Mollusks taken by Beam Trawl in the vicinity 

of Gray’s Reef National Marine Sanctuary on the

Continental Shelf off Georgia, Southeastern U.S.

NOAA Technical Memorandum NOS NCCOS 88
Mention of trade names or commercial products does not constitute 

endorsement or recommendation for their use by the United States 

government.

Citation for this Report

Wolfe, Douglas A. 2008. Mollusks taken by Beam Trawl in the vicinity of Gray’s Reef 

National Marine Sanctuary on the Continental Shelf off Georgia, Southeastern U.S.

NOAA Technical Memorandum NOS NCCOS 88. 40 pp.
Mollusks taken by Beam Trawl in the vicinity of 

Gray’s Reef National Marine Sanctuary on the 

Continental Shelf off Georgia, Southeastern U.S. 

Douglas A. Wolfe* 

NOAA, National Ocean Service

National Centers for Coastal Ocean Science

Center for Coastal Fisheries and Habitat Research

NOAA Beaufort Laboratory

101 Pivers Island Road

Beaufort, North Carolina 28516

*(retired) Present address: 109 Shore Drive, Beaufort, NC 28516

NOAA Technical Memorandum NOS NCCOS 88

December, 2008

Table of Contents

Abstract................................................................................ iii

Introduction............................................................................ 1

Methods................................................................................. 2

Results................................................................................... 4

Sample Characteristics......................................................... 4

Total Numbers of Taxa........................................................ 6

Species Associations and Community Composition....................... 7

Effect of Depth........................................................... 7

The “Atrina-Cymatium” Community................................. 10

Thee Nassarius acutus-Phascolion-Pythinella Connection...... 11

Rock-Dwelling Species................................................ 13

Range Extensions.............................................................. 14

Notes on Other Taxa.......................................................... 18

Gray’s Reef National Marine Sanctuary [Stations 2.1-2.4] Taxa....... 19

Potential Species Richness................................................... 21

Discussion............................................................................. 23

Acknowledgments.................................................................... 27

References............................................................................. 28

Appendix I. Station Locations and Depths for Beam Trawls.................. 34

Appendix II. Molluscan Taxa Recovered......................................... 35

i
ii
 

Abstract 

Mollusks were sorted from samples of shell hash (obtained as bycatch during 

NOAA-sponsored studies of larval and juvenile fish distribution), and analyzed to gain 

qualitative insights on species composition, distribution and habitat affinities of the 

molluscan fauna on the continental shelf off Georgia. Samples came from beam trawls at 

37 stations located in the immediate vicinity and offshore of the Gray’s Reef National 

Marine Sanctuary (GRNMS) at depths of 4.9 to 103 m. Two hundred sixty-three (263) 

taxa of mollusks (~58% as dead shells only) were collected, and nearly all (~99%) were 

identified to the species level. Ninety-seven of these taxa appeared in samples from one 

or more of the four stations established near the corners of the GRNMS. Samples were 

highly variable in terms of appearance, volume and species composition of mollusks, 

reflecting the extreme patchiness of benthic habitats within this region of the continental 

shelf. With very few exceptions, the mollusks were generally characteristic of either the 

Carolinian or Caribbean faunal provinces. The Georgia continental shelf, however, was 

outside the previously reported ranges for at least 16 of the species reported here. Most 

of these extralimital species were known previously from the East Coast of Florida, and 

represented northerly range extensions of 1-5° Latitude (110-560 km). One species 

represented a more significant range extension from the Bahamas and the southern 

Caribbean, and two represented southerly range extensions, known previously from only 

as close as off North Carolina. The high incidence of range extensions found in this 

study and the potential for discovery of additional species are discussed in the context of 

the diversity and patchiness of benthic habitats on the continental shelf of the region, and 

the sensitivity of species recruitment to variability in Gulf Stream patterns and global 

climate change. 

iii
iv
1

Introduction 

 

The Continental Shelf off the coast of Georgia and South Carolina supports a diverse 

assortment of demersal and benthic habitats. These range from flat bottoms of quartz and 

carbonate sands to high-relief, live-bottom rocky ledges that are densely colonized by 

invertebrates and fish communities (Parker et al., 1983; Nelson et al., 1999; Kendall et al., 

2006). Similar “patch reefs” occur along the continental shelf at least as far north as Cape 

Lookout, North Carolina (Menzies et al., 1966; Shoemaker, 1972; Van Dolah et al., 1994). 

With the establishment of the Gray’s Reef National Marine Sanctuary (GRNMS) in 1981, a 

17.5 square nautical mile (61 km2) portion of live-bottom patch reef habitat was set aside for 

research, protection, and management. Gray’s Reef contains extensive but discontinuous 

outcrops of limestone up to ten feet in height separated by flat-bottomed troughs with 

unconsolidated sediments, mainly sand. Parker et al. (1983, 1994) estimated that rocky reefs 

cover up to 30% of the Georgia shelf area in the vicinity of the GRNMS.

While it is known that live-bottom reef habitats support rich assemblages of 

invertebrates, the molluscan fauna of the continental shelf off Georgia has received little 

attention compared to similar areas both to the north and the south. The molluscan 

assemblages associated with patch reef habitats off North and South Carolina have been 

described in some detail (eg. Pearse and Williams, 1951, Wells et al., 1961; Menzies et al., 

1966; Shoemaker, 1972; Wenner et al., 1983), as have the mollusks associated with the 

calico scallop fishery off North Carolina (Wells et al., 1964; Porter and Wolfe, 1972). Ward 

and Blackwelder (1987) have suggested that these communities may have persisted relatively 

unchanged since the late Pliocene period (2.4 MYA). Wenner et al., (1983) examined 

diversity and biomass relationships among the epifauna (including mollusks) in the 

immediate region of the present study, but did not report details of species composition or 

occurrence. Lee (2006, pers. comm.) has compiled an extensive list of mollusks found off 

Northeast Florida. While the coastal and inshore molluscan fauna of Georgia has recently 

been described (Prezant et al., 2002), and the GRNMS has recently sponsored studies of the 

soft-bottom benthos (Vittor and Associates, Inc., 2004; Hyland et al., 2006) and demersal 

fish (Cooksey et al., 2004; Kendall et al., 2006; Walsh et al., 2006), the occurrence and 

distribution of mollusks on the continental shelf of Georgia remain largely undescribed.

The samples analyzed in this study were collected during February and April 2002 as 

bycatch on two NOAA cruises conducted in support of studies of the distribution and 

movements of larval and juvenile fish on the continental shelf off the coast of Georgia in the 

immediate vicinity of the GRNMS (Marancik et al., 2005; Walsh et al., 2006). These 

samples provided an opportunity for qualitative examination of the molluscan assemblages 

that occur on the patch reefs and associated habitats in this transitional subtropical shelf 

region.
2

Methods 

February samples were obtained on the NOAA Ship Oregon II; April samples on the 

NOAA Ship Ferrel. Sampling stations are mapped in Fig. 1; station coordinates and depths 

are shown in Appendix I.

Targeted juvenile fish were collected at each station using a standard 2-m beam trawl 

(Kuipers, 1975) with a 6-mm mesh body and a 3-mm mesh tail bag. The beam trawl had a 

tickler chain ahead of the trawl and a chain on the foot rope at the front of the trawl. Twokilogram weights were added to each skid of the trawl to ensure that it stayed on the bottom. 

Three 5-min bottom tows were made at each station. The catch was sorted on deck to 

remove larger fish and extraneous material, and the remaining fish, shell hash, and associated 

invertebrates were preserved in 95% ethanol for later separation of all fish. After removal of 

the fish, the residual materials (consisting mainly of shell hash) were further examined for 

mollusks. Procedures for sorting the samples were not rigorously consistent throughout the 

study, and varied with the volume and character of the sample (Table 1). No record was 

maintained of the details of sorting throughout the study. Initially (i.e., for most of the 

Oregon II samples), mollusks were sorted by visual examination– frequently with the naked 

eye and sometimes (if the proportion of fine-grained materials seemed to warrant it) at low 

magnification under a dissecting microscope. Sorting of large-volume samples of very 

coarse shell hash was facilitated by washing the material first through a 12.5-mm mesh 

hardware cloth to separate large shells and fragments, prior to examination of the finer hash 

retained on 3-mm mesh hardware cloth or (again, if the proportion of fine-grained materials 

seemed to warrant it) on fiberglass screen with a mesh size of approximately 1.1 mm. 

Toward the end of the study, use of the 1.1-mm mesh became more standard. Consequently, 

small taxa were more likely to have been lost or overlooked in earlier samples. 

Materials retained on the screens were sorted under a dissecting microscope. All livecollected mollusks (gastropods with opercula or visible animal material and paired bivalves) 

along with freshly dead specimens were sorted from the samples. Unusual or rare species 

were retained regardless of condition, including fresh fragments and representative single 

valves for bivalves not found paired. For the samples recovered from the Oregon II cruise 8-

13 February 2002, species occurrence (but not number of specimens) by station and total 

number of specimens for all stations were recorded. Except for the species recovered from 

only one or two stations, the number of specimens per station is not available from that 

cruise. By contrast, specimens were sorted and enumerated for the individual replicate tow 

samples at each station from the Ferrel cruise 13-16 April 2002. 

Mollusk specimens were identified to the lowest practical taxonomic level (LPTL) 

using published (e.g. Abbott, 1974; Abbott and Morris, 1995; Redfern, 2001; Williams, 

2006; Mikkelson and Bieler, 2008) and on-line sources for reference, and comparison with 

previously identified material in the author’s personal collection (e.g. Wolfe, 1968; Porter 

and Wolfe, 1972). Selected specimens were also sent to Harry G. Lee (Jacksonville, 

Florida), William G. Lyons (St. Petersburg, Florida), or Donn Tippett (U.S. National 

Museum, Washington, D.C.) for examination and further identification. 
3

Nomenclature generally follows Turgeon et al. (1998), with some exceptions based on 

Rosenberg (2005), or (for bivalves) Mikkelson and Bieler (2008). All specimens are 

archived in the author’s collection.

Figure 1. Station locations sampled during February 2002 from the Oregon II and 

during April 2002 from the Ferrel. Detailed station data are given in Appendix I.
4

Results 

Sample Characteristics 

Table 1 describes general characteristics of the samples obtained from the April cruise 

of the Ferrel (analogous records were not kept for the Oregon II samples). Volume of 

individual replicate samples ranged from only 7 ml to about 8 liters. Although samples 

consisted primarily of shell hash, they varied considerably both among stations and among 

replicates within station, in terms of their overall volumes, and their composition– especially 

with regard to the mix of large bivalve shells (most commonly including Argopecten gibbus, 

Anadara lienosa floridana, Glycymeris americana, Laevicardium serratum, Arcinella 

cornuta, and others) and the proportion of smaller-grained material. For example, the two 

smaller replicates from station 22 consisted largely of fragments of sanddollars (Mellita), 

seastars (Astropecten) and cordgrass (Spartina), while the third replicate was typical shell 

hash. Replicate #1 from station 25 consisted almost exclusively of shrimp and squid (Illex 

illecebrosus, five individuals); replicate #2 contained several shrimp along with seastar parts; 

and replicate #3 (1000 ml) was shell-hash; but all three replicates had a paucity of mollusk 

species. Samples from stations 38, 42 and 43 (and SC02 from the Oregon II cruise) 

contained high proportions of broken pen shells (mostly Atrina rigida). Replicates from 

stations 27 and 28 consistently had relatively small volumes and contained a glutinous mass 

of filamentous mucoid material of unknown origin (possibly bryozoans, coelenterate 

tentacles, or egg mass). 

Samples from three replicate tows of the beam trawl were sorted and tallied separately 

for stations from the April cruise of the Ferrel. Initial sample volumes varied widely among 

replicates at any given station (Table 1); leading to highly variable species numbers among 

replicates within a station, with each replicate exhibiting a highly skewed distribution of 

species with many “rare” taxa unique to that replicate. Coefficients of variation (standard 

deviation as percent of mean) for the mean number of species per tow at any station ranged 

from 16.5% to 110%, with an average of 59%. The ratio of total number of species 

recovered in all tows at a station to the mean number recovered per tow at that station ranged 

from 1.9 to 3.0 (average: 2.53). The variability was so great among replicates within station 

that the data from individual replicates could not be used meaningfully, and the data were 

therefore lumped by station. This also facilitated comparison with the data from the Oregon 

II cruise in February, where the data from different replicates had been lumped during initial 

sorting. Similar action was taken with data from those ten stations that were sampled on both 

cruises: data were combined by station. 
5

Table 1. Characteristics of Beam Trawl Samples from Ferrel cruise April 2002. ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

SAMPLE SAMPLE VOLUME (ml) 

STA. DESCRIPTION TOW1 TOW2 TOW3 NOTES ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

02.1 CSH-vCSH 1500 500 1000 Argopecten/Cardiidae/Glycymeris

02.2 FSH-CSH 400 350 600 Argopecten/Cardiidae/Glycymeris

02.3 CSH 250 500 800 Argopecten/Arcinella/Veneridae

02.4 CSH 500 1000 500 Argopecten/Cardiidae/Glycymeris

022 BioD-CSH 350 30 30 Mellita/seastar parts/Spartina

023 FSH-CSH 1000 1500 200 Anadara/Atrina/Neverita eggcases

024 FSH-CSH 1700 100 200 Argopecten/Cardiidae/Glycymeris

025 BioD-FSH-CSH 200 80 1000 starfish parts/shrimp/squid

027 BioD 100 300 350 filamentous mucous/bryozoa/wormtubes/

Neverita eggcases

028 BioD 120 120 200 filamentous mucous, bryozoa/eggmass?

029 BioD-CSH 400 400 250 tunicata,echinoid,holothuroid. 

035 BioD-FSH 30 400 60 Neverita eggcase& urchin fragments

036 FSH-CSH 100 2000 1300 Argopecten/Cardiidae/coral

037 CSH 1000 1500 1200 Argopecten/Glycymeris/Arcinella/ Chione

038 CSH 400 600 1800 Atrina rigidahash/bryozoa debris

039 BioD-FSH 125 200 30 Spartina/plant detritus/urchin spines & tests, 

040 vCSH 4500 4500 8000 Argopecten/Euvola/Anadara/Chione

041 CSH-vCSH 6000 1500 5000 Argopecten /Anadara/Glycymeris

042 BioD-CSH 800 1500 100 Atrina, Argopecten, bryozoa 

043 CSH 1200 1000 1000 Atrina rigida hash/bryozoa/anemones; 

044 FSH-CSH 20 7 30 flat circular (1 cm) concretions of 1mm 

black rocks ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

BioD = biodebris: plant and invertebrate parts

FSH = fine broken shell hash: nearly all passes through 12.5mm screen

CSH = coarse shell hash: about half retained on 12.5 mm screen

vCSH = very coarse shell hash: nearly all retained on 12.5 mm screen

 
6

Total Numbers of Taxa 

Examples of two hundred sixty-three (263) molluscan taxa were recovered from the 

beam trawl samples from all 37 stations (Appendix II). Of these, 120 were noted to be in live 

or very freshly dead condition from at least one of the stations. As “freshly dead” is a 

judgment call, many specimens/species may have been overlooked in making this 

assessment. All but two (Melanella sp. and Olivella sp.) of the taxa have been identified to 

species level. 

Figure 2 illustrates the highly skewed distribution of the taxa within and among 

stations. One hundred five (105) of the taxa made “single station” appearances (each was 

recovered from only one of the 37 stations), and nearly all of these were represented by only 

one (89) or two (14) specimens. Seventy-seven (77) taxa were recovered from five or more 

stations, while only eleven (11) were found at fifteen or more; Olivella mutica was the most 

widespread- found at twenty-seven stations. 

0

20

40

60

80

100

120

1357

9

11

13151719

212325

27

Station Occurrence

 

Figure 2. Species Occurrence among Sampling Stations. 
7

Numbers of taxa per station ranged from six (stations 22 & 35) to seventy-wo (station 

38); the mean was 28.4 ± 17.7 (SD) with a mode of 25. For those ten stations that were 

sampled on both cruises (Appendix I), the number of taxa ranged from 23 to 56 per station 

(37.0 ± 10.7 with a mode of 35.5). Diversity of recovered taxa was greatest, however, at four

stations that were sampled only once: station 38 with 72 species, 42 (65), 37 (64), and 41 

(59), 

Species Associations and Community Composition 

The data set from this study is suitable only for qualitative examination of community 

composition due to: (a) the highly skewed distribution in the numbers (and composition) of 

taxa at all stations, (b) inconsistencies of sample treatment and data recording, and (c) the 

inconsistency of sampling frequency among stations (ten stations were sampled twice as 

intensively as the other twenty-seven). Nonetheless, several features stood out and these are 

discussed in the following sections. 

To determine whether depth was a factor affecting species occurrence among stations, 

species distributions were compared qualitatively between two station clusters stratified by 

depth range. The “shallow” stratum (depth range 8.5 to 22 m) included all ten of the stations 

sampled twice, along with eight of the remaining twenty-seven, while the remaining nineteen

stations were 30 to 103 m deep (Appendix I). Rare species (those found at only one or two 

stations) were excluded for this comparison. However, the rare species were fairly evenly 

distributed between the two station strata: 55/105 (52%) of the single-station species came 

from the deep stations; while the two-station species were divided: 18/39 (46%) from deep 

stations, 7/39 (18%) from shallow, with the balance split between the two station clusters. 

Of the 120 taxa that occurred at three or more stations, 26 species were recovered nearly 

exclusively (>80%) from the “shallow” stratum (Table 2a), while 23 species were recovered 

nearly exclusively (>80%) from “deep” stations (Table 2b). The remaining 71 species were 

distributed more evenly and broadly across the range of station depth. Because of small 

samples and high sample/station variability, the depth affinities indicated for the species in 

Tables 2a and 2b cannot be viewed as limiting. 

With very few exceptions, the “shallow” species listed in Table 2a are well known 

species components of nearshore (intertidal to 10 m) habitats of the southeastern U.S. 

Possible exceptions may include Strombus alatus, Cancellaria reticulata, Epitonium 

championi, and Spondylus americanus, which typically occur farther offshore; but most of 

the species in Table 3a (except Spondylus americanus, Pythinella cuneata, and 

Nemocardium peramabile) are found regularly on beaches south and west of Cape Lookout, 

North Carolina. 

Effect of Depth 
8

Table 2a. Species Found Mainly in “Shallow” (4.9-22m) Stations. ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

 OCCURRENCE

 TOTAL FRACTION #RETRIEVED

SPECIES FAMILY # OF STA'S @ 4.9-22m ALL STA'S ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

SPECIES RECOVERED NEARLY EXCLUSIVELY FROM SHALLOW (≤22m) STATIONS

Angulus sybariticus (Dall, 1881) Tellinidae 15 0.80 39

Terebra dislocata (Say, 1822) Terebridae 10 0.80 18

Neverita duplicata (Say, 1822) Naticidae 7 1.00 7

Cerithium atratum (Born, 1778) Cerithiidae 7 0.86 9

Nassarius acutus (Say, 1822) Nassariidae 6 0.83 125

Terebra protexta (Conrad, 1846) Terebridae 6 0.83 9

Strombus alatus Gmelin, 1791 Strombidae 5 1.00 10

Cancellaria reticulata (Linnaeus, 1767) Cancellariidae 5 1.00 8

Eupleura caudata (Say, 1822) Muricidae 5 1.00 7

Epitonium championi Clench & Turner, 1952 Epitoniidae 5 1.00 6

Oliva sayana Ravenel, 1834 Olividae 5 1.00 6

Kurtziella atrostyla (Tryon, 1884) Turridae 5 0.80 15

Pandora trilineata Say, 1822 Pandoridae 5 0.80 10.5

Spondylus americanus (Hermann, 1781) Spondylidae 5 0.80 3.5

Strigilla mirabilis (Philippi, 1841) Tellinidae 4 1.00 8

Pyramidella suturalis H.C. Lea, 1843 Pyramidellidae 4 1.00 6

Spisula raveneli (Conrad, 1831) Mactridae 4 1.00 3.5

Pythinella cuneata (Verrill & Bush, 1898) Leptonidae 3 1.00 106

Kurtziella limonitella (Dall, 1884) Turridae 3 1.00 19

Ilyanassa obsoleta (Say, 1822) Nassariidae 3 1.00 7

Turbo castanea Gmelin, 1791 Turbinidae 3 1.00 7

Graptacme eborea (Conrad, 1846) Dentaliidae 3 1.00 4

Epitonium rupicola (Kurtz, 1860) Epitoniidae 3 1.00 4

Nemocardium peramabile (Dall, 1881) Cardiidae 3 1.00 3

Dinocardium robustum (Lightfoot, 1786) Cardiidae 3 1.00 2

Trachycardium muricatum (Linnaeus, 1758) Cardiidae 3 1.00 1.5 ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––
9

Table 2b. Species Found Mainly in “Deep” ( ̆30-103m) Stations. ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

 OCCURRENCE

 TOTAL FRACTION #RETRIEVED

SPECIES FAMILY # OF STA'S @ 4.9-22m ALL STA'S ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

SPECIES RECOVERED NEARLY EXCLUSIVELY FROM DEEP (≥30m) STATIONS

Polinices porcellanus (d'Orbigny, 1839) Naticidae 14 0.14 59

Naticarius canrena (Linnaeus, 1758) Naticidae 12 0.08 23

Nassarius albus (Say, 1826) Nassariidae 10 0.00 21

Caryocorbula contracta (Say, 1822) Corbulidae 8 0.13 13

Atrina rigida (Lightfoot, 1786) Pinnidae 7 0.14 100

Prunum hartleyanum (Schwengel, 1941) Marginellidae 7 0.14 21

Cerodrillia bealiana Schwengel & McGinty, 1942

Turridae 7 0.14 8

Nassarius consensus (Ravenel, 1861) Nassariidae 7 0.00 21

Euvola raveneli (Dall, 1898) Pectinidae 6 0.17 8.5

Xenophora conchyliophora (Born, 1780) Xenophoridae 6 0.00 11

Semicassis granulata (Born, 1778) Cassidae 6 0.00 6

Merisca aequistriata (Say, 1824) Tellinidae 5 0.20 3.5

Cirsotrema dalli Rehder, 1945 Epitoniidae 5 0.00 6

Aequipecten muscosus (W. Wood, 1828) Pectinidae 5 0.00 4.5

Cymatium cingulatum (Lamarck, 1822) Cymatiidae 4 0.00 38

Epitonium angulatum (Say, 1831) Epitoniidae 4 0.00 5

Busycon sinistrum Hollister, 1954 Melongenidae 4 0.00 3

Similipecten nanus (Verrill & Bush, 1897) Propeamussidae 3 0.00 12.5

Transenella stimpsoni Dall, 1902 Veneridae 3 0.00 5.5

Graptacme calamus Dall, 1889 Dentaliidae 3 0.00 4

Dentimargo aureocinctus (Stearns, 1872) Marginellidae 3 0.00 4

Astarte smithi Dall, 1886 Astartidae 3 0.00 3.5

Varicorbula limatula (Conrad, 1846) Corbulidae 3 0.00 2

Many of the “deep” species (Table 2b) are components of the calico scallop community 

off North Carolina (Porter and Wolfe, 1972), which occur there in depths of 20-40 m. Three 

of the species in Table 2b (Atrina rigida, Epitonium angulatum, and Busycon sinistrum) are 

known to occur intertidally or in the shallow subtidal throughout the southeastern U.S., so 

their appearance in this table is clearly an aberrant result. 
10

Stations 44 and 39, were substantially deeper than any others (at 103 and 90 m, 

respectively), and might have been expected to yield species more characteristic of deep 

habitats. These stations, however, had only three species in common, and all three were 

found commonly in the shallow stations as well. Replicate tows from both of these deep 

stations yielded very small sample volumes (Table 1), and relatively few taxa per station (12 

and 16, respectively). While station 39 yielded an unusually high proportion of single-station 

(7/16) or two-station (2/16) species, these categories accounted for only one species each of 

the 12 recovered from station 44. Seven of the twelve species recovered from station 44 and 

six of the sixteen from station 39 occurred at six or more stations, encompassing a broad 

range of depths as shallow as 8.5 m. Only two species recovered from these two stations 

were clearly recognizable as “deep-water species”: Cochlespira radiata (a freshly dead 

specimen), and Thracia conradi (two single valves). Samples from station 39 contained 

very little shell hash and notable amounts of biodebris, including urchin parts and Spartina

and seagrass fragments, suggesting that the station was in a depositional area that received 

material transported across the continental shelf from the coast. 

Stations 38 and 43 from the Ferrel cruise were characterized by large sample volumes 

with an unusual preponderance of broken pen shell (Atrina rigida) hash– mostly from shells 

100-125 mm long (Table 1). To a lesser degree, two stations from the Oregon II cruise 

(SC01 and SC02) shared this same trait. 

Three species of Cymatium (see cover photograph) appeared in samples from one or 

more of these four stations and at no other stations. Cymatium cingulatum occurred in 

replicate tow #3 from station 38 (two specimens, recently dead); in all three replicate tows 

from station 43 (1, 28, and 5 specimens, respectively- all either live or recently dead); and 

recently dead specimens were also obtained from stations SC01 and SC02. Cymatium 

parthenopeum also occurred in all three replicate tows from station 43 (2, 8, and 6 

specimens, respectively- all either live or recently dead); and a live specimen was obtained 

from station SC01. One recently dead specimen of Cymatium krebsii also appeared in tow 

#2 from station 43. Associated with and attached to fragments of the shell hash of Atrina 

rigida was an unidentified light brown organism with branched filamentous strands up to ~2-

3 cm long (referred to as “Bryozoa” in Table 1) that bore a striking resemblance to the 

periostracum of the Cymatiidae, and especially to that of C. parthenopeum. 

While most of the pen shell hash consisted of broken dead shells, many of the 

fragments had intact hinge ligaments, delicate unbroken spines, and interior surfaces free of 

fouling organisms; suggesting that the species is living at or very near the locations where the 

material was found. One recent and intact specimen of Atrina seminuda was picked out from 

among the abundant A. rigida. Rosenberg (2005) cites records of live occurrences for these 

two species at very shallow depth ranges (0.3-2 m and 1.5-2.4 m, respectively), while 

Mikkelson and Bieler (2008) indicate that both species usually occur at depths less than 30 

m. Stations 38, 43, SC01, and SC02 are clustered fairly closely together at depths of 43-47 

m (Fig. 1, Appendix I), so these records may represent unusually deep occurrences for both 

species.

The “Atrina-Cymatium” Community 
11

Typically, each of these Atrina-Cymatium stations had its own complement of other 

species unique to that station. The calico scallop Argopecten gibbus was the only other 

species recorded at all four of these stations. Twenty-five other species occurred at either 

two or three of these stations, but only one of these (Eulithidium thalassicola ) was found at 

no other station. The remaining species, were all found in addition at several other stations. 

Based on their relative occurrence at other stations and at similar depths, the following 

species were judged to be most closely associated with the Atrina-Cymatium community: 

Similipecten nanus, Depressiscala nautlae, Busycon sinistrum, Cirsotrema dalli, Aequipecten 

muscosus, Xenophora conchyliophora, Semicassis granulata, Arcinella cornuta, Calliostoma 

yucatecanum, Arene tricarinata, Naticarius canrena, Conus anabathrum, Polinices 

porcellanus, and Chama macerophylla. It is no surprise that several of these associated 

species appeared also in the cluster of “deep” species (Table 2b).

The Nassarius acutus—Phascolion-–Pythinella Connection 

Samples from stations 27 and 28 afforded a unique opportunity to examine and analyze 

the commensal relationship between a bivalve and a sipunculid worm that occupies vacant 

shells of small gastropods (Hampson, 1964; Gage 1966). The three tows from station 27 

yielded 86 specimens (29, 17, and 40, respectively) of Nassarius acutus. Only nine other 

specimens were recovered from four tows at three other stations (#’s 22, 23, and 42) from the 

Ferrel cruise. A total of thirty specimens were retrieved from the Oregon II cruise–– at 

stations 28, 23 and 24; and “live?” specimens were noted at station 28. About 75% of these 

N. acuta shells were occupied by hermit crabs; a few appeared quite freshly dead. At least 

twenty-two of these shells were occupied, however, by a sipunculid worm ––presumed to be 

Phascolion strombus (Montagu, 1804)–– and it was one or more of these specimens that I 

had erroneously judged to be “live”, mistaking the anterior end of the sipunculid (preserved 

with tentacles retracted) for a gastropod with an operculum. 

Of those shells containing sipunculids, nineteen (86%) had commensal bivalves 

[Galeommatoidea: Leptonidae: Pythinella cuneata (Verrill and Bush, 1898)] attached––

usually clustered around the tentacular ring of the sipunculid and half-buried in the mucoid 

mass filling the aperture of the gastropod shell (Fig. 3a-c). These bivalves probably graze- at 

least in part- on materials caught in the mucous secreted by the sipunculid. Specimens ranged 

in size from juveniles < 1 mm to adults just over 3 mm. The fifteen adult specimens were on 

13 different shells (maximum of two). The 39 mid-size (1-2.5 mm) specimens were attached 

on 15 shells, usually (on all but four shells) in conjunction with a full-sized individual. 

There were 75 small (< 1 mm) specimens on eight different Nassarius shells, accompanied in 

seven cases by at least one full-size and in six cases also by at least one mid-size specimen. 

The mean number of bivalves per sipunculid (including the three with no clam attached) was 

5.5 ± 6.8 (sd); without those three zeros the mean rose to 6.6 ± 7.0. While Nassarius acutus

was found at four other stations, Pythinella cuneata made only one other appearance (at 

station 31) where it was associated with a sipunculid in the aperture of Epitonium championi 

(Fig. 3d).
12

a

c

b

d

Figure 3. The bivalve Pythinella cuneata (Verrill & Bush, 1898) has a commensal relationship 

with sipunculids that occupy gastropod shells: a. here in Nassarius acutus. b. large Pythinella ~ 

3mm; sipunculid retracted. c. five Pythinella; sipunculid oral ring in view. d. Pythinella & 

sipunculid on Epitonium championi. 
13

The samples containing Pythinella cuneata also exhibited other distinctive 

characteristics. As mentioned previously, the Ferrel samples from stations 27 and 28 

exhibited the very unique character of small sample volumes consisting largely of a 

sticky mass of mucoid material (Table 1). Unusually large numbers of Neverita eggcases 

(sand collars) also were noted in these samples. The three tows from station 27 contained 

12 ± 2.6 (sd) species each with a total of 23 species combined. The three tows from 

station 28 contained 1.3 ± 0.6 species each with a total of only four species. The species 

totals from the three replicate tows for these same two stations in the Oregon II samples 

were essentially reversed– with only two species from station 27 and 22 from station 28. 

So– while there is considerable evidence that the two stations are very similar, the data 

also demonstrate the extreme variability and patchiness of species distributions on the 

continental shelf. When the two cruises are taken together the cumulative species totals 

and composition show high concordance: 25 and 24 total species for stations 27 and 28, 

respectively, with 15 species in common. Most notable among these (in terms of their 

scarcity at other stations) were two turrid species (Kurtziella limonitella and K. atrostyla), 

Pyramidella suturalis, Neverita duplicata, and Mitrella lunata (abundant in tow 3 from 

Ferrel station 27), Several gastropod species that were relatively rare in the overall 

dataset made appearances at either station 27 or 28, including Rictaxis punctostriatus, 

Sinum perspectivus, Cerodrillia simpsoni, Rubellatoma rubella, Sigatica carolinensis, 

Acteon candens, Epitonium candeanum, E. championi, E. krebsii, and E. rupicola. 

Epitoniids are well-known predators upon anemones and corals, and the turrids are also 

predators, feeding largely upon worms and other small benthic meiofauna. While the 

specific nature of the mucoid material in these samples remains a mystery, it bore some 

resemblance to the bryozoan Zoobotryon verticillatum, commonly known as animal 

grass. The absence of more typical shell hash and the presence of the diverse collection 

of gastropods, however, might reflect a habitat of relatively fine-grained sediment, 

populated by anemones and polychaetes, and loaded with organic detritus (and associated 

detritivores) at these stations.

Two species of rock-boring bivalves were represented by single valves in the beam 

trawl samples from two stations: Coralliophaga coralliophaga at 2.4 and Jouannetia 

quillingi at 33. Both species have been collected by the author from burrows in the 

limestone marl rock that forms the rocky ledges and reefs at 10-15 fms off Beaufort, 

North Carolina (Wolfe, 1968). I have also collected C. coralliophaga live from burrows 

in soft sandstone just below the intertidal zone at Myrtle Beach, South Carolina. Species 

found in the present study that were associated with J. quillingi on limestone boulders in 

North Carolina are: Lithophaga bisulcata, Diplodonta punctata (wedged into crevices 

and burrow-holes), Chama congregata, Chama macerophylla, Chama radians, and 

Vermicularia knorri, (Wolfe, 1968). Other species that are likely associated with the rock 

outcrops include: Spondylus americanus, Diodora cayenensis, Calliostoma pulchra, C. 

yucatecanum, the muricids Calotrophon ostrearum, Haustellum rubidum, Murexiella 

leviculus, and Stramonita haemastoma floridana, Erosaria acicularis, and Simnialena 

uniplicata (on gorgonian coral). Station numbers where these species were best 

represented in the samples were: 42, 40, 2.1, 2.4, 43, 41, 36, and 33. The occurrence of 

many of these species at low numbers in many different samples; however, suggests that 

suitable rock habitat is exposed in small patches over extensive areas of the shelf.

Rock-dwelling Species 
14

 

Range Extensions 

Prior records of distribution and occurrence were examined for each species on the 

list of taxa (Appendix II) in order to identify unusual records and possible range 

extensions. I started with Rosenberg’s (2005) online Malacolog database, which reports 

northern and southern range limits of records for most species known to occur in the 

Western Atlantic. Other authorities, including previous studies of molluscan fauna of 

similar continental shelf habitats off the coast of the southeastern United States (Wells et 

al., 1964; Porter and Wolfe, 1972; Shoemaker, 1972; Porter, 1974; Wenner et al., 1983) 

were also consulted to verify published distributions. Table 3 lists twenty-one (21) taxa 

found in this study that were flagged as possible range extensions, based strictly on their 

ranges of latitude as recorded by Rosenberg (2005). The species list (Appendix II) may 

contain numerous other additions to the molluscan fauna previously recorded on the 

Continental Shelf of Georgia, but that is not addressed here. As many of the species in 

Table 3 are represented only by dead specimens, the possibility clearly exists that they 

were physically transported to this location by oceanographic processes; but their 

presence here nonetheless indicates that they are living nearby. 

Exceptions to the ranges reported in Malacolog are noted here for five of the taxa 

listed in Table 3 (Lioberus castaneus, Orobitella floridana, Varicorbula philippii, 

Epitonium folaceicosta, Oliva reticularis). All five of these species have been previously 

reported to occur well to the north of Georgia. Mikkelson and Bieler (2008) described 

the range of L. castaneus as a typical extended Caribbean distribution from North 

Carolina to Florida, West Indies and Caribbean Central America to Brazil. Porter (1974) 

cited Coues (1871) for a North Carolina record of this species and a live specimen 

(DAW1667) was retrieved by the author in 1967 from a settling tank for a laboratory 

seawater system at Pivers Island, Beaufort, North Carolina. Mikkelson and Bieler (2008) 

described the range of Orobitella floridana, as a more nearly Carolinian distribution from 

North Carolina to Florida (and Bahamas), and Gulf of Mexico to Caribbean Central 

America to Brazil; Abbott (1974) noted it only from East and West Florida. Porter 

(1974) cited Jacot (1921) for a North Carolina record (Cape Lookout) of this species (as 

Montacuta floridana), but had not collected the species in North Carolina. In their 

review of Varicorbula, Mikkelson and Bieler (2001) provisionally recognized V. philippii

as distinct from V. limatula [synonyms: operculata (Philippi, 1848) and disparilis

d’Orbigny 1853], and gave its range (Mikkelson and Bieler, 2008) as North Carolina to 

Florida, Bermuda, Gulf of Mexico, West Indies, Caribbean Central America, and Brazil. 

Although not reported off North Carolina by Porter and Wolfe (1972) or Porter (1974), 

the species was included (with V. operculata) in material from the gut contents of starfish 

(Astropecten) on the calico scallop beds there (Mikkelson and Bieler, 2001). Porter 

(1974) cited Dall (1903) for a North Carolina record (Cape Hatteras) of Epitonium 

foliaceicosta (as Scala novemcostata), and a specimen (DAW925) was retrieved in 1966 

west of Cape Lookout, at a depth of 12-15 fathoms on calico scallop beds off Beaufort. 

Merrill and Petit (1969) reported Oliva reticularis living off McClellanville, South 

Carolina at 46m.
15

 

 

Table 3. Possible Range Extensions for Mollusks Found in this Study, Based Primarily on Ranges Reported by Rosenberg (2005). ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

DEPTH #RETRVD PREVIOUSLY RECORDED RANGE (MALACOLOG)

SPECIES STA #'s (M) ALL STA'S LATITUDE DEPTH REGIONS** ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

Ischnochiton hartmeyeri (Thiele, 1910) 2.4* 18 1 NR NR # EFL, FLK, WFL, BAH, YUC, BRZ

Lioberus castaneus (Say, 1822) 42* 40 1 28°N to 24°S 2-42 m EFL, WFL, YUC

Limatula hendersoni Olsson & McGinty, 1958 38* 43 1 26°N to 9°N NR EFL, BAH, PAN

Diplodonta notata Dall & Simpson, 1901 38,42 40-43 1.5 28°N to 18.2°N 4-7 EFL, WFL, FLK; PR 

Orobitella floridana (Dall, 1899) 29,34 15-15 2 28°N to 9°N 0-0.5 EFL,WFL,FLK

Ensis megistus Pilsbry & McGinty, 1943 40,42 22-40 1.5 30°N to 26.5°N 0-37 EFL, WFL

Pitar simpsonii Dall, 1889 40 22 0.5 27.7°N to 18°N 0-48 EFL, WFL

Cooperella atlantica Rehder, 1943 2.2*,34* 15-19 2 26°N to 34°S 3-3 EFL, FLK

Varicorbula philippii (E.A. Smith, 1885) 38 43 0.5 32.3°N ; 64.8°W NR BER

Caryocorbula cf. cymella (Dall, 1881) 6* 43 1 25°N to 6°S 36-125 FLK

Thracia cf. phaseolina Lamarck, 1822 33 13.4 0.5 25°N to 20°N 1170 FLK; YUC 

Thracia conradi Couthouy, 1838 39 90 2x 0.5 45.7°N to 41.2°N 274 NS; ME to NY

Periploma leanum (Conrad, 1831) 24 14 1 47°N to 35°N 0-45 QUE, NS; NY to NC

Barleeia cf. tincta Guppy, 1895 42 40 1 27°N to 10.5°N 0-8 EFL; TRD 

Caecum imbricatum Carpenter, 1858 37, SC06* 33-36 2 30°N to 24°S 0-183 EFL, WFL to YUC & BRZ

Epitonium foliaceicosta (d'Orbigny, 1842) 37 33 1 30°N to 21°S 0-219 EFL,WFL,FLK,BAH, CUB, PR, USVI, YUC, BRZ

Calotrophon ostrearum (Conrad, 1846) 2.1,24 14-20 3 30.3°N to 20°N 0-64 EFL WFL, FLK; YUC

Oliva reticularis Lamarck, 1811 39 90 1 32.3°N to 9.4°N 0-200 EFL; BER, BAH, CUB, PR, YUC to VEN

Inodrillia cf. avira Bartsch, 1943 a[8 sta’s] 11-45 13 26°N ; 80°W 183-382 EFL

Kurtziella dorvilliae (Reeve, 1845) 2.4 18 1 28°N to 21°S 0-50 EFL, WFL, USVI to COL, VEN & BRZ

Turbonilla krebsii (Mörch, 1875) 38 43 1 26.67°N to 12°S 0-0 BAH; CUB; USVI; BRZ ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

**To conserve space, some regions reported in Malacolog have been omitted. REGION CODES: QUE: Québec; NS: Nova Scotia; EFL: East Florida; WFL: West 

Florida; FLK: Florida Keys; BAH; Bahamas; BER: Bermuda; CUB: Cuba: PR: Puerto Rico; USVI: US Virgin Islands; DR: Dominican Republic; JAM; Jamaica; 

YUC: Yucatan; BRZ: Brasil; PAN: Panama; COL: Colombia; VEN: Venezuela; TRD: Trinidad

• live-taken specimen

NR = not reported in Malacolog (Rosenberg, 2005)

# (Lyons 1980 & pers. comm..)

a

[1,4*,6,32,36,37,42,43]
16

Three of the taxa noted in Table 3 (Ensis megistus, Caecum imbricatum, 

Calotrophon ostrearum) had previously been reported to occur off the coast of 

Florida north of Cape Canaveral (~28.4°N), and so their occurrence on the continental 

shelf off Georgia represents only a minor extension of reported range. Mikkelson 

and Bieler (2008) do not recognize Ensis megister as a distinct species, showing E. 

minor to occur from New Jersey to Florida and the Gulf of Mexico, and to reach a 

length of 146 mm. The Georgia specimens were broken single valves that appeared 

to have been at least 125 mm in length. Porter (1974) reported C. imbricatum from 

North Carolina (erroneously synonymizing the species with C. cooperi); Abbott 

(1974) gave its range as Florida and Bahamas to Texas and West Indies; and Redfern 

(2001) describes it as “fairly common” in Abaco. Abbott (1974) gave the range of C. 

ostrearum as the “west coast of Florida to the Florida Keys.” H. Lee (pers. comm.) 

describes it as rare off Jacksonville, Florida. None of these three taxa were found 

live.

Eight additional species (Ischnochiton hartmeyeri, Limatula hendersoni, 

Diplodonta notata, Pitar simpsoni, Cooperella atlantica, Barleeia cf. tincta, 

Inodrillia cf. avira, and Kurtziella dorvilliae) have the distinction of prior records on 

the East Coast of Florida south of Cape Canaveral. For most of these species the 

Georgia records represent range extensions greater than 325 km. The chiton (I. 

hartmeyeri), a single live specimen from station 2.4, had been previously recorded 

from Hutchinson Island, southeast Florida, Dry Tortugas and the Florida west coast 

(Lyons, 1980), although the species has subsequently been reported from additional 

east Florida locations (W. Lyons, pers. comm.). 

Two of the bivalves (L. hendersoni and C. atlantica) were also found alive. The 

Limatula was identified by H. Lee, and has also been found by him east of St. 

Augustine, Florida (pers. comm.). Cooperella atlantica was previously reported 

from SE Florida to the Greater Antilles and Brazil (Abbott, 1974; Mikkelson and 

Bieler, 2008), but has not been found in Jacksonville (H. Lee, pers. comm.). Pitar 

simpsoni (a somewhat worn single valve identified by H. Lee) has been previously 

recorded from S Florida and the Bahamas to the West Indies and the Gulf of Mexico 

(Abbott, 1974; Mikkelson and Bieler, 2008). The Diplodonta notata is a tenuous 

distinction of three single valves, based on the pattern of microscopic surface 

pustules, from other more common Ungulinids (Appendix 2). The species has been 

reported previously from the Florida Keys and Bahamas through the Gulf of Mexico 

and Caribbean to Colombia (Mikkelson and Bieler, 2008). A single specimen, dead 

and worn, resembling Barleia tincta, was retrieved from station 42 at 40 m. B. tincta

has been reported from Hutchinson Island, Florida (Lyons, 1989); and similar 

specimens have also been obtained off Jacksonville and other E Florida locations 

(H. Lee, pers. comm.). Taxonomic confusion surrounding Kurtziella dorvilliae is 

discussed by Rosenberg (2005) and Lee (pers. comm.). The species, previously 

reported off SE Florida (Abbott, 1974), has also been found off St. Augustine and at 

Jacksonville Beach (H. Lee, pers. comm.). 
17

Inodrillia cf. avira is only tentatively identified, based on its very close 

resemblance to the illustration of that species by Abbott (1974: p. 274, species 3095) 

and Bartsch’s (1943) very detailed original description. This species, previously 

recorded only in deep water off SE Florida (Bartsch, 1943; Abbott, 1974); was 

retrieved (thirteen specimens) from eight stations ranging from 11-45 m in depth; and 

at least one specimen was in very fresh condition (station 4*). These specimens were 

examined by H. Lee and by Donn Tippett, who concurred that they should be 

properly assigned to Viridrillia. Lee has found the same taxon among calico scallops 

trawled from at least two locations off E. Florida (specimens seen by author). 

Tippett (pers. comm.) indicated that Bartsch’s (1943) Inodrillia/Viridrillia material in 

the U.S. National Museum is in need of serious review, and that the Georgia 

specimens might even represent a new species. 

United States records in Malacolog (Rosenberg, 2005) for two of the Georgia 

species (Caryocorbula cf. cymella, Thracia cf. phaseolina) were restricted to the 

Florida Keys. A single live specimen of Caryocorbula cf. cymella was retrieved 

from station 6 at 43 m. The shell is distinguished from Caryocorbula contracta by a 

thinner, more transluscent shell with opaque white mottlings scattered over the 

surface, and lacking commarginal ridges near the umbones (Mikkelson and Bieler, 

2008). These same authors give its range as Florida Keys and West Indies to Brazil. 

According to Mikkelson and Bieler (2008), Thracia phaseolina is a European species, 

and the W Atlantic species is as yet undescribed. A single valve was found at station 

33.

Two taxa in Table 3 (Thracia conradi, and Periploma leanum) are northern 

species with reported ranges south only to Long Island Sound, New York and off 

North Carolina, respectively (Abbott, 1974). Although Thracia conradi is typically 

found in waters deeper than the stations sampled here, two (broken) left valves, the 

larger 70 mm long and retaining its color and gloss internally, were recovered from 

station 39 (90 m). It seems improbable that these specimens would have been 

transported upslope to this location; the species likely occurs live near here. The P. 

leanum (single valve only) appeared at a shallow (14 m) station (#24). Porter (1974) 

reported P. leanum from off Cape Fear (Southport), North Carolina.

The one remaining taxon in Table 3, Turbonilla krebsii, was previously known 

only from the Bahamas (Redfern, 2001) and the Netherlands Antilles (De Jong and 

Coomans, 1988). While this genus contains a bewildering array of species, the single 

specimen (~6.5 mm) found in excellent condition at station 38 (43 m) on the Georgia 

shelf is a dead ringer for those found in Abaco and illustrated by Redfern (2001: Pl. 

67, species 626)–– down to the orange-brown color and the incised spiral lines and 

faint cancellate sculpture on the body whorl behind the outer lip. The species occurs 

also in Bermuda (H. Lee, pers. comm.).

 
18

Notes on Other Taxa 

Five species listed in Appendix 2 (but not in Table 3) deserve special note here 

because of residual identification issues, and possible implications for further 

extensions of range. These taxa are discussed below:

Melanella sp., found at stations 2.2 and 38, strongly resembles Eulimostraca sp. B 

from Abaco (species 340: Redfern, 2001). Lee (pers. comm.) suggests that the species 

is one he has not seen before, and moreover, that the aperture is “that of a Melanella, 

not Eulimostraca”. He further likened the specimen to Polygireulima amblytera

(A.E. Verrill and Bush, 1900) and to Polygireulima sp. A (Pl. 6, fig. 6, in Lyons, 

1989)–– both recorded previously in southern Florida. 

Dentimargo aureocincta, represented by four specimens (dead, but in good 

condition) from three stations (33-103 m depth), correspond well to the illustration in 

Redfern (2001: Pl. 47, species 437A) found in Abaco and identified as Dentimargo

cf. macnairi (Bavay, 1922). Malacolog data (Rosenberg, 2005) report the range of 

Dentimargo macnairi as Martinique and Panama (14.5°N). Dall (1927) named 

several other species of Dentimargo from off Georgia for which I could locate no 

illustrations, but none of Dall’s written descriptions seemed to match these specimens 

exactly. These four specimens were examined by H. Lee and described by him as a 

“depauperate morph” of D. aureocincta; they are so identified in Appendix 2.

Olivella sp. is represented by two freshly dead, shiny creamy white 

specimens,11.5 and 12.9 mm in length, from stations 5 and SC10 at 33-37 m. The 

shells have 5.5-6 opaque whorls with a very deeply channeled suture; with heavy 

callus on the parietal area and lacking plicae on the columella and pillar structure; and 

the aperture is about 1/2 the total length. Based on Dall’s (1889) description and 

distribution data given by Abbott (1974) and Rosenberg (2005), I speculated that 

these might be Belloliva tubulata (Dall, 1889). Lee (pers. comm.), however, relates 

they are not that species and that they correspond very closely to an un-named species 

illustrated by Kaicher (1987: card no. 5019) that he treats as Olivella sp. aff. 

tunquina (Duclos, 1835), in his list of NE Florida Marine Mollusks (Lee, 2006). I 

have not yet seen an illustration that corresponds to this species.

The taxonomy of Platycythara elata is somewhat confusing in the literature: the 

species has variously been assigned to the genera Daphnella, Rubellatoma, 

Vitricythara, and Platycythara (Fargo, 1953; Lyons, 1989; Turgeon et al., 1998), and 

the discussion continues (Lee, pers. comm.). The single specimen (fresh but with a 

gaping hole in the body whorl) from the Georgia shelf corresponds extremely well to 

specimens illustrated by Fargo (1953) and Lyons (1989), and to a specimen recovered 

from a batfish taken off St. Johns County, Florida and illustrated (scanning electron 

micrograph, as Vitricythara auberiana) by Lee (2006). It is creamy white, however, 

with a light brown band emanating from the top of the aperture and encircling the 

body whorl around to the dorsum of the siphonal canal. Similar banding has not been 

reported on other specimens. The range of P. elata is reported (Rosenberg, 2005) as: 

34.53°N to 18.2°N; 82°W to 67.2°W, at depths of 11-73m. The type locality is 12 
19

miles east of Frying Pan Shoals, off North Carolina, at 12 fathoms. The species was 

found off North Carolina also by Wells et al., (1961), but was not found by Porter 

(1974).

The turrid Cryoturris fargoi also deserves special mention. It is a single 

specimen, collected dead but in good condition, whose identification was confirmed 

by Harry Lee. This specimen is represented quite well by an illustration in Lyons 

(1989: Plate XI, fig. 1), and described by him as having a range of “both coasts of 

Florida.” Three other specimens of Cryoturris fargoi , obtained from W of Cape 

Lookout, North Carolina (at 65 ft in seastar Astropecten stomach), Puerto Rico and 

W Florida, respectively, and verified by D. Tippett (USNM), reside in the author’s 

collection. Comparing the Georgia specimen with these, I suspect that two species 

may be involved here. The present specimen, about 5.2 mm long, is tan in color; its 

spire angle (corresponding with Lyon’s illustration) is greater; and it has more axial 

riblets than any of the other three specimens, all of which are white. All four of the 

specimens, however, possess fine spiral lines with microscopic beads, giving the 

shells a characteristic frosted appearance. Similar specimens have been obtained by 

Lee (pers. comm.) from scallop beds off eastern Florida, and he also noted variability 

of spire angle among specimens from different locations. Examination of more 

specimens may be required to ascertain the identification of this taxon, and the 

implications for range extension.

Gray’s Reef National Marine Sanctuary [Stations 2.1-2.4] Taxa 

Of the 263 taxa recovered in this study (Appendix II), ninety-seven (97) were 

retrieved from at least one of the four stations (2.1-2.4) bordering the GRNMS. 

These taxa are identified in Appendix II by an entry in the “STA 2?” column, 

indicating the actual number of stations (that is, of the four stations 2.x) where the 

species was found. These four stations all had heavily skewed distributions of 

species numbers similar to those found throughout the overall set of stations, and 

exhibited little similarity of species composition. The number of taxa occurring at 

stations 2.x (2.1 through 2.4) were 51, 32, 35, and 56, respectively, while the 

numbers of single-station (of the four 2.x stations) species were 19, 12, 6, and 20, 

respectively. Only eleven taxa occurred at all four of the stations 2.x; fifteen at three, 

fourteen at two, and 57 occurred at only one. Of the 57 taxa occurring at only one of 

the stations 2.x, sixteen (16) occurred at only one of the thirty-seven total stations, 

eight (8) were at 2; seven (7) were at 3; six (6) were at 4; eight (8) were at 5; three (3) 

were at each of 6 and 7; two (2) at 9; and one each was at 10, 12, 13, and 14. 

Since none of the stations 2.x actually lay within the GRNMS, and the species 

composition exhibited similar degrees of skewness and low concordance among these 

four stations as among the 37 stations at large (Fig. 2), I conclude that the station 2.x 

dataset is only slightly more likely to be representative of the GRNMS than the 

dataset from the overall set of 37 stations. One may treat the station 2.x dataset as 

reasonably positive evidence of a species’ occurrence within the GRNMS. Given the 

extreme patchiness of species occurrence noted among trawl replicates and among 
20

stations in this study, and the great diversity of habitats previously documented within 

the GRNMS, most of the species found in this study are nonetheless quite likely to 

occur also within the boundaries of the Sanctuary. This is particularly likely since the 

patch reef habitats characteristic of the GRNMS have been documented as 

widespread across the Georgia shelf, and extending to depths greater than within the 

GRNMS itself (Van Dolah et al., 1994). The absence of marked differences in 

species composition and occurrence with station depth, coupled with the fact that the 

GRNMS is located near the mid-range depths for this study, further supports this 

conclusion.

The list of taxa from the present study (Appendix II) was also compared with a 

list of benthos previously reported to occur within the Gray’s Reef National Marine 

Sanctuary (GRNMS, 2004), which was based on a benthic survey (Vittor and 

Associates, Inc., 2004). That survey used a Young dredge (sampling area of 0.04 

m

2

) and diver cores (area = 0.0071 m2), both of which are designed for quantitatively 

sampling benthic infauna. Either three or five replicate samples were taken at each of 

seven stations within the GRNMS. The beam trawl is more effective at collecting 

epifauna, and the 5-minute tows covered a much larger area, so the studies are 

complementary and one should not expect great similarity between the two lists. 

Benthic collections have also been made within the GRNMS by Hyland et al., (2006), 

and a few species are mentioned in that work that are not included on the GRNMS 

(2004) list.

The GRNMS benthos list includes 107 species of mollusks identified to the 

species level, 34 to genus, 37 only to family, and six only to a taxonomic level higher 

than family, for a total of 184 taxa. At least twenty-six species (including LPTL’s) of 

bivalves and twenty-eight species of gastropods on the GRNMS list were not found in 

the present study. The present study identifies one species of chiton, 81 species of 

bivalves, four species of scaphopods, 108 species of gastropods and one cephalopod 

not named explicitly on the GRNMS list. Some of these taxa may occur on the 

GRNMS list as indeterminate LPTL taxa, however, so the apparent discrepancy 

between study results is probably exaggerated to some extent. 

Potential Species Richness 

To assist the estimation of the potential total number of molluscan species that 

might ultimately be found in this region, I generated a “species discovery curve” 

(Keating et al., 1998; Solow and Smith, 2005) based on presence-absence data for the 

263 taxa from the 37 stations in this study (Fig. 4). Total species were cumulated 

over the 37 sampling stations twelve times, in randomized station order, and leastsquare regressions were performed on the resultant dataset, using logarithmic and 

negative exponential functions. In order to improve the fit of the regression curves to 

the upper portion of the dataset, the first five cumulative values were omitted from 

both regressions. This had the dual effects of: (a) forcing the curves through 263 

species at, or very near, 37 stations, and (b) increasing the projected number of 

species that would be discovered by additional sampling.
21

The negative exponential regression approaches an asymptotic maximum of 311 

species (compared to a value of 287 for the same regression on the complete dataset). 

The logarithmic function projects that 311 species would be discovered by sampling a 

total of 67 stations, and that further sampling would continue to reveal additional 

species (Fig. 4). 

Several more sophisticated extrapolative techniques have been developed and tested 

for estimating species richness (Chao, 1984; Foggo et al., 2003; Colwell et al., 2004), 

and Colwell (2006) provides downloadable software (EstimateS©: v. 8.0) enabling 

ready computation of additional estimates (Table 4). To generate a more robust 

database than the presence-absence data used for Fig. 4, I employed numbers of 

individuals for each species at each station, by spreading the species numbers 

collected during the Oregon II cruise, evenly among their stations of origin, back into 

the dataset. While this approach no doubt reduced the skewness of species numbers 

inherent among stations, it nonetheless provided a first order approximation of 

species distribution and relative abundance. Table 4 presents the estimates of species 

richness generated by EstimateS© (Colwell, 2006) for eight different extrapolative 

techniques applied to this reconstructed dataset, using 100 randomized cumulations 

without replacement.

Estimates of Smax ranged from 308 species (Bootstrap– similar to negative 

exponential regression of presence-absence data, Fig. 4) to 429 species (Jack2), with 

a median value in the range of 368-374 species (ACE/Chao 1). Given the limitations 

of this database, it is impossible to assess the relative merits of these estimates in 

quantitative terms, but there is a strong concensus among approaches that many 

species remain to be discovered. 
22

0

50

100

150

200

250

300

350

400

1 11 21 31 41 51 61 71 81 91 101

Cumulative stations

 

Figure 4. Species discovery curves based on 263 species at 37 Georgia Continental Shelf 

stations near Gray’s Reef National Marine Sanctuary, illustrating extrapolations based on 

logarithmic and negative exponential regression of twelve random cumulation curves. 

Table 4. Estimates of Mollusk Species Richness (Smax) on the Georgia Continental 

Shelf, using EstimateS©. See Colwell (2006) for additional references on estimators.

–––––––––––––––––––––––––––––––––––––––

Estimator Smax –––––––––––––––––––––––––––––––––––––––

ACE (Abundance-base coverage) 368

ICE (Incidence-base coverage) 383

Chao 1 (Chao, 1984) 374

Chao 2 (Chao, 1987) 404

Jack1 (first-order jackknife) 365

Jack 2 (second-order jackknife) 429

Bootstrap 308

Michaelis-Menton 343

–––––––––––––––––––––––––––––––––––––––

Y =(A)lnX - B

Y = Ymax – Be-kx
23

Discussion 

Most of the species retrieved in this study were recognized members of either 

the Caribbean or Carolinian provinces. The Carolinian province is the coastal and 

near-shore area that extends from Cape Hatteras (some species range north to New 

Jersey) to Cape Canaveral and on the West Coast of Florida from about Charlotte 

Harbor or Sarasota northward and around the Gulf of Mexico to the Yucatan 

Peninsula. The Caribbean province, centered in the West Indies, includes Southern 

Florida and the Bahamas, with extensions to Bermuda and (along the shelf break and 

Gulf Stream edge) to off Cape Hatteras. It includes the deeper portions of the Gulf of 

Mexico, the Greater and Lesser Antilles, Windward and Leeward West Indies, 

Caribbean Central America (eastern Yucatan to Panama), and coastal South America 

to about Sao Paolo, Brazil. The Carolinian and Caribbean faunas meet and mix on 

the Georgia continental shelf where this study was conducted. Provincial affinities of 

species have been discussed in some detail for the Florida Mixing Grounds (Hopkins 

et al., 1977) in the northeastern Gulf of Mexico where the Caribbean and Carolinian 

provinces also meet. Most of the northerly range extensions noted here involved 

Caribbean species, perhaps suggesting that the fauna of the nearshore region (i.e. the 

Carolinian Province) had received more thorough study. 

A number of species found at the shallower stations sampled here are wellknown components of inshore coastal and estuarine communities (Prezant et al., 

2002), and dead shells are probably transported onto the shelf by local circulation, 

including estuarine outwelling (Hanson et al., 1981), and by hermit crabs. The 

presence of dead specimens of common shallow species at the two deep stations (39 

and 44) also suggests that materials are transported across the continental shelf and 

onto the slope.

Although this study was necessarily qualitative in nature, some associations 

among taxa were still evident in the data. The over-riding conclusion that can be 

drawn from these samples, however, relates to the extreme patchiness of the various 

types of habitats and their associated fauna on the Continental Shelf of Georgia. This 

was reflected in the great variability of sample volume and sample character– even 

for replicate tows of the beam trawl at the same station, and in the high proportion of 

“rare” species within samples (one individual per tow), within stations (one individual 

per station), or within the entire data set from all stations. Taxa that would be 

expected to associate with a particular habitat type were frequently intermixed in the 

same tow with taxa expected from different habitats, demonstrating that the beam 

trawl was collecting from multiple habitat types during the tow. Each sample came 

from a 5-minute tow of the beam trawl, so the scale on which this habitat diversity 

occurs is small. The average bottom area that was swept by the beam trawl was 370 

m

2 (based on ship speed of 1.5 knots (0.77 m/sec) and a fishing width of 1.6 m for the 

2-m beam trawl). While the ship usually conducted the tows into the current and 

wind, the direction of the tows was haphazard (H. Walsh, pers. comm.). As a result 

some tows were parallel to the coast and shelf break (and to the general pattern of reef 

outcrops and troughs offshore), while others could have been more nearly 
24

perpendicular to these features. Part of the variability among replicate tows at 

individual stations probably arose from this variability of towing direction relative to 

the bathymetric contours and bottom features of the shelf. The effectiveness of the 

beam trawl no doubt varies considerably depending on the physical character of the 

bottom.

Since the mesh size of the beam trawl was 3-6 mm, fine material including 

small mollusks would likely have been excluded unless larger material was first 

entrained. Thus, the samples were biased against such small species. Nonetheless, 

many small species (1-5mm) did appear and they were probably entrained along with 

the large volumes of shell hash seen in some samples. Most of the species found in 

the soft-bottom infaunal studies (Vittor and Associates, 2004; Hyland et al., 2006) 

and not reported here were small species, and were probably missed because of the 

trawl design.

Using dredges, trawls, grabs and suction devices, Wenner et al., (1983) 

collected epifauna from eight hard-bottom habitats on the shelf off the coast between 

Jacksonville, Florida and Savannah, Georgia, and one southeast of Charleston, South 

Carolina (30°25’ to 32°29’ N). Their stations included three each on the inner shelf 

(17-22 m depth), the middle shelf (23-38 m) and the outer shelf (47-67 m), so their 

study area encompassed the area sampled in the present study except for the two deep 

stations. The focus of their study, however, was on biomass and diversity of 

numbers, not on details of species occurrence. Although they reported having 

collected 203 species of mollusks in all, mollusks comprised only about 6% of the 

total numbers of taxa collected by dredge and trawl in the study (slightly greater in 

the winter than in the summer). Only four species of mollusk occurred in their 

samples with sufficient frequency or density to warrant mention of species names. 

These were Pteria colymbus, which occurred in 35% of the 82 trawl samples; 

Crassinella lunulata, which was their 9th most abundant species (16/m2) on the outer 

shelf in winter; and Elliptotellina americana and Nassarius albus, which ranked 6th

(30/m2) and 9th (23/m2), respectively, in middle shelf samples in summer. Note that 

neither Pteria colymbus nor Elliptotellina americana was found in the present study. 

Their overall species list would no doubt be quite interesting to compare with that in 

Appendix II.

All of the estimates of potential mollusk species richness are predicated only on 

beam trawl samples, which are known to be biased against infaunal and small species, 

as well as species from high profile rocky areas or other habitats that may be poorly 

compatible with trawling. Because these samples included dead specimens gleaned 

from shell hash, the habitat bias is reduced, but the number of molluscan species 

occurring on the Georgia Continental Shelf most likely exceeds that estimated by any 

of the species discovery techniques (Fig. 4, Table 4).

Many of the species awaiting discovery on the Georgia Continental Shelf may 

be long-time resident species that have merely been missed or overlooked by the 

sampling effort to date because of their size, rarity or cryptic habits. Most certainly, 
25

however, new species have been, and will be, introduced to this region also as a result 

of changing environmental conditions or even human intervention. Seemingly small 

and subtle changes in either the circulation patterns or the temperature regimes of 

surface and bottom waters over the shelf could readily modify the recruitment success 

of planktonic molluscan larvae transported over great distances from upstream 

reproducing populations (Scheltema, 1989). The seasonal structure of water 

temperatures over the continental shelf of the South Atlantic Bight is strongly 

influenced by riverine flow from the landmass, onwelling of cool deep water onto the 

shelf edge, intrusions of the gulf stream and its eddies, and regional circulation 

patterns, which are largely wind-driven (Blanton et al., 2003). Tester and Steidinger 

(1997) described the circuitous paths taken by Gulf Stream eddies over this shelf 

region, in connection with transport of red tide organisms. Such complexity renders 

the detection of long-term sea-surface temperature trends very tenuous and uncertain, 

whether on a regional (Friedland and Hare, 2007) or global (Cane et al., 1997) scale. 

Nonetheless, the variations in sea-surface temperature and associated circulation 

patterns have strong implications for transport of eggs and larvae of offshore 

spawning fish (Stegmann and Yoder, 1996), planktonic veliger larvae of mollusks 

(Scheltema, 1989), and other organisms (Tester and Steidinger, 1997).

Examples of invasive “alien” marine species recently brought into the South 

Atlantic Bight by Gulf Stream circulation include: (1) the lionfish (Pterois volitans), 

presently distributed from Cuba to Cape Hatteras, and throughout the Bahamas to 

Bermuda (Whitfield et al., 2002; Schofield et al., 2008); (2) the green porcelain crab 

(Petrolisthes armatus), distributed northward as far as South Carolina (Hollebone and 

Hay, 2007) and Bermuda (Simmons, 2007); and (3) the Asian green mussel (Perna 

viridis), so far known only as far north as Georgia (Power et al., 2004; DeVictor and 

Knott, 2007). While these three examples represent truly “alien” species with historic 

distributions in the Pacific (lionfish and green mussel) and Brazil (porcelain crab), 

their rapid spread into the South Atlantic Bight region has been documented only 

within the past eight years from earlier locations on the west coast (lionfish and green 

mussel) or southeastern tip (porcelain crab) of Florida. 

Another recent molluscan range extension, reported here for the first time, is the 

zebra periwinkle [Echinolittorina ziczac (Gmelin, 1791)], which has been found by 

the author in colonies at two locations in North Carolina: in June 2005 on a concrete 

seawall near the Coast Guard Station at Fort Macon, inside Beaufort Inlet, and in 

August 2007 on the shore end of the rock jetty at Cape Lookout. That species was 

recently reported (Prezant et al., 2002) to occur in Georgia, but prior to that, was 

known only well south of Cape Canaveral, Florida (Rosenberg, 2005: Malacolog).

The foregoing examples further reinforce the present finding that of the 263 

taxa identified here, at least 16 (6.1%) were species not previously known to occur on 

the Georgia shelf. Moreover, numerous species associated with the calico scallop 

community off Beaufort, North Carolina (Wells et al., 1964; Porter and Wolfe, 1972) 

were not found in the present study. Taken together, the incidence of range 

extensions reported here, the documented patchiness and diversity of habitat in the 
26

region, the knowledge that additional molluscan members of the Caribbean province 

have been previously reported in more northerly waters (to Cape Hatteras), and the 

implications of global climate change, all serve to emphasize that the benthic and 

epibenthic molluscan fauna of the Georgia Continental Shelf remain only partially 

described and warrant continued study.
27

Acknowledgments 

The following personnel, many of whom at the time were at the NOAA Center for 

Fisheries and Habitat Research, assisted in the completion of this study. Don Hoss 

alerted me to the potential availability of shell hash from the NOAA cruises on the 

Georgia Shelf, and facilitated my access to the samples. Jon Hare (NOAA/NMFS, 

Narragansett, RI) and Harvey Walsh (WHOI, Woods Hole, MA) ensured that I 

received the samples after the target fish species were removed, and they (and Dave 

Meyer) also reviewed early drafts of the manuscript. Harvey also provided 

information on sampling procedures, and created the station location map. Mike 

Prager assisted in regression of species discovery data. Patti Marraro, librarian at the 

NOAA Center, helped me obtain molluscan literature needed for the study, and 

provided editorial review. James Morris formated the manuscript for publication. 

Jeff Govoni encouraged the completion of the study and facilitated preliminary 

scientific review of the manuscript. Nick Wolfe (Portland, OR) made the computer 

runs using EstimateS©. William G. Lyons (St. Petersburg, FL) kindly identified the 

chiton specimen and provided information on its prior distribution records. Dr. Harry 

G. Lee (Jacksonville, FL) examined numerous specimens and gave invaluable 

assistance in the identification of many taxa, along with his observations on their 

occurrence in NE Florida. Dr. Donn Tippett (Potomac, MD), volunteer at the 

Smithsonian Institution, assisted in the discussion and identification of several turrid 

species. Center support for the original sample collection was obtained from 

NOAA’s National Marine Sanctuary Office and Grays Reef National Marine 

Sanctuary. 
28

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34

APPENDIX I. STATION LOCATIONS AND DEPTHS FOR BEAM TRAWLS 

––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

Station Sample Sample LATITUDE LONGITUDE DEPTH

No. Date Date (M) ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

1 2/9/02 ––– 31°27.00'N 80°01.60'W 11

2.1 2/10/02 4/16/02 31°24.57'N 80°49.30'W 20

2.2 2/9/02 4/17/02 31°26.10'N 80°54.00'W 19

2.3 2/9/02 4/16/02 31°21.47'N 80°51.00'W 19

2.4 2/9/02 4/17/02 31°22.48'N 80°55.48'W 18

4 2/10/02 ––– 31°17.50'N 80°28.40'W 33

5 2/10/02 ––– 31°14.20'N 80°17.70'W 37

6 2/10/02 ––– 31°11.95'N 80°06.90'W 43

7 2/11/02 ––– 31°08.10'N 79°56.30'W 47

22 ––– 4/17/02 31°31.5 'N 80°04.6 'W 4.9

23 2/11/02 4/12/02 31°31.0 'N 81°00.0 'W 12.5

24 2/11/02 4/12/02 31°30.6 'N 80°55.3 'W 14

25 2/11/02 4/12/02 31°30.2 'N 80°50.6 'W 17.5

27 2/12/02 4/16/02 31°22.5 'N 81°09.9 'W 8.5

28 2/12/02 4/17/02 31°22.9 'N 81°03.8 'W 11.2

29 2/11/02 4/17/02 31°23.2 'N 80°58.3 'W 15

31 2/12/02 ––– 31°18.4 'N 81°11.5 'W 10

32 2/12/02 ––– 31°17.9 'N 81°06.2 'W 11.2

33 2/12/02 ––– 31°17.4 'N 81°01.3 'W 13.4

34 2/12/02 ––– 31°16.9 'N 80°56.4 'W 15

35 ––– 4/12/02 31°30.21'N 80°35.4 'W 22

36 ––– 4/12/02 31°32.18'N 80°20.4 'W 31

37 ––– 4/12/02 31°33.12'N 80°05.4 'W 33

38 ––– 4/13/02 31°34.03'N 79°50.4 'W 43

39 ––– 4/13/02 31°34.80'N 79°37.2 'W 90

40 ––– 4/16/02 31°23.36'N 80°39.74'W 22

41 ––– 4/14/02 31°23.36'N 80°25.67'W 30

42 ––– 4/14/02 31°23.36'N 80°11.65'W 40

43 ––– 4/13/02 31°23.36'N 79°57.60'W 45

44 ––– 4/13/02 31°23.36'N 79°44.28'W 103

SC01 2/8/02 ––– 31°32.00'N 79°47.40'W 47

SC02 2/8/02 ––– 31°35.00'N 79°55.00'W 44

SC06 2/9/02 ––– 31°28.70'N 80°06.50'W 36

SC07 2/13/02 ––– 31°17.48'N 79°51.85'W 47

SC08 2/12/02 ––– 31°20.37'N 80°02.89'W 43

SC09 2/12/02 ––– 31°23.12'N 80°15.56'W 37

SC10 2/12/02 ––– 31°26.67'N 80°25.35'W 33 ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––
35

APPENDIX II. MOLLUSCAN TAXA RECOVERED 

Taxa are presented in the phylogenetic sequence used by Turgeon et al., (1998). Data for 

each taxon includes:Station 2 occurrence (# of stations 2.1 to 2.4 represented), the total 

number of stations where it was collected (#), the depth range of those stations (meters), and 

the total number of specimens recovered (#). An asterisk (*) before the number of stations 

indicates that the taxon was collected live from at least one of those stations. 

_________________________________________________________________________________________________________________________

 STA STA'S DEPTH No. 

FAMILY GENUS & SPECIES 2? (#) (m) Rcvrd _________________________________________________________________________________________________________________________

Amphineura Ischnochiton hartmeyeri (Thiele, 1910) *1 *1 18 1

Bivalvia

Nuculidae Nucula proxima Say, 1822) 1 30 1

Mytilidae Lioberus castaneus (Say, 1822) *1 40 1

Mytilidae Lithophaga bisulcata (d'Orbigny, 1842) *1 *1 20 1

Mytilidae Modiolus squamosus Beauperthuy, 1967 *1 12.5 1

Mytilidae Musculus lateralis (Say, 1822) 1 *2 12.5-19 2

Arcidae Anadara floridana (Conrad, 1869) 4 12.5-33 1.5

Arcidae Anadara notabilis (Röding, 1798) *2 40-45 1.5

Arcidae Anadara transversa (Say, 1822) *6 12.5-43 14.5

Arcidae Arca zebra (Swainson, 1833) 1 40 0.5

Noetiidae Arcopsis adamsi (Dall, 1886) 1 1 19 0.5

Glycymerididae Glycymeris americana (DeFrance, 1829) 3 *9 4.9-40 4

Glycymerididae Glycymeris spectralis Nicol, 1952 1 45 0.5

Glycymerididae Glycymeris undata (Linné, 1758) *2 *9 18-47 13

Glycymerididae Tucetona pectinata (Gmelin, 1791) *1 *6 12.5-30 10

Pinnidae Atrina rigida (Lightfoot, 1786) *7 12.5-47 100

Pinnidae Atrina seminuda (Lamarck, 1819) *1 12.5 1

Limidae Limaria pellucida (C.B.Adams, 1846) 1 13.4 1

Limidae Limatula hendersoni Olsson & McGinty, 1958 *1 43 1

Pectinidae Aequipecten muscosus (W. Wood, 1828) *5 30-45 4.5

Pectinidae Argopecten gibbus (Linné, 1758) *3 *18 12.5-103 20

Pectinidae Euvola raveneli (Dall, 1898) *6 22-43 8.5

Pectinidae Spathochlamys benedicti (Verrill & Bush, 1897) *1 30 2

Propeamussidae Similipecten nanus Verrill & Bush, 1897 3 30-45 12.5

Plicatulidae Plicatula gibbosa Lamarck, 1801 *4 *11 14-43 23

Spondylidae Spondylus americanus (Hermann, 1781) *3 *5 15-30 3.5

Anomiidae Anomia simplex d'Orbigny, 1842 1 *6 13.4-45 12

Ostreidae Ostrea equestris Say, 1834 *1 *4 8.5-43 6

Lucinidae Callucina keenae Chavan, 1971 2 33-40 4

Lucinidae Cavilinga blanda (Dall, 1901) *3 *14 11-47 46

Lucinidae Ctena orbiculata (Montagu, 1808) 1 90 0.5

Lucinidae Divalinga quadrisulcata (d'Orbigny, 1842) 1 3 15-30 6.5

Lucinidae Lucinisca nassula (Conrad, 1846) 1 12.5 1

Lucinidae Parvilucina costata (d'Orbigny, 1845) 2 40-43 1

Lucinidae Parvilucina crenella (Dall, 1901) 1 15 0.5

Lucinidae Phacoides pectinatus (Gmelin, 1791) 1 1 19 0.5

Lucinidae Radiolucina amianta (Dall, 1901) *2 33-43 1.5

Ungulinidae Diplodonta notata Dall & Simpson, 1901 2 40-43 1.5
36

Ungulinidae Diplodonta nucleiformis(Wagner, 1840) 1 *5 15-43 5

Ungulinidae Diplodonta punctata (Say, 1822) 1 6 15-47 10.5

Ungulinidae Phlyctiderma semiasperum (Philippi, 1836) *2 *8 15-43 6

Chamidae Arcinella cornuta Conrad, 1866 *2 *10 12.5-47 8.5

Chamidae Chama congregata Conrad, 1833 *1 *5 20-40 6.5

Chamidae Chama macerophylla Gmelin, 1791 1 *7 15-45 12.5

Chamidae Chama radians Lamarck, 1819 *2 22-40 2.5

Lasaeidae Orobitella floridana (Dall, 1899) 1 15 0.5

Leptonidae Pythinella cuneata (Verrill & Bush, 1898) *3 8.5-11.2 106

Sportellidae Basterotia quadrata (Hinds, 1843) 2 15-15 2

Carditidae Glans dominguensis (d'Orbigny, 1853) *1 43 2

Carditidae Pleuromeris tridentata (Say, 1826) *4 *23 11-103 91

Carditidae Pteromeris perplana (Conrad, 1841) 1 *9 11.2-43 16

Astartidae Astarte crenata subequilatera G.B. Sowerby II 1 43 1.5

Astartidae Astarte nana Dall, 1886 3 33-43 3.5

Crassatellidae Crassinella dupliniana (Dall, 1903) 1 47 1

Crassatellidae Crassinella lunulata (Conrad, 1834) *3 *14 8.5-43 21.5

Crassatellidae Eucrassatella speciosa (A. Adams, 1852) *1 40 2.5

Cardiidae Trachycardium muricatum (Linnaeus, 1758) 3 4.9-22 1.5

Cardiidae Dinocardium robustum (Lightfoot, 1786) 1 3 4.9-18 2

Cardiidae Laevicardium laevigatum (Linnaeus, 1758) 3 *8 14-45 3.5

Cardiidae Laevicardium pictum (Ravenel, 1861) *4 *17 11.2-47 65

Cardiidae Nemocardium peramabile (Dall, 1881) 1 3 11-19 3

Mactridae Mulinia lateralis (Say, 1822) *5 8.5-40 16.5

Mactridae Spisula raveneli (Conrad, 1831) *4 11.2-15 3.5

Pharidae Ensis megistus Pilsbry & McGinty, 1943 2 22-40 1.5

Pharidae Ensis minor Dall, 1900 1 *5 15-37 5.5

Tellinidae Angulus sybariticus (Dall, 1881) *3 *15 8.5-40 39

Tellinidae Angulus tenellus (A.E. Verrill, 1874) *1 13.4 1

Tellinidae Angulus texanus (Dall, 1900) 1 11.2 1

Tellinidae Angulus versicolor (DeKay, 1843) 1 *4 14-33 3

Tellinidae Eurytellina alternata (Say, 1822) *1 8.5 1

Tellinidae Macoma brevifrons (Say, 1834) *2 13.4-15 1.5

Tellinidae Macoma tenta (Say, 1834) 1 15 1

Tellinidae Merisca aequistriata (Say, 1824) *5 15-40 3.5

Tellinidae Phyllodina squamifera (Deshayes, 1855) *1 90 1.5

Tellinidae Strigilla mirabilis (Philippi, 1841) *4 11.2-17.5 8

Tellinidae Tellinella listeri Roding, 1798 *2 *5 18-40 6

Semelidae Abra aequalis (Say, 1822) 2 22-30 1

Semelidae Ervilia concentrica (Holmes, 1860) *4 *17 8.5-103 114

Semelidae Semele bellastriata (Conrad, 1837) *3 *9 14-40 53

Semelidae Semele purpurascens (Gmelin, 1791) *1 15 1

Semelidae Semelina nuculoides (Conrad, 1841) 1 *5 15-43 13

Trapezidae Coralliophaga coralliophaga (Gmelin, 1791) 1 1 18 0.5

Veneridae Chione elevata Say, 1822 2 15-30 1.5

Veneridae Chione mazyckii Dall, 1902 1 30 0.5

Veneridae Cyclinella tenuis (Récluz, 1852) *1 43 1

Veneridae Gemma gemma (Totten, 1834) 2 15-47 64.5

Veneridae Gouldia cerina (C.B.Adams, 1845) *1 *12 11.2-43 45

Veneridae Lirophora latilirata (Conrad, 1841) 2 *5 18-43 9.5

Veneridae Macrocallista maculata (Linné, 1758) *3 *12 12.5-43 12.5
37

Veneridae Macrocallista nimbosa (Lightfoot, 1786) *1 15 1.5

Veneridae Mercenaria campechiensis (Gmelin, 1791) 1 33 0.5

Veneridae Pitar simpsoni (Dall, 1889) 1 22 0.5

Veneridae Puberella intapurpurea (Conrad, 1849) *4 *13 11.2-33 39.5

Veneridae Timoclea grus (Holmes, 1858) *2 30-33 1.5

Veneridae Transenella stimpsoni Dall, 1902 *3 33-43 5.5

Petricolidae Cooperella atlantica Rehder, 1943 *1 *3 15-33 3

Petricolidae Petricolaria pholadiformis (Lamarck, 1818) 1 17.5 1

Myidae Paramya subovata (Conrad, 1845) 1 15 0.5

Corbulidae Caryocorbula caribaea (d'Orbigny, 1853) *1 43 1

Corbulidae Caryocorbula chittyana (C. B. Adams, 1852) 1 31 0.5

Corbulidae Caryocorbula contracta (Say, 1822) *8 11.2-103 13

Corbulidae Caryocorbula cf. cymella (Dall, 1881) *1 43 1

Corbulidae Caryocorbula dietziana C.B.Adams, 1852 1 *3 11.2-43 3.5

Corbulidae Varicorbula limatula (Conrad, 1846) *3 33-43 2

Corbulidae Varicorbula philippii (E.A. Smith, 1885) 1 43 0.5

Pholadidae Jouannetia quillingi Turner, 1955 1 13.4 1

Lyonsiidae Lyonsia hyalina Conrad, 1831 *1 14 1

Pandoridae Pandora trilineata Say, 1822 *1 *5 10-40 10.5

Thraciidae Thracia cf. phaseolina Lamarck, 1822 1 13.4 0.5

Thraciidae Thracia conradi Couthouy, 1838 1 90 0.5

Periplomatidae Periploma leanum (Conrad, 1831) 1 14 1

Periplomatidae Periploma margaritaceum (Lamarck, 1801) 1 1 18 0.5

Verticordiidae Trigonulina ornata d'Orbigny, 1842 2 13.4-90 1.5

Cuspidariidae Cuspidaria obesa (Lovén, 1846) 1 43 0.5

Scaphopoda

Dentaliidae Antalis taphrium Dall, 1889 3 22-47 5

Dentaliidae Paradentalium americanum (Chenu, 1843) 1 10 2

Dentaliidae Dentalium laqueatum Verrill, 1885 1 30-90 4

Dentaliidae Graptacme calamus Dall, 1889 2 31-47 4

Dentaliidae Graptacme eborea (Conrad, 1846) 2 3 15-20 4

Gadilidae Polyschides tetraschistus (Watson, 1879) 2 40-43 2

Gadilidae Polyschides tetrodon (Pilsbry & Sharp, 1898) 2 30-33 2

Gastropoda

Fissurellidae Diodora cayenensis (Lamarck, 1822) 1 4 15-31 5

Fissurellidae Lucapinella limatula (Reeve, 1850) 1 30 1

Turbinidae Arene tricarinata (Stearns, 1872) 1 12 11-45 25

Turbinidae Eulithidium thalassicola (Robertson, 1958) 2 43-45 4

Turbinidae Turbo castanea Gmelin, 1791 1 *3 14-20 7

Calliostomatidae Calliostoma pulchrum (C.B.Adams, 1850) 1 14 1

Calliostomatidae Calliostoma yucatecanum Dall, 1881 1 *14 11.2-47 23

Litiopidae Litiopa melanostoma Rang, 1829 1 1 20 1

Cerithiidae Cerithium atratum (Born, 1778) 2 7 8.5-33 9

Modulidae Modulus modulus (Linnaeus, 1758) 1 4 18-43 5

Turritellidae Torcula acropora Dall, 1889 4 18 11-43 100

Turritellidae Vermicularia knorri (Deshayes, 1843) 1 47 1

Littorinidae Littoraria irrorata (Say, 1822) 1 11.2 1

Barleeiidae Barleeia tincta Guppy, 1895 1 40 1

Vitrinellidae Circulus multistriatus (A.E. Verrill, 1884) *1 *1 19 1
38

Vitrinellidae Cyclostremiscus beaui (P.Fischer, 1857) 1 3 19-43 4

Caecidae Caecum carolinianum Dall, 1892 1 43 2

Caecidae Caecum cooperi S. Smith, 1860 *1 43 2

Caecidae Caecum imbricatum Carpenter, 1858 *2 33-36 2

Caecidae Caecum pulchellum Stimpson, 1851 1 15 1

Strombidae Strombus alatus Gmelin, 1791 1 5 11.2-22 10

Calyptraeidae Bostrycapulus aculeatus (Gmelin, 1791) 1 31 1

Calyptraeidae Calyptraea centralis (Conrad, 1841) 4 *15 11-43 43

Calyptraeidae Cheilea equestris (Linnaeus, 1758) 1 14 1

Calyptraeidae Crepidula fornicata (Linné, 1758) 2 *6 12.5-40 15

Calyptraeidae Crepidula plana Say, 1822 *1 *4 13.4-90 6

Calyptraeidae Crucibulum striatum (Say, 1826) 1 40 2

Xenophoridae Xenophora conchyliophora (Born, 1780) *6 33-47 11

Vermetidae Petaloconchus erectus (Dall, 1888) 2 13.4-37 2

Cypraeidae Erosaria acicularis (Gmelin, 1791) 1 47 1

Ovulidae Simnialena uniplicata (G.B.Sowerby II, 1849) *2 13.4-17.5 2

Naticidae Naticarius canrena (Linnaeus, 1758) *12 11.2-47 23

Naticidae Neverita duplicata (Say, 1822) 1 7 8.5-22 7

Naticidae Polinices porcellanus (d'Orbigny, 1839) 2 14 18-40 59

Naticidae Sigatica carolinensis (Dall, 1889) 4 8.5-43 4

Naticidae Sinum maculatum (Say, 1831) 1 1 19 1

Naticidae Sinum perspectivum (Say, 1831) 1 11.2 1

Naticidae Tectonatica pusilla (Say, 1822) 3 *13 8.5-47 40

Cassidae Cassis madagascariensis Lamarck, 1822 3 8.5-40 3

Cassidae Cypraecassis testiculus (Linnaeus, 1758) 1 47 1

Cassidae Semicassis granulata (Born, 1778) *6 33-47 6

Ranellidae Cymatium cingulatum (Lamarck, 1822) 4 43-47 38

Ranellidae Cymatium krebsii (Mörch, 1877) 1 45 1

Ranellidae Cymatium parthenopeum (von Salis, 1793) *2 45-47 17

Cerithiopsidae Seila adamsii (H.C.Lea, 1845) *2 *5 15-43 6

Triphoridae Marshallora nigrocincta (C.B. Adams, 1839) 1 43 1

Epitoniidae Cirsotrema dalli Rehder, 1945 *5 37-45 6

Epitoniidae Depressiscala nautlae (Mörch, 1875) *3 14-47 3

Epitoniidae Epitonium albidum (d'Orbigny, 1842) *1 *1 19 1

Epitoniidae Epitonium angulatum (Say, 1831) 4 33-43 5

Epitoniidae Epitonium candeanum (d'Orbigny, 1842) 1 43 1

Epitoniidae Epitonium multistriatum (Say, 1826) 6 8.5-47 6

Epitoniidae Epitonium occidentale (Nyst, 1871) *2 11-12.5 2

Epitoniidae Epitonium championi Clench & Turner, 1952 1 *5 10-19 6

Epitoniidae Epitonium foliaceicosta (d'Orbigny, 1842) 1 33 1

Epitoniidae Epitonium krebsii (Mörch, 1875) 4 11.2-47 5

Epitoniidae Epitonium novangliae (Couthouy, 1838) 1 47 1

Epitoniidae Epitonium rupicola (Kurtz, 1860) 3 8.5-12.5 4

Epitoniidae Opalia pumilio (Mörch,1875) 1 31 1

Eulimidae Melanella sp. cf. Eulimostraca sp.B Redfern 340 1 2 20-43 2

Eulimidae Melanella hypsela (A.E. Verrill & Bush, 1900) *2 *2 18-19 2

Eulimidae Melanella jamaicensis (C.B. Adams, 1845) 1 12.5 1

Eulimidae Niso aeglees Bush, 1885 1 *9 17.5-43 11

Muricidae Calotrophon ostrearum (Conrad, 1846) 1 2 14-20 3

Muricidae Chicoreus florifer dilectus (A. Adams, 1855) *2 15-40 2

Muricidae Eupleura caudata (Say, 1822) *3 *5 12.5-20 7
39

Muricidae Haustellum rubidum (F.C.Baker, 1897) 1 43 1

Muricidae Murexiella levicula (Dall, 1889) 2 40-43 3

Muricidae Stramonita haemastoma floridana (Conrad, 1837) 2 22-45 2

Buccinidae Antillophos candeanus (d'Orbigny, 1842) 1 90 1

Buccinidae Hesperisternia multangulus (Philippi, 1848) 1 17.5 1

Melongenidae Busycon sinistrum Hollister, 1954 4 31-47 3

Melongenidae Busycotypus canaliculatus (Linnaeus, 1758) 1 90 1

Melongenidae Busycotypus spiratus (Lamarck, 1816) 1 47 1

Nassariidae Ilyanassa obsoleta (Say, 1822) 1 3 10-20 7

Nassariidae Ilyanassa trivittata (Say, 1822) 1 4 4.9-37 6

Nassariidae Nassarius acutus (Say, 1822) *6 4.9-40 125

Nassariidae Nassarius albus (Say, 1826) *10 30-47 21

Nassariidae Nassarius consensus (Ravenel, 1861) *7 33-47 21

Nassariidae Nassarius vibex (Say, 1822) *1 90 1

Fasciolariidae Fasciolaria lilium hunteria (G.Perry, 1811) *1 *2 13.4-20 2

Fasciolariidae Pleuroploca gigantea (Kiener, 1840) 1 11.2 1

Columbellidae Astyris raveneli (Dall, 1889) 1 43 1

Columbellidae Astyris lunata (Say, 1826) 2 *9 8.5-90 134

Columbellidae Costoanachis translirata (Ravenel, 1861) 4 *22 8.5-47 123

Columbellidae Suturoglypta iontha Ravenel, 1861 *1 14 1

Columbellidae Nassarina glypta (Bush, 1885) 2 33-43 5

Columbellidae Parvanachis obesa (C.B.Adams, 1845) 1 11 1

Olividae Oliva reticularis Lamarck, 1811 1 90 1

Olividae Oliva sayana Ravenel, 1834 *1 *5 11.2-18 6

Olividae Olivella floralia (Duclos, 1844) 3 *21 8.5-43 102

Olividae Olivella mutica (Say, 1822) 4 *27 8.5-90 216

Olividae Olivella nivea (Gmelin, 1791) 1 1 18 1

Olividae Olivella sp. aff. tunquina (Duclos, 1835) 2 33-37 2

Marginellidae Dentimargo aureocinctus (Stearns, 1872) 3 33-103 4

Marginellidae Prunum amabile authors non (Redfield, 1852) 1 *7 14-43 11

Marginellidae Prunum apicinum (Menke, 1828) *3 12.5-30 7

Marginellidae Prunum hartleyanum (Schwengel, 1941) 7 10-43 21

Marginellidae Prunum roscidum (Redfield, 1860) 1 3 20-33 4

Costellariidae Vexillum wandoense (Holmes, 1860) *1 *1 19 1

Cancellariidae Axelella smithii (Dall, 1888) 1 47 2

Cancellariidae Cancellaria reticulata (Linnaeus, 1767) 4 5 12.5-20 8

Cancellariidae Tritonoharpa lanceolata (Menke, 1828) 2 30-47 3

Drilliidae Cerodrillia simpsoni (Dall in Simpson, 1887) 2 11.2-43 2

Drilliidae Cerodrillia bealiana Schwengel & McGinty, 1942 1 7 20-45 8

Drilliidae Drillia cydia (Bartsch, 1943) 2 6 15-43 8

Drilliidae Splendrillia moseri (Dall, 1889) 2 45-103 4

Terebridae Terebra concava (Say, 1826) *3 *15 8.5-103 46

Terebridae Terebra dislocata (Say, 1822) *1 *10 4.9-47 18

Terebridae Terebra protexta (Conrad, 1846) *3 *6 14-103 9

Turridae Cochlespira radiata (Dall, 1889) *1 103 1

Turridae Crassispira fuscescens (Reeve, 1845) 1 1 20 1

Turridae Inodrillia cf. avira Bartsch, 1943 *8 11-45 13

Conidae Brachycythara biconica (C.B. Adams, 1850) 1 30 1

Conidae Conus anabathrum Crosse, 1865 1 *13 12.5-47 21

Conidae Conus stearnsii Conrad, 1869 4 *12 12.5-43 34

Conidae Cryoturris cerinella (Dall, 1889) 3 17.5-103 3
40

Conidae Cryoturris fargoi McGinty, 1955 1 1 20 1

Conidae Glyphoturris quadrata (Reeve, 1845) 1 30 1

Conidae Ithycythara lanceolata (C.B.Adams, 1850) 2 11-33 2

Conidae Kurtziella atrostyla (Tryon, 1884) 5 8.5-33 15

Conidae Kurtziella cerina (Kurtz & Stimpson, 1851) 1 17.5 1

Conidae Kurtziella dorvilliae (Reeve, 1845) 1 1 18 1

Conidae Kurtziella limonitella (Dall, 1884) 3 8.5-11.2 19

Conidae Platycythara elata (Dall, 1889) 1 43 1

Conidae Rubellatoma rubella (Kurtz & Stimpson, 1851) 3 18-25 11

Architectonicidae Heliacus bisulcatus (d'Orbigny, 1842) 2 37-43 3

Pyramidellidae Odostomia laevigata (d'Orbigny, 1841) *1 13.4 1

Pyramidellidae Petitilla crosseana (Dall, 1885) 1 90 1

Pyramidellidae Pyramidella suturalis H.C. Lea, 1843 *4 8.5-15 6

Pyramidellidae Turbonilla interrupta (Totten, 1835) 1 1 20 1

Pyramidellidae Turbonilla krebsii (Mörch, 1875) 1 43 1

Acteonidae Acteon candens Rehder, 1939 4 8.5-103 6

Acteonidae Rictaxis punctostriatus (C.B. Adams, 1840) 1 8.5 1

Cylichnidae Acteocina candei (d'Orbigny, 1841) 1 2 18-43 2

Cylichnidae Acteocina lepta Woodring, 1928 1 43 1

Philinidae Philine sagra (d'Orbigny, 1841) 1 43 1

Retusidae Pyrunculus caelatus (Bush, 1885) 1 43 1

Retusidae Retusa sulcata (d'Orbigny, 1841) 1 43 1

Cavoliniidae Diacavolinia longirostris (Blainville, 1821) *2 *10 15-103 25

Arminidae Armina mulleri (von Ihering, 1886) *1 14 1

Cephalopoda:

Ommastrephidae Illex illecebrosus (Lesueur, 1821) *1 17.5 5

––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––
41
42
United States Department of Commerce

Carlos M. Gutierrez 

Secretary

National Oceanic and Atmospheric Administration 

William J. Brennan, Ph.D.

Acting Under Secretary of Commerce for Oceans and Atmospheres 

Acting Administrator, National Oceanic and Atmospheric Administration

National Ocean Service

John H. Dunnigan

Assistant Administrator for Ocean Service and Coastal Zone Management
United States Department of National Oceanic and National Ocean Service

Commerce Atmospheric Administration

Carlos M. Gutierrez Conrad C. Lautenbacher, Jr. John H. Dunnigan

Secretary Administrator Assistant Administrator