reefs are now being given dictable recruitment and low survi­ estuaries, such as obsolete docks and serious scientific attention as habitat for vorship, although some encrusting piers in ports, can be preserved or epifauna and fishery resources (Leewis communities of bryozoans, com­ maintained because, besides attract­ and Waardenburg, 1991). But the char­ pound ascideans, colonial cnidaria, ing certain fishery resources or their acteristics and advantages of this habi­ and sponges are long-lived. (Day­ prey to submerged structures, they tat and its biological association are not ton, 1984), and little is known of can also be used by human urban considered in analyses and multiple re­ long-term community dynamics of dwellers to support subsistence or gression models of associations among subtidal reef communities, which low-income recreational fisheries mixed demersal species, which often can have several community states, (Hawkins et al., 1992; Able et al., use only bathymetric and thermal met­ some to the disadvantage of fishery 1998). Docks and piers also offer rics, but not benthic habitat structure resources, i.e. when coelenterates physically disabled people more op­ as a distribution variable (Colvocoress­ dominate (Steimle et al., 1999a). portunities to participate safely and es and Musick, 1984; Murawski and 7) To support the development of EFH comfortably in recreational fishing. Finn, 1988). The developers and man­ parameters for managed fishery re­ 6) Estuarine/coastal nutrient removal: agers of artificial reefs for recreational sources associated with structures Experiments with artificial reefs sug­ fishing have noted some of their general or reefs, an inventory of significant gest an abundant filter-feeding epi­ information needs (Steimle and Meier, structured habitats and concentra­ fauna supported by reef surfaces, 1997), and these include: feasibility of tions of habitats with limited struc­ such as mussel populations, can po­ estuarine applications, a better under­ tural components, e.g. shell fields tentially reduce and entrain eutro­ standing of noncoral reef community that may be important to juve­ phic phytoplankton production in a ecology, better information on the life niles, are needed, especially in the reef area and in some cases reduce histories of reef-associated species, best Bight. More side-scan sonar sur­ nutrients when attached algal colo­ reef designs for different applications veys are needed to define the dis­ nies are established (Laihonen et al., and situations, better information on the tribution and abundance of hard­ 1997). population dynamics of reef resources, bottom and reefs in estuarine and 7) Compensation for unavoidable hab­ more information on reef productivity marine waters, as per Smith and itat loss: In cases where aquatic hab­ compared to other habitats, more infor­ Greenhawk (1998), and to define, itat loss is unavoidable, and in-kind mation on the use of reefs to mitigate document, and monitor the shape habitat replacement is not available habitat loss, and socioeconomic data on and surface complexity of various or feasible, reef habitats can be used the value of reef fisheries. wrecks and reefs and link this with to enhance productivity of fishery Some other specific information needs LMR use. forage species, if not fishery species, for the Bight include, in no particular 8) Small scale spatial studies of the as out-of-kind mitigation (Burton et order: functional aspects (use) of reef hab­ al., 1999). itats are needed especially to relate 8) Provide opportunities for scientific 1) Diurnal and seasonal use of Bight the size and complexity of reef research: The ability to plan the de­ reef habitats by LMR species. habitats and interactions with hy­ ployment and design of artificial 2) Use of Bight reef habitats by the drographic environment with LMR reefs offers a range of opportuni­ early benthic phase of LMR species. use (Auster, 1988). ties for scientists to test hypotheses 3) Winter habitat requirements of south 9) The experimental approaches that about issues, including the behav­ and offshore migrating Bight LMR’s are conducted in warmer temperate ior of reef-associated species, the with strong summer reef-associa­ or tropical waters on the use and role of habitat size and complexity tions, e.g. black sea bass and scup. value of reef habitat, and the inter­ in species’ use, and a number of 4) Role of man-made reef habitat in relationships of habitat size, shape, other topics, such as those discussed conserving and enhancing fishing and complexity of fishery species below. resources is poorly known, includ­ use (Bohnsack et al., 1994) need to ing the use of these reefs as refuges be attempted in the cooler temper­ Information Deficiencies or reserves. ate waters of the Bight to address and Research Needs 5) Ecology and LMR species use of similar information issues. Because of the difficulty in surveying deep-water reefs, besides the sub­ 10) The generators of waste materials many reef habitats without the use of mersible studies associated with tile­ that are not readily or cost-effec­ divers or remote in situ cameras, much fish and submarine canyon habitat. tively recyclable or disposed of on

62(2), 2000 39 land, will continue to consider the vided location data on their current reef Bohnsack, J. A. 1992. Reef resource habitat pro­ tection: the forgotten factor. In R. A. Stroud sea as an disposal option and a sites. We also thank Annette Kalbach (Editor), Stemming the tide of coastal fish hab­ wide diversity of solid materials, for her assistance, and Bob Reid, Dave itat loss. Mar. Recreational Fish. 14:117Ð129. including nonsolids such as in­ Mountain, and Peter Auster for their ______, D. E. Harper, D. B. McClellan, and M. Hulsbeck. 1994. Effects of reef size on col­ cineration ash incorporated into helpful comments and suggestions. onization and assemblage structure of fishes at a solid matrix such as concrete, artificial reefs off southeastern Florida, U.S.A. Literature Cited are often proposed for use in de­ Bull. Mar. Sci. 55:796Ð823. Able, K. W., K. L. Heck Jr., M. P. Fahay, and C. Bombace, G. 1997. Protection of biological habi­ veloping artificial reef materials. T. Roman. 1988. Use of salt-marsh peat reefs tats by artificial reefs. In A. C. Jensen (Editor), Inadequate information is usually by juvenile lobsters on Cape Cod, Massachu­ European artificial reef research: proceedings available on the sustained value of setts. Estuaries 11(2):83Ð86. of the 1st EARRN Conference, Ancona, Italy, ______, J. P. Manderson, and A. L. Stud­ March 1996, p. 1Ð15. Southampton Ocean­ these diverse materials as non-tox­ holme. 1998. The distribution of shallow water ogr. Cent., U.K. ic and positive habitats for fishery juvenile fishes in an urban estuary: the effects Breitburg, D. L. 1999. Are three-dimensional of manmade structures in the lower Hudson structure and healthy oyster populations the resources. River. Estuaries 21:731Ð744. keys to an ecologically interesting and impor­ 11) Models are needed to explore the Adams, A. J. 1993. Dynamics of fish assemblages tant fish community? In M. W. Luckenbach, effects of changing the density or associated with an offshore artificial reef in R. Mann, and J. A. Wesson (Editors), Oyster the southern Mid-Atlantic Bight. Coll. Wil­ reef habitat restoration: a synopsis and syn­ distribution of artificial reef habi­ liam and Mary, Gloucester Point, Va., M.A. thesis of approaches. p 239Ð250. VIMS Press, tats on resident or seasonal fishery Thesis, 98 p. Coll. William and Mary, Gloucester Pt., Va. resource population dynamics and Alfieri, D. J. 1975. Organismal development on Briggs, P. T. 1975. An evaluation of artificial an artificial substrate 1 July 1972Ð6 July 1974. reefs in New York’s marine waters. N.Y. Fish. movements. Estuar. Coastal Mar. Sci. 3:465Ð472. Game J. 22:51Ð56. Allen, R. C., E. Gavish, G. M. Friedman, and J. ______and C. S. Zawacki. 1974. American Summary E. Sanders. 1969. Aragonite-cemented sand­ lobsters at artificial reefs in New York. N.Y. stone from outer continental shelf off Dela­ Fish Game J. 21:73Ð77. Man-made habitats have become a ware Bay: submarine lithification mechanism Bulloch, D. K. 1965. The development of the growing integral part of the coastal and yields product resembling beachrock. J. Sedi­ wreck, Pinta, as a marine habitat. Underwater ment. Petrol. 39:136Ð149. Nat. 3(1):17Ð19, 31Ð32. shelf ecosystem in the Bight over the last American Fisheries Society. 1997. Special issue Burton, W., J. S. Farrar, F. Steimle, and B. Conlin. 150 years and they support populations on artificial reef management. Fisheries 22(4): 1999. An artificial reef in Delaware Bay, of managed fishery resources, at least 4Ð36. U.S.A. II. assessment of out-of-kind mitiga­ Arve, J. 1960. Preliminary report on attracting tion success. Proceedings of the 7th Confer­ seasonally. Recognition of these habi­ fish by oyster-shell plantings in Chincoteague ence on Artificial Reefs and Related Aquatic tats, their availability, and their use by Bay, Maryland. Chesapeake Sci. 1:58Ð65. 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Sci. 8:67Ð75. Clark, S. H. (Editor). 1998. Status of fishery ways of shelter-associated species that ______. 1989. Species profiles: life histories resources off the Northeastern United States may benefit from the growing array of and environmental requirements of coastal for 1998. U.S. Dep. Commer., NOAA Tech. fishes and invertebrates (North Atlantic and Memo. NMFS-NE-115, 149 p. man-made reef habitats. Recognition is Mid-Atlantic)—tautog and cunner. U.S. Army Cobb, J. S. 1971. The shelter-related behavior a step toward conserving and making Corps Engr. Rep. TR-EL-82-4, 13 p. of the lobster, Homarus americanus. Ecology better use of this habitat to support appro­ ______and R. E. DeGoursey. 1983. Mass 52:108Ð115. aggregations of Jonah and rock crabs. Under­ Coen, L. D., M. W. Luckenbach, and D. L. Breit­ priate fishery management goals, by hab­ water Nat. 14(2):24Ð25. burg. 1999. The role of oyster reefs as essen­ itat manipulation or enhancement, and ______and R. W. Langton. 1999. The effects tial fish habitat: a review of current knowledge thus diversifying fisheries in many areas. of fishing on fish habitat. In L. R. Benaka and some new perspectives. In L. R. Benaka (Editor), Fish habitat: essential fish habitat (Editor), Fish habitat: essential fish habitat This summary is also a preliminary effort and rehabilitation, p. 150Ð187. Am. Fish. Soc. and rehabilitation, p. 438Ð454. Am. Fish. Soc. towards developing a baseline habitat in­ Spec. Symp. 22, Bethesda, Md. Spec. Symp. 22, Bethesda, Md. ______, R. J. Malatesta, and S. C. LaRosa. Colvocoresses, J. A., and J. A. Musick. 1984. ventory for the Northeast Region. 1995. Patterns of microhabitat utilization by Species associations and community compo­ mobile megafaunal on the southern New Eng­ sition of Middle Atlantic Bight continental Acknowledgments land (USA) continental shelf and slope. Mar. shelf demersal fishes. Fish. Bull. 82:295Ð313. Ecol. Prog. Ser. 127:77Ð85. Cooper, R. A., and J. R. Uzmann. 1971. 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