A guide to the Thalassinidea

(Crustacea: Malacostraca: Decapoda) 

of the South Atlantic Bight

Richard W. Heard

Rachael A. King

David M. Knott

Brent P. Thoma

Susan Thornton-DeVictor

NOAA Professional Paper NMFS 8

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NOAA Professional Paper NMFS 8

A guide to the Thalassinidea 

(Crustacea: Malacostraca: Decapoda) 

of the South Atlantic Bight 

Richard W. Heard

Rachael A. King

David M. Knott

Brent P. Thoma

Susan Thornton-DeVictor

October 2007

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Heard, Richard W., Rachael A. King, David M. Knott, Brent P. Thoma, and Susan 

Thornton-DeVictor. 2007. A guide to the Thalassinidea (Crustacea: Malacostraca: 

Decapoda) of the South Atlantic Bight. NOAA Professional Paper NMFS 8. 

30 p.

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iii

CONTENTS

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

 Overview of the Thalassinidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 Previous records of thalassinideans from the South Atlantic Bight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 Updated distributional records of thalassinideans from the South Atlantic Bight . . . . . . . . . . . . . . . . . . . . . . . . . 2

Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Key to the Thalassinidea from the South Atlantic Bight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Synopsis of Thalassinidea known from the South Atlantic Bight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

INFRAORDER THALASSINIDEA Latreille, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

SUPERFAMILY AXIOIDEA Huxley, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

FAMILY AXIIDAE Huxley, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Acanthaxius hirsutimanus (Boesch and Smalley, 1972) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Axius armatus S. I. Smith, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Calaxius jenneri (Williams, 1974) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Paraxiopsis gracilimana Kensley, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

FAMILY CALOCARIDIDAE Ortmann, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Calocaris templemani Squires, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

SUPERFAMILY CALLIANASSOIDEA Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

FAMILY Callianassidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Subfamily Callianassinae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Biffarius biformis (Biffar, 1971a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Biffarius cf. fragilis (Biffar, 1970) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Gilvossius setimanus (DeKay, 1844) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Necallianassa berylae Heard and Manning, 1998 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Subfamily Callichirinae Manning and Felder, 1991a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Callichirus major Say, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Subfamily Cheraminae Manning and Felder, 1991a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Cheramus marginatus (Rathbun, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

FAMILY LAOMEDIIDAE Borradaile, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Naushonia crangonoides Kingsley, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

FAMILY THOMASSINIIDAE de Saint Laurent, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Crosniera wennerae Heard and King, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

FAMILY UPOGEBIIDAE Borradaile, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Upogebia affinis (Say, 1818) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

A guide to the Thalassinidea 

(Crustacea: Malacostraca: Decapoda) 

of the South Atlantic Bight 

Richard W. Heard1, Rachael A. King,2 David M. Knott,2

Brent P. Thoma,

1

 and Susan Thornton-DeVictor2

1 The University of Southern Mississippi 

 Department of Coastal Sciences

 Gulf Coast Research Laboratory

 P.O. Box 7000

 Ocean Springs, Mississippi 39466-7000

2 Southeastern Regional Taxonomic Center (SERTC)

 South Carolina Department of Natural Resources

 P.O. Box 12559

 Charleston, South Carolina 29422-2559

Email (for R. King): KingR@dnr.sc.gov

Abstract. Ghost shrimp and mud shrimp 

in the decapod infraorder Thalassinidea are 

ecologically important members of many 

benthic intertidal and shallow subtidal infaunal communities, largely due to the sediment filtration and mixing that result from 

their burrowing and feeding behavior. These 

activities considerably modify their immediate environment and have made these cryptic 

animals extremely interesting to scientists 

in terms of their behavior, ecology, and 

classification. 

Over 20 years ago, seven species of thalassinideans were known from the South Atlantic Bight (Cape Hatteras, NC to Cape Canaveral, FL). During this study, the examination 

of extensive collections from the National 

Museum of Natural History (NMNH), the 

Southeastern Regional Taxonomic Center 

(SERTC), and regional institutions, resulted 

in the identification of 14 species of thalassinideans currently known to occur within 

this region. The family Axiidae is represented 

by three species: Axius armatus, Calaxius 

jenneri, and Paraxiopsis gracilimana; the Callianassidae by six: Biffarius biformis, B. cf. 

fragilis, Callichirus major, Cheramus marginatus, Gilvossius setimanus, and Necallianassa 

berylae; the Calocarididae by two: Calocaris 

templemani and Acanthaxius hirsutimanus; and 

the families Laomediidae, Thomassiniidae, 

and Upogebiidae are each represented by 

one: Naushonia crangonoides, Crosniera wennerae, and Upogebia affinis, respectively. An 

illustrated key is presented for species level 

identification and supplemental notes on the 

ecology, distribution, and taxonomy of the 

species are provided. 

Introduction

Despite the fact that they are commonly referred to as ghost shrimp 

or mud shrimp, thalassinideans 

belong to a decapod crustacean 

infraorder that is more closely related to anomurans and crabs than 

to true shrimp. These burrowing 

animals are an important benthic 

component of sandy or muddy intertidal and shallow subtidal habitats 

worldwide (Griffis and Suchanek, 

1991; Atkinson and Taylor, 2004). 

Although casual observers walking 

along the shore may notice the holes 

of thalassinidean burrows on tidal 

mud flats or sand bars, it is unlikely 

that many of them understand the 

significance of the animals within.

The actively burrowing lifestyle of 

thalassinideans and their filter feeding strategies result in high rates of 

sediment turnover. Using abdominal appendages that are specially 

modified, thalassinideans draw water into burrows that may be relatively simple “Y”- or “U”-shaped vertical 

tunnels, or much more elaborate 

systems of branching tunnels and 

chambers (Griffis and Suchanek, 

1991). The animals’ biofiltering activity disperses nutrients and oxygen 

throughout the surface sediments 

and also provides food that sustains 

them in their underground homes. 

Their burrowing and water circulation exert a large influence on the 

sediment structure and geochemistry of environments where they 

are abundant (Ziebis et al., 1996; 

Dworschak, 2000a; Dworschak et 

al., 2006). For that reason, these 

decapod crustaceans are fascinating to scientists, who are just beginning to understand their complex 

behavior, specialized morphology, 

and the evolutionary history of the 

infraorder. 

This guide comprises a broad 

taxonomic assessment and review 

of the thalassinidean fauna of the 

South Atlantic Bight (SAB), which 

is defined here as the coastal and 
2 

oceanic region of the southeastern United States between Cape Hatteras, North Carolina and Cape Canaveral, Florida. This guide is further restricted to depths 

ranging from the intertidal zone to 200 m. Since the 

last broad taxonomic treatment of SAB thalassinideans 

(Williams, 1984) is now over 20 years old and missing 

many of the species currently known to occur within 

the SAB, it was clear that a taxonomic revision of SAB 

thalassinideans was needed. 

The purpose of this study is to critically review previous information and studies, establish new regional 

records, document distributional records for the SAB, 

summarize the taxonomic information available on 

each species treated, and present an illustrated key for 

the identification of the Thalassinidea currently known 

from the inshore, nearshore, and offshore shelf waters 

of the SAB. 

Overview of the Thalassinidea

Currently, the Thalassinidea contains three superfamilies (Axioidea Huxley, 1879; Callianassoidea Dana, 

1852; Thalassinoidea Latreille, 1831), whose members 

are widely distributed in estuarine and marine habitats 

throughout much of the world from the intertidal to 

depths exceeding 2500 m (Lemaitre and Ramos, 1992; 

Poore, 1997; Dworschak, 2000a; Dworschak, 2005).

Members of the infraorder Thalassinidea are highly 

specialized decapod crustaceans that construct and live 

in permanent burrows in sediment. These burrows can 

be highly complex, with multiple chambers, and the 

burrows of some species can exceed a depth of 2 m 

(Weimer and Hoyt, 1964; Griffis and Suchanek, 1991; 

Nickell and Atkinson, 1995; Tamaki and Ueno, 1998; 

Dworschak, 2000b; Kinoshita et al., 2003). Thalassinideans possess an elongate and flexible sub-cylindrical body 

(partly sclerotized in callianassids and upogebiids, more 

so in axiids); chelipeds, legs, and mouthparts adapted 

for locomotion, burrowing, and feeding; broad setose 

pleopods for pumping water through burrows; and 

broad uropods for sealing burrows (Bishop and Williams, 2005). Figure 1 shows the general morphology 

of a thalassinidean.

Several different feeding strategies have been recorded within the Thalassinidea: suspension feeding, 

deposit feeding, and scavenging (Griffis and Suchanek, 

1991; Nickell and Atkinson, 1995; Abed-Navandi and 

Dworschak, 2005). Predation or carnivorous feeding 

behavior by the callianassid Corallianassa longiventris (A. 

Milne-Edwards, 1870) on other crustaceans has been 

observed during field studies in the Turks and Caicos 

(Heard, pers. observ.) Under laboratory conditions, 

however, this species appeared to exhibit deposit and 

herbivorous feeding behavior (Dworschak, 2005). The 

feeding and burrowing activity of thalassinideans has 

a great impact on the environment surrounding the 

burrows, given the high densities of many species and 

the large amounts of sediment re-working they cause. 

Callianassa species can filter as much as 2.59kg/m2

of sediment per day (Suchanek, 1983), and Biffarius 

biformis Biffar, 1971a has been recorded at densities of 

483 individuals/m2 in St. Catherines Sound, Georgia 

(Bishop and Bishop, 1992). 

Detailed treatments of the classification of Thalassinidea have been presented by Manning and Felder 

(1991a), Poore (1994), Tudge et al. (2000), Martin and 

Davis (2001), and Sakai (2005). While it is the most 

recent, Sakai’s (2005) classification of the Infraorder 

Thalassinidea Latreille, 1831, has not yet been critically 

evaluated, and so we follow the classification presented 

by Martin and Davis (2001), which largely followed 

Poore (1994). 

Members of the thalassinidean superfamilies Axioidea and Callianassoidea are found in the SAB. The 

Axioidea is composed of four families, two of which are 

represented in the SAB: the Axiidae Huxley, 1879 and 

the Calocarididae Ortmann, 1891. The Callianassoidea 

contains six families, four of which are represented in 

the SAB: Callianassidae Dana, 1852, Laomediidae Borradaile, 1903, Thomassinidae de Saint Laurent, 1979, 

and Upogebiidae Borradaile, 1903. The Thalassinoidea, 

a small group containing a single genus with a few species, is largely restricted to the western Pacific (Poore 

and Griffin, 1979; Dworschak, 1992) and is not known 

from the SAB.

Previous records of thalassinideans 

from the South Atlantic Bight

The last comprehensive review of regional thalassinideans was in A. B. Williams’ (1984) guide to shrimp, 

lobsters, and crabs of the Atlantic coast of the eastern 

United States. Williams (1984) recorded seven species 

from the South Atlantic Bight (Callianassa major Say, 

1818; Callianassa atlantica Rathbun, 1926; Callianassa 

biformis Biffar, 1971a; Axiopsis jenneri (Williams, 1974); 

Calocaris templemani Squires, 1965; Naushonia crangonoides Kingsley, 1897; Upogebia affinis (Say, 1818)). Nizinski 

(2003) expanded the list to eight species with the addition of Gilvossius setimanus (De Kay, 1844) (=Callianassa 

atlantica) and Necallianassa berylae Heard and Manning, 

1998, based on an extensive review of taxonomic literature published since 1984. 

Updated distributional records of 

thalassinideans from the South Atlantic Bight

Since Williams’ (1984) comprehensive monograph on 

the Decapoda of the U.S. Atlantic coast, three new species have been described within the SAB (Paraxiopsis 
3

Figure 1

The general morphology of thalassinideans: (A) dorsal view; (B) lateral view. Modified from Biffar, 1971b. P1-5: pereopods 

1–5; A1-6: abdominal segments 1–6.

gracilimana Kensley, 1996, Necallianassa berylae Heard 

and Manning, 1998, and Crosniera wennerae Heard and 

King, 2007), and a range extension of Axius armatus S. 

I. Smith, 1881 was reported (Kensley, 2001). Range extensions of four more species, Acanthaxius hirsutimanus 

(Boesch and Smalley, 1972), Calaxius jenneri (Williams, 

1974), Cheramus marginatus (Rathbun, 1901), and Necallianassa berylae are recorded here. Thus, we report here 

that 14 species of thalassinideans occur within the SAB.

Table 1 gives a systematic listing of the Thalassinidea 

known from the SAB.

There have been a number of taxonomic changes 

within the Thalassinidea since Williams (1984) that have 

been adopted here. A taxonomic listing of the species 

within the Thalassinidea currently known from the SAB 

is given in Table 1. 

Materials and methods

We examined scientific publications, “gray literature” 

(e.g. regional agency technical reports, contract reports, etc.), and collections from the region (National 

Museum of Natural History, Smithsonian Institution, 

Washington, DC; Duke Marine Laboratory, Beaufort, 

NC; Grice Marine Laboratory, Charleston, SC; South 

Carolina Department of Natural Resources, Charleston, 

SC; and University of Louisiana at Lafayette, LA), to produce a comprehensive list of thalassinidean species from 

the SAB. Wherever possible, we verified these records by 

re-examining the specimens reported.

The following abbreviations have been used: 

USNM—United States National Museum, now known 

as the National Museum of Natural History (NMNH), 
4 

Table 1

A systematic listing of Thalassinidea currently known 

from the inshore, shelf, and slope waters of the SAB. 

(*), new taxon reports since Williams (1984); (+), 

range extensions; (=), synonyms. 

Order Decapoda Latrielle, 1802

Infraorder Thalassinidea Latreille, 1831

Superfamily Axioidea Huxley, 1879

Family Axiidae Huxley, 1879

 Genus Axius Leach, 1815

 Axius armatus S. I. Smith, 1881*+

 Genus Acanthaxius Sakai and de Saint Laurent, 1989

 Acanthaxius hirsutimanus (Boesch and Smalley, 1972)* +

 Genus Calaxius Sakai and de Saint Laurent, 1989

 Calaxius jenneri (Williams, 1974)* +

 =Axiopsis jenneri (Williams, 1974)

 Genus Paraxiopsis DeMan, 1905

 Paraxiopsis gracilimana Kensley, 1996*

Family Calocarididae Ortmann, 1891

 Genus Calocaris Bell, 1853

 Calocaris templemani Squires, 1965

Superfamily Callianassoidea Dana, 1852

Family Callianassidae Dana, 1852

 Subfamily Callianassinae Dana, 1852

 Genus Biffarius Manning and Felder 1991a

 Biffarius biformis (Biffar, 1971a)

 =Callianassa biformis Biffar, 1971a

 Biffarius cf. fragilis (Biffar, 1970)

 Genus Gilvossius Manning and Felder, 1991b

 Gilvossius setimanus (DeKay, 1844)

 =Callianassa atlantica Rathbun, 1926

 Genus Necallianassa Heard and Manning, 1998

 Necallianassa berylae Heard and Manning, 1998*+

 Subfamily Callichirinae Manning and Felder, 1991a*

 Genus Callichirus Stimpson, 1866 

 Callichirus major (Say, 1818)

 Subfamily Cheraminae Manning and Felder, 1991a*

 Genus Cheramus Bate, 1888

 Cheramus marginatus (Rathbun, 1901)* +

 =Callianassa marginata Rathbun, 1901

Family Laomediidae Borradaile, 1903

 Genus Naushonia Kingsley, 1897 

 Naushonia crangonoides Kingsley, 1897 

Family Thomassiniidae de Saint Laurent, 1979

 Genus Crosniera Kensley and Heard, 1991

 Crosniera wennerae Heard and King, 2007*

Family Upogebiidae Borradaile, 1903

 Genus Upogebia Leach, 1814

 Upogebia affinis (Say, 1818)

Smithsonian Institution, Washington DC; SERTC—

Southeastern Regional Taxonomic Center, South Carolina Department of Natural Resources, Charleston, SC; 

GML—Grice Marine Laboratory, College of Charleston, SC; MARMAP— Marine Resources Monitoring 

Assessment and Prediction Program, South Carolina 

Department of Natural Resources, Charleston, SC; and 

ULLZ—University of Louisiana at Lafayette, Zoological 

Collection. 
5

1. Pereopod 1 strongly subchelate (dactylus articulated to fold back against the ventral margin of the propodus). Uropodal endopod and exopod each with distinct distal suture. Pereopod 2 simple (not chelate) 

(Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Naushonia crangonoides Kingsley, 1897

Figure 2

Naushonia crangonoides: (A) dorsal view of carapace; (B) inner surface of pereopod 1 (chela); (C) telson and uropods.

A–C: modified from Williams, 1984.

— Pereopod 1 strongly or weakly chelate (dactylus forming pincer with propodus). Uropodal endopod lacking 

distal suture; exopod with distal suture present or absent. Pereopod 2 chelate or simple (Fig. 3). . . . . . . . . . 2

Figure 3

(A) Acanthaxius hirsutimanus: pereopod 1 (chela); (B) Paraxiopsis gracilimana: distal end of pereopod 1 (chela); (C) 

Callichirus major: distal end of pereopod 1 (chela); (D) Paraxiopsis gracilimana: telson and uropods; (E) Gilvossius setimanus: 

telson and uropods; (F) Acanthaxius hirsutimanus: telson and uropods. A, F: modified from Boesch and Smalley, 1972; C, E: 

Modified from Williams, 1984; B, D: modified from Kensley, 1996.

Results

Key to the Thalassinidea from the South Atlantic Bight
6 

2. Rostrum well developed, spinose, either rounded or acutely tipped. Eyestalks rounded. Pereopod 3 not highly 

modified: similar to pereopods 4 and 5. Pleopods 2–5 setose, well developed (Fig. 4). . . . . . . . . . . . . . . . . . . . 3

— Rostrum small, usually triangular, flattened, dorsally smooth. Eyestalks flattened dorsoventrally. Pereopod 3 

highly modified, propodus usually broadly expanded and distinctly different from pereopods 4 and 5. Pleopods 

1–2 reduced; pleopods 3–5 well developed with broad, highly setose rami (Fig. 5). . . . . . . . . . . . . . . . . . . . . . 8

Figure 4

(A) Upogebia affinis: dorsal view of carapace showing rostrum; (B) Acanthaxius hirsutimanus: dorsal view of anterior carapace 

showing rostrum; (C) Upogebia affinis: lateral view of abdomen showing pleopods; (D) Acanthaxius hirsutimanus: lateral 

view of anterior carapace showing eyestalks; (E) Paraxiopsis gracilimana: lateral view of cephalothorax showing pereopods 

1–5. A: modified from Williams, 1984; B, D: modified from Boesch and Smalley, 1972; C: modified from Williams, 1993; E: 

modified from Kensley, 1996.

Figure 5

(A) Necallianassa berylae: dorsal view of anterior carapace showing rostrum; (B) Callichirus major: dorsal view of anterior 

carapace showing rostrum; (C) Gilvossius setimanus: dorsal view of anterior carapace showing rostrum; (D) Biffarius 

biformis: pereopod 3; (E) Biffarius biformis: lateral view of anterior carapace showing flattened eyestalks; (F) lateral view of 

generalized thalassinidean showing pleopods 1 and 2 reduced. A: modified from Heard and Manning, 1998; B, C: modified 

from Williams, 1984; D, E: original illustrations by R.W. Heard; F: modified from Biffar, 1971b.
7

3. Pereopod 1 strongly subchelate, superficially appearing chelate, but having short fixed finger (distal extension 

of propodus). Pereopod 2 simple (not chelate), similar to pereopods 3–4. Rostrum large, spinose, broadly 

rounded to a blunt tip (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Upogebia affinis (Say, 1818) 

— Pereopod 1 strongly chelate. Pereopod 2 chelate. Rostrum triangular, laterally spinose, acutely tipped 

(Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Figure 6

Upogebia affinis: (A) lateral view of whole animal; (B) detail of dactyl of pereopod 2; (C) dorsal view of carapace; (D) 

pereopod 1 (chela) showing weakly chelate form. A, B: modified from Williams, 1993; C, D: modified from Williams, 1984.

Figure 7

(A) Calocaris templemani: pereopod 1 (chela); (B) Acanthaxius hirsutimanus: pereopod 1 (chela); (C) Acanthaxius hirsutimanus: 

dorsal view of anterior carapace showing rostrum; (D) Calaxius jenneri: dorsal view of anterior carapace showing rostrum; 

(E) Paraxiopsis gracilimana: dorsal view of anterior carapace showing rostrum; (F) Acanthaxius hirsutimanus: pereopod 2. A, 

D: modified from Williams, 1984; B, C, F: modified from Boesch and Smalley, 1972; E: modified from Kensley, 1996.
8 

4. Eyes with pigment indistinct (diffuse or appearing absent); eyestalk rudimentary, flattened obliquely on distal surface. Uropodal exopod with weakly defined distal suture, recognizable part of suture 

margin lacking spines; endopod with medial carina unarmed (lacking spines). Family Calocariidae 

(Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Calocaris templemani Squires, 1965

Figure 9

(A) Acanthaxius hirsutimanus: dorsal view of anterior carapace; (B) Calaxius jenneri: dorsal view of anterior carapace; (C) 

Paraxiopsis gracilimana: dorsal view of anterior carapace; (D) Acanthaxius hirsutimanus: telson and uropods; (E) Calaxius 

jenneri: telson and uropods. A, D: modified from Boesch and Smalley, 1972; B, E: modified from Williams, 1984; C: modified 

from Kensley, 1996.

Figure 8

Calocaris templemani: (A) pereopod 1 (chela); (B) lateral view of anterior carapace; (C) telson and uropods. A–C: modified 

from Williams, 1984.

— Eyes with eyestalk well developed, nearly reaching and sometimes overreaching tip of rostrum. Uropodal 

exopod with well-defined distal suture, suture margin armed with spines; endopod with medial carina armed 

with several distinct spines. Family Axiidae (Fig. 9). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
9

5. Pereopod 1 major or minor chela with dactylus distinctly longer than the palm (proximal part of propodus), dorsal margin of dactylus with a few sharp spines. Eyes extending slightly beyond rostrum. 

Carapace dorsal surface distinctly setose, granulose, and spinose; lateral carina with 5 or more spines 

(Fig. 10). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthaxius hirsutimanus (Boesch and Smalley, 1972)

Figure 10

Acanthaxius hirsutimanus: (A) dorsal view of anterior carapace; (B) pereopod 1 major (left) chela; (C) pereopod 1 minor 

(right) chela; (D) telson and uropods; (E) lateral view of anterior carapace; (F) lateral view of posterior part of carapace 

and first three abdominal segments; (G) pereopod 2. A–G: modified from Boesch and Smalley, 1972.

— Pereopod 1 major and minor chelae with dactylus a similar length to or shorter than the palm (proximal 

part of the propodus), dorsal margin of dactylus smooth, or nearly so. Eyes not extending beyond rostrum. 

Carapace dorsal surface weakly to moderately setose; lateral carina with 4 or fewer spines (Fig. 11). . . . . . . . . 6

Figure 11

(A) Calaxius jenneri: pereopod 1 (chela); (B) Paraxiopsis gracilimana: pereopod 1 (chela); (C) Paraxiopsis gracilimana: dorsal 

view of anterior carapace; (D) Axius armatus: dorsal view of anterior carapace; (E) Calaxius jenneri: dorsal view of anterior 

carapace. A, E: modified from Williams, 1984; B, C: modified from Kensley, 1996; D: modified from Kensley, 2001.
10

6. Carapace dorsal surface with indistinct lateral and submedial carinae; lateral carinae indicated by a single strong 

spine (not noticeably coalescing to form rostral margin). Pereopod 1 major chela masive, densely setose. Telson 

without medial spine on distal margin (Fig. 12). . . . . . . . . . . . . . . . . . . . . . . . . Calaxius jenneri (Williams, 1974) 

— Carapace dorsal surface with distinct lateral, submedial, and medial carinae; lateral carinae extending anteriorly to coalesce and form rostral margin. Pereopod 1 major chela not massive or exceptionally setose. Telson 

with medial spine present on distal margin (Fig. 13). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Figure 12

Calaxius jenneri: (A) lateral view of whole animal; (B) pereopod 2; (C) dorsal view of anterior carapace; (D) pereopod 1 

major (right) chela; (E) telson and uropods. A–E: modified from Williams, 1984.

Figure 13

(A) Axius armatus: dorsal view of anterior carapace; (B) Paraxiopsis gracilimana: dorsal view of anterior carapace; (C) Axius 

armatus: pereopod 1 major chela; (D) Paraxiopsis gracilimana: pereopod 1 major chela; (E) Paraxiopsis gracilimana: telson and 

uropods; (F) Axius armatus: telson and uropods. A, C, F: modified from Kensley, 2001; B, D, E: modified from Kensley, 1996.
11

7. Carapace medial carina lacking spines; submedial carinae each with distinct spine anteriorly; each lateral 

carina with an anterior spine (situated posterior to, and merging with, the rostrum); rostrum with 4–6 progressively smaller lateral spines and associated setae, anterior half smooth or with inditinct spines and small setae 

(Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paraxiopsis gracilimana Kensley, 1996

— Carapace medial carina armed anteriorly with 1–3 spines; submedial carinae lacking spines; lateral 

carinae lacking spines; rostrum with 3–6 pairs of distinct similarly sized lateral spines, continuing to tip 

(Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axius armatus S. I. Smith, 1881

Figure 14

Paraxiopsis gracilimana: (A) lateral view of whole animal; (B) pereopod 1 (chela); (C) dorsal view of anterior carapace; (D) 

telson and uropods. A–D: modified from Kensley, 1996.

Figure 15

Axius armatus: (A) lateral view of carapace; (B) telson and uropods; (C) dorsal view of anterior carapace; (D) pereopod 1 

major chela; (E) pereopod 1 minor chela. A–E: modified from Kensley, 2001.
12

8. Pereopod 3 propodus distal margin with single spiniform seta. Maxilla 2 with long distinctive seta on posterior lobe of scaphognathite. Uropodal exopod with broadly rounded margin, not appearing bilobed. Family 

Thomassiniidae (Fig. 16). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crosniera wennerae Heard and King, 2007

— Pereopod 3 propodus distal margin lacking spiniform seta. Maxilla 2 lacking long distinctive seta on posterior 

lobe of scaphognathite. Uropodal exopod appearing bilobed, with subdistal fringe of setae. Family Callianassidae (Fig. 17). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Figure 16

Crosniera wennerae: (A) lateral view of carapace; (B) uropods and telson; (C) pereopod 3 showing spiniform seta including 

enlargement with fine setae removed; (D) maxilla 2 with long seta. A–D: original illustrations by R.W. Heard. 

Figure 17

(A) Biffarius biformis: pereopod 3; (B) Gilvossius setimanus: telson and uropods; (C) Cheramus marginatus: telson and uropods; 

(D) Callichirus major: telson and uropods; (E) Biffarius biformis: maxilla 2 without seta. A, E: original illustrations by R.W. 

Heard; B, D: modified from Williams, 1984; C: modified from Biffar, 1971b.
13

9. Uropodal exopod at least twice as long as wide and nearly twice as long as telson; subdistal plates indistinct, appearing absent, represented by row of stout spiniform setae immediately behind distal fringe of plumose setae 

on distal upper margin. Pleopods 3–5 with finger-like appendices internae on endopod. Pereopod 1 (major 

chela) merus lower margin convex, smooth, lacking hook-like process or small spines. Subfamily Cheraminae 

(Fig. 18). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cheramus marginatus (Rathbun, 1901)

— Uropodal exopod distinctly less than twice as long as wide; subdistal plates distinct, with distal fringe of setae. 

Pleopods 3–5 having short stubby-like or embedded appendices internae on endopod. Pereopod 1 (major chela) 

merus lower margin with hook-like process or spine. Subfamilies Callianassinae and Callichirinae) (Fig. 19). . . 10

Figure 18

Cheramus marginatus: (A) dorsal view of carapace; (B) male pleopod 3 with appendix interna; (C) pereopod 1 major chela; 

(D) telson and uropods; (E) pereopod 1 minor chela. A–C: modified from Manning and Felder, 1991a; D–E: modified 

from Biffar, 1971b.

Figure 19

(A) Callichirus major: telson and uropods; (B) Necallianassa berylae: telson and uropods; (C) Biffarius biformis: telson and 

uropods; (D) tip of pleopod showing embedded appendix interna for Callichirus species; (E) tip of pleopod 3 showing stubby 

appendix interna for Callianassa species; (F) Biffarius biformis: pereopod 1 major chela; (G) Callichirus major : pereopod 1 

major chela. A, C, F, G: modified from Williams, 1984; B: modified from Heard and Manning, 1998; D, E: modified from 

Manning and Felder, 1991a.
14

10. Uropodal endopod lateral margin with one large spine. Telson lateral margins with 2 spines (Fig. 20). 

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Necallianassa berylae Heard and Manning, 1998 

— Uropodal endopod lateral margin without spines. Telson lateral margins without spines (Fig. 21). . . . . . 11

Figure 20

Necallianassa berylae: (A) dorsal view of anterior carapace; (B) lateral view of anterior carapace; (C) pereopod 1 major chela 

of male; (D) pereopod 1 minor chela of male (similar to major chela of female); (E) telson and uropods; (F) pereopod 1 

minor chela of female. A–F: modified from Heard and Manning, 1998.

Figure 21

Telson and uropods: (A) Biffarius biformis; (B) Gilvossius setimanus; (C) Callichirus major. A–C: modified from Williams, 

1984.
15

Figure 22

Callichirus major: (A) dorsal view of carapace: (B) pereopod 1 major (right) chela; (C) telson and uropods. A–C: modified 

from Williams, 1984.

11. Uropodal endopod narrow, length four times width. Telson with distinct medial groove (Fig. 22).

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callichirus major (Say, 1818)

— Uropodal endopod rounded, length less that two times width. Telson lacking distinct medial groove 

(Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Figure 23

Telson and uropods: (A) Gilvossius setimanus; (B) Biffarius biformis. A, B: modified from Williams, 1984.
16

12. Eyestalks flattened and pointed, curving laterally at apex. Telson posterior margin with small acute spine medially (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gilvossius setimanus (DeKay, 1844) 

Figure 24

Gilvossius setimanus: (A) dorsal view of carapace; (B) pereopod 1 major (right) chela; (C) pereopod 1 minor (left) chela; 

(D) telson and uropods, with enlargement of spine on telson. A–D: modified from Williams, 1984.

— Eyestalks rounded at apex. Telson posterior margin lacking spine (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . 13

Figure 25

(A) Biffarius cf. fragilis: dorsal view of anterior carapace showing rounded eye stalks; (B) Biffarius cf. fragilis: telson and 

uropods; (C) Biffarius biformis: telson and uropods. A: original illustration by R. W. Heard; B: modified from Biffar, 1971b; 

C: modified from Manning and Felder, 1991a. 
17

13. Antenna 1 peduncle distinctly shorter than antenna 2 peduncle. Pereopod 1 minor chela of male similar in size 

and shape to major and minor chelae of female, merus with weak hook-like spine; carpus about three times 

longer than wide at greatest width. Telson posterolateral corners with two, small, short spiniform setae; posterior 

margin fringed with row of closely spaced, plumose setae (Fig. 26) . . . . . . . . . .Biffarius biformis (Biffar, 1971a)

Figure 26

Biffarius biformis: (A) dorsal view of carapace; (B) pereopod 1 major chela; (C) pereopod 1 minor chela; (D) telson, with 

enlargement of posterolateral corner of telson. A: modifed from Williams, 1984; B, C: modified from Manning and Felder, 

1991a; D: original illustration by R. W. Heard.

— Antenna 1 peduncle equal to, or slightly longer than, antenna 2 peduncle. Pereopod 1 major chela of female 

distinctly larger and more developed than minor chela; minor chela of male and female markedly elongate; 

merus lacking weak hook-like spine; carpus narrow, 5–6 times longer than greatest width. Telson posterolateral corners lacking setae or with a single, short, spiniform seta; posterior margin sparsely setose (Fig. 27).

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biffarius cf. fragilis (Biffar, 1970)

Figure 27

Biffarius cf. fragilis: (A) dorsal view of anterior carapace; (B) pereopod 1 major chela; (C) pereopod 1 minor chela; 

(D) telson and uropods, with enlargement of posterolateral corner of telson. A and enlargement of telson: original 

illustrations by R. W. Heard; B–D modified from Biffar, 1971b.
18

Synopsis of Thalassinidea known from 

the South Atlantic Bight

INFRAORDER THALASSINIDEA Latreille, 1831

SUPERFAMILY AXIOIDEA Huxley, 1879

Family Axiidae Huxley, 1879

Acanthaxius Sakai and de Saint Laurent, 1989

Acanthaxius hirsutimanus (Boesch and Smalley, 1972)

(Fig. 10)

Calocaris (Calastacus) hirsutimana Boesch and Smalley, 

1972: 45–52. 

Axiopsis hirsutimana.—Williams, 1984: 187.

Acanthaxius hirsutimanus.—Sakai and de Saint Laurent, 1989: 73.

Material examined. SERTC #S780, one female, off Tybee 

Island, GA, 32°00.0ʹN, 80°11.6ʹW, 137 m, coll. MARMAP, 

5 April 1975. ULLZ #1474, one female, 40 miles SSW of 

Grand Isle, LA, 40 fathoms, coll. D.L. Felder, July 1975, 

identification by D.L. Felder.

Diagnosis. (Modified from Boesch and Smalley, 1972.) 

Carapace with five gastric carinae (one medial, two 

submedial, and two lateral); medial carina with eight 

spines; submedial carinae with five spines; lateral carinae with six or seven spines; medial carina continues 

posteriorly for length of carapace after short trench 

extending from cervical groove. Eyes with eyestalks well 

developed and overreaching rostrum. First pereopods 

(chelae) unequal in size, robust, heavily setose. Uropodal exopods broad, sub equal in length to telson.

Known range. Gulf of Mexico, tropical Atlantic, off 

Guyana, Costa Rica (Boesch and Smalley, 1972; Vargas 

and Cortés, 1999), South Atlantic Bight (Wenner and 

Read, 1982), off Santee Rivers, SC (Wenner et al., 1984), 

off Kiawah Island, SC (Wenner et al., 1984), off Tybee 

Island, GA (present study). 

Habitat. Soft bottom substrata, 11 to 137 m.

Remarks. This species was originally described based 

on specimens from off northeastern South America 

and the northeast Gulf of Mexico (Boesch and Smalley, 

1972). The first SAB record of the species was reported 

by Wenner and Read (1982) in a study of the shelf and 

upper slope Crustacea. During the present investigation a single female specimen was found in the marine 

invertebrate collection of the College of Charleston’s 

Grice Marine Laboratory (GML) from off Tybee Island, 

Georgia.

There appear to be distinctive differences, especially 

in size, shape, and ornamentation of the first pair of 

chelae, between the specimen of A. hirsutimanus from 

Georgia and the original description. These differences 

could be ecophenotypic, the result of re-growth after 

the individual was damaged, or representative of specific differences. Further study of additional material is 

needed before any conclusions can be made.

Axius Leach, 1815

Axius armatus S. I. Smith, 1881

(Fig. 15)

 Axius armatus S. I. Smith 1881: 433.

Material examined. SERTC #S783, two females, off St. 

Helena, SC, 31°41ʹN, 79°25ʹW, 232 m, coll. R/V Dolphin, 

21 September 1976. USNM 174450, one ovig. female, 

off South Carolina, 32°30ʹN, 78°29ʹW, Sta. 2G, 218 m, 

coll. by Texas Instruments, Inc. for BLM/MMS, SABP, 

20 November 1977. USNM 174451, one male, off South 

Carolina, 32°30ʹN, 78°29ʹW, Sta. 2G, 218 m, box corer, 

coll. by Texas Instruments, Inc. BLM/MMS, SABP, 12 

May 1977. 

Diagnosis. (Modified from Kensley, 2001.) Carapace 

with submedial carina unarmed, without supraocular 

spines. Rostrum longer than eyes, with 3–6 pairs of 

spines along lateral edges. Antenna 2 with scaphocerite 

well developed, simple. Pereopod 1 major chela fixed 

finger of propodus with single spine; merus armed with 

3–4 ventral spines, dorsal margin strongly carinate; 

minor chela lacking spine on fixed finger of propodus; 

merus dorsal margin with single spine, ventral margin 

with 4 spines. Pereopod 2 merus ventral margin setose, 

with 4 spines. Pereopod 4 sternal plate with lateral 

flange rounded. Telson with 4–6 dorsal spines at mid 

length. Uropodal exopod transverse suture armed with 

7–8 spines. 

Known range. U.S. Atlantic Seaboard from Massachusetts to South Carolina (Kensley, 2001; present study).

Habitat. Continental shelf, 108–260 m.

Remarks. Prior to this report, this species was known 

from only two records in the SAB from off the coast of 

South Carolina (Kensley, 2001). There are few records 

of this species in the literature, all of which are female 

(Kensley, 2001). Two specimens from off St. Helena, SC 

examined in this study (SERTC S#783) represent the 

third report of this species in the SAB.

Axius armatus appears to be primarily a resident of 

the waters off the northeast U.S. coast (Kensley, 2001). 

In the northern portions of its range, it is sympatric 
19

with the morphologically similar and apparently closely 

related species Axius serratus Stimpson, 1852. For more 

information on these two species refer to Kensley 

(2001).

Calaxius Sakai and de Saint Laurent, 1989

Calaxius jenneri (Williams, 1974)

(Fig. 12)

Calocaris (Calastacus) jenneri Williams, 1974: 451.

Axiopsis jenneri.—Williams, 1984: 185.

Calaxius jenneri.—Sakai and de Saint Laurent, 1989: 

86. 

Material examined. SERTC #S778, 10 specimens, off 

Charleston, SC, 32°06.5ʹN, 79°12.5ʹW, 79 m, coll. by 

GML staff, 23 April 1977, housed within the College of 

Charleston’s Grice Marine Laboratory marine invertebrate collection. USNM 174448, one male specimen, off 

Sapelo Island, GA, 30°57.08ʹN, 79°57.85ʹW, 62 m, coll. 

by Texas Instruments, Inc. for BLM/MMS, SABP, 30 August 1977. USNM 214904, one specimen, off Savannah, 

GA, 31°32.0ʹN, 79°44.4ʹW, 56 m, coll. by South Carolina 

Marine Resources for BLM/MMS, R/V Oregon, Sta. 

OS01, fly net, 10 March 1981. 

Diagnosis. (Modified from Williams, 1974.) Carapace 

nearly smooth, with scattered tufts of setae; lateral carinae with a small spine, sub medial carinae with four 

spines. Rostrum acute, extending slightly beyond eyes; 

tip slightly upturned with 1–3 spines on lateral margins. Eyes with cyclindrical stalks; cornea rounded, well 

developed, pigmented. Antenna 2 with scaphocerite 

long and slender. First pereopods (major and minor) 

sub equal, asymmetrical, densely setose. Major chela 

strongly developed, fingers with distinct gap proximally; 

propodus (including fixed finger) about same length as 

carapace, fixed finger with spine; dactylus (movable finger) with distally inflated cutting edge. Telson rounded 

posteriorly; posterior margin fringed with setae, lacking 

posterio-medial spine. Uropodal exopod with transverse 

distal suture with short, stout spines.

Known range. Southwest of Cape Lookout, NC (Williams, 1974), South Atlantic Bight (Wenner and Read, 

1982), and off Savannah River, GA (Wenner et al., 1984, 

as Axiopsis hirsutimana; present study).

Habitat. Deep continental shelf and upper slope, 62 to 

232 m (Williams, 1974; Wenner and Read, 1982; present study).

Remarks. This species was transferred to the new genus 

Calaxius by Sakai and de St. Laurent (1989), which then 

contained eight species. Kensley and Hickman (2001) 

subsequently recognized 10 species and divided the 

genus into two groups: species “with strongly triangular 

and ventrally acute abdominal pleura,” and “those with 

ventrally rounded pleura.” Felder and Kensley (2004) 

added an eleventh species to this genus based on material from chemosynthetic (seep) communities off the 

Louisiana continental slope.

This species was previously recorded in the literature solely from North Carolina (Williams, 1974; Williams, 1984). Wenner and Read (1982) recorded two 

specimens from 232 m in the South Atlantic Bight that 

they identified as “Calocaris jenneri” but we have been 

unable to locate the material to confirm the identification. Wenner et al. (1984) recorded one specimen of 

“Axiopsis hirsutimana” from 56 m, near the shelf edge off 

Savannah, Georgia, that we found in the USNM collections (USNM 214904) and re-identified as Calaxius 

jenneri. In our examination, 10 additional specimens 

from South Carolina of C. jenneri were located in the 

collections of the GML.

Paraxiopsis De Man, 1905

Paraxiopsis gracilimana Kensley, 1996

(Fig. 14)

Paraxiopsis gracilimana Kensley, 1996: 719–722, 

figs. 6, 7.

Material examined. USNM 169669 (Holotype), female, 

Bonaire Harbor, Netherlands Antilles, 2 m, coll. R/V 

Harrison, 25 August 1976. USNM 211454 (Paratypes) 

one female, one juvenile, Looe Key Reef, FL, sta FLK21, 6 m, coll. B. Kensley and M. Schotte, 29 September 1982. USNM 174452, off South Carolina, 3245N, 

7856W, Sta 2D, 27 m, coll. by Texas Instruments, Inc. 

for BLM/MMS, SABP, 25 August 1977.

Diagnosis. (Modified from Kensley, 1996.) Carapace 

with surface smooth, lateral carina more or less continuous with lateral rostral margin, armed with a single pair 

of anterior spines, submedial carina with a single pair 

of anterior spines, medial carina lacking spines; rostrum 

extending beyond eyes, with four pairs of lateral spines 

(each with associated seta) that get progressively smaller 

(in smaller specimens there may also be 1–2 pairs of 

very small spines towards the rostrum tip). Abdominal 

pleura ventrally rounded, with small spine on anterior 

margin of pleura 3–5. Telson lateral margins with four 

pairs of spines and a pair of distal robust setae, dorsal 

surface with three pairs of spines, distal margin with 

small medial spine.

Known range. South Carolina southward and westward 

to Gulf of Mexico; throughout Caribbean as far south 

as Tobago (Kensley, 1996).
20

Habitat. Live bottom habitats, 1 to 27 m (Kensley, 1996).

Remarks. This species is known in the SAB from a single 

record off South Carolina (Kensley, 1996). Examination 

of the South Carolina specimen led to questions regarding the rostrum and maxilliped morphology that could 

not be answered in Kensley’s (1996) description and it 

was apparent that the types for this species would need 

to be examined. Subsequently, it was found that smaller 

specimens have 1–2 pairs of much smaller spines on the 

anterior portion of the rostrum, in addition to the four 

pairs of spines described. Also, on maxilliped 3 the mesial edge of the merus possesses around 17 spines along 

its entire length. While the spines on the mesial edge of 

maxilliped 3 were not illustrated by Kensley (1996), it was 

given as a general diagnostic character for the genus.

Paraxiopsis gracilimana is the type-host for the bopyrid 

isopod, Gigantione uberlackerae Adkison, the type locality of 

which is the northern Gulf of Mexico (Adkison, 1984). At 

present this isopod parasite is not known from the SAB. 

Family Calocarididae Ortmann, 1891

Calocaris Bell, 1853

Calocaris templemani Squires, 1965

(Fig. 8)

Calocaris templemani Squires, 1965: 2, figs. 1, 2A, 2B, 

CtC, CtD, 3Ct, 4, 5Ct, 6.

Material examined. USNM 294422, one female, off 

Cape Lookout, NC, 34°14.86ʹN, 075°43.78ʹW, 1013 

m, R/V Columbus Iselin, coll. by Battelle New England 

Marine Lab. for BLM/MMS, ASLAR, Sta. SA1:02, box 

corer, Core 7, 12 Nov 1983. 

Diagnosis. (Modified from Williams, 1984.) Carapace having medial carina extending full length, continuing on to 

rostrum. Eyes with pigment indistinct (diffuse or appearing absent), eyestalk rudimentary, flattened obliquely on 

distal surface. Antenna 2 peduncular article 2 with small 

lateral spine and lateral scale reduced to a short spine. 

Telson rounded distally, lateral margin with a small pair of 

anterior spines. Uropodal exopod with a weakly defined, 

partially fused distal suture, suture lacking spines. 

Known range. Newfoundland to Virginia; off Cape 

Lookout, NC (Squires, 1965; Williams, 1984).

Habitat. Soft sediments, 200–1013 m (Williams, 1984).

Remarks. Although the single specimen from off Cape 

Lookout represents the known southern limit for this 

species, it is possible that this species occurs in deeper 

waters further south (Williams, 1984).

SUPERFAMILY CALLIANASSOIDEA Dana, 1852

 Family Callianassidae Dana, 1852

 Subfamily Callianassinae Dana, 1852

Biffarius Manning and Felder, 1991a

Biffarius biformis (Biffar, 1971a)

(Fig. 26)

Callianassa biformis Biffar, 1971a: 225, fig. 1. 

Biffarius biformis.—Manning and Felder, 1991a: 769.

Material examined. SERTC #S777, one male, six females, Edisto Beach, near entrance to Big Bay Creek, 

SC, 32°29.6ʹN, 80°20.7ʹW, intertidal, coll. by GML staff,

20 August 1972. SERTC #S1072, one male, one ovigerous female, two undetermined, on protected beach 

near Cummings Point, Morris Island, SC, 32°44.48ʹN, 

79°52.38ʹW, intertidal, coll. By R. Heard, 13 May 

2004. USNM 42401 (labeled as Gilvossius setimanus) 

off Beaufort, NC. USNM 150246, two female, three 

males, Sapelo Island Beach, GA, intertidal, coll. by 

M. Gray, 9 February 1961. USNM 155371, 10 females, 

16 males, south end of Sapelo Island, GA, intertidal, 

coll. by R. Heard, 6 April 1969. USNM 266218, 50+ 

males and females, north end of Saint Catherines Island, GA, intertidal, coll. by GA Bishop, yabby pump, 

1 June 1988. 

Diagnosis. (Modified from Biffar, 1971a.) Antenna 1 

peduncle shorter than antenna 2 peduncle. Eyes flattened proximally, rounded distally, cornea central; 

eyestalks shorter than first antennular segment. In 

males, the first pair of chelae are greatly unequal in 

size, with two forms of major chelae (a “strong” form 

and a “weak” form); ischium dorsal margin with small 

spur-like projection; merus dorsal margin with fringe 

of setae, ventral margin with hook-like projection. In 

females, the first pair of chelae are more or less symmetrical, resembling the minor chela of the male, not 

noticeably elongate; merus ventral margin with small 

curved hook-like projection; carpus narrower than 

propodus, length about three times greatest width. 

Telson distal margin bluntly rounded and lacking 

medial projection, lateral margins each with pair of 

small distal spiniform setae present. Uropodal exopod 

irregularly shaped, basically bilobed distally; endopod 

rounded.

Known range. Massachusetts to Georgia (Sapelo Island); northern Gulf of Mexico, including Bald Point, 

Franklin County, FL; Louisiana, Corpus Christi, TX, and 

the south Texas outer continental shelf (Biffar, 1971a; 

Rabalais et al., 1981).
21

Habitat. Lower intertidal in the vicinity of the mouths of 

sounds, bays, and inlets to about 10 m depths offshore 

in fine to moderate sand and sand-silt substrata; salinity 

10–30‰ (Biffar, 1971a).

Remarks. In the SAB, Biffarius biformis has been reported from both lower intertidal (Biffar, 1971a) and 

subtidal habitats (Dörges, 1972, 1977; Frankenberg 

and Lieper, 1977; Prezant et al., 2002; Van Dolah et 

al.1; Jutte and Van Dolah2; Jutte et al.3; Van Dolah et 

al.4). In past studies this species has been confused with 

juveniles of C. atlantica (=Gilvossius setimanus) (Frey and 

Howard, 1969; Biffar, 1971a). Populations are known 

to occur seaward to depths of 7–8 m over 3 km off the 

mouth of Doboy Sound, Georgia (Smith, 1971). Dörges 

(1977) considered it one of the dominant organisms 

in high salinity estuaries and the inner shelf outside 

the wave zone in Georgia. One population with densities exceeding 50 individuals per m2

 was discovered 

on the north shore of Tybee Inlet, near the mouth on 

the south end of Tybee Island, Georgia (Heard, pers. 

observ.). See Frey and Howard (1969) for a description 

of B. biformis burrows.

Biffarius biformis has been reported off the Texas coast 

in depths of 10–15 m (Rabalais et al., 1981); however, 

the specific status of the Texas and other northern Gulf 

of Mexico populations needs to be verified by analysis of 

morphological and molecular data (Felder5).

Biffarius cf. fragilis (Biffar, 1970)

(Fig. 27)

?Callianassa fragilis Biffar, 1970: 45, fig. 3.

?Biffarius fragilis.—Manning and Felder, 1991a: 769.

Material examined. USNM 174404, 1 male, off South 

Carolina, 32°54.1ʹN, 79°11.98ʹW, 17 m, TI/MMS, Sta. 

2B, 17 November 1977.

Diagnosis. (Modified from Biffar, 1970.) Eyestalks with 

apices rounded (lacking acute, elongate tips). Antenna 

1 peduncle equal to or extending slightly beyond antenna 2 peducle. Third maxilliped with ischium and 

merus subquadrate, longer than wide. Major chela of 

male and female with hook-like projection on inner 

margin of merus; major chela of male much larger and 

strongly developed (nearly as long as entire body) than 

that of female. Minor chelae of both male and female 

similar, distinctly elongate with merus lacking hooklike spine; carpus narrow, 5–6 times greatest width. 

Pleopods 3–5 having short stubby-like appendices 

internae. Telson lacking distinct medial grove, each 

posterior lateral corner with single, short, spiniform 

seta; posterior margin sparsely setose, lacking medial 

projection. Uropodal endopod rounded, length less 

that two times width. 

Known range. South Carolina; Florida; Puerto Rico; Antigua; Venezuela (Biffar 1970, 1971b; present study).

Habitat. Intertidal to shallow subtidal; sand, clay or mud 

substrates (Biffar, 1970).

Remarks. The single damaged specimen from South 

Carolina is similar in most respects to the description 

of the species given by Biffar (1970, 1971b); however, 

it differs slightly in the dentition of the cutting edges 

of the major chela and the presence of a single, short 

spiniform seta on each posterior lateral margin of the 

telson. The presence of this spiniform seta was not 

noted by Biffar (1970, 1971b) and may represent a 

variable characters or evidence for a yet undescribed 

species.

Gilvossius Manning and Felder, 1991b

Gilvossius setimanus (DeKay, 1844)

(Fig. 24)

Gonodactylus setimanus DeKay, 1844: 34. 

Callianassa stimpsoni S. I. Smith, 1873: 549, pl. 2, 

fig. 8.

Callianassa atlantica Rathbun, 1926: 107.

Callianassa setimanus.—Manning, 1987: 388.

Gilvossius setimanus.—Manning and Felder, 1991b: 558.

1

 Van Dolah, R. F., R. M. Martore, A. E. Lynch, M. V. Levisen, P. H. 

Wendt, D. J. Whitaker, and W. D. Anderson. 1994. Environmental 

evaluation of the Folly Beach nourishment project. Final Report to 

the U.S. Army Corps of Engineers, Charleston District. Charleston, 

SC. 100 p. and appendices. Marine Resources Research Institute, 

South Carolina Department of Natural Resources, P.O. Box 12559, 

Charleston, SC 29422.

2

 Jutte, P. C., and R. F. Van Dolah. 1999. An assessment of benthic 

infaunal assemblages and sediments in the Joiner Bank and Gaskins 

Banks borrow areas for the Hilton Head beach renourishment project. Final Report – Year 1 to Olsen Associates, Inc. and the Town 

of Hilton Head Island from the Marine Resources Division, South 

Carolina Department of Natural Resources. 33 p. Marine Resources 

Research Institute, South Carolina Department of Natural Resources, PO Box 12559, Charleston, SC 29422.

3

 Jutte, P. C., R. F. Van Dolah, G. W. Eason, Jr., and M. V. Levisen. 

1999. An assessment of benthic infaunal assemblages and sediments 

in the vicinity of the Port Royal Ocean Dredged Material Disposal 

Site. Final Report to the U. S. Army Corps of Engineers, Charleston 

District. Charleston, SC. 55 p. Marine Resources Research Institute, 

South Carolina Department of Natural Resources, PO Box 12559, 

Charleston, SC 29422.

4

 Van Dolah, R. F., P. C. Jutte, G. H. M. Riekirk, M. V. Levisen, L. 

E. Zimmerman, J. D. Jones, A. J. Lewitus, D. E. Chestnut, W. McDermott, D. Bearden, G. I. Scott, and M. H. Fulton. 2002. The 

condition of South Carolina’s estuarine and coastal habitats during 1999–2000: Technical Report No. 90. South Carolina Marine 

Resources Division. 132 p. Marine Resources Research Institute, 

South Carolina Department of Natural Resources, P.O. Box 12559, 

Charleston, SC 29422.

5

 Felder, D. Pers. commun. University of Louisiana at Lafayette, Department of Biology, PO Box 42451, Lafayette, LA 70504.
22

Material examined. USNM 51007, fishing grounds 

off Beaufort, NC, 34°22ʹN, 076°57ʹW, 29 m, R/V Fish 

Hawk, Sta. 7958, 7 Sept. 1913. USNM 174405, off 

South Carolina, 32°57.0ʹN, 79°17.0ʹW, 12.5 m, coll. 

by Texas Instruments, Inc. for BLM/MMS, SABP, Sta. 

2A, box corer, 12 February 1977. USNM 68386, one 

female, Wassaw Sound, Cabbage Island, Chatham 

County, GA, coll. by I. R. Tomkins, 01 April 1933.

Diagnosis. (Modified from Williams, 1984.) Antennular 

peduncle extending beyond distal tip of antenna 2 peduncle. Eye with cornea laterally positioned; eyestalks 

extending beyond first antennular segment, narrowing distally and curving outward. Pereopod 1 chelae 

unequal and showing sexual dimorphism; major chela 

merus ventral margin with strong hook-like projection 

in males (smaller in females); minor chela with carpus 

about as wide as propodus merus, ventral margin entire. 

Telson posterior margin bluntly rounded, with small 

medial spine. Uropodal exopod distally rounded, two 

distinct areas defined on dorsal surface by distal fringes; 

endopod distally rounded. 

Known range. Nova Scotia to southern Georgia (Williams, 1984; Manning, 1987).

Habitat. Lower intertidal and shallow subtidal to depths 

of 134 m, firm mud-sand substrates, salinity 25–32‰ 

(Frankenberg and Leiper, 1977; Rabalais et al., 1981; 

Williams, 1984; Van Dolah et al.6). Deeper records of 

“Callianassa atlantica” have been recorded within the 

South Atlantic Bight (collected at 218 m and 401 m and 

recorded by Tenore7 in the South Atlantic Benchmark 

Program 1977 Report) (Knott and Wendt8). Despite our 

efforts, we have not been able to locate these specimens 

for examination, therefore the maximum depth record 

for this species within the South Atlantic Bight remains 

at 134 m.

Remarks. Individuals of this species are intermediate in 

body length (59 mm males, 68 mm females) between 

Callichirus major (Say, 1818) (95 mm males, 80–92 mm 

females) and the smaller Biffarius biformis (32 mm 

males, 27 mm females). Gilvossius setimanus is easily 

distinguished from the superficially similar B. biformis 

by the shape of the third maxilliped, the tail fan, and 

the outwardly curved apices of the eyestalks. Gilvossius 

setimanus has been collected irregularly and may be 

more characteristically found subtidally in the higher 

salinity waters of the sounds and bays of the SAB; however, a well-established intertidal population was present 

on an exposed mud-sand bar with associated oysters on 

the Bull River side of Cabbage Island, Wassaw Sound, 

Georgia (Heard, pers. observ.).

During September and October of 1968, specimens 

were found in the stomachs of large specimens of the Atlantic stingray Dasyatis sabina (Lesueur) near the mouth 

of St. Catherines Sound (Heard, pers. observ.). An additional adult specimen was also found in the stomach of 

a large American stingray, D. americana Hildebrand and 

Schroeder, in the mouth of Calibogue Sound, South 

Carolina (Heard, pers. observ.). The species has also 

been found to be occasional prey of tomtate, Haemulon 

aurolineatum Cuvier 1830, and whitebone porgy, Calamus 

leucosteus Jordan and Gilbert 1885, in studies of food 

habits of fish on the continental shelf between Santee 

River, South Carolina and Jacksonville, Florida (MRRI)9.

The bopyrid isopod, Ione thompsoni Richardson, 1904, 

was originally described from the branchial chamber of 

G. setimanus in New England (Markham, 1995). This 

isopod parasite was found on specimens of G. setimanus

collected in the lower intertidal zone of Cabbage Island 

(Wassaw Sound), Georgia (Heard, pers. observ.) and it 

was found infesting this host in an intertidal area just 

south of St. Augustine, Florida (Maturo10).

Necallianassa Heard and Manning, 1998

Necallianassa berylae Heard and Manning, 1998

(Fig. 20)

Necallianassa berylae Heard and Manning, 1998: 884.

Material examined. USNM 260881 (Holotype), one 

male, off South Carolina, 32°00.95ʹN, 79°31.05ʹW, 

43 m, TI/MMS Pierce Sta. 9737, sample 0177-2 (3E), 

August 1977. USNM 260882 (Paratype), one female, 

6

 Van Dolah R. F., D. R. Calder, D. M. Knott, and M. S. Maclin. 1979. 

Effects of dredging and unconfined disposal of dredged material 

on macrobenthic communities in Sewee Bay, South Carolina. South 

Carolina Marine Resources Center Technical Report 39. 54 p. Marine Resources Research Institute, South Carolina Department of 

Natural Resources, P.O. Box 12559, Charleston, SC 29422.

7

 Tenore, K. R. 1978. Macroinfaunal benthos. In South Atlantic 

Benchmark Program 1977 Report, Volume 3, p. 255–288 and appendices. Prepared for the Bureau of Land Management, Outer 

Continental Shelf Office, New Orleans, LA by Texas Instruments, 

Inc., Dallas, TX, 440 p.

8

 Knott D. M., and P. H. Wendt. 1985. Special literature analysis study: 

Final report on benthic communities in certain slope areas of the 

South Atlantic Bight. Prepared for Minerals Management Service, 

Washington, D.C., Contract 14-12-0001-29185, OCS study MMS 85-

0051, 75 p. Marine Resources Research Institute, South Carolina 

Department of Natural Resources, P.O. Box 12559, Charleston, SC 

29422.

 9 Marine Resources Research Institute. 1984. Final Report, South 

Atlantic OCS Area Living Marine Resources Study, Phase III. Vol. 

II: Appendices. Prepared for Minerals Management Service, Washington, D. C., Contract 14-12-0001-29185. 82 p. Marine Resources 

Research Institute, South Carolina Department of Natural Resources, P.O. Box 12559, Charleston, SC 29422.

10 Maturo, F. J. S. 2006. Pers. commun. University of Florida, Box 

118525, Gainesville, FL 32611-8525.
23

off South Carolina, 32°00.95ʹN, 79°31.05ʹW, 43 m, TI/

MMS Pierce Sta. 9737, sample 0177-2 (3E), August 1977. 

USNM 260886 (Paratype), off Georgia, 31°33.63ʹN, 

79°39.02ʹW, 75 m, 6 August 1963. USNM 260887 (Paratype), off Georgia, 31°33.06ʹN, 79°40.35ʹW, 65 m, 6 

August 1963. USNM 260888 (Paratype), off Georgia, 

31°33.05ʹN, 79°40.63ʹW, 53 m, 6 August 1963. USNM 

174408, one specimen, off North Carolina, 32°12.00ʹN, 

77°36.00ʹW, 44 m, TI/MMS Sta. 1E, 10 May 1977; USNM 

174411, one specimen, off South Carolina, 31°27.00ʹN, 

79°46.00ʹW, 64 m, TI/MMS Sta. 4F, 28 August 1977.

Diagnosis. (Modified from Heard and Manning, 1998.) 

Rostrum acute, sharp, reaching about 3/4 the length of 

the eyestalks. Eyestalks dorso-ventrally flattened. Pereopod 1 chelae sexually dimorphic; major chela of males 

with ischium narrow, ventral margin crenulate; merus 

ventral margin with hook-like projection (projection 

smaller in females). Telson lateral margins with 1–2 

pairs of spines, distal margin with medial spine. Uropodal endopod with lateral spine.

Known range. Georgia, North Carolina, and South 

Carolina (Heard and Manning, 1998).

Habitat. Sub tidal soft sand substrata, depths of 35 to 

75 m (Heard and Manning, 1998).

Remarks. The species was described by Heard and Manning (1998) based on material from off Georgia and 

South Carolina. This species appears to be the western 

Atlantic cognate of Callianassa acanthura Caroli, from 

the eastern Atlantic waters of the Mediterranean. At 

the time N. berylae was described, two eastern Atlantic 

species were transferred to Necallianassa: Callianassa 

acanthura Caroli, 1946 and C. truncata Giard and Bonnier, 1890. However, in 1999, Sakai synonymized Necallianassa with Callianassa. Ngoc-Ho (2003) re-examined 

the group and declared Necallianassa a valid monotypic 

genus, containing only N. berylae, based on the large 

rostral spine, the lateral spines on the telson, the morphology of the uropods, and female with pleopod 2 

lacking an appendix interna. We follow Ngoc-Ho (2003), 

in accepting Necallianassa as a valid genus. 

 Subfamily Callichirinae Manning and 

 Felder, 1991a

Genus Callichirus Stimpson, 1866

Callichirus major (Say, 1818)

(Fig. 22)

Callianassa major Say, 1818: 238.

Callichirus major.—Hay and Shore, 1918: 407, pl. 29, 

fig. 10.

Material examined. SERTC #S779, one specimen, Folly 

Beach, Charleston, SC, 32°41.5ʹN, 79°53.4ʹW, intertidal, coll. by GML staff, 1 August 1985. SERTC #S1969, 

six males, five females, Folly Beach, Charleston, SC, 

32°38.49ʹN, 79°57.91ʹW, intertidal, coll. by SERTC staff, 

March 2005. USNM 78348, one female, one male, Edisto Beach, SC, coll. by H. G. Rutledge, 3 August 1937. 

USNM 150248, two females, Sapelo Island Beach, GA, 

intertidal, coll. by M. Gray, 20 November 1961. USNM 

266228, 45 females, 5 males, north beach of Saint Catherines Island, Georgia, intertidal, coll. by G. A. Bishop, 

yabby pump, 1988.

Diagnosis. (Modified from Williams, 1984.) Rostrum 

minute. Eyes with cornea reduced, near middle of lateral margin of ocular peduncles. Antennular peduncles 

approximately 2/3 as long as carapace, densely setose 

ventrally, terminating in two sub equal flagella. Chelae 

unequal, sexual dimorphism present; in major chela 

of males ischium and merus ventral margins granular; merus with ventral spine; carpus proximal ventral 

margin granular, dactylus hooked over fixed finger of 

propodus and with a proximal granular process on the 

ventral margin; the ischium, merus, and carpus ventral 

margins are smooth; merus does not have process on 

ventral margin uropods and telson lacking lateral processes; uropodal endopod narrow (approximately four 

times long as wide), with only terminal setae. 

Known range. Cape Lookout, North Carolina southward 

to Florida, the Gulf of Mexico; northern South America 

(Williams, 1984; Heard, pers. observ.).

Habitat. In the SAB this species occurs on open beaches, 

primarily in the intertidal zone (Dörges, 1972; KirbySmith and Gray, 1977; Prezant et al., 2002) but also in 

shallow subtidal depths (2–3 m). It occurs in salinities 

ranging from 15–36‰ and can tolerate salinities as low 

as 10‰ for short periods. It constructs deep burrows 

with some approaching depths of 2 m (Weimer and 

Hoyt, 1964; Heard, pers. observ.).

Remarks. This large callianassid is one of the most 

common, characteristic and bionomically important 

species on the open beaches of the SAB, but because 

it occurs in deep burrows C. major is seldom observed 

or collected. As with all callianassids, it is a permanent 

burrow dweller with the only indication of its presence 

being a small hole fringed with fecal pellets. Ovigerous 

females were observed in June and July near Beaufort, 

South Carolina (Pearse et al., 1942) and in August on 

Sapelo Island, Georgia (Heard, pers. observ.). The 

burrow morphology on Sapelo Island and surrounding areas have been studied and described by Weimer 

and Hoyt (1964) as tube shaped, from 1 to 2 meters 
24

in depth, with walls 1/4 of an inch thick, and bound 

together by a brown cementing agent consisting of 

callophanite (amorphous calcium phosphate). The 

fecal pellets of this organism were used in studies of 

the biological significance of coprophagy in estuarine 

animals by Frankenberg et al. (1967) and Frankenberg 

and Smith (1967).

Two commensal crustaceans are often associated 

with C. major; a bright orange copepod, Clausidium 

dissimile Wilson, 1921, which is found on the body 

surface, branchial cavity, and burrow wall; and a small 

elongate, pinnotherid crab Austinixa cristata (Rathbun), 

which occurs in the upper constricted part of the burrows (Williams, 1984; Heard, pers. observ.). The larval 

stage of the tapeworm Prochristianella hispida (Linton, 

1890) occurs in the digestive gland of C. major where 

its oval-shaped whitish-colored blastocysts can often be 

observed through the translucent exoskeleton of its 

host. The adult stage of this marine tapeworm matures 

in the spiral valve of the blunt nose stingray, Dasyatis sayi 

(Lesueur), which represents one of the few predators 

capable of extracting the C. major host from its burrows 

(Overstreet, 1983; Heard, pers. observ.).

 Subfamily Cheraminae Manning and 

 Felder, 1991a

Cheramus marginatus (Rathbun, 1901) 

(Fig. 18)

Callianassa marginata Rathbun, 1901: 92, fig 15.—Borradaile, 1903: 547.—Biffar, 1971b: 689–694, figs 15, 16.

Cheramus marginatus.—Manning and Felder, 1991a: 

780, fig. 14.

Material examined. USNM 174406, one ovigerous female, off the coast of Georgia, 30°53.98ʹN, 79°43.96ʹW, 

383 m, coll. by TI/MMS, 17 May 1977, Id by: B. Boothe. 

USNM 174407, one specimen, off Florida, 29°35.98ʹN, 

80°11.00ʹW, 241 m, coll. by TI/MMS, Sta.7E, 20 May 

1977.

Diagnosis. (Modified from Manning and Felder, 1991a.) 

Rostrum acute, reaching beyond eyes. Eyes with cornea 

lateral, subterminal. Pereopod 1 major chela ischium inner margin with short spines; merus with inner margin 

unarmed. Pleopods 3–5 with finger-like appendices internae. Telson distal margin bilobed, with medial spine, 

with 3 robust seta on the distolateral margins. Uropodal 

exopod subquadrate, more than twice as long as wide 

and nearly twice as long as telson.

Known range. Puerto Rico, Mexico, Texas, southeast 

Florida and Georgia (Rathbun, 1901; Biffar, 1971b; 

present study).

Habitat. Soft substrata on the deep shelf and upper 

slope, 40 to 383 m. 

Remarks. This species was originally described as Callianassa marginata by Rathbun (1901) based on specimens 

from Puerto Rico, from depths of 40 m to more than 314 

m (Rathbun, 1901). In 1991, Manning and Felder transferred this species to the genus Cheramus Bate, 1888. 

Confirmation of the presence of this species within 

the SAB has been elusive. Biffar (1971b) reported the 

species from off Miami Beach. Abele and Kim (1986) 

included the species (as Callianassa marginata) in their 

illustrated key to the marine decapod crustaceans of 

Florida, but did not indicate the species distribution. 

The two specimens examined from the USNM collections constitute a documented range extension of the 

species.

We compared the ovigerous female from off Georgia 

with specimens collected off Puerto Rico, which were 

made available to us by Barry A. Vittor & Associates 

(SERTC #S2805). Several differences were observed 

including (1) differences in type, size, and number 

of setae on the uropods and telson, (2) the shape 

and length/width ratios of the uropodal endopods 

and exopods, and (3) the shape and dentition of the 

chelae. One of the more distinctive differences was the 

length/width ratios of the uropodal exopods (Georgia 

length/width: 1.8 vs. Puerto Rico length/width: 2.5). 

Whether or not these differences are clinal or indicate 

the possibility of a cryptic species being present in 

the SAB can only be resolved by study of additional 

specimens from the waters off Georgia and possibly 

the Carolinas.

 Family Laomediidae Borradaile, 1903

Naushonia Kingsley, 1897

Naushonia crangonoides Kingsley, 1897

(Fig. 2)

Naushonia crangonoides Kingsley, 1897: 96, pl. 3, figs. 

8–10.—Goy and Provenzano, 1979: 339–359, figs 1–8.

Material examined. USNM 170634, one male, two females, off Cobb Island, Virginia, 37°15ʹN, 075°40ʹW, 14 

m, Sta. 66, R/V Delaware II, trawl, 2 October 1977.

Diagnosis. (Modified from Williams, 1984.) Carapace 

cylindrical. Rostrum flattened, broadly rounded. Eyes 

not visible from above; pigment reduced. First leg 

strongly subchelate; dactyl articulated to fold back 

against the distoventral margin of the elongate palm. 

Second leg simple, not chelate. Uropod with distinct 

suture on both endopod and exopod. Telson broadly 

rounded.
25

Known range. Newfoundland to Gulf of Maine; off Cape 

Lookout, NC (Williams, 1984).

Habitat. Soft bottom in sand and mud, near shore to 

14 m (Williams, 1984).

Remarks. This species is represented in the SAB by a 

single post-larva collected from Bogue Sound in North 

Carolina in a surface plankton sample identified by Williams (1984). The present authors did not examine this 

specimen; nonetheless the species is included within 

this guide given that it may be present in the northern 

region of the SAB.

 Family Thomassiniidae de Saint Laurent, 1979

Crosniera wennerae Heard and King, 2007 

(Fig. 16)

Material examined. USNM 174403, one male, off North 

Carolina, 33°00.96ʹN, 77°20.20ʹW, 318 m, TI/MMS Station 1F, 10 February 1977.

Diagnosis. (Modified from Heard and King, 2007.) Female: Carapace with linea thalassinica not extending into 

posterior half; rostrum short, acutely tipped, extending 

to, or just short of tips of eyes. Eye stalks with subacute tip, 

dorsally rounded and flattened ventrally; eyes poorly developed, each with two ommatidia and some defuse pigment. Maxilla 2 posterior lobe of scaphognathite bearing 

single long distinctive seta. Maxilliped 2 and 3 each with 

well-developed exopod, extending beyond articulation 

of merus and propodus. Maxilliped 3 lacking spine on 

distal, flexor margin of merus. Pereopod 1 (first chelae) 

sub equal (left chela slightly more robust); merus lower 

margin convex, smooth except for few small simple setae; 

left (major) chela with cutting edge of fixed finger bearing strong blunt subdistal spine and small proximal spine 

near articulation with movable finger; right (minor) chela 

with fixed finger having single, relatively small, low spine 

midway along cutting edge. Pereopod 3 with propodus 

having distal margin armed with single spiniform seta. 

Uropods broadly rounded; exopod not bilobed, entire 

(no lateral spine). Telson dorsal surface smooth with shallow lobes proximally on each margin, posterior margin 

broadly rounded and fringed with plumose setae.

Known range. Collected from off North Carolina.

Habitat. The single nonovigerous female specimen was 

obtained at a depth of 318 m in soft substratum on the 

continental slope.

Remarks. Crosniera wennerae is distinguished from the 

other four nominal species of the genus by having maxillipeds 2 and 3 with relatively long exopods that extend 

well past the merus of the endopod. It appears to have 

closer affinities with C. corindon Poore, 1997, a western 

Pacific species known only from Indonesia (Poore, 

1997), than with C. minima, the only other member of 

the genus currently known from the western Atlantic. 

Crosniera wennerae differs from C. minima in several characters, including: (1) the shape and dentition of the 

first chelae, (2) the absence of a lateral spine on both 

the uropodal endopod and exopod, (3) the absence of 

one or two submarginal, elongate, spiniform setae near 

distal margin of each uropodal endopod, and (4) the 

absence of lateral spines on the margins of the telson. 

 Family Upogebiidae Borradaile, 1903

Upogebia Leach, 1814

Upogebia affinis (Say, 1818)

(Fig. 6)

Gebia affinis Say, 1818: 241.

Upogebia affinis.—Stebbing, 1893: 185.

Material examined. SERTC #S611, two females, Southern Bass Hole Bay, north of Cooks Creek, SC, intertidal, 

coll. by GML staff, 13 September 1970. SERTC #S776, 

seven specimens, tide pool, on west side of Wood Creek, 

SC, 33°18.2ʹN, 79°10.6ʹW, intertidal, coll. by GML staff, 

21 July 1970. SERTC #S781, two females, north bank 

of Crabhaul Creek, North Inlet, Georgetown, SC, 

33°21.0ʹN, 79°11.3ʹW, intertidal, coll. by GML staff, 7 

July 1971. SERTC #S782, one female, east bank of Old 

Man Creek, 500 m north of Town Creek, SC, 32°20.5ʹN, 

79°10.5ʹW, 0–4 m, coll. by GML staff, 18 August 1970. 

SERTC #S784, one female, Charleston Harbor, SC, 

32°45.0ʹN, 79°51.6ʹW, 12–18 m, coll. by GML staff, 17 

May 1974. USNM 174414, one female, off Georgia, 

31°13.02ʹN, 81°13.05ʹW, Sta. 5A, 11 m, box corer, coll. 

by Texas Instruments, Inc. for BLM/MMS, SABP, 21 

February 1977. USNM 174415, two females, off Florida, 

29°27ʹN, 081°03ʹW, 20 m, coll. by Texas Instruments, 

Inc. for BLM/MMS, SABP, Sta. 7A, 4 September 1977. 

USNM 174416, one female, one male, off Florida, 

29°27ʹN, 081°03ʹW, 20 m, coll. by Texas Instruments, 

Inc. for BLM/MMS, SABP, box corer, Sta. 7A, 4 September 1977. 

Diagnosis. (Modified from Williams, 1993.) Carapace 

dorsal surface anteriorly flattened, coarsely tuberculate 

and covered with stiff setae; postocular spine present. 

Rostrum well developed, large, triangular, slightly 

downturned, thickly covered with stiff setae dorsally, 

0–8 spines ventrally; lateral projections terminating in 

acute spines. Eye stalks rounded, shorter than rostrum. 

First pereopods of approximately same size, chelate but 
26

with greatly reduced fixed finger; carpus with two strong 

spines on anteromesial margin; propodus fixed finger 

small, distinct. Pereopod 2 simple, merus with proximal 

mesoventral spine, single subdistal spine dorsally. Pereopod 3 merus with cluster of spines ventrolaterally. Pereopod 4 merus unarmed. Abdominal sternites unarmed. 

Pleopods somewhat broad and flattened.

Known range. Massachusetts to southern Texas (Williams, 1993).

Habitat. Firm mud or mud-sand substrates; lower intertidal to 20 m (Williams, 1993).

Remarks. Often called the coastal mud shrimp, U. affinis

is a common resident of the sounds and mesohaline 

tidal rivers systems of the SAB (e.g. see Prezant et al., 

2002; Van Dolah et al.11). Zoea were collected in nearshore plankton samples at Folly Beach, South Carolina 

(DeLancey, 1984), but adults are obligate burrow dwellers. Frey and Howard (1969) picture a cast of a burrow 

and describe the burrows in the Sapelo Island area as 

typically 1–2 cm wide, 1–2 m long extending to a depth 

of 50 cm. They found that the burrow walls “lack the pelletoidal exterior characteristic of Callianassa burrows.” 

The size of the individuals in the populations occurring 

along the Atlantic Seaboard is considerably larger than 

those in the populations known from the Gulf of Mexico 

(Heard, pers. observ.).

In Georgia waters, mud shrimp are a common 

food item of the Southern stingray, Dasyatis americana

(Hildebrand and Schroeder, 1928), and the Atlantic 

stingray, Dasyatis sabina (Lesueur, 1824) (Dahlberg and 

Heard, 1969; Howard et al., 1977; Heard, pers. observ.). 

The hakes, Urophycis regius (Walbaum, 1792) and Urophycis floridanus (Bean and Dresel, 1884), also feed heavily 

on this crustacean (Sikora et al., 1972).

Upobebia affinis serves as host for a variety of symbionts. 

Two parasitic bopyrid isopods, Progebiophilus upogebiae

(Hay, 1917) and Orthione furcata (Richardson, 1904), 

occur in the branchial chamber of U. affinis (Markham, 

2001). In addition, the alpheid shrimp, Leptalpheus 

forceps Williams, was described from the burrows of U. 

affinis (Williams, 1965). Numerous specimens of U. affinis, along with specimens of L. forceps, were recovered 

from the stomach of a large American stingray (D. 

americana) caught in Wassaw Sound, Georgia (Heard, 

pers. observ.). McCloskey and Caldwell (1965) reported 

the presence of the symbiotic fungus, Enteromyces callianassae, in the hindgut of specimens from the Beaufort, 

North Carolina area. Ruppert and Fox (1988) listed additional symbionts of U. affinis, including the polychaete

Parahesione luteola (Webster), the flatworm Stylochus 

ellipticus (Girard), the copepod Hemicyclops adhaerens 

(Williams), and the decapod Pinnixia sayana Stimpson. 

An unidentified tapeworm plerocercus attributed to 

the tetrarhynch genus Prochristianella Dollfus, was found 

in the digestive gland of specimens examined from 

the vicinity of Beaufort, North Carolina (Heard, pers. 

observ.). For a review of what is known about U. affinis, 

see Williams (1993).

Acknowledgments

We extend our gratitude to J. M. Foster, S. E. LeCroy, and 

J. N. Thoma (Gulf Coast Research Marine Laboratory, 

Ocean Springs, Mississippi) for their untiring assistance 

and support of this project. We thank Barry Vittor, Barry 

A. Vittor and Associates, Mobile, Alabama, for providing 

specimens of Crosniera minima and Cheramus marginatu

from Puerto Rico. This manuscript was completed with 

funding to the Southeastern Regional Taxonomic Center through the NOAA National Marine Fisheries Service grant NA16FL1490. This document is also Special 

Scientific Report No. 18 of the Marine Resources Division, South Carolina Department of Natural Resources.

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30

Appendix

Figure sources

The illustrations used in this document were obtained 

from a variety of sources and include published figures and original drawings. In most cases, the illustrations have been modified from the original to better 

illustrate the character being described. The sources 

for all illustrations are cited in the figure captions 

and in full in the literature cited section. Permission 

has been obtained for the use of illustrations taken 

from copyrighted sources, and those are credited as 

follows:

 Illustrations from the Proceedings of the Biological Society of Washington (citation: Heard and Manning, 1998) 

are reproduced with permission of the editor.

 Illustrations from The Bulletin of Marine Science (citations: Biffar, 1971a, 1971b; Boesch and Smalley, 1972; 

Kensley, 1996; Manning and Felder, 1991a) are reproduced with permission of the editor.

 Illustrations from Crustaceana (citation: Kensley, 

2001) are reproduced with permission of the editor 

and publisher (Brill Academic Publishers).
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