Site by Bennett Web & Design Co.
2008 SOUTHEASTERN NATURALIST 7(2):191–206
Reef Morphology and Invertebrate Distribution at
Continental Shelf Edge Reefs in the South Atlantic Bight
Sarah B. Fraser1,2,* and George R. Sedberry3
Abstract - Video footage recorded from 14 submersible dives on the continental shelf
edge was used to describe and categorize reef morphology and quantify density and
number of morphotypes of large sponges and corals. Significant variation in number
of morphotypes and density of three dominant species among temperature classes,
depth classes, and reef morphology categories was tested using a multiple response
permutation procedure. The greatest densities of Ircinia campana, Stichopathes sp.,
and Muricea pendula, and the largest numbers of morphotypes were found between
18.1 and 21.0 °C and at depths between 51.0 and 60.9 m. Among reef morphology
types, those that contained unconsolidated sediments such as “sand” and “large boulders
with sand” exhibited the lowest densities and richness of morphotypes, while
“block-shaped boulders,” “buried block-shaped boulders,” and “low-relief bioeroded”
reefs had the greatest densities and largest numbers of coral and sponge morphotypes.
Rocky reefs along the shelf edge with rough texture, complexity, and relief provide
favorable conditions for epibenthic invertebrates. The warming and stabilizing effect
of the Gulf Stream along the continental shelf edge allows some sessile macrofauna to
inhabit deeper waters and more northern latitudes.
Rocky bottom formations are found throughout the South Atlantic Bight
(SAB) between Cape Canaveral, FL and Cape Hatteras, NC among large expanses
of sand (Miller and Richards 1980, Struhsaker 1969). Rocky outcrops
and hard bottom reefs at the continental shelf edge are important habitats
for sessile marine invertebrates and they are often densely encrusted with
sponges, corals, tunicates, algae, hydroids, and bryozoans because they provide
stable attachment surfaces (Barans and Henry 1984, Miller and Richards
1980, Struhsaker 1969, Van Dolah et al. 1994, Wenner et al. 1983). Variations
in rock morphology and amount of relief (Barans and Henry 1984), as well as
differences in density and diversity of epibenthic fauna (Wendt et al. 1985;
Wenner et al. 1983, 1984), have been observed among hard bottom habitats of
the shelf edge. However, the relationship between reef morphology and density
and richness of sessile invertebrates has not been examined.
The shelf edge is composed primarily of limestone, sandstone with calcareous
cement, or carbonate sediments at depths of 45–110 m (MacIntyre
and Milliman 1970, Parker 1986, Struhsaker 1969). Shelf-edge reefs vary
from low-relief (<0.5 m) outcrops with rounded or gentle slopes to highrelief
(2.0–15.0 m) scarps and ridges with steep walls (Barans and Henry
1Environmental Studies Program, College of Charleston, 66 George Street, Charleston,
SC 29424. 2Current address - 11 Jett Court, Asheville, NC 28806. 3Gray’s Reef
National Marine Sanctuary, 10 Ocean Science Circle, Savannah, GA 31411. *Corresponding
author - email@example.com.
192 Southeastern Naturalist Vol.7, No. 2
1984). Temperature and salinity on the shelf edge remain relatively stable
throughout the year due to infl uence from the Gulf Stream (Matthews and
Pashuk 1986, Miller and Richards 1980, Struhsaker 1969). However, lateral
meanders in the path of the Gulf Steam at specific locations allow cold upwellings
and intrusions towards the coast (Atkinson and Targett 1983, Bane
and Brooks 1979, Blanton et al. 1981, Mathews and Pashuk 1986, Menzel
et al.1993). Hydrographic stability, warmth provided by the Gulf Stream,
and hard substrate produce favorable conditions for tropical and subtropical
species to permanently inhabit the continental shelf as far north as North
Carolina (Cerame-Vivas and Gray 1966, MacIntyre and Pilkey 1969, Miller
and Richards 1980, Struhsaker 1969, Wenner et al. 1983).
Previous studies investigating hard bottom reefs within the SAB concentrated
on general descriptions of habitats (Barans and Henry 1984), fish
assemblages (Sedberry and Van Dolah 1984), fish spawning habitats and behavior
(Sedberry et al. 2006), occurrence of invasive species (Meister et al.
2005), and ichthyoplankton assemblages (Powles and Stender 1976). Early
studies of epibenthic invertebrates primarily focused on cataloging the number
and type of species found (Cain 1972, Cerame-Vivas and Gray 1966, MacIntyre
and Pilkey 1969, Pearse and Williams 1951). More recent work has described
assemblages of SAB invertebrates in terms of depth, latitude, and seasonality
(Wenner et al. 1983, 1984). Association with artificial reefs (Wendt et al. 1989)
and evaluation of damage to sessile biota from trawling (Van Dolah et al. 1987)
have also been investigated. This research project describes reef habitat and assesses
density and richness of sessile megafauna in relation to reef morphology.
Large sessile invertebrates include sponges and corals, which anchor to the sea
fl oor substrate as larvae. As they grow, these organisms contribute to reef complexity
and support a variety of other invertebrate and fish species.
Field site description
Hard bottom reef sites were selected for sampling based on known areas of
shelf-edge reef (Barans and Henry 1984, Wenner et. al. 1984, Van Dolah et al.
1994), the occurrence of reef fish species (Miller and Richards 1980, Sedberry
and Van Dolah 1984, Sedberry et al. 2006), and historical data on snappers
(Family Lutjanidae) and groupers (Family Serranidae) obtained by the Marine
Resources Monitoring, Assessment, and Prediction (MARMAP) program, a
fishery-independent monitoring survey conducted in the SAB by the South
Carolina Department of Natural Resources (SCDNR) since 1973. Sites were
located along the shelf edge between north Florida and northern South Carolina
(Fig. 1). Dive locations were accessed by the R/V Seward Johnson II and
the Clelia submersible during 2001, and the R/V Seward Johnson and Johnson
Sea Link II (JSL II) submersible during 2002 and 2003. In 2001, three dives
were conducted on the Savannah Scarp, a shelf-edge reef east of Savannah, GA
(Fig. 1). In 2002, eight dives were made at five shelf-edge locations: St. Augustine
Scarp, Jacksonville Scarp, Julians Ridge, Scamp Ridge, and Georgetown
Hole (Fig. 1). In 2003, two dives were made, one each at Tattler Town and Razorback
2008 S.B. Fraser and G.R. Sedberry 193
The Clelia and JSL II submersibles were equipped with a SeaBird Electronics
(SBE) 25 Sealogger CTD to record environmental variables, and a
video camera with two fixed distance lasers for measurements of fish length.
Depth, temperature, salinity, and the laser points were visible on the videotape.
The submersible conducted four-minute transects along the length
of the main reef at a distance of 1–2 m off the bottom. Transect width was
Figure 1. Location of submersible dive sites 2001–2003 on the continental shelf between
north Florida and northern South Carolina. Dives were conducted by Johnson
Sea Link II and Clelia submersibles.
194 Southeastern Naturalist Vol.7, No. 2
approximately 7 m, and average transect length was 76.7 m. Location fixes
of the submersible’s position were taken using the ship’s ORE Trackpoint II
system. Currents and bottom topography caused variations in submersible
speed and altitude. During each transect, the video camera mounted on the
sub was zoomed out to the furthest extent and pointed down towards the seafl
oor at approximately a 45-degree angle. The video camera recorded visual
observations and audio notes onto mini digital video (DV) tapes.
Analysis of videotapes
Some transects were excluded from analysis because of poor video quality
or changes in camera zoom during the transect. Examination of invertebrate
fauna was limited to large, erect sponges and corals because it was not possible
to accurately distinguish other organisms and quantify them from the
videotape. Encrusting sponges were not included. Large sponges and corals
were identified to the lowest possible taxon based on known species
distributions, assistance from marine invertebrate experts, and published
identification guides (Hooper and van Soest 2002, Smith 1971, Veron 2000).
Some organisms seen on videotape could be compared to reference material
collected by the submersible during the dives. Without specimens for microscopic
examination, it was impossible to make accurate identification to the
species level of many sponges and corals. Therefore, morphotypes were counted,
and each assigned a unique identification code (sequential number) with a
description of the external morphological characteristics used to differentiate
it from others. This unique identification code, along with descriptions and
potential classifications, were useful for making richness estimates. Species
richness is a measure of diversity, and richness of morphotypes in this study is
presumably a refl ection of species richness (Kikkawa 1986).
Using a Sony DV Recorder, each transect was viewed three times on a
14-inch fl at-screen color monitor. A count was made of large sponge and
coral morphotypes. Each transect was again reviewed and assigned a reef
morphology category based on size and shape of rocks, amount of bioerosion,
presence of sand or crevices, and occurrence of holes in the reef. When
transects crossed multiple morphology categories, each category type was
noted; however, these transects were not used for statistical analyses. Eleven
reef morphology categories describe transect morphology and habitat in this
study (Table 1).
Transects were also grouped into the following relief and sediment
categories: high relief (>2 m) = “block-shaped boulders” and “high-relief
bioerosion;” moderate relief (1–2 m) = “buried block-shaped boulders,”
“bioeroded rocky fl at,” “large boulders with sand,” and “moderate-relief
bioerosion;” low-relief (<1 m) = “slab pavement,” “buried rocky fl at,” “sand
with scattered rocks,” “low-relief bioerosion,” and “sand;” high sediment =
“sand,” “sand with scattered rocks,” “buried rocky fl at,” and “large boulders
with sand;” moderate sediment = “buried block-shaped boulders,” “bioeroded
rocky fl at,” “moderate relief bioerosion,” and “low-relief bioerosion;” and
low sediment = “block-shaped boulders,” “slab pavement,” and “high-relief
bioerosion.” Total numbers of organisms were compiled and graphed for
each relief and sediment grouping.
2008 S.B. Fraser and G.R. Sedberry 195
Table 1. Reef morphology categories describing transect habitat. Relief categories: High relief = >2 m, moderate relief = 1–2 m, and low relief = <1 m.
Reef morphology Dive location Description Relief Sediment
Block-shaped boulders (BSB) St. Augustine Scarp Large square boulders with fl at edges along a continuous reef ridge High Low
Buried-block shaped boulders (BBB) Jacksonville Scarp Continuous reef ridge, fl at on top with partially buried square shaped Moderate Moderate
Slab pavement (SP) Jacksonville Scarp Large slabs of fl at rock with narrow fractures filled with sand Low Low
Large boulders with sand (LBS) Razorback Large boulders mixed with patches of sand, some boulders partially Moderate High
Bioeroded rocky fl at (BiRF) Savannah Scarp Hard bottom, low relief rubble, and rocky pavement. Rocks appear Moderate Moderate
pitted and eroded
Buried rocky fl at (BuRF) Savannah Scarp Similar to Bioeroded rocky fl at, but with sand partially covering the Low High
High relief bioeroded (HRB) Julians Ridge and Complex reef heavily pitted with crevices, holes, overhangs, and High Low
Scamp Ridge rock ledges (>2m)
Moderate relief bioeroded (MRB) Julians Ridge and Rough texture with depressions and small crevices (1–2m) Moderate Moderate
Low relief bioeroded (LRB) Georgetown Hole Low mounds of exposed bioeroded hard bottom with moderate sand Low Moderate
and Julians Ridge and sediment (<1m)
Sand with scattered rocks (SSR) Tattler Town Flat expanses of coarse sand with widely scattered rocks Low High
Sand (S) Tattler Town, Savannah Coarse unconsolidated sediment with no visible rock Low High
196 Southeastern Naturalist Vol.7, No. 2
Submersible tracks and fixes were plotted in ArcView v 3.2 (ESRI, Redlands,
CA), and transect lengths were estimated by measuring the distance
between each transect start and end point. Width of the visible transect field
was estimated at a random point within each transect using the 25-cm distance
between lasers for measurement. Temperature, depth, and salinity were
recorded from the on-screen data logger display, and mean values for these
variables were calculated for each transect.
Ircinia campana Nardo (stinking vase sponge), Stichopathes sp. (whip
coral), and Muricea pendula Verrill (pinnate spiny sea fan) were chosen to
determine whether significant differences in species densities existed among
temperatures, depths, or reef morphology categories. These organisms were
selected because they were identified with confidence, were found throughout
many of the dive locations, and typically had high abundances within most transects
at each dive location. Density data were tested for normality in JMP v 5.0
(SAS Institute Inc., Cary, NC) by applying a Shapiro-Wilks goodness-of-fit test
(p ≥ 0.05). This test revealed that the data were not normally distributed. Both
log and square root transformations were attempted, but failed to normalize the
data, so a non-parametric multiple response permutation procedure (MRPP)
was used to evaluate significance. The MRPP reports an overall p value to
evaluate significant difference, and an agreement statistic, A, that describes
within-group homogeneity compared to random expectation. In addition, the
test provides pairwise comparisons that allow assessments between groups. All
MRPP significance tests were conducted at the 0.05 significance level using
PC-Ord v. 5.0 (MJM Software, Glen Eden Beach, OR). Sorensen (Bray-Curtis)
distances were employed as the distance metric.
Water temperatures ranged from 13.7–22.6 °C and were divided into
four classes: 12.1–15.0 °C, 15.1–18.0 °C, 18.1–21.0 °C, and 21.1–24.0 °C.
Similarly, depth ranged from 44.5–78.3 m and was divided into four classes:
41.0–50.9 m, 51.0–60.9 m, 61.0–70.9 m, and 71.0–80.9 m. Median densities
for I. campana, Stichopathes sp., and. M. pendula were calculated for each
temperature class, depth class, and reef morphology category. The median
was selected over the mean as the best measure of the center of the data
because of the large number of zero values in the data set. Significant differences
in median densities for I. campana, Stichopathes sp., and M. pendula
were tested using a MRPP.
Richness of sponge and coral morphotypes and mean numbers of morphotypes
were evaluated within each transect by each temperature class,
depth class, and reef morphology category in terms of number of morphotypes
observed. Significant variation among morphotypes was tested in
Minitab v. 13.0 (Minitab Inc., State College, PA) using a non-parametric
Kruskal-Wallis analysis of variance on ranks.
A total of 143 transects from 14 submersible dives at eight different
locations were analyzed. Approximately 29,500 m2 were surveyed with a
2008 S.B. Fraser and G.R. Sedberry 197
mean transect area of 225 m2. Transects conducted at Scamp Ridge covered
the largest area, 6790 m2, while only 1308 m2 were surveyed at Georgetown
Hole (Fig 1.). A total of 8241 organisms from 32 different morphotypes
were counted in this study. The largest numbers of organisms (4590) were
recorded at St. Augustine Scarp, the most southerly site, while the lowest
numbers of organisms (17) were observed at Tattler Town. While only 10
of the morphotypes represented were corals, they accounted for 6224 individuals,
while the remaining 22 morphotypes were identified as sponges
and totaled 1989 individuals. I. campana, Stichopathes sp., M. pendula,
and “sponge morph 4” accounted for 91.2% of all individuals counted.
Most transects (104) covered only one of the 11 reef morphology categories.
Video clips of examples of each reef morphology can be viewed
at http://www.csc.noaa.gov/seageofish/. Total area observed on those transects
was approximately 21,000 m2. The remaining 39 transects crossed
multiple habitat types and were therefore eliminated from statistical
analysis comparing biota among reef morphology categories. The largest
number of transects were categorized as “low-relief bioeroded” habitat,
and the fewest were classified as “large boulders with sand.” St. Augustine
Scarp and Jacksonville Scarp had a well-defined ridge with a noticeable
drop-off, while the reefs at the more northern dive sites appeared less distinct.
When reef morphology categories were grouped by amount of relief
and sediment, those transects characterized by large vertical relief (>2 m)
and low amounts of sediment had the greatest numbers of sessile invertebrates
Ircinia campana and Stichopathes sp. were not found at temperatures
colder than 15.0 °C or at depths greater than 70.0 m, while M. pendula was
found in all temperature and depth classes. The greatest densities of I. campana
and Stichopathes sp. were found at temperatures within the 18.1–21.0
°C class, while M. pendula densities were greatest within 21.1–24.0 °C
(Figs. 3, 4, 5). All three organisms had the greatest densities within the
51.0–60.9 m depth class (Figs. 3, 4, 5). Significant differences in densities
of I. campana, M. pendula, and Stichopathes sp. were found among all
temperature classes (A = 0.246, p = 0 .000), and among all depth classes (A
= 0.449, p = 0.000).
The greatest densities of I. campana and Stichopathes sp. were found
within reef morphology categories with moderate to high relief, and moderate
to low sediment (“buried block-shaped boulders” and “block-shaped
boulders”) at St. Augustine Scarp (Fig. 6). The largest densities of M.
pendula were found on transects that crossed reef morphologies of low
relief and moderate sediment (“low-relief bioerosion”) at Julians Ridge
(Fig. 6). All three organisms were found in low densities or were absent
from transects that crossed low-relief and high-sediment reef morphology
categories such as “sand,” “sand with scattered rocks,” and “buried
198 Southeastern Naturalist Vol.7, No. 2
Figure 3. Mean density of I. campana, by temperature and depth. The four bar shades
represent the four temperature categories.
Figure 2. Number of organisms by relief and sediment classes. High relief (>2 m):
block-shaped boulders and high-relief bioerosion; moderate relief (1–2 m): buried
block-shaped boulders, bioeroded rocky fl at, large boulders with sand, and moderate-
relief bioerosion; and low-relief (<1 m): slab pavement, buried rocky fl at,
sand with scattered rocks, low relief bioerosion, and sand. High Sediment: sand,
sand with scattered rocks, buried rocky fl at, and large boulders with sand; moderate
sediment: buried block-shaped boulders, bioeroded rocky fl at, moderate relief bioerosion,
and low relief bioerosion; and low sediment: block-shaped boulders, slab
pavement, and high relief bioerosion. See Table 1 for summary of reef morphology
2008 S.B. Fraser and G.R. Sedberry 199
rocky flat” (Fig. 6). No I. campana or Stichopathes sp. and few M. pendula
were found at the Tattler Town dive site. MRPP tests indicated significant
differences in species densities among reef morphology categories (A =
0.999, p = 0.000). However, pairwise comparisons revealed that densities
between the “sand with scattered rocks” and “large boulders with sand”
and between the “moderate” and “high-relief bioeroded” reef morphology
categories were not significantly different.
Figure 4. Mean density of Stichopathes sp. by temperature and depth. The four bar
shades represent the four temperature categories.
Figure 5. Mean density of M. pendula by temperature and depth. The four bar shades
represent the four temperature categories.
200 Southeastern Naturalist Vol.7, No. 2
Among temperature classes, the number of morphotypes was lowest
within 15.1–18.0 °C and highest within 18.1–21.0 °C (Fig. 7). The number of
morphotypes was also highest within the 51.0–60.9 m depth classes and lowest
within the 71.0–80.9 m depth class (Fig. 7). The greatest number of morphotypes
was found on “low-relief bioeroded” reef morphology and the fewest on
“sand” (Fig. 8). Significant differences in mean numbers of morphotypes were
Figure 6. Mean density of I. campana, Stichopathes sp., and M. pendula by reef
morphology category. See Table 1 for summary of reef morphology categories and
Figure 7. Number of morphotypes by temperature and depth.
2008 S.B. Fraser and G.R. Sedberry 201
found among temperature classes (σ = 1.794, p = 0.000), depth classes (σ =
1.756, p = 0.001), and reef morphology categories (σ = 1.925, p = 0.000).
Each dive location site was nearly distinct from all others in its reef morphology.
“Block-shaped boulders” were found only at St. Augustine Scarp,
“buried block-shaped boulders” and “slab pavement” only at Jacksonville
Scarp, and “bioeroded rocky fl at” and “buried rocky fl at” only at Savannah
Scarp. “Large boulders with sand” were found only at Razorback, and “sand
with scattered rocks” occurred only at Tattler Town. “High-relief” and “moderate-
relief bioerosion” were both found at Julians Ridge and Scamp Ridge,
and “low-relief bioerosion” was found at Julians Ridge and Georgetown Hole.
Generally, the southern dive sites had the most well-defined ledge or ridge features,
while reefs north of Florida exhibited less distinct forms.
Bioerosion refers to the mechanical and chemical activity of algae, sponges,
polychaetes, mollusks, barnacles, fish, and other marine organisms that
penetrate and break down the substrate (Wilkinson 1983). This process appears
to be a significant force controlling reef morphology among continental
shelf edge reefs within the SAB, with the more northern reefs off the coast of
South Carolina exhibiting greater amounts of bioerosion than reefs off Florida
and Georgia. The organisms responsible for the erosion could not be identified
or quantified among sites, and the observed erosion may have been caused by
communities that are no longer present on the reefs.
High A values reported by the MRPP test refl ect strong within-group
homgeneity and suggest that reef morphology categories are distinct from
Figure 8. Number of morphotypes observed within each reef morphology category.
See Table 1 for summary of reef morphology categories and abbreviations.
202 Southeastern Naturalist Vol.7, No. 2
one another. The p values for the pairwise comparisons clarify the exceptions.
“Large boulders with sand” and “sand with scattered rock” could be lumped
together, as could “moderate” and “high-relief bioeroded” reef morphology
categories. “Low,” “moderate,” and “high-relief bioeroded” sites as well as
“bioeroded rocky fl ats” and “block-shaped boulders” generally exhibited the
greatest densities and diversity of large sessile invertebrates in this study, and
had far greater densities than sandy habitats. Hard substrates such as these
provide an environment more suitable for attachment than sandy sites (Barans
and Henry 1984, Miller and Richards 1980, Powles and Barans 1980, Wenner
et al. 1983). Near-bottom currents create migrating sand waves, scouring, and
re-suspension of sediments that can bury fl at hard bottom or the organisms
associated with that habitat. Relief of 2–3 m (such as found at some of these
sites) offers protection from these types of disturbances.
Other research has shown greatest densities and diversities of sessile
invertebrates to be associated with hard surfaces, rough texture, and relief.
Wenner et al. (1983) and Bright et al. (1984) noted greater diversity of epifaunal
macro-invertebrate communities on rocky bottom than on soft, sandy
seafl oors. Bell and Smith (2004) observed that diversity of sponges on an
Indonesian reef was higher on vertical and inclined surfaces than horizontal
planes. Goldberg (1973) described three terrace-like reefs off the coast
of southeast Florida and noted that the terrace with the most vertical relief
and rugged texture exhibited the greatest diversity and density of gorgonian
and scleractinian corals. Erwin (1976) discovered at Strangford Lough in
Northern Ireland that distribution of epifaunal and infaunal organisms was
closely related to bottom type and water movement.
Densities of I. campana, Stichopathes sp., and M. pendula were greatest
within the 51.0–60.9 m depth class and at temperatures between 18.1 and
21.0 °C. The deepest (71.0–80.9 m) and coldest (12.1–15.0 °C) transects had
relatively low abundances and diversities of sessile invertebrates. This finding
may be explained by the fact that these transects were conducted over fl at
sand and sand with scattered low-relief rock morphologies, providing relatively
little suitable habitat for attachment and growth of sessile organisms.
Previous sampling in the SAB (Cerame-Vivas and Gray 1966, Pearse and
Williams 1951, Wenner et al. 1983) indicates I. campana and Stichopathes
sp. are typically found at shallower and more southern sites. It is likely that
the warming and stabilizing infl uence of the Gulf Stream along the continental
shelf edge allows these organisms to exist in deeper waters and at more
Studies conducted at depths of 0–30 m have shown that richness, evenness,
and density of sponges and coral in reef habitats often increase with
depth up to 30 m as disturbance and temperature fl uctuations decrease and
available space increases with light attenuation (Andres and Witman 1995,
Bell and Barnes 2000, Dinesen 1983, Huston 1985, Liddell and Ohlhorst
1987, Schmal 1990). In the present study, the number of sponge and coral
morphotypes increased with depth until 60.9 m. At greater depths, transects
sampled mostly “sand” and “large boulders with sand,” habitats not suitable
2008 S.B. Fraser and G.R. Sedberry 203
In recent years, recorded video footage has been used to investigate marine
fauna and habitat. Observations made from video recordings provide the
unique and complementary opportunity to note variation in bottom topography,
texture, and behavior of fish and invertebrates in relation to bottom
features. Video recordings made from submersibles offer several advantages
over remotely operated underwater televisions and diver-held cameras.
Submersibles can operate in deeper waters and for longer periods of time
than divers, and are often equipped to accommodate sample collection. Finally,
video footage recorded on these dives provides permanent documentation
without destroying fauna and habitat (Parker 1986).
One of the primary goals of recent submersible operations off the southeastern
US coast has been to explore spawning locations of commercially or
recreationally important reef fishes, such as our sampling sites (Sedberry et
al. 2006). Declining catch per unit effort (CPUE) of several of these species
indicates overfishing and collapsing populations, despite efforts to manage
these fisheries (McGovern et al. 1998, Miller and Richards 1980, Sedberry
1988, Sedberry and Van Dolah 1984). Spawning locations on shelf edge reefs
represent important habitat essential for fish life cycles. Large sponges and
corals may be important components of these habitats because they enhance
structural complexity of the environment and therefore contribute shelter and
hiding places attractive to fishes (Van Dolah et al. 1987). Additionally, these
large sessile organisms provide microhabitats for various smaller invertebrate
species that may provide food for a variety of reef and pelagic fishes (McGovern
et al. 1998, Miller and Richards 1980, Sedberry 1988, Sedberry and Van
Dolah 1984, Weaver and Sedberry 2001). Results of this study suggest differences
in relief, sediment, and density and diversity of invertebrate macrofauna
on shelf edge reefs in the SAB. These factors may be important considerations
in natural resource management
Fishery management has recently shifted from single species management
to an ecosystem approach. In response to this change, studies utilizing
submersible video have included analyses of fish composition, density, and
behavior (Schobernd 2005). The present study and others focused on lower
food-web components and abiotic elements help provide a more ecosystemwide
understanding. Implementation of ecosystem-based fishery management
relies on evaluation of biodiversity, genetic variability, and ecological functionality
(Lubchenco et al. 2003, Roberts et al. 2003). Management actions
include development of marine protected areas (MPAs) to protect bottom
habitat. Reefs along the continental shelf edge off the southeastern US are potential
candidates for MPA designation (SAFMC 2004, Sedberry et al. 2006).
Effective positioning of reserves will maximize protection of important species
and habitat and, hopefully, serve as biotic sources for neighboring waters.
Fish biomass and production are thought to be related to the presence of
sessile invertebrates and reef morphology, but that remains to be demonstrated.
Quantitative measurements of reef morphology, angle of the reef slope,
position of the organisms on the reef, and fl ow regime could help explain
distribution of sessile megafauna and fish because these variables infl uence
current velocity, sedimentation rates, extent of benthic structure, and available
204 Southeastern Naturalist Vol.7, No. 2
light for benthic photosynthesis. For example, bioerosion rates may affect
habitat relief and structure available for fish and benthic invertebrates. Verification of substratum age could explain differences in reef morphology and
shed light on why the northern dive sites exhibited greater amounts of bioerosion
and less relief.
This research was funded by NOAA Ocean Exploration grants NA16RP2697,
NA03OAR4600097, and NA0ROAR4600055, G.R. Sedberry, Principal Investigator.
We thank Leslie Sautter, Rick DeVoe, Cass Runyon, Christina Schobernd, Matt Fraser,
and Dana Griffin for constructive criticism and advice; Jessica Stephens for database
assistance; Susan Thornton-DeVictor, David Knott, Stacie Crowe, and Cara Fiore for
invertebrate identification; Josh Loefer, Tim Snoots, and Karen Swanson for technical
help; and Martin Jones and Jodi Slapcinsky for statistical advice.
Andres, N.G., and J.D. Witman. 1995. Trends in community structure on a Jamaican
reef. Marine Ecology Progress Series 118:305–310.
Atkinson, L.P., and T.E. Targett. 1983. Upwelling along the 60-m isobath from Cape
Canaveral to Cape Hatteras and its relationship to fish distribution. Deep-Sea Research
Bane, J.M., and D.A. Brooks. 1979. Gulf Stream meanders along the continental
margin from the Florida Straits to Cape Hatteras. Geophysical Research Letters 6:
Barans, C.A., and V.J. Henry, Jr. 1984. A description of the shelf edge groundfish habitat
along the southeastern United States. Northeast Gulf Science 7:77–96.
Bell, J.J., and D.K.A. Barnes. 2000. A sponge diversity centre within a marine “island.”
Bell, J.J., and D. Smith. 2004. Ecology of sponge assemblages (Porifera) in the Wakatobi
region, southeast Sulawesi, Indonesia: Richness and abundance. Journal of
Marine Biology Association of the United Kingdom 84:581–591.
Bright, T.J., G.P. Kraemer, G.A. Minnery, and S.T. Viada. 1984. Hermatypes of the
Flower Garden Banks, northwestern Gulf of Mexico: A comparison to other western
Atlantic reefs. Bulletin of Marine Science 34:461–476.
Blanton, J.O., L.P. Atkinson, L.J. Pietrafesa, and T.N. Lee. 1981. The intrusion of Gulf
Stream water across the continental shelf due to topographically induced upwelling.
Deep Sea Research 28A:393–405.
Cain, T.D. 1972. Additional epifauna of a reef off North Carolina. Journal of the Elisha
Mitchell Scientific Society 88:79–82.
Cerame-Vivas, M.J., and I.E. Gray. 1966. The distributional pattern of benthic invertebrates
of the continental shelf off North Carolina. Ecology 47:260–270.
Dinesen, Z.D. 1983. Patterns in the distribution of soft corals across the central Great
Barrier Reef. Coral Reefs 1:229–236.
Erwin, D.G. 1976. A diving survey of Strangford Lough: The benthic communities and
their relation to substrate–a preliminary account. Pp. 215–224, In B.F. Keegan,
P.O. Ceidigh, and P.J. Boaden (Eds.). Biology of Benthic Organisms. Pergamon
Press, Oxford, UK. 630 pp.
Goldberg, W.M. 1973. The ecology of the coral-octocoral communities off the southeast
Florida coast: Geomorphology, species composition, and zonation. Bulletin of
Marine Science 23:465–488.
2008 S.B. Fraser and G.R. Sedberry 205
Hooper, J.N.A., and R.V.M. van Soest (Eds.). 2002. Systema Porifera: A Guide to the
Classification of Sponges. Kluwer Academic/Plenum Publishers, Amsterdam, The
Netherlands. 1810 pp.
Huston, M. 1985. Patterns of species diversity in relation to depth at Discovery Bay,
Jamaica. Bulletin of Marine Science 37:928–935.
Kikkawa, J. 1986. Complexity, diversity, and stability. Pp. 41–62, In J. Kikkawa and
D.J. Anderson (Eds.). Community Ecology: Pattern and Process. Blackwell Scientific Publications, Oxford, UK. 432 pp.
Liddell, W.D., and S.L. Ohlhorst. 1987. Patterns of reef community structure, north
Jamaica. Bulletin of Marine Science 40:311–329.
Lubchenco, J., S.R. Palumbi, S.D. Gaines, and S. Andelman. 2003. Plugging a hole in
the ocean: The emerging science of marine reserves. Ecological Applications 13(1)
MacIntyre, I.G., and J.D. Milliman. 1970. Physiographic features on the outer shelf
and upper slope Atlantic continental margin, southeastern United States. Geological
Society of America Bulletin 81:2577–2598.
MacIntyre, I.G., and O.H. Pilkey. 1969. Tropical reef corals: Tolerance of low temperatures
on the North Carolina continental shelf. Science 166:374–375.
Mathews, T.D., and O. Pashuk. 1986. Summer and winter hydrography of the US South
Atlantic Bight (1973–1979). Journal of Coastal Research 2:311–336.
McGovern, J.C., G.R. Sedberry, and P.J. Harris. 1998. The status of reef fish stocks off
the southeast United States, 1983–1996. Proceedings of the Gulf and Caribbean
Fisheries Institute 50:871–895.
Meister, H.S., D.M. Wyanski, J.K. Loefer, S.W. Ross, A.M. Quattrini, and K.J. Sulak.
2005. Further evidence for the invasion and establishment of Pterois volitans (Teleostei:
Scorpaenidae) along the Atlantic coast of the United States. Southeastern
Menzel, D.W., L.R. Pomeroy, T.N. Lee, J.O. Blanton, and C.R. Alexander. 1993. Introduction.
Pp. 1–8, In D.W. Menzel (Ed.). Ocean Processes: US Southeast Continental
Shelf; A Summary of Research Conducted in the South Atlantic Bight under the
Auspices of the US Department of Energy from 1977 to 1991. Skidaway Institute
of Oceanography, Savannah, GA. Prepared by the US Department of Energy Office
of Scientific and Technical Information.
Miller, G.C., and W.J. Richards. 1980. Reef fish habitat, faunal assemblages, and factors
determining distributions in the South Atlantic Bight. Proceedings of the Gulf
and Caribbean Fisheries Institute 32:114–130.
Parker, R.O. 1986. Observing reef fishes from submersible off North Carolina. Northeast
Gulf Science 8:31–49.
Pearse, A.S., and L.G. Williams. 1951. The biota of the reefs of the Carolinas. Journal
of the Elisha Mitchell Scientific Society 67:133–161.
Powles, H., and C.A. Barans. 1980. Groundfish monitoring in sponge coral areas off
the southeastern United States. Marine Fisheries Review 42(5):21–35.
Powles, H., and W. Stender. 1976. Observations on composition, seasonality and distribution
of ichthyoplankton from MARMAP cruises in the South Atlantic Bight in
1973. Technical Report 11. South Carolina Marine Resources Center, Charleston,
SC. 46 pp.
Roberts, C.M., S. Andelman, G. Branch, R.H. Bustamante, J.C. Castilla, J. Dugan, B.S.
Halpern, K.D. Lafferty, H. Leslie, J. Lubchenco, D. McArdle, H.P. Possingham, M.
Ruckelshaus, and R.R. Warner. 2003. Ecological criteria for evaluating candidate
sites for marine reserves. Ecological Applications 13(1) supplement:S199–S214.
Schmal, G.P. 1990. Community structure and ecology of sponges associated with four
southern Florida coral reefs. Pp. 376–387, In K. Rutzler (Ed.). New perspectives in
Sponge Biology. Smithsonian Institution Press, Washington, DC. 533 pp.
206 Southeastern Naturalist Vol.7, No. 2
Schobernd, C. 2005. Submersible observations of fish assemblages in deep reef
habitats off the southeastern United States: Implications for management. M.Sc.
Thesis. College of Charleston, Charleston, SC.
Sedberry, G.R. 1988. Food and feeding of Black Sea Bass, Centropristis striata, in
live bottom habitats in the South Atlantic Bight. Journal of the Elisha Mitchell
Scientific Society 104:35–50.
Sedberry, G.R., and R.F. Van Dolah. 1984. Demersal fish assemblages associated
with hard bottom habitat in the South Atlantic Bight of the USA. Environmental
Biology of Fishes 11:241–258.
Sedberry, G.R., O. Pashuk, D.M. Wyanski, J.A. Stephen, and P. Weinbach. 2006.
Spawning locations for Atlantic reef fishes off the southeastern US. Proceedings
of the Gulf and Caribbean Fisheries Institute 57:463–514.
Smith, F.G.W. 1971. Atlantic Reef Corals; A Handbook of the Common Reef and
Shallow-Water Corals of Bermuda, the Bahamas, Florida, and the West Indies
and Brazil. University of Miami Press, Coral Gables, FL. 164 pp.
South Atlantic Fishery Management Council (SAFMC). 2004. Informational public
hearing document on marine protected areas to be included in Amendment 14
to the fishery management plan for the snapper grouper fishery of the South Atlantic
region. Unpublished report. South Atlantic Fishery Management Council,
Charleston, SC. 46 pp.
Struhsaker, P. 1969. Demersal fish resources: Composition, distribution, and commercial
potential of the continental shelf stocks off southeastern United States.
Fishery Industrial Research 4:261–300.
Van Dolah, R.F., P.H. Wendt, and N. Nicholson. 1987. Effects of a research trawl on
a hard-bottom assemblage of sponges and corals. Fisheries Research 5:39–54.
Van Dolah, R.F., P.P. Maier, G.R. Sedberry, C.A. Barans, F.M. Idris, and J.J. Henry.
1994. Distribution of bottom habitats on the continental shelf off South Carolina
and Georgia. Final Report to Southeast Area Monitoring and Assessment
Program (SEAMAP) South Atlantic Committee Charleston, SC, NOAA Award
Veron, J.E.N. 2000. Corals of the World. Australian Institute of Marine Science,
Townsville, Australia. 1382 pp.
Weaver, D.C., and G.R. Sedberry. 2001. Trophic subsidies at the Charleston Bump:
Food-web structure of reef fishes on the continental slope of the southeastern
United States. Pp. 137–152, In G.R. Sedberry (Ed.). Island in the Stream: Oceanography
and Fisheries of the Charleston Bump. American Fisheries Society,
Symposium 25, Bethesda, MD. 240 pp.
Wendt, P.H., R.F. Van Dolah, and C.B. O’Rourke. 1985. A comparative study of the
invertebrate macrofauna associated with seven sponge and coral species collected
from the South Atlantic Bight. 1985. Journal of the Elisha Mitchell Scientific
Wendt, P.H., D.M. Knott, and R.F. Van Dolah. 1989. Community structure of the
sessile biota on five artificial reefs of different ages. Bulletin of Marine Science
Wenner, E.L., D.M. Knott, R.F. Van Dolah, and V.G. Burrell, Jr., 1983. Invertebrate
communities associated with hard bottom habitats in the South Atlantic Bight.
Estuarine Coastal Shelf Science 17:143–158.
Wenner, E.L., P. Hinde, D.M. Knott, and R.F. Van Dolah. 1984. A temporal and spatial
study of invertebrate communities associated with hard bottom habitats in the
South Atlantic Bight. NOAA Technical Report NMFS 18. 104 pp.
Wilkinson, C.R. 1983. Role of sponges in coral reef structural processes. Pp. 263–
274, In D.J. Barnes (Ed.). Perspectives on Coral Reefs. B. Clouston Publishers,
Manuka, ACT, Australia. 288 pp.