From the Upper Pliocene Yorktown Formation of Southeastern Virginia

Home , Libinia dubia

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116( 1): 168—189. 2003. New brachyuran crabs (Crustacea: Decapoda) from the Upper Pliocene Yorktown Formation of southeastern Virginia

Warren C. Blow

Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A.

Abstract.—Six new species of crabs, representing three families and one new genus, are described and illustrated from the Upper Pliocene Yorktown For- mation of southeastern Virginia. They are remarkable for their state of preservation and represent the first fossil record for three of these living genera in Virginia's rich Neogene marine deposits. The presence of Stenocionops along with the common occurrence of Persephona in these deposits suggests that warm temperate waters covered southeastern Virginia during the deposition of late Yorktown sediments.

Introduction Stimpson, 1859, the remaining species— Callianassa suffolkensis Rathbun, 1935; C. In 1935, Mary J. Rathbun published the atlantica Rathbun, 1926; Persephona punc- first comprehensive paper on the fossil tata (Linne, 1758); Callinectes sapidus crustaceans, principally decapods, of the Rathbun, 1896; Panopeus herbstii Milne- Atlantic Gulf Coastal Plain of Eastern Edwards, 1834; and Libinia dubia Milne- North America. In this landmark paper she Edwards, 1834—are all listed from locali- describes or mentions all of the fossil deca- ties now regarded as Upper Pliocene. Cal- pod, stomatopod and isopod crustaceans linectes sapidus Rathbun, 1896 is the only from this region made available to her at species listed from the Pleistocene. that time from a host of sources (Rathbun In the present paper six additional taxa, 1935:1). Of the 25 families, 67 genera, and Hepatus bottomsi, n. sp., Pterocarcinus 167 species covered, and for the most part baileyi, n. gen, n. sp., Persephona niemey- illustrated, in her paper only eight species eri, n. sp., P. rodesae, n. sp., Stenocionops representing six genera and six families dyeri, n. sp., and Euprognatha ricei, n. sp. were listed and treated from Virginia from are described as new and illustrated from deposits of Miocene, Pliocene and Pleisto- deposits of late Pliocene age in the same cene age. All but one were listed as occur- general geographic area as that treated by ring in deposits of the Yorktown Formation Rathbun 67 years ago. Unlike Rathbun's which at that time was regarded as Mio- material, which is very fragmentary, often cene. However the Neogene stratigraphy of consisting only of fingers, the taxa present- southeastern Virginia has evolved consid- ed in this paper are remarkable for their erably since 1935 (Hazel 1971a, Ward & state of preservation and completeness. All Blackwelder 1980, Ward & Gilinsky 1993), are represented by their original exoskele- and consequently only Cancer borealis ton, and many even exhibit a uniform color Stimpson, 1859, Panopeus herbstii Milne- similar to that of their living relatives. Edwards, 1834, and Libinia dubia Milne- The material studied in this paper was Edwards, 1834, are listed from localities collected from the Moore House Member that are now regarded as old as Miocene. of the upper part of the Yorktown Forma- With the exception of Cancer borealis tion from three localities in southeastern VOLUME 116, NUMBER 1 169

Virginia. They are: The Lone Star Lakes, water mollusks. Locally large offshore bars near Chuckatuck, Suffolk; Riddick Pit, bor- were the site of rapid, large-scale, cross- row pit east side Virginia Routes 10/32, 4.4 bedded sand deposition." All of the speci- km southeast of Benns Church; and Rices mens studied in this paper and the strata at Pit, a now flooded borrow pit in Hampton the localities from which they were derived (Ward & Blackwelder 1980). Detailed lo- are now regarded by L. W. Ward as be- cality data for the these localities are pro- longing to the Moore House Member and vided in Appendix 1. of late Pliocene age (Ward, pers. comm., 26 With the exception of one female para- November 2002). Kier (1972), in a study of type of Stenocionops, the gender of the re- the echinoids from two (Chuckatuck and maining specimens described in this paper Rices Pit) of the three localities covered in could not be determined. this paper, gives a late Miocene age for his material. This age, though now regarded as Stratigraphy & Paleoenvironment incorrect, was based on the ostracode zones of Hazel (1971a) and the work of Gibson Mansfield (1943) expanded on his earlier (1967). A late Miocene age for strata found biostratigraphic division of the Yorktown at these localities was also the consensus of Formation into Zones I and II and regarded Neogene molluscan workers at that time. both units as entirely Miocene. His zona- Kier's interpretation of the life habits of the tion, though flawed, was very workable in Yorktown echinoids is a valuable resource the field for many years. Hazel (1971a) di- for understanding the paleoenvironmental vided the Yorktown into three ostracode conditions that existed during the deposi- zones of which the uppermost or Puriana tion of the Yorktown at Rices Pit and par- mesacostalis zone, he regarded as Pliocene. ticularly at the Chuckatuck Bar. (See John- Ward & Blackwelder (1980) refined the son & Coch 1969, Johnson 1969 and stratigraphy of the Yorktown Formation by Campbell 1993 for a detailed description of redefining and dividing it into two forma- the structure, extent and mollusks of the tions: the Upper Miocene Eastover Forma- Chuckatuck Bar, which includes the depos- tion overlain by the Lower Pliocene York- its at the Riddick Pit). town Formation. The latter was further di- Kier (1972:3) states, "The Yorktown vided into four members which in ascend- echinoids, like the ostracodes, indicate a ing order were named: Sunken Meadow, past climate warmer than now" . . . "Al- Rushmere, Mogarts Beach, and Moore though some of these living species range House. Ward & Blackwelder (1980:44), in into cooler waters, they all occur in sub- defining the uppermost member state, "the tropical regions, suggesting that the fossil Moore House Member, representing the re- echinoids they resemble lived in waters gressive phase of the Yorktown Formation, warmer than the mild-temperate waters now reflects a renewal of higher current and occurring off the coast of Virginia" Earlier, wave energy conditions. The member con- Hazel (1971b) in a detailed study of the sists of sandy shell beds and cross-bedded Yorktown ostracodes coupled with his own shell hash and locally is cemented to form detailed studies of Western Atlantic Recent a very indurated rock." They further (p. 45, ostracodes concluded that the climate of the 47) state, "The Moore House Member is Yorktown Sea was much warmer than Vir- found only east of the Surry Scarp," and ginia's coastal waters are today. Kier (1972: "The Moore House Member reflects a pro- 2) in summarizing Hazel (1971b) states, gressively shallowing, regressive sea. Mol- "This equable thermal regime is markedly luscan assemblages indicate normal salini- different from that of any province and con- ties, but some of the highest beds in the comitant climate zone now extant along the Williamsburg area contain a few brackish- Atlantic coast of the United States. The 1S 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON temperatures varied from about 12°C to has its structure and function been de- 15°C in the winter to about 17.5°C to 20°C scribed previously." These structures have, and finally to 20°C to 25°C in summer." however been known for some time as not- Ward & Gilinsky (1993:26) state, "The ed above, and both their development and Yorktown Sea supported a large, warm- function studied in great detail (Serene temperate to subtropical molluscan assem- 1954, Guinot 1979). Given the vast range blage." Western Atlantic species of the spi- of shapes suggested by the term Fungi, the der crab Stenocionops are not reported term bolitimorph is more specific and is north of Cape Hatteras, North Carolina but preferred here to denote mushroom-shaped inhabit warmer waters further south, some structures. to Brazil (see Rathbun 1925; Williams Repositories.—Primary and secondary 1965, 1984 for geographic ranges). The types are deposited in the National Museum purse crab, Persephona mediterranea of Natural History, Smithsonian Institution, (Herbst, 1794) though reported from as far Washington, and The Virginia Museum of north as New Jersey and the southern Ches- Natural History, Martinsville, Virginia as apeake Bay, is not commonly found there indicated. but is commonly found in warmer more Abbreviations.—NMNH, National Mu- southern waters from off Cape Hatteras to seum of Natural History, Smithsonian In- Brazil. The crabs of the Yorktown Sea, like stitution; S.I., Smithsonian Institution; the mollusks and other invertebrates found USGS, U.S. Geological Survey (when used at the localities cited here, are representa- with number indicates a locality; see local- tive of similar taxa that would be found to- ity register at end of paper); USNM, abbre- day in warm-temperate to subtropical, open viation for catalogue numbers of the former marine waters of normal salinity. U.S. National Museum now the National Museum of Natural History; VMNH, Vir- Terms and Conventions ginia Museum of Natural History.

The minute mushroom-shaped structures Measurements found on the surfaces of numerous brach- yurans, and referred to by French authors Measurements are expressed in millime- for more than 100 years as "champignons" ters. Abbreviations are as follows: cl, car- (mushrooms) in reference to their "mush- apace length, maximum longitudinal mea- room-like" appearance, are here referred to surement; cw, maximum transverse mea- as bolitimorphs after the Greek bolites surement; prl, propodus length, length of (mushroom) in combination with the Greek palm or combined length of palm and fixed morphe (form, shape). Bolitimorph as here finger; prh, propodus height; prt, propodus defined is intended to replace the phrases thickness. "mushroom-like structure" or "mushroom- shaped structure". At least one living spe- Systematic Paleontology cies, Merocryptus boletifer Milne-Edwards Family Calappidae & Bouvier, 1894, from the Azores was Subfamily Matutinae named for this common feature. Genus Hepatus, Latreille, 1802 Haj & Feldmann (2002) observed similar Hepatus bottomsi, new species structures in members of the family Ran- Fig. 1 inidae and introduced the term "fungi- form" for their mushroom shaped struc- Diagnosis.—Carapace arcuate (shaped tures that form a "pebbled surface" and like a drawn bow), front prominent, strong- stated, "This pebbled surface has not been ly elevated, produced well beyond orbits recognized in any other decapod taxon, nor and anterolateral margin; dorsal surface VOLUME 116. NUMBER 1 171

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Fig. 1. Hepatiis bottoinsi Holotype (USNM 520705) dorsal \iew. Scale = 10 mm.

with eight distinct, pustulate protuberances; width between one-fourth and one-third suborbital flank deeply depressed above de- carapace width. Front prominent, thick, bi- scending, weakly sinuous anterolateral mar- lobate, strongly projecting beyond orbits gin; surfaces everywhere pitted, particularly and anterolateral margin (and placed well anteriorly where appearing eroded. above anterolateral continuation of borders Description. —Carapace arcuace, strongly of carapace); lobes truncate with lateral narrowing posteriorly, broad, length about margins rounded anteriorly, dorsally divid- three-fourths width, widest just forward of ed by deep sulcus, ventrally fused or closed anterior-lateral angle, strongly convex lon- along entire length. Antennules very gitudinally and highly arched transversely, obliquely set. Orbit small, subcircular. de- appearing tripartite in frontal view, gastric fined by slightly raised wide rim; rim com- and branchial regions distinctly swollen, re- posed of three lobes, base of eye in orbit gions moderately well defined, greatly ele- level with front; orbits directed obliquely vated in younger individuals; dorsal surface downward parallel to slope of thick, round- with eight prominent, pustulate protuber- ed dorsal margin above suborbital depres- ances. Surfaces everywhere pitted, giving sion. Anterolateral margin comprised of an eroded appearance; pits most abundant about 40 blunt denticles situated on from on anterior one-third of carapace particular- 12 to 13 or more teeth; teeth separated by ly on front, eyes, orbits, post orbital and closed incisions, most of which begin with anterolateral margins, and adjacent surfac- a deep pit; teeth bidentate anteriorly other- es. Surfaces somewhat smoother in larger, wise most often tridentate with median den- more mature individuals. Fronto-orbital ticle strongest. Margin, where descending 1S 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON anteriorly to buccal cavity, weakly sinuous, when wet. Recent species in alcohol are comprised of single row of denticles of sub- usually gray. equal strength. Posterolateral margin Etymology.—This species is named for straight, strongly convergent posteriorly; E. Edward Bottoms, paleoanthropologist of anterior portion of margin, just posterior to Portsmouth, Virginia, in recognition of his strong tooth defining anterolateral angle, numerous writings on Paleo Indians of the with a long low-angle tooth directed Eastern United States, for his years of obliquely forward, tooth with double or tri- teaching regional history on all student lev- ple close set rows of low tubercles; tuber- els in the Tidewater Area, and for stimulat- cles continue as a single or double row to ing the author's early interest and develop- distinct, though small, subacute tooth defin- ment in paleontology. ing posterolateral angle; single row of tu- Holotype.—The Lone Star Lakes, near bercles continue transversely across poste- Chuckatuck, Suffolk, Virginia. USGS rior margin of carapace where it eventually 26891; (USNM 520705) cl 42.88 mm, cw continues along the opposing lateral mar- 54.92 mm. Paratypes: (USNM 520706), cl gins until returning to the opposite side of 49.97 mm, cw 66.18 mm. USGS 26891 as the buccal cavity from which it began. Pos- above; (USNM 520707), cl 33.63 mm, cw terior margin high, narrow, width about 42.77 mm. USGS 26891 as above. (VMNH one-seventh to one-fifth carapace width; 13546), cl 47.24 mm, cw 64.19 mm. Same margin defined above by slightly convex geographic locality as USGS 26891 as row of tubercles continuous with row of above. posterolateral margin, and submarginal row Occurrence.—The Lone Star Lakes, near of tubercles below. Postfrontal surface with Chuckatuck, Suffolk, Virginia. medial, broad, very shallow sulcus leading Remarks.—Recent comparisons: The to base of frontal lobes. Epigastric region greatly produced front of H. bottomsi im- defined by minute pustules on either side of mediately separates it from all living spe- base of postfrontal sulcus. Hepatic region cies of Hepatus but may confuse it with depressed, bordered anteriorly by thick, species of the genus Hepatella, Smith, in rounded carapace margin leading to orbit. Verrill, 1869, type species H. arnica, from Remaining dorsal regions defined by eight which it differs as follows. Adult specimens prominent, pustulate protuberances distrib- of Hepatus bottomsi have eight low dorsal uted as follows: paired protogastric, one protuberances compared to the six greatly mesogastric, one cardiac, paired epibran- elevated dorsal protuberances found in chial, and paired mesobranchial. Protogas- adults of Hepatella, and as Rathbun, 1937 tric and mesogastric regions large, of sub- notes, Hepatella lacks the suborbital de- equal size, subcircular. Epibranchial regions pression found in members of the genus developed into oblique ridges directed to- Hepatus. This feature is distinct in H. bot- ward marginal tooth of anterolateral angle. tomsi. Mesobranchial regions developed into ridg- Fossil comparisons: Hepatus bottomsi es aligned roughly with anterior ridge of differs from the middle Miocene H. nodo- cardiac region to form a transverse line of sus Collins & Morris, 1976, of Trinidad, the protuberances aligned with first posterolat- only fossil species with which it might be eral tooth. Anterior face of ridges, smooth, confused, in having a greatly produced steep. Intestinal region weakly inflated, in- front and obliquely directed epibranchial distinct. Posterior gastric pits minute, close- ridges. In H. nodosus the front is marginal ly spaced. and the epibranchial ridges are transverse. Color.—Rust or yellowish brown to Individuals of H. bottomsi become brownish gray when dry, gray or dark gray smoother and their dorsal protuberances VOLUME 116, NUMBER 1 173 lower and much less pronounced as they this new species, differs in having very grow larger and older. thick or rotund branchial "wings" unlike the thin carinate wings of Pterocarcinus Leucosiidae and in having the cardiac and urogastric re- Subfamily Ebaliinae gions joined and not completely separated Pterocarcinus, new genus by a deep sulcus as in Pterocarcinus (Fig. 2A). In the absence of the first right pleo- Diagnosis.—Carapace suboctagonal, pod of a male and other features present in wing like; "wing" or branchial region ex- living material, the true relationship of this tending laterally well beyond basis of legs; new fossil genus to other leucosiids may margins very thin, carinate; front narrow never be fully understood, but its form is with two very weak rostral "horns"; car- quite different from all known genera. Frag- diac region most prominent, a raised, sub- ments of this new genus are fairly common circular platform or "mesa" surrounded by in strata of Pliocene age along the Atlantic a deep, wide sulcus; posterior margin Coastal Plain. Given this situation it seems broad, bilobate. best to give this unusual leucosiid a distinct Type species.—Pterocarcinus baileyi, n. generic name. sp. by present designation and monotypy. Related species.—lEbalia rotundata (A. Pterocarcinus baileyi, new species Milne-Edwards, 1880); See A. Milne-Ed- Figs. 2-4 wards & E. L. Bouvier, 1902 for figures. Generic status uncertain. Diagnosis.—As for genus. Etymology.—Ptero from the Greek, pter- Description.—Carapace suboctagonal, on, wing, in combination with the generic wing-like, length about eight-tenths width, name Carcinus derived from the Greek, broadest at anterolateral angle; regions karkinos, or crab. Gender masculine. moderately well defined; dorsal ridge from Remarks.—In outline Pterocarcinus re- front along midline to and including inter- sembles a number of species currently secting diagonal branchial ridges; hepatic placed in Ebalia Leach, 1817; Lithadia and cardiac regions and posterior lobes dis- Bell, 1855 and Speloeophorus A. Milne- tinct to prominent, ridges greatly elevated; Edwards, 1865, but it differs dramatically irregular inner margin of elevated hepatic in overall shape and form from the type region deeply excavated. Cardiac region species of all of these genera. It does not most prominent (Figs. 2A, 3B, F), a raised, possess the subhemispherical form of the subcircular or oval free-standing "mesa- type species of Ebalia, E. tuberosa (Pen- like" platform completely surrounded by a nant, 1777). Nor does it exhibit the outline, deep, wide sulcus filled with scattered, tall "thick" wings, and caverns of the type spe- bolitimorphs (Fig. 3B, D); "mesa" slightly cies of Speloeophorus, S. nodosus (Bell, elevated above ridge of branchial region 1855), and it does not exhibit the highly and strongly produced posteriorly. Most elevated and excavated form represented by surfaces of carapace covered with pavement the type of Lithadia, L. cumingii Bell, 1855. of low, closely spaced or abutting boliti- (See Rathbun 1937, for figures of North and morphs (Fig. 2A); bolitimorphs separate or South American species placed in these free standing in depressions adjacent to genera). In overall form and outline P. bail- ridge leading to front, along and anterior to eyi most closely resembles Ebalia rotun- diagonal ridge of branchial region, and in data (A. Milne-Edwards, 1880), a species sulcus completely surrounding cardiac clearly not in the genus Ebalia, but one "mesa-like" region as well as ventral sur- which might be its closest living relative. faces of pterygostomian and branchial re- Ebalia rotundata, though very similar to gions where they are highest and most dis- 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Pterocarcinus balleyi Holotype (VMNH 13547) A. dorsal view; B. frontal view, close up. Scale

1 mm. VOLUME 116, NUMBER 1 175

Fig. 3. Pterocarcinus baileyi Holotype (VMNH 13547) A. anterior view; B. posterior view; C. left lateral view; D. oblique posterior view. Scale = 1 mm. E. left lateral view of ventral surface beneath branchial "wing".

Scale = 0.5 mm. F. oblique frontal view to show height of cardiac "mesa". Scale = 1 mm.

tinct. Front truncate, elevated, narrow, Pterygostomian region developed into width slightly less than one-sixth width of long ridge subparallel to anterolateral mar- carapace, lateral margins defined by very gin (Fig. 3A); ridge composed of free short, rounded, blade-like "horns" (Fig. standing rows of bolitimorphs of median 2A, B); front weakly projecting. Horns ap- length that cluster to form a downward, pearing more distinct in younger individu- obliquely directed tooth before terminating als. Fronto-orbital width about one-fourth posteriorly. Buccal cavity triangular, ante- carapace width. Orbit sub-circular, margin- rior margins of afferent channels with mi- al, with three sutures, slightly produced at nute pustules, margin curving downward ex-orbital margin. Orbit and antennule fos- laterally to a narrow notch or sulcus. Bran- sae openly connected; basal antennal seg- chial "wings" extending laterally well be- ment not present (not preserved); antennule yond basis of ambulatory legs and chela fossae oblique. Bolitimorphs on fronto-or- (Fig. 3A-C); legs probably not visible in bital surfaces closely spaced forming pave- dorsal view. "Wings" ventrally covered ment (Fig. 2B). with distinctly separated tall bolitimorphs 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Pterocarcinus baileyi Holotype (VMNH 13547) A., B. stereo pair in dorsal view. Scale = 1 mm. Pterocarcinus baileyi Paratype (USNM 520708) C, D. close-up of bolitimorphs in post-hepatic depression. Scale = 200 |xm. Pterocarcinus baileyi Holotype (VMNH 13547) E. close-up of stalk of bolitimorph to show dagger-like structures that might function to hold sediment in place. Scale = 10 [xm. Pterocarcinus baileyi Paratype (USNM 520708) F high magnification to show unique structure of crown of bolitimorph seen in figures C. and D. above. Scale = 20 ixm.

(Fig. 3E); bolitimorphs much shorter and pearing beaded below where bordered by a abutting around margins at insertion of am- single row of minute bolitimorphs. Antero- bulatory legs and chela. Anterolateral, lat- lateral margin longest, convergent anteri- eral, and posterolateral margins very thin, orly, set at 40 degrees to longitudinal axis; carinate (Fig. 3A-C); leading edge of each margin interrupted medially by a very dis- curving downward like an airfoil; edge ap- tinct fissure between hepatic and branchial VOLUME 116, NUMBER 1 177 regions; margin anterior to fissure thickened Co., Virginia, and The Lone Star Lakes, and posterior to fissure sinuous; anterolat- near Chuckatuck, Suffolk, Virginia. eral angle rounded to subacute. Lateral mar- Remarks.—The function of the boleti- gin about two-thirds length of anterolateral morphs found on Pterocarcinus baileyi and margin, distinctly concave in dorsal view other crabs probably differs with each of and raised medially in lateral view; anterior the taxa on which they occur. Given the dif- half weakly divergent anteriorly; posterior ficulty of cleaning specimens covered with half subparallel to longitudinal axis and ter- these boletimorphs, it is obvious to this au- minating in a subacute angle. Posterolateral thor that their primary function is most margin perpendicular to longitudinal axis, probably one of concealment, in that they length less than one-half length of antero- very effectively trap sand particles and hold lateral margin; margin separated from pos- them in the grooves and excavations of the terior lobes by deep sulcus. Posterior dis- carapace, thus obscuring the crab's outline tinctly bilobate, width slightly less than and allowing it to blend into the substrate. one-half width of carapace; lobes thick, Such concealment would allow it to hide separated by a deep fissure; dorsal surface from predators and ambush prey as well. of each, smooth; lobes moderately pro- These sometime very ornate, even flower- like, surface structures might also aid in an- duced posteriorly, margin of each terminat- choring the crab in the substrate. They also ing in a thin, raised, beaded rim. undoubtedly have a structural advantage in Apparent posterior gastric pits distinct, their reinforcement of the carapace. Serene very widely spaced, set on branchial ridges (1954) elaborates on their development and just anterior to gastro-cardiac sulcus (Fig. Guinot (1979) discusses their development 2A). and presents the hypothesis that they serve Color.—Brown, buff, or sometimes to channel water to the bases of the legs weathered bluish grey. and that they aid in concealment as well. Etymology.—This species is named for My own observations have indicated that Richard H. Bailey, geologist and paleontol- their individual form can be species specific ogist, Professor, Northeastern University, and therefore of particular use to paleon- Boston, Massachusetts, for his research tologists in the identification of crab frag- contributions to Atlantic Coastal Plain ge- ments. ology and paleontology, for his many years of superb teaching, and especially for his years of encouragement and field assistance Subfamily Iliinae to the author. Genus Persephona Leach, 1817 Persephona niemeyeri, new species Holotype.—Riddick Pit, about 4.4 km Figs. 5A, C, E, F southeast of Benns Church, Isle of Wright County, Virginia. Same geographic locality Diagnosis.—Carapace hemispherical as USGS 26892 (VMNH 13547), cl 10.04 with three posterior spines; front bilobate, mm, cw 12.75 mm. Paratypes: (VMNH elevated, produced with orbits well beyond 13548), cl 10.77 mm, cw 13.36 mm esti- anterolateral margin; post frontal, hepatic mate, incomplete. Same geographic locality and all other dorsal surfaces thickly covered as USGS 26892 above. (USNM 520708), with relatively high, subacute tubercles of cl 9.16 mm, cw 10.22 mm, USGS 26891, various sizes; tubercles strongest along lat- The Lone Star Lakes, near Chuckatuck, eral and posterior margins, and highly con- Suffolk, Virginia; (USNM 520709), cl 8.79 centrated on front, orbits and adjacent areas mm, cw 10.95 mm, USGS 26891 as above. where developed into bolitimorphs. Occurrence.—Riddick Pit, about 4.4 km Description.—Carapace hemispherical, southeast of Benns Church, Isle of Wright width about nine-tenths of length, regions 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 116, NUMBER 1 179 very weakly defined, front and orbits ad- bercles, sometimes indistinct or appearing vanced well beyond anterolateral margins; as double row where intermixed with dorsal posterior with three equally spaced re- tubercles; area below plural suture covered curved spines, two marginal, and one me- with closely spaced low tubercles, surface dian above margin. Dorsal surfaces thickly appearing very rough, bordered below by covered with relatively high, subacute tu- distinct row of strong tubercles just above bercles of various sizes; tubercles strongest bases of ambulatory legs. Posterior margin along lateral and posterior margins, most slightly more than one-third carapace concentrated below plural suture, where width, produced posteriorly, laterally de- very low and closely spaced, and highly fined by a broad, stout, triangular, acute re- concentrated on front, orbits and adjacent curved spine at each corner; margin be- areas where developed into bolitimorphs. tween spines comprised of from seven to Front very narrow, width about 0.09 width 10 large recurved tubercles; intestinal mar- of carapace, bilobate, lobes separated by gin above with a single medially placed, distinct v shaped sulcus, laterally defined by stout, recurved spine. Spines coarsely gran- raised clusters of tubercles; anterior mar- ulate. Posterior gastric pits not evident. gins of lobes blunt, denticulate. Orbit very Color.—Reddish orange to yellow when small, divided by three deep, narrow, open dry. sutures creating four lobes. Minute, elon- Etymology.—This species is named for gate tubercles span sutures from both sides, Antonio B. Niemeyer, Jr., science teacher, creating a pectinate appearance. Supraorbit- retired, of Chesapeake, Virginia, in recog- al eave and median lobe meet at a steep nition of his contribution to science through angle to form a broad v as seen from above. his many years of teaching science in the Large distinct bolitimorphs concentrated on public schools of tidewater Virginia and for outer lobe. Suborbital lobe with afferent the fundamentals of general science that he channel below; channel deeply notched at instilled in this author so many years ago. anterolateral angle, inner angle rounded dis- Holotype.—The Lone Star Lakes, near tally, margin pustulose and channel pro- Chuckatuck, Suffolk, Virginia. USGS duced beyond front and orbits as seen in 26891, (USNM 520710) cl 52.30 mm, cw dorsal view. Hepatic region slightly swol- 48.13 mm; Paratypes: (USNM 520711) cl len, weakly defined, thickly covered with 43.69 mm, cw 38.85 mm, USGS 26891 as tubercles. Superior margin of subhepatic re- above; (USNM 520712) cl 30.06 mm, cw gion with low ridge of concentrated tuber- 25.91 mm, USGS 26891 as above; (USNM cles; ridge not joining but rather descending 520713) cl 23.02 mm, cw 18.07 mm. Frag- just below tubercles of anterolateral margin. ment of anterior portion of carapace only. Anterior and posterior lateral margins of USGS 26891 as above; (VMNH 13549) cl carapace continuous, defined by a row of 36.83 mm, cw 33.27 mm. Same geographic larger tubercles beginning at orbit and con- locality as USGS 26891 above. tinuing along upper edge of plural suture to Occurrence.—The Lone Star Lakes, near a point just above tooth of posterolateral an- Chuckatuck, Suffolk, Virginia. gle; row poorly separated from dorsal tu- Remarks.—Recent comparisons: Perse-

Fig. 5. Persephona niemeyeri Paratype (USNM 520711) A. closeup of left anterior surface in dorsal view. Scale = 4 mm. Persephona rodesae Holotype (USNM 520714) B. closeup of right anterior surface in dorsal view. Scale = 5 mm. Persephona niemeyeri Paratype (USNM 520711) C. dorsal view. Scale = 10 mm. Per- sephona rodesae Holotype (USNM 520714) D. dorsal view. Scale = 10 mm. Persephona niemeyeri Holotype (USNM 520710) E. dorsal view of geronic specimen, bare areas are a result of wear and abrasion. Scale = 10 mm. Persephona niemeyeri Paratype (USNM 520712) F. dorsal view. Scale = 10 mm. 1S 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON phona niemeyeri is most similar to the Re- bercles of subequal size; tubercles strongest cent western Atlantic purse crab, P. medi- along lateral and posterior margins, and terranea (Herbst, 1794) (see: Williams very small to minute where highly concen- 1984 for synonymy) from which it can be trated on distal surface of front, and around immediately separated on the basis of its orbits. dense tuberculate covering alone. Description.—Carapace hemispherical, Fossil comparisons: Persephona niemey- width about nine-tenths of length, regions eri most closely resembles P. rodesae de- very weakly defined, front elevated, pro- scribed below from which it can be sepa- duced with orbits well beyond anterolateral rated as follows: 1.) Persephona niemeyeri margin; posterior with three equally spaced has far more dorsal tubercles than P. ro- recurved spines, two marginal, and one me- desae, especially anteriorly on the postfron- dian above margin. Dorsal surfaces sparsely tal, frontal, and orbital surfaces; 2.) these covered with relatively low, blunt to sub- tubercles are much higher and more acute acute tubercles of subequal size; tubercles than those of P. rodesae\ 3.) the densely strongest along lateral and posterior mar- tuberculate hepatic region of P. niemeyeri gins, most concentrated below plural suture, differs considerably from that of P. rodesae where very low, rounded and closely which possesses only a few tubercles; and spaced; very small to minute tubercles 4.) the lateral row of tubercles just above highly concentrated on front and orbits the plural suture of P. niemeyeri is poorly where some are developed into small boli- separated from the dorsal tubercles with timorphs; post frontal surfaces and adjacent which it inner grades, where as this row in areas behind orbits nearly barren, void of P. rodesae is well separated from the dorsal tubercles. Front very narrow, width about tubercles. 0.11 width of carapace, bilobate, lobes sep- Specimens of isolated "arms" referred to arated by distinct v-shaped sulcus, laterally P. punctata (Linne, 1758) by Rathbun defined by low ridge formed by raised clus- (1935:106), from her Virginia localities will ters of tubercles; anterior margins of lobes probably, in time, be shown to represent P. blunt, denticulate. Orbit very small, divided niemeyeri and/or P. rodesae. The right pos- by three deep, narrow, open sutures creating terior spine of paratype (USNM 520712) four lobes. Minute, elongate tubercles span was broken off prior to photography but has sutures from both sides, creating a pectinate since been restored. The barren surfaces appearance. Supraorbital eave and median seen on the figure of the holotype (USNM lobe meet at a steep angle to form a broad 520710) of P. niemeyeri are the apparent v as seen from above. A few very small result of abrasion on this gerontic specimen. bolitimorphs are present along margin of Remnants of the dense covering of these outer lobe and just posterior to orbit along tubercles and bolitimorphs can be seen on anterolateral margin. Hepatic region slight- the actual specimen. ly swollen, weakly defined, surface with only a few tubercles. Subhepatic region not Persephona rodesae, new species preserved in type series. Anterior and pos- Fig. 5B, D terior lateral margins of carapace continuous, defined by a distinct row of larger tu- Diagnosis.—Carapace hemispherical bercles beginning at orbit and continuing with three posterior spines; front bilobate, along upper edge of plural suture to a point elevated, produced with orbits well beyond just above tooth of posterolateral angle; row anterolateral margin; post frontal surface of tubercles usually well ordered and well barren of tubercles, hepatic region and all separated from dorsal tubercles, particularly other dorsal surfaces sparsely covered with anteriorly; posteriorly row may inner grade relatively low, small, obtuse to subacute tu- with some dorsal tubercles and appear as a VOLUME 116, NUMBER 1 181 double row; area below plural suture cov- patic region; 4.) fewer, though stronger tu- ered with pavement of closely spaced low bercles along its lateral margins; 5.) a much tubercles, surface appearing very rough. rougher surface below its lateral margins; Posterior margin slightly more than one- and 6.) far fewer and much stronger tuber- third carapace width, produced posteriorly, cles along its posterior border. laterally defined by a broad, stout, triangu- Fossil comparisons: Persephona rodesae lar, acute recurved spine at each corner; most closely resembles P. niemeyeri de- margin between spines comprised of from scribed above. See remarks for P. niemeyeri seven to 11 large recurved tubercles; intes- for comparison. tinal margin above with a single medially Persephona rodesae and P. niemeyeri placed, stout, recurved spine. Spines have not been found at the same locality or coarsely granulate. Posterior gastric pits not in similar sediments. Persephona rodesae evident. has been found in the shelly sands of Rices Color.—Pale yellow when dry. Pit, a much calmer, much less violent en- Etymology.—This species is named for vironment during its deposition than that Mary Betty Rodes, science teacher, de- seen at the Chuckatuck Bar where P. nie- ceased, formerly of Portsmouth, Virginia, in meyeri is found. The morphological differ- recognition of her contribution to science ences between these two species therefore through her many years of teaching science appear directly related to these environmen- in the public schools of tidewater Virginia. tal differences. In addition Miss Rodes provided the author with his first formal introduction to pale- Family Majidae ontology. Subfamily Inachinae Holotype.—Rices Pit, Hampton, Virgin- Genus Euprognatha Stimpson, 1871 ia. USGS 26893, (USNM 520714) cl 27.05 Euprognatha ricei, new species mm, cw 23.86 mm. Paratypes: (USNM Fig. 6 520715) cl 32.05 mm, cw 28.39 mm, USGS 26893 as above; (USNM 520716) cl 32.60 Diagnosis.—Carapace pyriform in out- mm, carapace incomplete, cw 35.50 mm, line, surface with five unusually long nar- USGS 26893 as above; (USNM 520717) cl row blunt spines, mesogastric and cardiac 15.47 mm, cw 16.41 mm, fragment of an- spines longest, surface between base of me- terior portion of carapace only, USGS sogastric spine and protogastric regions ex- 26893 as above; (VMNH 13550) cl 27.88 hibiting a distinct, pentagonal array of gran- mm, cw 24.95 mm, same geographic local- ulations mimicking the shape of a royal ity as USGS 26893 above. crown; intestinal spine present, well devel- Occurrence.—Rices Pit, Hampton, Vir- oped; interantennular spine absent. ginia. Description.—Carapace pyriform in out- Remarks.—Recent comparisons: Perse- line; fronto-orbital region strongly project- phona rodesae appears most similar to the ing, narrow, occupying less than one-fourth Recent Western Atlantic purse crab, P. width of carapace; regions well defined, mediterranea (Herbst, 1794) (see: Williams moderately swollen. Front bidentate, pro- 1984 for synonymy) from which it differs jecting; projections represent superior sur- in having: 1.) relatively high, obtuse to sub- face of "shoe-shaped" antennual fossae be- acute tubercles, compared to the very low low; projections broadly triangular or ob- rounded tubercles of P. mediterranea; 2.) tuse distally, divided by a shallow sulcus small to minute tubercles concentrated on extending posteriorly where terminating at its front and orbits unlike P. mediterranea an inverted v-shaped row of eight granules. where these areas are all but barren; 3.) Orbit circular, margins vertical, subparallel; fewer, though stronger tubercles on its he- supraorbital eave covered with triangular 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Euprognatha ricei Holotype (USNM 520718) A. right lateral view. Scale = 2 mm. B. ventral closeup of antennule fossae, and long basal segment of antenna. Scale = 1 mm. C. dorsal view. Scale = 2 mm.

field of granules posteriorly; point of field nal, and branchial regions each surmounted distal where defined by a strong tubercle on with a single prominent cylindrical blunt orbital margin. Eyes apparently on long spine; mesogastric spine longest, vertically stalks, resting within depression of postor- directed; cardiac spine subequal to meso- bital spine. Postorbital spine very strong; gastric spine, slightly directed posteriorly; broad basally, tapering to a point distally, intestinal and branchial spines much shorter obliquely directed forward. Basal antennal and of subequal length; the former oblique- article with inner and outer rows of gran- ly directed posteriorly; latter obliquely di- ules of which several are developed into rected laterally. Spines smooth laterally short, obtuse spines (tubercles); article di- with their distal ends granulated. Surface of rected only slightly obliquely forward, ter- carapace noticeably punctate, otherwise rel- minating in a granulated spine subequal in atively smooth except for concentrations of advancement to the front. Antennal sternum low tubercles or rounded granules along weak, not produced; interantennular spine frontal margins, supraorbital eaves, postor- absent. Outer margin of pterygostomian re- bital spines, hepatic regions, lateral mar- gion with rows of coarse granules; superior gins, and bases of prominent spines. Irreg- row with large blunt tooth posteriorly. Buc- ular concentrations of tubercles and gran- cal cavity broadest anteriorly, lateral mar- ules appearing as a distinct anteriorly di- gins divergent, width about one-third width rected row on each branchial region; row of carapace. Hepatic region conical, sparse- beginning just posterior to prominent spine ly covered with rounded granules or small of branchial region and converging anteri- tubercles, terminating obliquely in a strong orly where tapering to a point at branchial- blunt spine; spine produced laterally to a hepatic junction. Surface between base of distance subequal to advancement of post mesogastric spine and protogastric regions orbital spine. Mesogastric, cardiac, intesti- defined by a very distinct pentagonal array VOLUME 116, NUMBER 1 183 of granulations arranged in the shape of a of E. ricei are much longer than those of E. royal crown. Tubercles strongest on or near gracilipes and E. ricei unlike E. gracilipes hepatic and anterolateral margins. Posterior possesses an intestinal spine; and 2.) the gastric pits distinct, close set at base of me- dorsal surface of E. ricei is much less gran- sogastric region, each bounded posteriorly ulate than the evenly, coarsely granulated by cluster of three granules. Distinct broad, surface of E. gracilipes. The presence of smooth, groove separating cardiac and in- long mesogastric, branchial, and cardiac testinal regions from branchial region. spines, and in particular a long intestional Branchial region posterolateral^ pro- spine in combination with its sparsely gran- duced into a relatively wide, flattened, granulated surface, the absence of an interanten- ulated margin. Posterior margin moderately nular spine and the presence of a pentago- produced posteriorly, width about one-third nal array of granules in the shape of a royal carapace width. crown on its mesogastric region easily sep- Color.—Ash white when dry, or gray arate E. ricei from all other living species when wet. of Euprognatha. Etymology.—This species is named in Fossil comparisons: Rathbun (1935) honor and appreciation of Mr. William M. identified two left dactyls from the Miocene Rice, deceased, of Hampton, Virginia, who of Liberty County, Florida as Euprognatha allowed the author unrestricted access to his sp., p. 112, pi. 24, figs. 16-19. Lacking as- borrow pit and who, with his wife, Made- sociated chela, the type specimens of E. ri- line, and family built a museum on his cei cannot be compared with Rathbun's ma- property for the purpose of educating stu- terial. dents in the Hampton Roads area. This The distal end of the left basal antennal small museum, known as The Kenneth E. article was broken and lost by the author in Rice Memorial Museum (after his youngest handling the specimen after the photograph son), the borrow pit behind it, and the tu- was completed. torage of Mr. Rice served more than five thousand students of all ages each year. His Subfamily Mithracinae positive impact on the paleontology of the Genus Stenocionops Desmarest, 1823 area is unsurpassed. Stenocionops dyeri, new species Holotype.—Rices Pit, Hampton, Virgin- Fig. 7 ia. USGS 26893, (USNM 520718) cl 10.31 mm, cw 9.01 mm. Paratypes: (USNM Diagnosis.—Carapace oblong-ovate, 520719) cl 4.41 mm, cw 3.46 mm, USGS strongly arched anteriorly; superior lateral 22209 same geographic locality as USGS marginal spines three; frontal depression 26893 above; (VMNH 13551) cl 7.75 mm, laterally bordered by 10 to 18 small spines; cw 6.83 mm, USGS 26893 as above. cardiac region greatly elevated, armed with Occurrence.—Rices Pit, Hampton, Vir- an irregular pentagonal array of five short ginia. spines. Remarks.—Recent comparisons: Of the Description.—Carapace oblong-ovate, known living species of Euprognatha, E. strongly arched anteriorly, surface uneven, ricei appears most similar in overall form, regions well defined. Length from rostral number and placement of prominent dorsal notch along middorsal line from 1.33 to spines and distribution of tubercles and 1.41 times greatest width, broadest across granules to E. gracilipes A. Milne-Ed- middle of branchial regions; greatest height wards, 1878b (Florida Keys to Barbados, approximately one-half that of width. Ros- Rathbun 1925). Euprognatha ricei can tral horns basally divergent, laterally sub- however be easily separated from E. gra- parallel, short, flattened, slightly upturned cilipes as follows: 1.) the prominent spines distally. Frontal depression deep, ovate, lat- 184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Stenocionops dyeri Holotype (USNM 520720) A. dorsal view and B. left lateral view. Scale 10 mm.

erally bordered by 10 to 18 small spines mian region triangular, armed with two decreasing in height anteriorly. Fronto-or- small outer marginal spines; anterior lon- bital region broad, more than one-half times gest. Inner margins raised, rim like, anteri- that of width; orbits strongly projecting. orly terminated by an anvil-like process. Eyes small, retractable within deep tubular Buccal frame broader than long, widest an- superior, orbits; preocular spine a small dis- teriorly; width about one-third width of car- tinct spine or tubercle medial to its base; apace width. Superior lateral marginal postocular spine smaller, acute. Orbit com- spines three; one hepatic and two branchial. pleted below by a blade-like lateral expan- Secondary lateral marginal spines common, sion of basal antennal article. Basal anten- may exceed ten, their number apparently in- nal article broader than long, armed with 3 creasing with age. Hepatic region swollen, spines, one anterior-external, short, acute, very distinct, reaching beyond fronto-orbit- barely visible dorsally; another, posterior- al width, armed laterally with a cluster of external, usually very weak; and last, an- five or more spines; superior inner margin terior-internal, strong, acute, extending for- with spines, ward and obliquely downward from basal two posteriormost longest and strongest. insertion of second antennal article. Second Mid-dorsal line slightly raised, antennal article not extending forward to or ridge-like. Gastric region greatly arched an- beyond rostral notch. Opening of green teriorly; armed with from eight to 10 short gland oval, noticeably raised, a prominent, spines, of which four are usually mesogas- obliquely downward-directed, acute spine tric, two metagastric, and three urogastric. near its external-lateral border. Pterygosto- Posterior gastric pits faint, bordered by VOLUME 116, NUMBER 1 185 curved row of spines just distal to cervical species of Stenocionops are dark red, such groove. as Stenocionops far cat a coelata (A. Milne- Cardiac region greatly elevated, usually Edwards, 1878a). See Williams (1984:339). armed with a characteristic irregular pen- Etymology.—This species is named for tagonal array of five small spines, posteri- Brian J. Dyer, microbiologist. Old Domin- ormost two, often close set. Cardio-intesti- ion University, Norfolk, Virginia for his nal and metabranchial regions separated by contribution to science through his years of deep furrows. Intestinal region armed with research and teaching and especially for his two distinct spines along midline, posteri- years of encouragement and field assistance ormost largest; spines aligned perpendicular to the author. to a lower horizontal row of from two to Holotype.—The Lone Star Lakes, near five smaller spines and/or tubercles, their Chuckatuck, Suffolk, Virginia. USGS number apparently decreasing with age; 26891, (USNM 520720) cl including ros- row of spines bordered below by wide, trum 58.38 mm, excluding rostrum 54.47 shallow sulcus. Posterior margin rounded, mm, cw including spines 44.65 mm, ex- narrow, slightly projecting, width less than cluding spines 40.03 mm. Paratypes: one-half fronto-orbital width. Branchial re- (USNM 520721) Female: cl including ros- gions uneven, moderately spinous. A dis- trum 53.91 mm, excluding rostrum 49.96 tinct, armed, broadly conical prominence on mm, cw including spines 41.70 mm, ex- each epibranchial, mesobranchial and me- cluding spines 37.72 mm, right propodus: tabranchial region, which when roughly prl 21.63 mm, prh 6.09 mm, prl 3.15 mm, aligned with rostral notch, forms an invert- left propodus, prl 20.96 mm, prh 5.13 mm, ed V of approximately 30°. Epibranchial prt 3.22 mm, USGS 26891 as above; prominence usually with two secondary (VMNH 13552) cl including rostrum 42.95 spines or tubercles. A prominent ridge mm, excluding rostrum 39.21 mm, cw in- armed with three low spines, posterior lon- cluding spines 32.22 mm, excluding spines gest, occupying inter-lateral margin of me- 27.75 mm, same geographic locality as tabranchial region. USGS 26891 above. Lower margin of entire carapace termi- Occurrence.—The Lone Star Lakes, near nating in a rounded rim. Entire surface of Chuckatuck, Suffolk, Virginia. body (chelipeds—excepting distal half of Remarks.—Recent comparisons: Steno- fingers) closely covered with minute, cir- cionops dyeri appears most similar in shape cular pores. and overall character to S. spinosissima Coxa, basis, and ischium of chelipeds (Saussure, 1857) from which it can be eas- and ambulatory legs, unarmed. Merus of ily separated as follows: 1.) Stenocionops cheliped quadrangular in section, armed dyeri has three strong lateral spines com- with four distinct lateral rows of spines, the pared to the five found in S. spinosissima; dorsal row being most prominent with four 2.) S. dyeri has short, flattened rostral horns superior spines and as many as five smaller compared to the much longer, more acute spines or tubercles. Carpus smooth. Hand horns of S. spinosissima; and 3.) S. dyeri elliptical in section, highest proximally, has between 13 to 15 short median spines, smooth to finely granulate. Fingers less than of which those on the cardiac region are one-half length of palm, not gaping. Am- arranged in an irregular pentagonal array in bulatory legs circular in section; merus of contrast to S. spinosissima which exhibits first and second pair moderately tuberculate 10 median spines of which only two are above; third and fourth pairs smooth. Legs found on the cardiac region. Stenocionops decreasing in length posteriorly. dyeri also superficially resembles a much Color.—Ash white with faint patches of smaller species of Stenocionops, S. trian- pink when dry, gray when wet. Some living gulate. (Rathbun, 1892) which also possess- 1S 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON es three lateral spines. These spines in S. ing washed about and buried in the Chuck- dyeri, however, are shorter and much less atuck Bar deposits by strong currents dur- acute. Stenocionops dyeri also differs from ing storms. These same barnacles also this species in having: 1.) much flatter, strengthened the dorsal aspect of the cara- shorter and less divergent rostral horns; 2.) pace allowing it to be preserved in this high 13 to 15 median spines compared to about energy environment of deposition. nine on S. triangulata; and 3.) an irregular pentagonal array of five spines on its car- Acknowledgments diac region compared to only one or two on this region in S. triangulata. In reference to Mr. H. B. Roberts, deceased, formerly of S. triangulata, Rathbun (1925:461) states: the Department of Invertebrate Zoology, "Known only from small and immature NMNH, SI, correctly identified Hepatus specimens". The pentagonal array of spines bottomsi, Persephona rodesae, Euprogna- on the cardiac region of S. dyeri alone sets tha ricei and Stenocionops dyeri as new in it apart from all other known species of a letter to the present author, dated 13 Oc- Stenocionops. tober 1964. Given the incomplete and often Fossil comparisons: Rathbun (1935) de- fragmentary nature of the material present- scribed Stenocionops primus from the Up- ed to him at that time, his identifications are per Cretaceous Brownstown Formation, of remarkable for their accuracy, a testament Howard County, Arkansas and Stenocion- to his knowledge and ability. Due to his de- ops suwanneeana from the Eocene Ocala clining health, Mr. Roberts was unable to Limestone of Suwannee County, Florida. pursue descriptions of these and other new The chela of these two species may indeed taxa (see: Manning & Blow 1980, Blow & resemble those of adult males of Stenocion- Bailey 1992) and passed that task on to this ops or other mature majid chela but in size author before his retirement in June 1973. and coarse granulation they are very differ- He died 14 March 1979. ent from the nearly smooth diminutive fe- I thank E. E. Bottoms for introducing me male chela of S. dyeri. The chela in both to a number of fossil decapod localities in adult male and female species of some liv- the Hampton Roads area in the fall of 1962, ing representatives of Stenocionops are rel- and for years of cooperative fieldwork and atively small, like those of S. dyeri, but in mutual assistance. I particularly thank my reaching maturity the chela in gerontic parents, R. V. and C. D. Blow, for years of males can become quite large. The geologic logistical support, field assistance, and for time difference between S. dyeri and these their own collecting efforts which have geologically much older taxa is such that it benefitted and continue to benefit my re- would be very unlikely that they represent search these past 40 years. Mr. W. A. Stay- the same species. lor, Mr. H. F. Saunders and the other men The barnacle, Balanus sp., is often found and women of the Lone Star Cement Co. completely covering the dorsal surface of kindly allowed the author unrestricted ac- the carapace of S. dyeri where it apparently cess to their diggings. R. H. Bailey, B. A. lived as a symbiont during the life of the Bedette, B. J. Dyer, L. W. Ward, and the crab. This barnacle has not yet been found late D. Wilson helped the author in the field covering the orbital openings of this crab, on a number of occasions, adding their ex- thus revealing that the crab was indeed pertise and collections to the research and alive and able to keep its orbits clear during writing of the present paper. L. D. Campbell this relationship. The massive numbers of and Svein Nielsen kindly made their col- this barnacle found on some specimens of lections available to the author for study. S. dyeri greatly increased the crab's surface I thank S. Whittaker, of the SEM lab, area and undoubtedly contributed to its be- Smithsonian Institution, for providing the VOLUME 116, NUMBER 1 187 photomicrographs of P. baileyi, M. G. Har- Geological Survey Professional Paper 704:1- asewych for providing the digital images of 13. . 1971b. Paleoclimatology of the Yorktown the remaining taxa except for the dorsal and Formation (upper Miocene and lower Pliocene) lateral views of E. ricei, which were taken of Virginia and North Carolina: Pp. 361-375 in by Mr. T. F. Phelan. My sincere thanks to J. H. J. Oertli, ed., Paleoecologie Ostracodes, Pau, A. Sanner for enhancing the images in Ado- 1970.—Bulletin Centre Recherche Pau-SNPA. be Photoshop and constructing the finished 953 pp., 3 tab. digital plates. In addition B. A. Bedette read Herbst. J. F W. 1794. Versuch einer Naturgeschichte der Krabben und Krebse, nebst einer systema- the manuscript and offered numerous sug- tischen Beschreibung ihrer verschiedenen Arten gestions and corrections. T. R. Waller pro- 2(5): 147-162 + pis. 37-40. vided space in his laboratory, and scientific Johnson, G. H. 1969. Guidebook to the geology of the and editorial advice throughout the prepa- York-James Peninsula and south bank of the ration of this manuscript. James River.—College of William and Mary Department of Geology Guidebook No. 1. At- lantic Coastal Plain Geological Association. Literature Cited Tenth Annual Field Conference and First An- nual Geological Field Conference: 1-33. Bell, T. 1855. A Monograph of the Leucosiadae, with , & N. K. Coch. 1969. A Coquina facies in the observations on the relations, structure, habits, Yorktown Formation near Chuckatuck, Virginia and distribution of the family: a revision of the and its geological implications.—(Abs.) Geo- generic characters; and descriptions of new gen- logical Society America Special Paper 121: era and species, Horae Carcinologicae, or Notic- 448. es of Crustacea, I.—Transactions of the Linnean Kier. P. M. 1972. Upper Miocene echinoids from the Society 21:277-314 + pis. 30-34. Yorktown Formation of Virginia and their en- Blow, W. C., & R. H. Bailey. 1992. Chasmocarcimts vironmental significance.—Smithsonian Contri- robertsi, a new crab species from the Miocene butions to Paleobiology 13:1-41. Smithsonian of Virginia, with notes on the genus Fcilcono- Institution Press, Washington. plax (Crustacea, Decapoda, Goneplacidae).— Latreille, P. A. 1802. Histoire naturelle generale et par- Tulane Studies in Geology and Paleontology ticuliere des Crustaces et des Insects 3:13-467. 25(4):175-185. Paris. Campbell, L. D. 1993. Pliocene molluscs from the Leach, W. E. 1817. Malacostraca Podophthalma Bri- Yorktown and Chowan River Formations in tanniae; or descriptions of the British species of Virginia.—Virginia Division of Mineral Re- crabs, lobsters, prawns, and of other Malacos- sources Publication 127:1-259. traca with pedunculated eyes. James Sowerby. Collins, J. S. H„ & S. F. Morris. 1976. Tertiary and London, XIV, 5 p., unpaged + pis 16, 25 and Pleistocene crabs from Barbados and Trini- 44. dad.—Palaeontology 19( 1): 107-131. Linnaeus, C. 1758. Systema naturae per regna tria na- Desmarest, A. G. 1823. Malacostraces. Pp. 158-425 turae, secundum classes, ordines, genera, spe- in Dictionnaire des Sciences Naturelles, vol. 28. cies, cum characteribus. differentii, synonymis. Strasbourg. locis, 10th edition. Laurentius Salvius. Stock- Gibson, T. G. 1967. Stratigraphy and paleoenviron- holm 1:1-824. ment of the phosphatic Miocene strata of North Manning, R. B„ & W. C. Blow. 1980. Henry B. Rob- Carolina.—Geological Society of America Bul- erts 1 September 1910-14 March 1979.—Crus- letin 78(5):631-650, pis. 1, 2. taceana 39(1): 104-107. pi. 1. Guinot, D. 1979. Morphologie et phyogenese des Mansfield, W. C. 1943 [1944], Stratigraphy of the Mio- Brachyoures.—Memoires du Museum national cene of Virginia and the Miocene and Pliocene d'Histoire naturelle (Paris), new series A (zo- of North Carolina. Pp. 1-19 in J. Gardner, ed.. ology) 112:1-354. Mollusca from the Miocene and Lower Pliocene Haj, A. E.. & R. M. Feldmann. 2002. Functional mor- of Virginia and North Carolina: Pt 1, Pelecy- phology and taxonomic significance of a novel poda.—U.S. Geological Survey Professional cuticular structure in Cretaceous raninid crabs Paper I99-A:l-178. (Decapoda: Brachyura: Raninidae).—Journal of Milne-Edwards, A. 1865. Description de quelques Paleontology 76(3):472-485. Crustaces nouveaux on peu connus de la famille Hazel, J. E. 1971a. Ostracode biostratigraphy of the des Leucosiens.—Annales Societe Entomolo- Yorktown Formation (upper Miocene and lower gique de France 4(5): 148-159. pi. 6. Pliocene) of Virginia and North Carolina.— . 1878a. Note sur quelques Crustaces nouveaux 1S 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

appartenant au groupe des Oxyrhynques.—Bul- . 1937. The Oxystomatous and allied crabs of letin de la Societe Philomathique de Paris 7(2): America.—United States National Museum 222-225. Bulletin 166:1-278, pis. 1-86. . 1878b. Etudes sur les Xiphosures et les Crus- Saussure, H. de. 1857. Diagnoses de quelques Crus- taces de la Region Mexicaine.—In Mission taces nouveaux de l'Amerique tropicale.—Re- Scientifique au Mexique et dans l'Amerique vue et Magzin de Zoologie Pure et Appliquee, centrale part 5(4): 121-184, pis. 21-27, 29-30. 2(9):501-505. . 1880. Etudes preliminaries sur les Crustaces, Serene, R. 1954. Sur quelques especes rares de Brach- 1 ere partie. In reports on the results of dredging yures (Leucosidae) de l'lndo-Pacifique.—Treu- under the supervision of Alexander Agassiz, in bia 22(3):453-499, pis. 7-10. the Gulf of Mexico, and in the Caribbean Sea, Stimpson, W. 1859. Notes on North American Crus- 1877, '78, '79 by the U.S. Coast Survey Steam- tacea, No. 1.—Annals of the Lyceum of Natural er "Blake," Lieut. Commander C. D. Sigsbee, History of New York 7(1862)(2):49-93, 1 plate. U.S.N., and Commander J.R. Bartlett, U.S.N., . 1871. Preliminary report on the Crustacea Commanding.—Bulletin of the Museum of dredged in the Gulf Stream in the Straits of Comparative Zoology at Harvard College 8(1): Florida, by L. F. de Pourtales, Assistant United 1-68, pis. 1, 2. States Coast Survey. Part 1. Brachyura.—Bul- , & E. L. Bouvier. 1894. Brachyures et An- letin of the Museum of Comparative Zoology omoures. In Crustaces decapodes provenant des at Harvard College 2(2): 109-160. campagnes du yacht l'Hirondelle (1886, 1887, Verrill, A. E. 1869. On the parasitic habits of Crusta- 1888), Premiere Partie. Resultats des Campag- cea.—American Naturalist 3:239-250. [.Hepa- nes Scientifiques accomplies sur son yacht par tella arnica Smith described as footnote p. 250] Albert 1 er, Prince Souverain de Monaco 7:1- Ward, L. W„ & B. W. Blackwelder. 1980. Stratigraphic 112, pis. 1-11. revision of Upper Miocene and Lower Pliocene , & E. L. Bouvier. 1902. Les Dromiaces et Ox- beds of the Chesapeake Group, Middle Atlantic ystomes. In reports on the results of dredging, Coastal Plain.—Contributions to Stratigraphy, under the supervision of Alexander Agassiz, in Geological Survey Bulletin 1482-D: 1-61, pis. the Gulf of Mexico (1877-78), in the Caribbean 1-5. Sea (1978-79), and along the Atlantic Coast of , & N. L. Gilinsky. 1993. Molluscan assem- the United States (1880), by the U. S. Coast blages of the Chowan River Formation, Part A, Survey Steamer "Blake", Lieut. Com. C. D. Biostratigraphic analysis of the Chowan River Sigsbee, U. S. N., and Commander J. R. Bart- Formation (Upper Pliocene) and adjoining lett, U. S. N., Commanding, XXXIX.—Mem- units, The Moore House Member of the York- oirs of the Museum of Comparative Zoology at town Formation (Upper Pliocene) and the James Harvard College 27(1): 1-127, pis. 1-25. City Formation (Lower Pleistocene).—Virginia Museum of Natural History Memoir 3, Part A: Milne-Edwards, H. 1834. Histoire naturelle des Crus- 1-32, plate 1. Tables 1, 2, & 4-6 & fig. 6 not taces, comprenant l'anatomie, la physiologie et paginated (in pocket). la classification de ces animaux 1:1-468. Paris. Williams, A. B. 1965. Marine decapod crustaceans of Pennant, T. 1777. Crustacea, Mollusca, Testacea. Brit- the Carolinas.—Fishery Bulletin 65(1): 1-298. ish Zoology, edition 4, 4:London, 1-154, pis. . 1984. Shrimps, lobsters, and crabs of the At- 1-93. lantic Coast of the eastern United States, Maine Rathbun, M. J. 1892. Catalogue of the crabs of the to Florida. Smithsonian Institution Press, Wash- family Periceridae in the U.S. National Muse- ington, D.C., 550 pp. um.—Proceedings of the United States National Museum 15(901):231-277, pis. 28-40. . 1896. The genus Callinectes.—Proceedings of Appendix 1 the United States National Museum 18(1070): Locality register 349-375, pis. 12-28. . 1925. The spider crabs of America.—United USGS 26891 The Lone Star Lakes, formerly Lone States National Museum Bulletin 129:1-613, Star Cement Company open-pit mine, about 1 km pis. 1-283. north of Chuckatuck, Suffolk, Virginia. USGS . 1926. The fossil stalk-eyed Crustacea of the Chuckatuck, 71/i-minute quadrangle map, 1965 pho- Pacific slope of North America.—United States torevised 1979. National Museum Bulletin 138:1-155, pis. 1- USGS 26892 Open-pit mine, locally known as Rid- 39. dick Pit, about 4.4 km southeast of Benns Church, . 1935. Fossil Crustacea of the Atlantic and on east side of Va. route 10/32, Isle of Wright Coun- Gulf Coastal Plain.—Geological Society of ty, Virginia. USGS Benns Church 71/4-minute quad- America, Special Papers 2:1-160, pis. 1-26. rangle map, 1965 photorevised 1986. VOLUME 116, NUMBER 1 189

USGS 26893 Rices Pit belonging to Mr. William M. USGS 22209 Same geographic locality as 26893 Rice, about 0.4 km north of intersection of Fox Hill above but collected by a large group of Smithsonian Road (Rte. 167) and Harris Creek Road, Hampton. scientists lead by G. A. Cooper in 1966. Virginia. USGS Hampton, 7%-minute quadrangle map. 1965 photorevised 1986.