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UC Berkeley PaleoBios

Title Late () elasmobranchs from the Dry Branch Formation (Barnwell Group) of Aiken County, South Carolina, USA

Permalink https://escholarship.org/uc/item/0fh2f5n3

Journal PaleoBios, 36(0)

ISSN 0031-0298

Authors Cicimurri, David J. Knight, James L.

Publication Date 2019-06-04

DOI 10.5070/P9361043964

License https://creativecommons.org/licenses/by-nc-sa/4.0/ 4.0

Peer reviewed

eScholarship.org Powered by the California Digital Library University of California PaleoBios 36:1–31, June 4, 2019 PaleoBios

OFFICIAL PUBLICATION OF THE UNIVERSITY OF CALIFORNIA MUSEUM OF PALEONTOLOGY

DAVID J. CICIMURRI & JAMES L. KNIGHT (2019). Late Eocene (Pri- abonian) elasmobranchs from the Dry Branch Formation (Barnwell Group) of Aiken County, South Carolina, USA.

Cover: Various late Eocene and ray teeth from the Dry Branch Formation, Aiken County, South Carolina. Citation: Cicimurri, D.J. and J.L. Knight. 2019. Late Eocene (Priabonian) elasmobranchs from the Dry Branch Formation (Barnwell Group) of Aiken County, South Carolina, USA. PaleoBios, 36. ucmp_paleobios_43964. Late Eocene (Priabonian) elasmobranchs from the Dry Branch Formation (Barnwell Group) of Aiken County, South Carolina, USA

DAVID J. CICIMURRI¹* and JAMES L. KNIGHT²

¹South Carolina State Museum, 301 Gervais Street, Columbia, SC 29201 [email protected]

[email protected] ²Department of Biology and Geology, University of South Carolina Aiken, 476 University Parkway, Aiken, SC 29801

A survey of the Eocene (Priabonian) Dry Branch Formation exposed in Aiken County, South Carolina, resulted in the collection of thousands of fossil teeth and bone fragments. Two sites located near the city of Aiken proved to be particularly productive, and 24 of elasmobranchs, 11 osteichthyans, and -

threeof daggernose reptiles (oneshark, crocodilian Isogomphodon and twoaikenensis turtles) n. have sp. Cicimurri been identified. and Knight. Herein The we fossils focus are on derived the elasmo from branchthe upper species part of (17 the Dry Branch and seven Formation, rays) that and are the part fossiliferous of the assemblage, strata accumulated which includes within a a new high species energy - nearshoreture was at marine least 22 depositional° C . Elasmobranch environment species that composition was influenced is similar by a river to other system. late Based Eocene on elasmobranchthe assemblagesand invertebrate reported fossils from we identified,the Gulf and the Atlantic water depth Coastal was plains, less thanparticularly 40 m, and Georgia, surface and water several tempera of the

taxa indicate affinities to the Tethyan region. Keywords: fossil, elasmobranch, Eocene, Dry Branch Formation, South Carolina

INTRODUCTION of elasmobranch fossils from the Dry Branch Formation Although Eocene strata within the South Carolina of South Carolina. More than 3,000 shark and ray teeth coastal plain have been extensively mapped and studied were collected from two sites in Aiken County (Fig. 1A), lithologically, only scattered accounts of the elasmo- but only approximately 2,100 of these were complete branch fossils these units contain have been published ( , , , Leriche 1942, White discovery of a new species of described , Müller 1999, ). hereenough as toIsogomphodon be identified toaikenensis or species,n. sp. Cicimurri and included and SeveralGibbes reports1848 1850 on theLeidy elasmobranch 1877 species occurring Knight. In addition to the taxonomic discussions about 1956in the Barnwell GroupCicimurri of Georgia, and which Ebersole, includes 2015 the Aiken assemblages to each other, as well as to those have been published ( , Case and Borodin 2000, previouslythe elasmobranchs reported wefrom identified, the Barnwell we compareGroup of Georgia.the two ParmleyClinchfield, and Dry Cicimurri Branch, 2003 and ),Tobacco but such Road fossils Formations, occurring We also present our interpretation of the depositional within equivalent strataCase in South1981 Carolina have been only environment as inferred from the lithostratigraphy, casually mentioned in the literature ( , taphonomic indicators, and paleoecology of the extant , ). Cicimurri and Ebersole ( ) reported a new speciesZullo of ray, et Pseudaeal. 1982- GEOLOGICAL CONTEXT Zullotobatus and undulatus Kite 1985, fromSteele the et Dry al. Branch1986 Formation in representatives of fossil taxa we have identified. 2015 Stratigraphic framework of Myliobatidae . Aiken County, but only briefly noted other coeval species The fossils described herein were all collected from Bonaparte, 1838 the Eocene Dry Branch Formation in Aiken, Aiken County, *authorOur paper for correspondence presents the first comprehensive analysis Citation: Cicimurri, D.J. and J.L. Knight. 2019. Late Eocene (Priabonian) elasmobranchs from the Dry Branch Formation (Barnwell Group) of Aiken County, South Carolina, USA. PaleoBios, 36. ucmp_paleobios_43964. Permalink: Copyright: Published under Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International (CC-BY-NC-SA) license. https://escholarship.org/uc/item/0fh2f5n3

LSID urn:lisd:zoobank.org:pub:3F95876E-933FF-48AF-9CF0-A840A333220B 2 PALEOBIOS, VOLUME 36, JUNE 2019

South Carolina ( , Falls and Prowell 2001), in central Aiken County the Dry Branch Forma- A Fallaw and Price 1995 North Carolina tion disconformably overlies kaolinite deposits of the lower-to-middle Eocene Huber Formation (Fig. 1B). South The contact between these two formations has been re- Carolina ported as a sequence boundary, with the base of the Dry Columbia Branch Formation consisting of a lag deposit (Harris et Aiken al. 2002, Schroeder et al. 2002) that formed during the Georgia initial transgression of the Jackson Sea into the region ( , Huddlestun 1993) at the start of the Tejas A4.2 cycle (Harris and Zullo 1991). The Atlantic DryHuddlestun Branch Formation and Hetrick is disconformably 1986 overlain by the Ocean Tobacco Road Formation (Fig. 1B), and these two units Florida have been studied extensively for economically impor- N tant occurrences of kaolinite ( ). The base of the Tobacco Road Formation may be very Gulf of Mexico pebbly, but it is also relativelyBuie easy and to identifySchrader because 1982 of its indurated nature, compared to the unconsolidated 0 240 sands of the Dry Branch Formation. In some areas the Dry Branch Formation is absent and the Tobacco Road Forma- km tion disconformably overlies Huber Formation kaolinite. The Dry Branch Formation extends laterally into central Georgia, and within this region the formation B has been subdivided into three members, including the Age Group South Carlolina Stratigraphy NP Zone Stage ( ). The stratigraphic rela-

e l Tobacco Road Formation tionshipsGriffins Landing between Sand, the members Twiggs Clay, may and not beIrwinton as simple Sand as w NP Huddlestun and Hetrick 1979 19/20 Priabonian sub- and superjacent units, as Huddlestun and Hetrick

U p e r Dry Branch Formation

B a r n ( ) observed intertonguing lateral facies changes. t ) Additionally, Huddlestun and Hetrick ( ) and Eversull Bartonian (1986 and Irwinton sands that were similar 1979to those occurring

o c e n ( p a r 2005) reported clay beds within the Griffins Landing E within Georgia is facilitated by exposures of relatively M i d l e thickwithin and the fossiliferous Twiggs Clay. sections, Identification but identifying of Eocene and units cor- O’Burg NP (part) Huber Formation (part) relating strata in South Carolina has been hampered by lack of thick exposures, and those that could be exam- Figure 1. A. Outline map of southeastern US coastal states ined are often highly weathered and devoid of fossils showing South Carolina, Aiken County (shaded region), and the ( ). city of Aiken. B. Generalized stratigraphic section of Barnwell Group strata in the Aiken, South Carolina area. The fossilifer- In Aiken County, the Dry Branch Formation is at least ous horizon at the South Aiken Site occurs approximately 2 Huddlestun 1982 m below the base of the Tobacco Road Formation. A and B , ). Abbreviations: NP=calcareous nannoplankton zone, O’burg=Orangeburg. 28 m thick, and Twiggs Clay, Griffins, Landing Sand, and). modified from Cicimurri and Ebersole (2015 InvertebrateIrwinton Sand fossils have may been be identified common within(Mittwede calcareous 1982 Nystrom and Willoughby 1982 Zullo and Kite 1985 South Carolina. The Dry Branch Formation is part of South Carolina, and species include various types of the Barnwell Group, which also includes the subjacent barnacles,sediments benthonicof the Griffins foraminifera, Landing a Sand few echinoderm of Georgia and crustacean remains, and the ostreid Crassostrea gigantis- sima ( ) may be locally abundant (Huddlestun inClinchfield the subsurface Formation of the and western superjacent part of TobaccoAiken County, Road ). Zullo and Kite ( ) reported that Formation. Although the Clinchfield Formation occurs Finch, 1824 and Hetrick 1979 1985 CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 3

been interpreted as representing inner neritic (sands) Landing Sand was excellent, and that C. gigantissima oc- and bay or lagoonal environments (clays), and scour curspreservation in Allendale of invertebrateCounty but not fossils in Aiken within County. the FurtherGriffins structures could represent tidal channels (Huddlestun to the south in Allendale County, strata attributed to the , ). Overall, the formations within the Barnwell Group of South Carolina Formation ( ), whereas in Aiken County, representand Hetrick a transgressive-regressive 1986 Fallaw and Price 1995sequence, with the Griffins Landing Sand disconformably overlie the McBean lower portion of the Dry Branch Formation represent- Huber FormationSteele (et al. 1986 ). Both Zullo and ing a high-energy mixed siliciclastic-carbonate shelf that KiteGriffins ( Landing) and Steele deposits et al. disconformably ( ) noted occurrences overly the periodically received input from a shoreward source, the Zullo and Kite 1985 upper part of the formation being deposited in a shallow but no1985 details were provided. 1986 siliciclastic shelf environment, and the Tobacco Road of sharkStrata andassigned ray teeth to the within Twiggs the Clay Griffins in the Landing Graniteville Sand, Formation accumulating in a lower delta plain environ- and Hollow Creek quadrangles (west and southwest of ment, with occasional shoreward transport of marine Aiken, respectively) typically occur as well-layered se- sediments (Segalla et al. 2000). quences of clay varying in color from gray, green, brown, Age of deposits quartz sand ( ). Further to The late Eocene (Priabonian) age of the Dry Branch theand southpurple, in and the clay Jackson beds quadrangle, are separated Mittwede by fine-grained ( ) Formation in South Carolina has been demonstrated Nystrom and Willoughby 1982 by the various barnacle and clypeasteroid echinoderm medium-grained quartz sand containing thin, discontinu1982- species that are known to occur ( , ousreported clay beds mustard-yellow that were attributed to orange-yellow to the Irwinton loose, fine- Sand, to , ). Huddlestun and Hetrick and he noted that the unit thickened markedly towards ( ) placed the Twiggs Clay of theZullo Dry andBranch Kite Forma 1985- the Aiken area. The fossiliferous Dry Branch Formation Steeletion within et al. 1986calcareousCarter nannofossil 1987 Zone NP19/20, and deposits we examined are consistent with Mittwede’s tektites1986 in the Dry Branch Formation that formed dur- ( ) description of the Irwinton Sand. Non-quartz ing the Chesapeak Bay Impact have a laser fusion age of grains that we recovered along with the fossils include Albin and Wampler 1996, Albin zircon,1982 kyanite, and muscovite mica, and several types 1999, Povenmire and Povenmire 2002, Schroeder et al. of igneous and metamorphic rocks occuring in the Aiken approximately2002, Harris et 34.5 al. 2002 Ma (, 2004). area could have been source rocks for these Dry Branch sediments ( , ). METHODS The fossils in this report were collected from two loca- DepositionalSnoke setting and Secor 1982 Speer 1982 tions in the Aiken city area, Aiken County, South Carolina. During the Eocene, deposition of marine strata in South Carolina and Georgia occurred within the Gulf The first is the North Aiken Site (NAS), a clay pit south from the Gulf of Mexico (Huddlestun 1990, Eversull overof the the Aiken course Municipal of several Airport years (33.624867, and are included -81.681713 within Trough). Atas thethis Suwanee time, siliciclastic Current flowed deposition northeastward (i.e., Dry Wat the eastern entrance). Fossils were recovered as float Branch Formation) dominated on the northern side of as a housing subdivision and our recent exploration of the2005 Gulf Trough, whereas carbonate deposition (i.e., Ocala itaccession yielded SC96.97.only a few This additional site is currently shark being teeth developed near the Limestone) dominated to the south (Huddlestun, 1990, southern limit of the pit. The second is the South Aiken ). Site (SAS) located in a housing subdivision south of the Zullo and Kite ( EversullLanding 2005Sand in Aiken County represented subtidal to require entry through private property, and permission inner shelf deposition,1985 whereas) concluded Fallaw thatand Pricethe Griffins ( ) mustAiken be city obtained limit (33.504444, before visiting -84.742778). them. Both sites suggested a bay or lagoon to open ocean environment The SAS is the type locality for Pseudaetobatus undula- based on foraminifera they studied. Kirby (20001995) be- tus ( ). Its primary exposure lieved that the occurrence of Crassostrea gigantissima consists of a 0.6 m-thick section covering a six-square- - meterCicimurri area. We and collected Ebersole more 2015 than 100 kg (dry) of in situ ments as opposed to brackish environments. In Georgia, bulk matrix from a very fossiliferous horizon within the the(Griffins Irwinton Landing Sand Sand)Member indicated is locally fully fossiliferous marine environ and has section, and recovered macrofossils that had weathered 4 PALEOBIOS, VOLUME 36, JUNE 2019 out of place. A second, mostly covered and nearly vertical Notorynchus sp. cf. No. kempi Ayres, 1855 section, occurs in an E-W trending wall approximately 5 Ward, 1979a Although no fossils were recovered from this section, we Referred specimens m to the north, where up to 5 m of thickness is preserved. (Fig. 2A–D) were able to determine that the fossiliferous horizon right tooth; SC2001.1.46, lower tooth fragment; within the Dry Branch Formation occurs approximately —SC96.97.4, incomplete lower 2.0 m below the base of the Tobacco Road Formation. Remarks—The incompleteness of the three specimens In the laboratory, Dry Branch Formation matrix was inhibitsSC2013.38.1, our ability incomplete to accurately lower left identify tooth. the Dry Branch disaggregated in water and gently rinsed through U.S.A. from Eocene deposits elsewhere, we found that the acro- Matrix that passed through this screen was also saved, hexanchid. However, when compared to taxa identified Standard Testing Sieves down to 0.25 mm (#60 screen). and the resulting concentrates were dried and sorted numerous but much smaller than those of Notidanodon under a binocular microscope. The SAS matrix sample cone mesial cusplets of SC96.97.4 (Fig. 2C–D) are more yielded about 30 specimens per kilogram, and these (Fig. 2A, B) also conspicuously diminish in size away from fossils are included under accessions SC2001.1 and theCappetta, acrocone 1975 (Gurr, and 1962the distal, cusplets ,of SC2013.38.1). Lower teeth of agassizi are In order to more accurately identify the many broken Casier 1967 Ward 1979a SC2013.38. teeth of Myliobatidae we recovered (particularly those than six on the Dry Branch taxon ( Cappetta, 1976, Ward in median positions), we repaired as many as possible smaller). andIn overall have 7–12 morphology, distal cusplets, our material as opposed compares to less favorably to lower teeth of NotorynchusCappetta, particularly1976 No.1979a serratissimus ( ) and No. kempi. Early to using thin butvar (B-76 in acetone). Specimens were attached to a Wild compound microscope. The teeth are middle Eocene teeth of No. serratissimus have a smaller photographed with a Nikon Coolpix 995 digital camera- Agassiz, 1843 cult to discern, but brightness and contrast were adjusted mesial serrations on the acrocone increase in size apically white and details in unretouched photographs were diffi slightly for clarity using Adobe Photoshop software. (crown (less than, 17 mm width),, fewer distal, Van cusplets, den Eekhaut and and De Schutter 2009). In contrast, middle to late Eocene beyond the generic level was hampered due to small No.Cappetta kempi teeth1976 areWard larger 1979a thanNolf the 1988 aforementioned taxon In many cases, identification of the elasmobranch teeth sample size and diagenetic alteration of the remains. (greater than 30 mm width) and the mesial serrations are Overall, teeth have a chalky preservation, enameloid large and of relatively uniform size ( ). The has been partly or completely leached away, and roots Dry Branch hexanchid appears to have been as large as are eroded and/or incomplete. Comparison of the Dry No. kempi and the evenness of the acroconeWard serrations 1979a on Branch material to a sample of several thousand teeth at the South Carolina State Museum, and to specimens SC96.97.4 (Fig.SQUATINIFORMES 2C, D) is also consistent with this species. from the Clinchfield Formation of central Georgia, housed- SQUATINIDAE lina, housed at the McKissick Museum at the University SQUATINA Buen, 1926 ofrecovered South Carolina, from the allowed Griffins us Landing to more Sand accurately of South identify Caro Squatina Sq. primaBonaparte, ( 1838 ) some of our Dry Branch Formation material. The features Dumeril, 1806 that we used to identify specimens and distinguish be- sp. cf. Winkler, 1874 Referred specimens(Fig. 2E) tween similar morphologies are provided in the Remarks section for each taxon. Higher level largely incomplete tooth. —SC96.97.9, tooth (Fig. 2E), follows Naylor et al. (2012). SC96.97.10,Remarks—The tooth; four SC2001.1.40, Dry Branch tooth; Formation SC2013.38.2, teeth are Institutional abbreviations—SC, South Carolina morphologically similar to those of Eocene Squatina State Museum, Columbia , South Carolina; MSC, McWane prima reported elsewhere ( , Kemp et al. 1990, Science Center, Birmingham, Alabama. Parmley and Cicimurri 2003 - tion is tentative because of theCase small 1981 and imperfectly pre- SYSTEMATIC PALEONTOLOGY served sample. Additionally,), Squatina and our toothspecific morphology identifica has remained conservative and relatively stable since the HEXANCHIDAE Huxley, 1880 elongated lateral shoulders and low, roughly triangular Buen, 1926 , largely consisting of a narrow cusp flanked by Gray, 1851 CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA

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Figure 2. Shark teeth from the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–D. Notorynchus sp. cf. No. kempi teeth. Lower left tooth in lingual (A) and labial (B C) and lingual (D) views, E. Squatina sp. cf. Sq. prima F, G. Nebrius sp. cf. Neb. thielensi. Antero- lateral tooth in labial (F) and occlusal (G ) views, SC2013.38.1. Lower. H. . right tooth in labial (Tooth in labial view, SC96.97.4. I–O. sp. Third. lowerAntero-lateral right anterior tooth toothin labial in labialview, (SC96.97.9.I in lingual (J) and labial (K ) views, SC2013.38.5. Labial at top in G L SC2013.38.7.anterior tooth in labial (M ) view, SC2001.1.55.N Third upper right anterior tooth tooth in lingual (O ) views, SC2001.1.57.P–S. Alopias First sp. lower Tooth right in labial anterior (P) and tooth lingual in labial (Q ( ) view, SC2001.1.59. Second lowerR right) and lingual (S) views, SC2001.1.66.) view, Scale SC96.97.44.1. bars=1 cm Second in A–D, upper I–K, N right, anterior –toothG, L– inM, lingualP–S; 1 mm( ) view,in H. SC96.97.42. Lower left lateral ) view, SC96.97.43. ) views, SC2001.1.65. Tooth in labial ( O; 5 mm in E 6 PALEOBIOS, VOLUME 36, JUNE 2019

(in basal view) root ( ). Carolina coastal plain deposits is unknown, and the fos- sils could be as old as Eocene or as young as Pleistocene ORECTOLOBIFORMESCappetta 1987 (see : pl. 34). Although Neb. obliquus was GINGLYMOSTOMATIDAE based on a single tooth from the Eocene of New Jersey, Applegate, 1972 NEBRIUS it hasLeidy been reported1877 as being common in lower Eocene Nebrius Neb. thielensi (Gill, 1862 ) Rüppell, 1837 () strata elsewhere (Noubhani and Cappetta sp. cf. Winkler, 1874 , Cappetta 2012). Referred specimens(Fig. 2F, G) Morphologically, Neb. thielensi was differentiated from - 1997Neb. blackenhorni in having a crown that is wide but —SC96.97.1, three incomplete rather low, a larger and less distally inclined main cusp, teeth; SC2001.1.41, incomplete tooth; SC2013.38.3, an fewer lateral cusplets, and a shorter but conspicuously teeth.terior tooth; SC2013.38.4, posterior tooth; SC2013.38.5, , anteriorRemarks tooth—We (Fig. identify 2F, G); SC2013.38.6,these nine orectolobid two incomplete teeth ). In contrast, Neb. blackenhorni has a as Nebrius because they have a wide crown bearing a ratherflat or slightlyhigh crown bifid in labial proportion protuberance to width, (Winkler there are 1874 ten orArambourg more cusplets 1952 on the mesial side of the main cusp, and cusplets. In contrast, teeth of morphologically similar the labial protuberance is more elongated and rounded Ginglymostomacentral cusp that is flanked by up to seven have pairs up of tolateral two basally. These generalities are seen in the suite of teeth il- Herman lustrated in Arambourg ( , pl. 22), as well as Stromer et al. 1992, PurdyMüller et al. and 2001 Henle,). Unfortunately, 1837 identify- Ginglymostoma ingpairs fossil of lateral Nebrius cusplets teeth flanking is often the hampered main cusp by ( the fact fourtaui ( ) (1952 , 1909). that original descriptions of fossil species are based on 1905Noubhani, and teeth and originally Cappetta identified ( ) used as features like relatively few specimens, and the degree of morphologi- Priem, 1905 Priem 1907 - cal variation within true biological species is unknown. tuberance, and concavity of the1997 basal attachment surface There is a lack of agreement as to species occurrences totooth separate size, labial Neb. thielensicrown profile, from Neb. shape obliquus of the . labialSome prolate within Eocene strata of North America, with Neb. thielensi Eocene records of Nebrius (Case and Borodin 2000, Parmley and Cicimurri 2003), as Neb. thielensi (Case and Borodin 2000, Parmley and Neb. blackenhorni (Stromer, 1903) (Thurmond and Jones Cicimurri 2003) and Neb. obliquus, including ( teeth identified), appear ), Neb. serra ( ), and Neb. obliquus ( ) (Ginglymostoma obliquum Case 1981 1981of ) beingManning recognized and Standhardtin the literature. 1986 Kent to be conspecific, as all have a large and rather erect main (1994) reportedLeidy Neb. 1877thielensi Neb. Drycusp Branch flanked Nebrius by 7–9 teeth cusplets, lack andenameloid, a basal makingprotuberance direct blackenhorniCase 1981 - that is moderately elongated and often weakly bifid. The jemoy Formation of Maryland and (p. 34, Virginia fig. 8.3b) (Kent and 1999 ), comparable to Neb. thielensi (note that the name has and he considered (p. 34, fig.the 8.3a) possibility from the that lower the Eocene morpholo Nan- alsocomparisons variously to been other published specimens as Neb. difficult, thielensis but andthey Neb. are gies represented heterodonty within a single species. thielense; see Cappetta [2012] for further discussion on Woodward ( spelling) in that they are larger than Neb. obliquus sensu Eocene of Alabama as Gi. serra ( ), but Leriche Leidy ( (1942 1889: pl. 16, fig. 9) identified:146) synonymized teeth from the record with Neb. obliquus. However,Leidy Thurmond 1877 and Jones protuberance.1877: pl. 34, fig. 14), have a larger main cusp, ( :44):27, reported28) and White that the (1956 middle Eocene Gosport Sand fewer lateral cusplets, and shorter and weakly bifid labial of Alabama contained a continuous morphological series between1981 the obliquus and serra morphologies, and they synonymized obliquus with serra. Unfortunately, Thur- GINGLYMOSTOMATIDAE gen. et sp. indet. mond and Jones ( ) only provided an illustration of Referred specimen(Fig. 2H) - Leidy’s ( ) original material, not specimens from terior tooth. the Gosport Sand.1981 Noubhani and Cappetta ( ) con- Remarks—This specimen—SC2013.38.7, differs from incomplete Nebrius sp. an cf. 1877 Neb. serra Neb. thielensi (see above) in having only a single pair of to be incorrect and preferred instead to identify1997 teeth large lateral cusplets, and the labial crown foot is dis- assidered Neb. obliquusthe identification (also Darteville of Eocene and teeth Casier as 1943 ). The precise geologic age of Neb. serra, collected from South tinctively bifid but not drawn out into an elongate basal protuberance. SC2013.38.7 is similar to the anterior teeth CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA

7 of Plicatoscyllium , tooth. but there is no indication of labial crown ornamenta- Remarks—Specimens bear a broadly triangular, dis- tion. The single pair of lateral cuspletsCase and also Cappetta, distinguishes 1997 tally curving crown. Unfortunately, the enameloid is lack- ing due to taphonomic processes, and the dentine cores of Ginglymostoma ( ). In are exposed. There is no indication that lateral cusplets SC2013.38.7 from the various early species ProtoginglymostomaNoubhani ypresiensis and (Casier, Cappetta 1946 )1997 from the cores of what appear to be two mesial basal cusplets termsearly to of middle gross Eocenemorphology, of Belgium SC2013.38.7 and Morocco is similar (Casier to andwere an present incipient on distal SC2001.1.65 cusplet are (Fig. visible 2P, Q),on SC2001.1.66but dentine 1946, , ), but the poor (Fig. 2R, S). Unfortunately, the enameloid on the two condition of the only available tooth makes a more pre- specimens is imperfectly preserved, and the shape of Herman 1977 Tabuce et al. 2005 the very limited sample size (n=2), we could therefore cise identificationLAMNIFORMES difficult. notthe crowneffectively may notcompare be accurately the Dry reflected. Branch AlopiasCoupled teeth with ODONTASPIDIDAE to those of any of the several Eocene species that have Berg, 1958 CARCHARIAS been described, which include Al. crochardi , CarchariasMüller and Henle, 1838 Rafinesque, 1810 Al. leeensis , Al. denticulatus , cf. sp. and Al. alabamensis . The crownWard, foot of 1978 both Referred specimens(Fig. 2I–O) Dry Branch Ward,Formation 1978 teeth is very convexCappetta, and obviously 1981 overhangs the root (Fig.White, 2Q, 1956 S), and the lingual side of the —SC96.97.42, second upper root is bisected by a shallow nutritive groove (Fig. 2P, R). right anterior tooth (Fig. 2N); SC96.97.43, lower left These features are consistent with teeth of Al. crochardi lateral tooth (Fig. 2O); SC96.97.44.1, second lower and Al. denticulatus, but our specimens appear to have right anterior tooth (Fig. 2M); SC96.97.44.2, four teeth;- lacked cusplets like those seen on Al. denticulatus. Teeth SC96.97.45, 236 incomplete teeth; SC2001.1.55, third of Al. leeensis and A. alabamensis lack nutritive grooves lower right anterior tooth (Fig. 2I); SC2001.1.56, an and crowns appear much wider than seen on the Dry terior tooth; SC2001.1.57, third upper right anterior Branch Formation specimens. Our two specimens are 2L);tooth SC2001.1.60, (Fig. 2J, K); SC2001.1.58, three posterior six incompleteteeth; SC2001.1.61, anterior similar to teeth we observed within the jaws of Al. super- teeth; SC2001.1.59, first lower right anterior tooth (Fig. ciliosus ( ) housed at SC, and perhaps the Dry Branch Formation taxon is related to this “group” (sensu 40 incomplete teeth; SC2013.38.8, anterolateral tooth; Cappetta,Lowe, 2012 1841). A larger and better preserved sample SC2013.38.9, four partial teeth; SC2013.38.10, three twopartial incomplete teeth; SC2013.38.11, anterior teeth. lateral tooth; SC2013.38.12, 14Remarks tooth fragments;—The 332 SC2013.38.13, teeth in the tooth; sample SC2013.38.14, are simi- is needed for a more specific determination. lar to extant Carcharias taurus , and GALEOCERDIDAE examination of two Recent dentitions of this species Galeocerdo Compagno, 1973 Rafinesque, 1810 HermanGaleocerdo and van den Eeckhaut, 2010 Genus Müller and Henle, 1837 various anterior and lateral positions in the fossil spe- sp. (SC2001.120.6 and SC86.62.2) allowed us to identify cies. Unfortunately, diagenetic alteration of the teeth Referred specimens(Fig. 3E, F) - of various species that have been reported from Eocene SC2001.1.9, lateral tooth;—SC96.97.11, SC2001.1.10, threethree incompartial makes confidently assigning the fossil material to any plete teeth; SC2001.1.8, incomplete anterior tooth; lateralstrata difficult,cusplets andas characteristics crown ornamentation, that have are been generally used teeth; SC2013.38.16, anterolateral tooth (Fig. 3E, F); notfor speciesevident identification, or imperfectly like preserved. the number and shape of SC2013.38.17,Remarks incomplete lateral tooth; SC2013.38.18, areincomplete morphologically tooth; SC2013.38.19, consistent with four Galeocerdo incomplete, having teeth. ALOPIIDAE an elongated—All and broadly 15 specimens convex lackmesial enameloid edge, short but distal they ALOPIAS edge that forms a triangular cusp with the mesial edge, AlopiasBonaparte, 1838 Rafinesque, 1810 and elongate, serrated distal heel. In terms of gross mor- sp. phology, the Dry Branch Formation specimens differ from Referred specimens (Fig. 2P–S) Eocene Galeocerdo eaglesomei White, 1926 in having a

—SC2001.1.65, tooth; SC2001.1.66, PALEOBIOS, VOLUME 36, JUNE 2019

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Figure 3. Carcharhiniform shark teeth the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–D. Physogaleus sp. cf. Ph. latus. Upper left lateral tooth in lingual (A) and labial (B C) and lingual (D E, F. Galeocerdo sp. Anterolateral tooth in lingual (E) and labial (F G–I. Abdounia sp. cf. Ab. enniskilleni. Anterior tooth in lingual (G) and labial) views, (H SC2013.38.65. Lower left anterior tooth in labialI) view, ( J–L) .views, SC96.97.26. ?ganntourensis . Anterolateral tooth in labial (J) and lingual (K ) views, SC2013.38.16. in lingual view (L M–P. Hemipristis sp. cf. He. curvatus. Upper) views, left lateralSC96.97.30. tooth Lateralin lingual tooth (M )in and lingual labial ( (N) views, SC96.97.31.SC2001.1.21. Lower right anterior tooth in lingual (O) and labial (P Q, R). Negaprionviews, SC2013.38.98. gilmorei. Upper Lateral right tooth anterior tooth in lingual), SC2013.38.99. (Q Lower right anterolateral tooth in lingual (R ) views, SC2013.28.35. ) view, SC2013.38.61. ) view, SC2013.38.64. Scale bars=1 cm in A–F, M–N, Q–R; 5 mm in G–L, O–P. CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 9 lower and more uniformly convex mesial cutting edge, Physogaleus Ph. latus ( ) as opposed to a rather high and conspicuously convex sp. aff. Storms, 1894 apical portion to the mesial edge. We believe that the Referred specimens(Fig. 3A–D) Dry Branch Formation Galeocerdo teeth are comparable —SC96.97.26, anterior tooth;- Ga. alabamensis Leriche, 1942 by Parmley SC96.97.27, lateral tooth; SC96.97.28, lateral tooth; to specimens from the Clinchfield Formation of Georgia and Cicimurri (2003 - SC96.97.29, 18 incomplete teeth; SC2001.1.51, ante identified as tion made by those authors appears to be in error, as rior tooth; SC2001.1.52, lateral tooth; SC2001.1.53, ). However, the specific identifica examination of Leriche’s (1942) alabamensis holotype posterior tooth; SC2001.1.54, five incomplete teeth; leads us to conclude that it is actually Physogaleus Cap- SC2013.38.56, lateral tooth; SC2013.38.57, anterior tooth;Remarks SC2013.38.58,—Dignathic lateral heterodonty tooth; SC2013.38.59, is evident among tooth; by Parmley and Cicimurri (2003) was Ga. alabamensis theSC2013.38.60, 39 Dry Branch six incomplete specimens, teeth. with upper teeth having petta, 1980. The assignment of the Clichfield material to based on White’s ( a broader central cusp and better developed serrations from South Carolina as Ga. alabamensis, which are not on the lateral shoulders (Fig. 3A, B) compared to lower 1956) identification of specimens 1942) taxon. teeth. Lower teeth have a very narrow, sigmoidal cusp In addition to Alabama ( , Thurmond and Physo- conspecific), with Leriche’s ( was also documented from Ga. alabamensis teeth are similar in gross morphology to those of Arkansas ( ) andWhite Louisiana 1956 (Manning and andGaleocerdo significantly, but they thickened are smaller root (Fig. in overall 3C, D). size The and de- Jones 1981 ). Another species, Ga. clarkensis, was velopment of serrations (especially on the mesial edge) Westgate 1984 founded by White ( ) based on a small number of is much reduced. For example, only the basal half of the Standhardt 1986 teeth from the late middle Eocene of Alabama, but Man- mesial cutting edge of Physogaleus teeth is serrated, 1956 ning and Standhardt ( ) synonymized this taxon with whereas the apical half is smooth. This phenomenon Ga. alabamensis, considering the former to represent up- can be discerned even on teeth lacking enameloid, as the 1986 per teeth and the latter lower teeth in the same dentition. dentine core preserves small denticulation basally, but However, as noted above, the “Galeocerdo” alabamensis the edge is smooth apically. Physo- Several of the species that White (1926) described galeus. In Georgia, Case ( Ga. clarkensis from the Eocene of Nigeria, including Eugaleus falconeri, morphology is more appropriately identified as - Sphyrna itoriensis, Sph. tortilis, and niger- 1981) identified ber) Formations, but Case and Borodin (2000) later iensis, have been placed within Physogaleus (Cappetta, from the Clinchfield and Dry Branch (Twiggs Clay Mem reported Ga. latidens from the Dry Branch 2006), but it is as yet unclear if the morphologies rep- Formation (Irwinton Sand Member). It is apparent that resent multiple species or heterodonty within a single Agassiz, 1843 Eocene records of Galeocerdo from North America are in species (monognathic, dignathic, and gynandric heter- further need of evaluation. odonty). The teeth we tentatively identify as Ph. latus Our examination of Ga. clarkensis teeth from the compare favorably to teeth described by Storms ( ) middle Eocene (Bartonian) Gosport Sand (Claiborne from the lower of Belgium, and several of the Group) of Alabama, housed at MSC, showed that the Dry Branch teeth (i.e., Fig. 3A, B) are practically identical1894 mesial cutting edge bears coarse, compound serrations (largest serrations bear smaller serrations). Additionally, particular teeth differ from other Eocene and Oligocene examination of Ga. eaglesomei teeth from the middle speciesto the specimen of Physogaleus he illustrates in that in the plate cusp 6, figureis broad 17c. with These bi- Eocene (Lutetian) Lisbon Formation (Claiborne Group) convex cutting edges, and the cusp is well differentiated of Alabama, also at MSC, revealed that serration on from the coarsely serrated distal heel and basal half of the the mesial edge is simple (large serrations do not bear mesial cutting edge ( , Reinecke and others smaller serrations). As our specimens are incompletely , , ). Teeth that preserved, we cannot determine the precise shape of Winkler 1876 the original crown or if the serrations were simple or are2005 comparableHaye et al. to2008 the Reineckespecimen et illustrated al. 2008 by Storms compound (serrations on serrations), and we refrain (we identify as being from anterior files (i.e., Fig. 3C, D) Case ( ) named Galeorhinus huberensis (=Ph. hube- rensis1894); frompl. 6, the fig. Dry 18). Branch Formation of Georgia, and we from CARCHARHINIDAEmaking a specific identification. PHYSOGALEUS 1981 Phy. secundus Jordan and Evermann, 1896 Cappetta, 1980 consider that taxon to be conspecific with 10 PALEOBIOS, VOLUME 36, JUNE 2019

( ) as reported by Parmley and Cicimurri is generally cuspidate ( , Herman et al. (2003 1991, Case et al. 1996, , TheWinkler, South 1876 Carolina Dry Branch taxon appears to differ ), butArambourg our specimens 1952 lack enameloid from the) from Georgia the Clinchfield material Formationin that the of teeth central can Georgia. attain and this feature is not preserved.Noubhani andHowever, Cappetta we exam1997- larger size, the cusp is wider, distal serrations are smaller Mustafained a tooth et al. of 2005 Rhiz. sp. cf. Rhiz. ganntourensis that was and more numerous, and the mesial serrations may be recovered by Zullo and Kite ( larger or more numerous. Landing Sand (see Appendix 1), and found that it is very similar to the material listed above.1985 The) from well the preserved Griffins ABDOUNIA Abdounia enniskilleni ( ) heel, but the translucent enameloid allows us to view Cappetta, 1980 theGriffins unifromly Landing convex Sand dentine tooth exhibits core of thea cuspidate heel, as can distal be White, 1956 seen on the specimens in our sample. Referred specimens(Fig. 3G–I) SC2001.1.43, anterior tooth;—SC96.97.30, SC2001.1.44, three anterior teeth; Negaprion gilmorei ( ) tooth; SC96.97.31, lateral tooth; SC96.97.32, 43 teeth; Whitley, 1940 Remarks—Gradient monognathic heterodonty is Leriche 1942 SC2013.38.15, incomplete tooth. Referred specimens(Fig. 3Q, R) - anterior teeth having tall vertical main cusps (Fig. 3G, H),evident but cuspsamong becoming the 50 specimens lower, broader, we examined, and distally with —SC96.97.37, upper right ante curved (Fig. 3I) in lateral and posterior positions. Fine rior tooth; SC96.97.38, upper right antero-lateral tooth; vertical enameloid ridges are preserved on specimens upperSC96.97.39, tooth; upper right lateral, tooth; tooth; SC2001.1.49, SC96.97.40, lower 101 from the NAS, and this feature, combined with the gross tooth;upper teeth; SC96.97.41, 24 lower teeth; SC2001.1.47,- morphology of teeth, leads us to assign them to Abdounia SC2001.1.48 enniskilleni. This species has been reported from several SC2001.1.50, 75 teeth; SC2013.38.61, upper an upper Eocene localities in the Gulf and Atlantic coastal terior tooth; SC2013.38.62, upper right lateral tooth; enniskilleni prior SC2013.38.63, upper right tooth; SC2013.38.64, lower Manning and Standhardt tooth; SC2013.38.65, seven lower teeth; SC2013.38.66, plains), (often Arkansas identified ( as ), Alabama (White sevenRemarks upper anterior teeth; SC2013.38.67, 29 teeth;- to 1987),), North including Carolina Louisiana ( ( ), mensSC2013.38.68, that exhibit 14 lower dignathic teeth. heterodonty, with upper and1986 Georgia ( Westgate, Case and 1984 Borodin 2000, Parmley —This species is represented by 267 speci and1956 Cicimurri 2003). WithinTimmerman the Dry and Branch Chandler sample, 1995 Ab. lateral shoulders (Fig. 3Q), whereas lower teeth have a enniskilleni canCase be differentiated1981 from small odontaspidid shorterteeth having and narrowera broader cusp,cusp flankedand smooth by elongate, shoulders oblique that teeth in that the lateral cusplets are large in relation to are nearly perpendicular to the cusp (Fig. 3R). Although the enameloid is often missing, the remaining dentine are shorter and more triangular in outline. core of the lateral shoulders on some upper teeth shows the main cusp, the lingual root face is flat, and root lobes indications that they were at least weakly serrated. RHIZOPRIONODON Leriche (1942) erected Sphyrna gilmorei based on Rhizoprionodon ?ganntourensis ( ) Whitley, 1929 teeth occurring in upper Eocene deposits of Alabama. Arambourg, 1952 White ( ) assigned the morphology to the subspe- Referred specimens(Fig. 3J–L) cies Negaprion gibbesi gilmorei, and Müller (1999) and Adnet et1956 al. ( ) used the name Carcharhinus gilmorei. —SC96.97.2, four teeth; Underwood et al. (2011) and Underwood and Gunter SC2001.1.18, tooth; SC2001.1.19, 40 teeth; SC2013.38.98, (2012 2007 anterior tooth (Fig. 3J, K); SC2013.38.99, lateral tooth (Fig. such as those reported here, as Negaprion rather than eight3L); SC2013.38.100, distal lateral teeth. 18 teeth; SC2013.38.101, 32 teeth; Carcharhinus) identified. If our unserrated association to is weakly correct, serrated the dignathic teeth, SC2013.38.102,Remarks—One eight hundred posterior thirteen teeth; Dry SC2013.38.103, Branch Forma- heterodonty of Neg. gilmorei is more pronounced than tion teeth are similar to those of extant Rhizoprionodon, in a dentition of Recent Neg. brevirostris ( ) we and to Eocene Rhiz. ganntourensis, in particular. The dis- examined (SC uncurated). tal heel of Rhizoprionodon, including Rhiz. ganntourensis, We believe that Eocene material from Poey,North 1868 Carolina CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 11 and Georgia that was referred to Neg. eurybathrodon of the extant species are weakly serrated and edges of ( Neg. lower anterior teeth are often limited to the upper half gilmorei ( , Case and Borodin 2000, Parmley and of the crown. In addition, the Eocene teeth have shorter CicimurriBlake, 1862 2003) is). moreTeeth appropriately of Neg. identified eurybathrodon as cusps, many exhibit very convex lateral shoulders (espe- are up toCase 2 cm 1981 in height and width, much larger than cially more lateral positions), and the root lobes are more Eocene specimens attributed to this species. The main elongated and divergent (Herman et al. 1991, Compagno cusp of Neg. eurybathrodon upper teeth is also much ). taller but narrower than the Eocene specimens, and the The new species differs from the fossil species I. acuar- lateral shoulders are more evenly serrated ( ). iuset al. ( 2005 ), I. lerichei (Darteville and Casier 1943), Negaprion gilmorei is common in the Dry Branch Forma- I. caunellensis ( ), and I. gracilis (Jonet tion, and it is the dominant elasmobranch Whitetaxon within1955 1966Probst) in being 1879 smaller in overall size. Additionally, the Dry Branch speciesCappetta has narrower 1970 upper anterior teeth thousand teeth within two SC accessions (SC2004.34 and and complete cutting edges on all teeth when compared SC2013.44;the Clinchfield also Formation Parmley and of Georgia,Cicimurri based 2003 on). several to I. caunellensis, and the convexity of the lateral shoul- ders of anterior teeth appears to be more pronounced ISOGOMPHODON than on I. acuarius ( , , Müller Isogomphodon aikenensis 1999). The transition from main cusp to lateral shoulders Genus Gill 1862 appears to be slightlyCappetta more angular 1970 onCase teeth 1980 of I. lerichei Cicimurri and Knight n. sp. (Darteville and Casier 1943). (Fig. 4E–P) Holotype Description—These 393 distinctive teeth generally tooth. Paratypes—SC2013.38.110 (Fig. 4E), upper anterior- narrowmeasure cusp, less and than enameloid 5 mm in shoulderstotal height, that although extend nearly some —SC2013.38.111 (Fig. 4P), upper antero toanterior the tips teeth of the are root up lobes. to 7 mm. The cuttingAll teeth edges have of a alltall teeth and lateral tooth; SC2013.38.119 (Fig. 4H, I), lower anterior are smooth and extend to the tips of lateral shoulders. tooth;Referred SC2013.38.112 specimens (Fig.—SC2001.1.62, 4J, K), upper upperlateral tooth; SC2013.38.115 (Fig. 4N, O), posterior tooth. that is slightly asymmetrical (Fig. 4E). The crown consists - ofThe a tall,holotype, narrow SC2013.38.110, and erect cusp, is anwith upper cutting anterior edges tooth that SC2001.1.63, lower tooth; SC2001.1.64, 97 teeth; diverge slightly towards the crown base. Elongate lat- SC2013.38.113, symphyseal tooth; SC2013.38.114, pos eral shoulders extend obliquely onto the root lobes. The terolateral tooth (Fig. 4L, M); SC2013.38.116, upper tooth; root is bilobate, with rather short and sub-rectangular SC2013.38.117, 15 symphyseal teeth; SC2013.38.118, lobes that are separated by a narrow but deep U-shaped anterior tooth; SC2013.38.120, four anterior teeth; interlobe area. The mesial lobe is slightly smaller than SC2013.38.121, six anterior teeth; SC2013.38.122, 52 the distal lobe, and a deep but narrow nutritive groove teeth; SC2013.38.123.1 lower lateral tooth (Fig. 4F, G), SC2013.38.123.2,Occurrence—Type 84 teeth; locality: SC2013.38.124, South Aiken 112Site teeth;(SAS) teeth (Fig. 4P) are wider than those near the jaw sym- SC2013.38.125, 16 posterior teeth. bisects the flat lingual root face. Upper anterolateral- yellow, orange and red variegated sand, Upper Eocene terior positions, but slightly distally inclined. The lateral (Priabonian)33.504444, -84.742778, Dry Branch AikenFormation, County, approximately South Carolina; two shouldersphysis. The are cusp more is narrowelongated and and flat nearly as seen horizontal. in more Thean meters below the contact with the overlying Tobacco sub-rectangular root lobes are lower, more elongated and Road Formation. divergent than on anterior teeth, and the interlobe area Etymology—The species name recognizes the city of is wider but shallower. Aiken and Aiken County, South Carolina, the only known Lower anterior teeth (Fig. 4H, I) have a very tall and area from which Eocene Isogomphodon have thus far narrow cusp, with sub-parallel cutting edges. Lateral been documented from North America. shoulders are short and nearly horizontal. The bilobate Diagnosis—Nearly 400 specimens are referred to the root is bisected by a deep, narrow nutritive groove, and new species, which differs from Recent Isogomphodon the short lobes are widely separated by a U-shaped oxyrhynchus ( ) in that all teeth interlobe area. Lower lateral teeth (Fig. 4F, G) are simi- have smooth cutting edges extending from the apex to lar to those in the upper jaw, but the cusp is narrower, the very base Müllerof the crown. and Henle In contrast, 1841 the upper teeth lateral shoulders are shorter, the transition from cusp 12 PALEOBIOS, VOLUME 36, JUNE 2019

Figure 4 . Carcharhiniform shark teeth from the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–D. Carcha- rhinidae indeterminate. Anterolateral tooth in labial (A) and lingual (B C) and lingual (D) views, SC2001.1.43. Labial at right in C. E–P. Isogomphodon aikenensis n. sp. Upper anterior tooth in lingual view (E), F) and lingual) ( views,G SC2013.38.105. Anterolateral tooth in labial ( (H) and lingual (I J) and lingual (K - type).SC2013.38.110 Posterolateral (Holotype). tooth in Lower labial lateral (L) and tooth lingual in labial (M ( ) views, SC2013.38.123.1. LowerN) and anterior lingual tooth (O) inviews, labial ) views, SC2013.38.119 (Paratype). Upper lateralP tooth in labial ( ) views, SC2013.38.112– (Para in E–P. ) views, SC2013.38.114. Posterior tooth in labial ( SC2013.38.115 (Paratype). Upper anterolateral tooth in lingual ( ) view, SC2013.38.111 (Paratype). Scale bars=1 cm in A D; 5 mm to shoulder is more angular, and the mesial root lobe is a centrally located narrow but deep nutritive groove. shorter than the distal one. The root is low and bilobate, Remarks—A number of fossil species have been as- with rounded and diverging lobes that are separated by signed to Aprionodon , a junior synonym of a wide and shallow U-shaped interlobe area. A narrow Isogomphodon, including Ap. acuarias ( ), but deep nutritive groove is located on the lingual face. Ap. amekiensis White, Poey,1926 ,1868 Ap. caunellensis Cappetta, Posterolateral and posterior teeth (Fig. 4L–O) are , Ap. collata (Eastman, 1904), Ap. elongatusProbst, (Leriche, 1879 small with a T-shaped outline. The cusp is triangular 1910), Ap. gibbesi ( ), Ap. gracilis (Jonet, but sharply tapering apically, rather low compared to 19661970), Ap. frequens ( ), Ap. lerichei Darteville teeth in more anterior positions. The cusp is vertical to and Casier, 1943, Ap.Woodward, macrorhiza 1889 Jonet, 1966, Ap. mar- çaisi Dames,, and a1883 subspecies, Ap. lerichei var. are short to elongate, perpendicular to the cusp, and the minuta Jonet, 1966. Although some of these species are transitionslightly distally from inclined, cusp to shoulderflat in profile. is more Lateral angular shoulders than at leastArambourg, in part based 1952 on specimens correctly assigned seen on more anterior jaw positions. The lateral shoul- to the genus, most are appropriately referred to other ders in these positions are very convex at their distal genera or are perhaps nomina dubia. extremities, resulting in a cusp-like appearance in lingual For example, Carcharhinus collata Eastman, 1904 view. The root is bilobate with short, rounded, highly was referred to Aprionodon by Powlowska (1960), but diverging lobes, which are separated by a broad, shallow illustrations of three teeth in Eastman (1904 to deep U-shaped interlobe area. The lingual face bears Carcharhinus and, possibly, : pl. 22, figs. 3-5) show them to represent CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 13

Negaprion. Antunes and Jonet ( ) concluded that are larger than I. aikenensis, have wider upper anterior Jonet’s (1966) Ap. macrhorhiza represents symphyseal teeth, and lower anterior teeth with incomplete cutting teeth of indeterminate Carcharhinidae.1970 The teeth that edges. Probst ( White (1926) illustrated as Ap. amekiensis I. acuarius from his sample of approximately 11-26) largely appear to be Carcharhinus (with some pos- 1879) illustrated a specimen (pl. I.1, aike figs.- sibly representing Negaprion), and Arambourg’s (pl. (8, figs.) 76-77)nensis. However,of his description of the material (p. 140) Ap. marçaisi Carcharhinus 50is generic teeth that for isthe not genus, dissimilar but the to maximumanterior teeth tooth of size he marçaisi by Noubhani and Cappetta ( 1952 reported (10 mm) is larger than that of I. aikenensis of the Ap. gibbesi morphology ( was identified) and as Ap. elongatus mm). (Leriche 1910) species have weakly to1997 coarsely: 151). serrated Teeth Isogomphodon lerichei was based on a suite of Miocene (7 heels, particularly theWoodward uppers, and 1889 both species have been teeth described by Darteville and Casier (1943). However, placed within Carcharhinus (Reinecke et al. 2001, , this taxon also appears to be a mixture of multiple spe- , Cicimurri and Knight 2009). cies, with teeth of Isogomphodon and non-Isogomphodon With regard to Jonet’s (1966) Ap. lerichei var. minuta2005 species included in the illustrated suite of teeth provided subspecies,Haye et al. 2008 Cappetta ( ) considered the material to by Darteville and Casier (1943 represent the lateral teeth of Isogomphodon acuarius, I. lerichei, but but he later (2006) treated1970 the morphology as a valid examination of the daggernose: i.e., shark pl. 13, teeth figs. 41,shown 42, 49,by species distinct from I. acuarius without providing sup- Darteville50). This calls and intoCasier question (1943 the validity of porting evidence. However, our examination of the teeth show that they are much larger and the angle between illustrated in Jonet’s (1966) plate II leads us to conclude the cusp and lateral shoulders: i.e., pl.may 13, be figs.37, sharper 39, 43-46)than I. that many of them, including the type specimen shown aikenensis. Antunes et al. ( ) tentatively reported Isogomphodon. Specimens shown in I. lerichei from the Miocene of Portugal, but the two 1966) plate II appear to 1981 bein figureIsogomphodon 1, are not more appropriately referred to Carcharhinus. offigures further 7, 11evaluation. and 13 in Jonet’s ( referredIsogomphodon teeth (pl. oxryrhynchus2, figs. 13 and is17) the are, only in our extant opinion, spe- Dames’ ( ), butCarcharhinus the specific (Aprionodon attribution) is frequens in need cies of daggernose shark, and it is largely restricted to from the late Eocene of Egypt contains more than one coastal waters of northeastern (Lessa and species of shark.1883 His description of the teeth (pp. 143, others 1999, Lessa et al. 2016). Teeth of fossil Isogomph- odon, particularly I. acuarias, are predominantly known from Mio-Pliocene deposits of the Tethyan regions of are144) Isogomphodon and accompanying. However, illustrations the frequens (pl. morphology 3, figs. 7a–p) is Europe and Africa, but an increasing number of reports consideredindicate that valid some but specimens has been attributed he examined to Carcharhinus (figs. 7b, f) document the taxon from temporally equivalent strata in (i.e., ) and, more recently, Negaprion (Un- South America (i.e., Carrillo-Briceño et al. 2016). Some derwood et al. 2011). As is the case with Jonet’s (1966) of these latter records, however, are based on inaccurate gracilisCappetta morphology, 1970 the Isogomphodon teeth reported by Dames ( ) are in need of further study. (i.e., Mora 1999). Two reports of I. acuarius from North Cappetta ( ) believed that Jonet’s (1966) Ap. graci- Carolinaidentifications in the easternof teeth belongingUnited States to other have Carcharhinidaebeen published lis species was1883 actually the anterior teeth of I. acuarius. ( ; Müller 1999). The strata yielding the fossils However, Cappetta1970 (2006) later treated the species as were considered to be of Miocene age, but the deposits valid, but in doing so provided no comment supporting areCase actually 1980 of Oligocene age (Harris and Zullo 1991; the conclusion. We concur with Cappetta ( ) that Zullo et al. 1992; Denison et al. 1993). Considering the teeth illustrated by Jonet (1966 - limited geographic distribution and habitat preference sent anterior teeth, and they are not dissimilar1970 from those of extant I. oxyrhynchus, it may be prudent to re-evaluate of I. acuarius. Regardless of whether: pl. 2, Jonet’s figs. 14–21) (1966 repre) spe- Oligo-Miocene records of the genus. Discovery of I. I. acuarius, the teeth are aikenensis in the Dry Branch Formation unequivocally both larger and have more elongated cusps than I. aike- extends the temporal range of the genus back to the late nensiscies is n.valid sp. Caseor conspecific and Cappetta with ( 1990) placed Cappetta’s Eocene, and it is the only Eocene record in North America. ( ) Ap. caunellensis species within Carcharhinus, but later Cappetta (2006, 2012 1970 Isogomphodon caunellensis. If valid, I. caunellensis teeth CARCHARHINIDAE genus indeterminate ) supported identification as (Fig. 4A–D) 14 PALEOBIOS, VOLUME 36, JUNE 2019

Referred specimens - rior tooth; —SC2013.38.104, tooth; two juvenile upper tooth; SC2013.38.23, lower ante SC2013.38.105,Remarks—Unfortunately, tooth; SC2013.38.106, the tooth; eight SC2013.38.107, teeth lack incomplete upperSC2013.38.24, teeth. lower anterolateral tooth; enameloid.tooth; SC2013.38.108, However, they tooth; are SC2013.38.109, distinguished by three the teeth.pres- SC2013.38.25,Remarks—Monognathic lower lateral and tooth; dignathic SC2013.38.26, heterodonty five ence of a single pair of cusplets that mark the base of can be discerned within the sample of 39 teeth. Upper the main cusp and beginning of short enameloid shoul- anterior teeth are narrow and rather erect, whereas lat- eral teeth are broadly triangular with a crown apex that to Abdounia enniskilleni, but whereas the cusplets of is distinctively distally curved. The mesial edge of upper Ab.ders enniskilleni (Fig. 4A, C). form These the teethmesial are and superficially distal ends similar of the lateral teeth is completely smooth or weakly serrated crown, cusplets of Carcharhinidae indet. are compara- on its basal half, but the distal edge is coarsely serrated tively smaller and located near the middle of the crown nearly to the apex (Fig. 3M). Lower anterior teeth are (compare Fig. 3H, I to Fig. 4A, C). Some of the specimens comparatively narrower, erect, and serrations on cutting are very similar to White’s (1926) Hypoprion overricus edges are restricted to the crown base (Fig. 3P). (reassigned to Abdounia overrica by Cappetta 2006), but Hemipristis teeth are distinctive in having very large the teeth in the Dry Branch sample differ in that cusplets cusplets along the distal crown edge, a characteristic evi- appear to have been smaller. The Dry Branch teeth also dent even on teeth lacking enameloid (Fig. 3N). Hemipris- bear similarities to Case’s ( ) Negaprion furmiskyi tis curvatus He. wyattdurhami (Cappetta [2006] considers this species to belong within in older literature) appears to have been widespread in Abdounia 1980 the Eocene Atlantic (identified and as Gulf coastal regions, White,having 1956been morphologies is that the diminutive cusplets occur at the reported from North Carolina (Timmerman and Chan- ends of the), butlateral a significant shoulders difference on Ab. furimskyi between, whereas the two ), Georgia ( , Case and Borodin 2000, they are between the end of the shoulder and the base Parmley and Cicimurri 2003), Alabama ( , of the cusp of the Dry Branch teeth. dler 1995 Case 1981), Louisiana (Manning and Comparison of the Dry Branch material to some of the ), and Arkansas ( White ).1956 This Thurmondtaxon is easily and separated Jones 1981 from the temporally younger as Carcharhinus frequens StandhardtHe. serra 1986 in that teethWestgate are much 1984 smaller late Eocene teeth identified by Case and Cappetta (1990) in size and the mesial cutting edges of lateral teeth are similarities. Case and Cappetta’s (pl. 5, figs. (1990 100–103,) assignment 106–107; of nearly or completelyAgassiz, 1835 devoid of serrations. thepl. 7, Egyptian figs. 143–144, material 146–148, to Carcharhinus 151–159) frequensrevealed strikingis in er- ror because the teeth differ from most of the specimens SCYLIORHINIDAE illustrated by Dames ( FOUMTIZIA Foumtizia Gill, 1862 possess a broader, lower cusp and exhibit serrated lateral Noubhani and Cappetta, 1997 shoulders. The frequens1883 morphology: pl. 3, fig. was 7), consideredwhich clearly to sp. be a species of Negaprion by Underwood et al. (2011), Referred specimens(Fig. 5A–G) and they concluded that the sample examined by Case and Cappetta (1990) contained teeth of Negaprion and —SC96.97.5, incomplete- an undescribed species of Abdounia. tooth (Fig. 5D); SC96.97.6, incomplete tooth (Fig. 5E); SC96.97.7, incomplete tooth (Fig. 5F); SC96.97.8,–C) in; complete tooth (Fig. 5G); SCSC2001.1.42, incomplete

HEMIPRISTIS tooth; SC2013.38.27, incomplete tooth (Fig. A Hemipristis H. curvatusHasse, 1879 lateral tooth. Agassiz, 1835 SC2013.38.28,Remarks—These two incomplete scyliorhinid teeth; teeth areSC2013.38.159, large enough sp. cf. Dames, 1883 Referred specimens(Fig. 3M–P) naked eye. Tooth crowns consist of a tall, erect, basally (up to 4 mm in height) to be found in the field with the —SC96.97.33, upper lateral pair of large lateral cusplets. The labial crown foot is SC2001.1.20,tooth; SC96.97.34, incomplete upper upper lateral tooth; tooth; SC2001.1.21, SC96.97.35, up- thick,broad overhangs but sharply the tapering root, and cusp bears that a conspicuous is flanked by me a- lower lateral tooth; SC96.97.36, 13 incomplete teeth; dial concavity. A blunt cutting edge extends across the main cusp and generally only onto the inner side of the per tooth; SC2001.1.22, five upper teeth; SC2001.1.23, lateral cusplets. In terms of overall morphology these four incomplete teeth; SC2013.38.20, lower anterior tooth; SC2013.38.21, lower lateral tooth; SC2013.38.22, CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA

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Figure 5. Scyliorhinid shark teeth from the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–G. Foumtizia sp. Anterior tooth in labial (A), lingual (B C D An- terior tooth in labial (E F Anterior tooth in labial view (G H–N. Premontreia (Oxyscyllium) sp. cf. Pr.) ( andO.) gilberti profile. (Anterior) views, tooth SC2013.38. in labial 27. (H Anterior) and distal tooth (I in labial ( ) view, SC96.97.5. in I. Anterior? tooth in labial) view, ( JSC96.97.6.), lingual ( KAnterior) and basal tooth (L in labial view ( ), SC96.97.7. M) and lingual), ( NSC) views,96.97.8. O–CC. Scyliorhinus sp. Lateral tooth in labial (O), lingual (P), occlusal (Q), and basal) views, (R SC2013.38.29. Labial at left- terolateral tooth in basal (S ) views, SC2013.38.30.T), lingual Lateral (U), tooth and occlusal in labial ( (V SC2013.38.31.Lateral tooth in labial (W), lingual (X), and occlusal (Y Z)), views, lingual SC2013.38.36. (AA), and occlusal An (BB ) view, SC2013.38.35. LateralCC tooth in labial ( ) views, SC2013.38.39. Scale bars=1mm. ) views, SC2013.38.37. Lateral tooth in labial ( ) views, SC2013.38.38. Anterior? tooth in occlusal ( ) view, SC2013.38.40. Labial at top in L, R, S; at bottom in Q, V, Y, BB, CC. teeth compare well to Foumtizia, but the Dry Branch , Malyshkina 2006). The preservation of our mate- teeth are larger than any species previously attributed rial and small sample size (n=9) leads us to refrain from to that genus ( , Noubhani and Cappetta 1997

Cappetta 1976 making a more specific identification. The morphological 16 PALEOBIOS, VOLUME 36, JUNE 2019 features noted above, along with larger size and lack of height of the main cusp and lateral cusplets cannot be enameloid ornamentation, distinguish the teeth from two determined. Teeth are distinctive in that the labial crown other scyliorhinids occurring in the Dry Branch sample foot is highly concave so that two lobes, a short mesial and (see below). a more elongated distal lobe, are developed. One or two cusplets are found on the mesial lobe, but only one large PREMONTREIA cusplet occurs on the distal lobe. The specimens exhibit ( Premontreia Oxyscyllium) crown ornamentation and lateral cusplets, and they are Pr. (O.) gilbertiCappetta, ( 1992 ) sp. of similar size to other scyliorhinid teeth in our sample cf. Casier, 1946 that we identify as Premontreia (see above). However, Material examined Fig. 5H–N the root morphology and strongly asymmetrical nature of the crown are not typical of species within the genus —SC2013.38.29, tooth (Fig. 5H, Premontreia ( , Malyshkina I); SC2013.38.30, tooth (Fig. 5J–L); SC2013.38.31, tooth 2006, ). (Fig.Remarks 5M, N);—The SC2013.38.32, root morphology tooth; SC2013.38.33, of our six specimens tooth; The discontinuousNoubhani nature and Cappetta of the nutritive 1997 groove on leadsSC2013.38.34, us to assign incomplete the teeth tooth. to Premontreinae Cappetta, our teethMollen is consistent 2008 with the morphology of Scyliorhi- 1992, which currently contains the genera Premontreia nus Noubhani and Cappetta and Pachyscyllium . Noubhani ). The preservation of the Dry Branch teeth inhibits and Cappetta ( ) have since divided Premontreia into our speciesability identifiedto compare elsewhere them to ( known species, but in two subgenera: PremontreiaReinecke and and others, Oxyscyllium 2005 . Although terms1997 of overall morphology they are similar to only the dentine1997 core of the Dry Branch Formation Sc. entomodon of Morocco specimens is preserved, we can discern that the tooth and Sc. joleaudi from the lower Miocene crowns bore coarse basal longitudinal ridges and they of France. However,Noubhani the Dry and Branch Cappetta, teeth 1997 differ from the were distally inclined, which is in contrast to what is seen Moroccan speciesCappetta, in that the 1970 crown lobes are much more on teeth of the various Oligocene to Pliocene species of widely separated, and all specimens exhibit enameloid Pachyscyllium that have been described ( , ornamentation ( ). The main , , Reinecke et cusp of Sc. joleaudi appears to have been larger than the ). With respect to crown shape, Cappetta,the Dry Branch 1970 Dry Branch taxon,Noubhani the crown and lobesCappetta are 1997closer together, Reinecke and others 2005 Haye et al. 2008 Premon- and cusplets are also larger ( ). A larger treiaal. 2008 (Oxyscyllium) because sample size and more complete material will help with ofFormation the large teeth and are well identifiable differentiated to the subgenus lateral cusplets. Cappetta 1970 Additionally, we believeNoubhani that and the Cappetta, teeth compare 1997 favor- ably to Pr. (O.) gilberti, a species formerly placed within making RHINOPRISTIFORMESa more specific determination. Scyliorhinus (Casier 1946, , , Kemp RHINOBATIDAE Naylor et al., 2012 et al. 1990, Case et al. 1996, also ). RHINOBATOS RhinobatosBonaparte,. 1835 We also consider specimensBor that 1985 ParmleyNolf and 1988 Cicimurri Linck, 1790 (2003 Scyliorhinus Reineckegilberti and et al. material 2008 sp Scy. distans ( ) by Manning and Referred specimen(Fig. 6A–D) Standhardt) identified ( as Remarks—The single tooth can be distinguished from Dryidentified Branch as taxon. Probst, 1879 teeth of Rhynchobatus —SC2013.38.45, tooth. (see be- 1986:fig. 1, no. 2) to be conspecific with the low) in lacking crown ornamentation, the medial lingual SCYLIORHINUS Müller and Henle, 1837 - Scylliorhinus Blainville, 1816 berances are present (Fig. 6A). The crown apex is worn, sp. andprotuberance it is unclear is veryif this narrow, tooth was and low-crowned flanking lateral (female) protu Referred specimens(Fig.—SC2001.1.42, 5O–CC) incomplete or cuspidate (male). The tooth is identical to Rhinobatos

central Georgia (DJC, unpublished data). Although repre- tooth; SC2013.38.35, tooth; SC2013.38.36, tooth; sentedteeth from by a the single temporally specimen, older the Clinchfield Dry Branch Formation Rhinobatos of tooth.SC2013.38.37, tooth; SC2013.38.38, tooth; SC2013.38.39, exhibits features in common with middle to late Eocene tooth;Remarks SC2013.38.40,—Unfortunately, tooth; SC2013.38.160, our eight teeth anterior lack Rhinob. steurbauti , including an enameloid and/or have broken cusps, and the true Cappetta and Nolf, 1981 elongated central lingual protuberance flanked by much CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA

17

Figure 6. Batoid teeth from the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–D. Rhinobatos sp. Tooth in occlusal (A), labial (B C), and basal (D E–H. Rhynchobatus sp. cf. Rhy. pristinus. Tooth in occlusal (E), labial (F G), and basal (H I–P. Dasyatis sp. cf. D. tricuspidatus. Male anterior tooth in labial (I) and distal (J ), profile ( ) views,K) and SC2013.38.45. occlusal (L M), labial (N), pro- O), and ),basal profile (P ( ) views,Q–Y SC2013.38.46.. Dasyatis Q) and distal (R) views. Tooth in occlusal) views, (S SC2013.38.42. Tooth Toothin labial in (occlusal (T), labial )( Uviews, SC2013.38.43.V Tooth in occlusal ( file(W ),( labial (X ) views,Y SC2013.38.41. sp. SC2013.38.49. Male anterior–T, W;tooth at left in labial in C, G, ( J, O, R, V, Y. Scale bars=1 mm. ) view, SC2013.38.55. ), and profile ( ) views, SC2013.38.52. Tooth in occlusal ), and profile ( ) views, SC2013.38.53. Labial at top in A, D, E, H, L, M, P, S smaller and slightly diverging lateral protuberances used the presence of Rhinobatos in a paleofauna as an ( , Case et al. 1996). Rhinobatos indicator of a middle shelf-depth environment. teeth occurring in upper Eocene strata of Georgia and Cappetta and Case (2016 Cappetta and Nolf 1981 teeth from middle Eocene strata of Alabama as Pristis Formation species, and we believe that those records . However, those teeth) recently clearly identified have an elongat similar- Louisiana appear to be conspecific with the Dry BranchRhi- Pristis nob. casieri ( , Manning teethLinck, lack1790 lateral uvulae ( , were erroneously identified). Manning as the and Cretaceous Standhardt taxon ( ) Carrillo-Briceñoed medial uvula flanked et al. 2016 by lateral). Negative uvulae, evidence whereas within Cappetta and Case, 1975 Case 1981 Carrillo-Briceño et al. 2015 and Standhardt 1986 1986 PALEOBIOS, VOLUME 36, JUNE 2019

18 the Dry Branch Formation sample, which lacks Pristidae Referred specimens not present at the time of deposition. For the purposes —SC2013.38.41, tooth; Bonaparte, 1838 rostral spines, indicates sawfishRhinobatos were, SC2013.38.42,Remarks—Nine male large tooth; Dasyatis SC2013.38.43, teeth are characteristic male tooth; but our generic assignment could change with a larger inSC2013.38.44, their lack of six enameloid teeth. ornamentation, a feature that sampleof this report size that we willidentify allow SC2013.38.45 for better comparisons as to readily distinguishes them from most other species that the tooth morphologies of other extant rhinopristiform have been reported from Eocene strata ( , genera. The Dry Branch Formation tooth bears some ). The intersection of the similarity to Glaucostegus typus (Under- transverse crest with a sagittal lingual crestWard results 1979b in a ), but its labial face appears to be more tripartiteNoubhani division and Cappetta of the crown 1997 (i.e., Fig. 6M). This species convex than seen on male and femaleBennett, teeth 1830 of Zapteryx was originally reported from lower Eocene strata of Bel- brevirostriswood et al. 2015( ) (Rangel et al. 2014). gium (Casier 1946), and the taxon has subsequently been documented in middle Eocene strata of Europe (Kemp et RHINIDAEMüller and Henle, 1841 al. 1990, Van den Eeckhaut and de Schutter 2009). RHYNCHOBATUS Dasyatis exhibits gynandric heterodonty (male teeth Rhynchobatus MüllerRhy. pristinus and Henle, ( 1841 ) Müller and Henle, 1837 sp. cf. Probst, 1877 male and female teeth of a fossil species may not be as Referred specimens(Fig. 6E–H) cleardiffer cut from as previously those of females), thought. Kajiurabut the and identification Tricas (1996 of) found that male and female teeth of Dasyatis sabina Remarks—The three teeth—SC2013.38.46, are easily distinguished tooth; as (=Hypanus sabinus) ( ) may be nearly indis- RhynchobatusSC2013.38.47, in tooth; having SC2013.38.48, granular crown tooth. ornamentation tinguishable from each other, with both sexes having low- and a central lingual protuberance that is long and wide, crowned teeth. However,Lesueur, as mating 1824 season approaches, male teeth exhibit a transition (Fig. 6K) to taller, highly fossil Rhynchobatus species have been reported, Rhy. vin- cuspidate crowns that are effective at grasping pectoral centiand flanking protuberances from Eocene are deposits absent ((Fig. 6E). Two) Kajiura et al. 2000). and Rhy. pristinus from Oligo-Miocene strata (Cappetta Assuming this phenomenon was developed in Eocene ,Jaekel, Cicimurri 1894 and Knight 2009). Overall,Leriche teeth of 1905 these fossilfins of species, females isolated during copulationlow-crowned ( teeth (Fig. 6M–P) two species are comparable in size and morphology, but in the Dry Branch sample could belong to males or fe- Rhy.1970 vincenti has a wider, shorter lingual protuberance males, whereas high-crowned cuspidate teeth belonged than Rhy. pristinus ( , Müller 1999). Jaekel to males (Fig. 6I–J). ( Rhy. vincenti, and the JaekelDry Branch 1894 specimens have a long Dasyatis 1894:76–77) makes no mention of ornamentation on and narrow protuberance similar to Rhy. pristinus. We sp. tentatively refer the Dry Branch teeth to this latter spe- Referred specimens(Fig. 6Q–Y) cies due to limited sample available to us. Müller (1999) reported a damaged lower Eocene Rhynchobatus tooth —SC2013.38.49, male tooth; that is similar to Rhy. vincenti, but he did not believe the SC2013.38.51, 12 male teeth; SC2013.38.52, tooth; Pristis SC2013.38.53,Remarks tooth; SC2013.38.54, 39 teeth; SC2013.38.55,Dasyatis differ from those of Rhynchobatus in lacking crown sp.tooth; cf. D.SC2013.38.161, tricuspidatus four (see teeth. above) in that they are smaller ornamentation,taxa were conspecific. but having Although a sharp similar, and teeth conspicuous of in overall size—These and are 59 easily teeth distinguished differ from those by their of highly transverse crest. Additionally, Pristis teeth have a shorter lingual face but much more elongated medial uvula (Cap- make accurate comparisons to other species with highly petta 2012, , Carrillo-Briceño ornamented teethlabial ( face (Fig. 6S,, Noubhani U, W). It andis difficult Cappetta to et al. 2016). ) due to the lack of enameloid on the Dry Branch Carrillo-Briceño et al. 2015 specimens. However,Ward the 1979b teeth of D. jaekeli (Leriche, MYLIOBATIFORMES 1997) have ornamentation that is generally restricted DASYATIDAE to the apical region of the labial face ( ), Compagno 1973 DASYATIS and1905 ornamentation on Jordan 1888 D. wochadunensis Dasyatis D. tricuspidatus Leriche 1905 Rafinesque 1810 extends well onto the lingual side of the crown. Highly sp. cf. Casier 1946 cuspidate teeth bearing weak labial ornamentationWard, 1979b (i.e., (Fig. 6I–P) CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 19

and the latter generic name is applied to the Dry Branch associate them with Dasyatis sp. due to their small size sample listed above. Fig.and 6Q–R)ornamented are herein labial identified face, in comparison as male teeth, to Dasyatis and we sp. cf. D. tricuspidatus discussed above. PSEUDAETOBATUS Pseudaetobatus undulatus MYLIOBATIDAE Cappetta, 1986 AETOMYLAEUS Cicimurri and Ebersole, 2015 AetomylaeusBonaparte, 1838 Referred specimens(Fig. 7F–I) Garman, 1908 sp. —SC96.97.52, 22 medial tooth Material examined (Fig. 7A–E) - fragments; SC96.97.53, lateral tooth; SC2001.1.3,- 118 partial medial teeth; SC2001.1.4, three lateral tooth; SC2001.1.1, eight—SC96.97.49, partial median 17 teeth; incomplete SC2001.1.2, me teeth; SC2001.1.5, 10 lateral teeth; SC2001.1.6, lat incompletedian teeth; SC96.97.50, median tooth; lateral SC2001.1.29, tooth; SC96.97.51, lateral lateraltooth; eral tooth; SC2001.1.7, lateral tooth; SC2013.38.84, SC2001.1.30, three lateral teeth; SC2001.1.31, two in- lower medial tooth; SC2013.38.85, upper medial tooth; SC2013.38.86, upper medial tooth; SC2013.38.87, upper medial tooth; SC2013.38.88, incomplete upper medial complete median teeth; SC2001.1.32, five partial median- tooth; SC2013.38.89, incomplete upper medial tooth; teeth; SC2013.38.80, partial median tooth; SC2013.38.81, SC2013.38.90, lower medial tooth; SC2013.38.91, lower twoRemarks incomplete median teeth; SC2013.38.82, four in distal-most lateral tooth; SC2013.38.92, upper distal- thickcomplete crowns median with teeth; slightly SC2013.38.83, lingually inclined two lateral labial teeth. and most lateral tooth; SC2013.38.93, five lower distal-most lingual faces.—These The labial 48 teethface appears are distinguished highly pitted by their(Fig. laterals; SC2013.38.94, 12 upper distal-most lateral teeth;Remarks SC2013.38.95, lateral tooth; SC2013.38.96, five a very thin and sharp transverse ridge is located at the lateral teeth; SC2013.38.97, 87 incomplete medial teeth. 7A), whereas the lingual face has a granular texture and Pseudaetobatus. Upper median teeth of Pseudaetobatus —We assign 275 teeth in our sample to be distinguished from those in the lower dentition in be- crown-root junction (Fig. 7B). Upper median teeth can from the lower dental battery are more arcuate (Fig. ing more convex overall, with a straight to curved crown are wide and straight (Fig. 7F), whereas median teeth base that parallels the occlusal surface. In contrast, lower of (see above) in that the labial and lingual - Aetomylaeus 7G). Median teeth are easily distinguished from those face, and the crown foot is parallel to the basal root face. crown faces are only weakly ornamented (as opposed to Lateralmedian teeth teeth arehave longer a straight than crown wide and with bear flat labialocclusal pitting sur heavily pitted labially and with granular texture lingually) and the lingual ridge at the crown/root junction is thick and rounded instead of thin and sharp. These features and lingual tuberculation on crown faces (Fig. 7D, E). - can also be used to distinguish the lateral teeth of both mationThe Dry of central Branch Georgia material (DJC appears unpublished to be conspecific data). In species, and the distal-most lateral teeth of Pseudaeto- termswith a of myliobatoid overall morphology, occurring this in taxon the Clinchfield compares quite For batus favorably to Miocene teeth Cappetta ( ) identi- Pseudaetobatus has only recently been formally re- Pteromylaeus Garman, 1913, and to Myliobatis ported are from also the sinuous United (Fig. States, 7I). with two new species meridionalis ( ). Cappetta (20061970) synony- described by Cicimurri and Ebersole ( ). Pseudae- mizedfied as M. meridionalis with Pteromylaeus, and earlier tobatus belli is an early Cappetta ( Gervais, 1852 Eocene species that occurs in Mississippi2015 and Alabama, material he described in could represent old indi- whereas Ps. undulatusCicimurri isand thus Ebersole, far only 2015known from the viduals of 1987Aetomylaeus:171) considered. Hovestadt the and possibility Hovestadt-Euler that the upper Eocene Dry Branch Formation of South Carolina. (1999 1970 myliobatidae teeth and stated that Pteromylaeus is only RHINOPTERA ) noted the difficulty of identifying isolated fossil Rhinoptera known from extant species. However, Hovestadt and Cuvier, 1829 Hovestadt-Euler (2013) later assigned several fossil spe- sp. Myliobatis to Pteromylaeus Referred specimens(Fig. 7K–R) and Aetomylaeus, thereby extending the record of the lattercies previously genera back identified into the as Eocene. White (2014) more fragments; SC2001.1.11,—SC96.97.54, 233 partial medial tooth;teeth; recently synonymized Pteromylaeus with Aetomylaeus, SC2001.1.12,SC96.97.55, 8lateral lateral tooth; teeth; SC2001.1.13, SC96.97.56, lateral 38 tooth;tooth 20 PALEOBIOS, VOLUME 36, JUNE 2019

Figure 7. Myliobatidae teeth from the Dry Branch Formation (Priabonian) of Aiken County, South Carolina. A–E. Aetomylaeus sp. Incomplete lower median tooth in labial (A), lingual (B), and occlusal (C) and views, SC2001.1.2. Lateral tooth in occlusal (D E) views, SC2001.1.29. F–I. Pseudaetobatus undulatus. Upper median tooth in occlusal (F Lower median tooth in occlusal (G H) and labial (I) views, ) and profile ( J K–R. Rhinoptera sp. Lateral tooth in labial) view, (K), lingualSC2013.38.86. (L), occlu - sal (M), and basal (N) views, SC2001.1.12.) view, SC2013.38.84. Median tooth Upperin occlusal right ( Odistal-most), lingual ( Plateral), and toothbasal in(Q occlusal) views, (SC2001.1.16. Lateral tooth inSC2013.38.91. lingual (R Lateral tooth in occlusal ( ) view, SC2013.38.96.2.–H, J, M–O, Q; at right in E. Scale bars=1 cm in A–C, F–I, K– , E, J.

) view, SC2013.38.70. Labial at top in C, D, F R; 5 mm in D

Description—Rhinoptera is represented in our material SC2001.1.14, lateral tooth; SC2001.1.15, lateral tooth; lateral tooth; SC2013.38.79, 100 incomplete teeth. SC2001.1.16, medial tooth; SC2001.1.17, 19 medial teeth; to 1 cm in thickness, but in cases of extreme wear crowns SC2013.38.69, medial tooth; SC2013.38.70, lateral tooth; areby 435 only teeth. 2 mm Unworn thick. Theteeth labial have and a crown lingual measuring faces of upall SC2013.38.71, lateral tooth; SC013.38.72, lateral tooth; SC2013.38.73, lateral tooth; SC2013.38.74, four medial vertical wrinkles which grade apically into a granular teeth; SC2013.38.75, three medial or proximal lateral textureteeth are on vertical the lingual and face. flat, Aand large ornamented rounded transverse with fine teeth; SC2013.38.76, ten lateral teeth; SC2013.38.77, eight distal lateral teeth; SC2013.38.78, distal-most CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 21 ridge is located at the lingual crown/root junction (Fig. Bearing these differences in mind, assigning a spe- cies name to the Dry Branch Rhinoptera is somewhat there is a shallow groove on the ventral surface of the problematical. The available material compares very 7L, P, R). The labial crown face overhangs the root, and well to Leidy’s ( ) Zygobates dubius from the “Ash- ley Phosphate beds” region of South Carolina (Leidy did longcrown (4.4:1 just ratio).anterior We to observed, the root as (Fig. did 7K).White Median (1926 ),teeth that not illustrate teeth1855 until median(i.e., Fig. teeth 7O–Q) are measure evenly wornup to and35 mmmay wide be arched and 8 (Fig.mm Leidy ( Z. dubius 1877Trygon; see pl.carolinensis 31, figs. 21–33). named by Emmons1877 :248)( mentioned) from the Eocenethe possibility of North that Carolina. his 7P). The largest teeth that we consider to have been from was conspecific with easythe first to identifylateral row because measure the betweencrown is 23 obviously mm and thicker25 mm appear to be undulating1858 rather than straight or weakly in width and 5 mm to 9 mm in length. These teeth are arched.However, Cappetta Emmons’ ( illustrated2006) syonymized teeth (p. 243, Z. dubius figs. 91, with 92) and root lobes are often oblique to the long axis of the Mio-Pliocene Rhinoptera studeri , but un- on the mesial side than on the distal side (Fig. 7K, L, R), fortunately the stratigraphic position and age of Leidy’s from the outermost row of lateral teeth are symmetrical fossils are unknown. Due to a lack ofAgassiz, stratigraphic 1843 control tooth (Fig. 7N). Other specimens that we believe to be and more limited knowledge of Cenozoic stratigraphy in are distinguished by having an angular mesial margin Leidy’s time, material purportedly from the “Ashley Phos- thator only articulates slightly wider with than the longremainder (1:0.7 ratio). of the These dentition, teeth phate beds” of South Carolina could range in age from the but the distal margin is rounded. At this time we cannot Oligocene through Pleistocene epochs (Sanders 2002). ascertain if intermediate lateral positions were part of Leriche ( ) illustrated a partial Rhinoptera denti- the dentition. Remarks—Unworn median teeth we refer to Rhi- 1927 noptera have a square cross section, whereas those of Rhinop.tion (p. 44,studeri fig. 6) and that it clearlymay shed shows some that light a median on the issue.tooth Pseudaetobatus are rectangular. The labial and lingual rowThat andspecimen, at least from three the lateral Swiss Miocene,tooth rows was were identified present. as crown faces of Rhinoptera vertical wrinkling, whereas the labial face of Aetomylaeus of the width of the median tooth, but the second median is concave with extensive pitting, sp. are andvertical the lingualand bear face fine is The first lateral tooth of the Swiss specimen is only 86% convex with granular ornamentation. In addition, the root lobes of Rhinoptera do not extend lingually past tooth is only 29% of the width of the medial tooth. We the crown base as they do on Pseudaetobatus and Ae- can only confirm two lateral tooth rows in the dentition tomylaeus. The lingual crown/root ridge on Rhinoptera theof the width Dry of Branch the median taxon tooth. (first The and Swiss distal-most Rhinop. studerilateral is thick and rounded as opposed to thin and sharp on dentitionpositions). appears Additionally, to have the been first disjunct lateral toothwith respectis 63% toof Aetomylaeus. tooth size reduction towards the commissure, but size The Dry Branch Rhinoptera is easily distinguished reduction within the Dry Branch appears to have been from Paleocene Rhinop. prisca and Rhi- more gradual and similar to Recent Rhinop. brasiliensis nop. raeburni White, 1934 in that the crown is not nearly ( ). Leidy’s Z. dubius appears as thick, labial and lingual facesWoodward, are nearly 1907 vertical and to be identical to the Dry Branch taxon, and his fossils couldBigelow be andolder Schroeder than Miocene. 1953 Unfortunately, we can- or scattered nodes. Occlusal ornamentation appears to not corroborate these hypotheses, and at this time we bestraight, absent, and and ornamentation nutritive grooves is reduced are more to numerous fine wrinkles and identify the Dry Branch taxon only to the generic level. more closely spaced. Although of similar morphology, Dry Interestingly, Darteville and Casier ( ) illustrated Branch Rhinoptera teeth are larger than Rhinop. sherborni White, 1926 from the African Eocene, with the crown Myliobatis1959 sp., that are very of the largest complete tooth in our sample measuring similartwo teeth to fromthe Dry the Branch Eocene Rhinoptera of western. Africa (pl. 34, fig. White 2a–c. figs. 7, 8), identified as 1926 ). Even DISCUSSION 3.6 cm in width as opposed to only 1.7 cm (seeMyliobatis by White: pl. (10,1926 figs. 16–26; also Arambourg 1952 Stratigraphic position of the Dry Branch Arambourgconsidering (specimens) to be originally Rhinop. sherborniidentified medialas teeth, elasmobranch assemblage : pl. 10, fig. 5–7) but later considered by In the Aiken area, the Huber, Dry Branch, and Tobacco 1952 the largest specimen measures only 2.5 cm. 22 PALEOBIOS, VOLUME 36, JUNE 2019

Road Formations are generally visible at the surface as thin outcroppings along roadways and in housing sub- sandy clay, and overlies a loose sand of more uniform divisions. Limited exposure, combined with a great deal yellow-orangeleast 15 cm of thin color. alternating beds of red sand and white of lithologic variability and inconsistent local thicken- ing and thinning due in part to highly irregular contacts Composition of the Dry Branch elasmobranch assemblage atbetween the NAS these are assignableformations, to makes the Dry identification Branch Formation, of units teeth to the species level was sometimes hampered by the and correlationa thick grayish-white of sections clay difficult. bed occurs All of the at the exposures base of preservationAs noted earlier, of the identification material. The of enameloid the Dry Branch was sharkoften the unit. Unfortunately, this site has been reclaimed as partly or completely leached, and the presence of crown a housing development, and only thin patches of strata ornamentation, height of lateral cusplets, and the nature are currently exposed. Of the strata preserved in the E-W trending wall at the SAS, only the sediment within specimens. For example, Figure 3A, B and 4L–M show teethof cutting on which edge serrations the enameloid was difficult is partially to discern lost, whereason many (we also dug a trench up to 1.3 m below ground level but Figure 3 provides a contrast between teeth with (3O–P) onlythe lower encountered 2.0–2.5 morem belong unconsolidated to the Dry Branch sand). FormationThe upper and without (3M–N) enamleoid. However, it can be said part of the section consists of indurated medium- to very that carcharhiniform sharks dominate the Dry Branch coarse-grained quartz sands (grains are sub-rounded to elasmobranch assemblage, with 11 taxa having been rounded) that are poorly sorted and contain abundant interstitial clay, which we assign to the Tobacco Road hexanchid, one squatinid, two orectolobiforms, and two Formation. A basal pebble lag is not obvious, but the lamniforms.identified. The The remaining batiod portion component of the fauna consists is composed of one contact with the Dry Branch Formation is irregular, and Twenty-one taxa have presumed benthic/epibenthic pinkish to pale yellow quartz sand containing little in- habitsof two rhinopristiform(including Squatina rays ,and most five of myliobatiform the charcharhini rays.- terstitialthere is aclay sharp (soft change and friable=Dry from fine- Branch to coarse-grained, Formation), forms, batoids). These far outnumber the three with to medium- to coarse-grained, poorly sorted quartz presumed pelagic habits that include Galeocerdo, Alopias, sand containing abundant orange interstitial clay (more and Carcharias. The varied tooth morphologies in our indurated=Tobacco Road Formation). sample are indicative of clutching, cutting, tearing, and Consistent with previous observations of elasmo- crushing dentitions ( ), and a larger array branch teeth occurring in the Dry Branch Formation of prey species was probably available, both vertebrate ( ), the sample from the SAS is derived and invertebrate, thanCappetta is presently 1987 indicated by the fos- from the upper part of the formation, and the lithology sil record. isZullo consistent et al. 1982 with the Irwinton Sand as it is composed predominantly of golden-yellow, brownish-yellow, and Depositional environment pinkish to purple medium- to very coarse-grained quartz Vertebrate species occurrences at the NAS and SAS are sand that is moderately well-sorted and generally con- very similar, and we believe that the depositional settings tains very little interstitial clay. at the two sites were comparable. For both sites, evidence suggests shallow, subtropical, high-energy environments. cm thick and has variegated red and white color, with The lithology and presence of terrestrial/brackish water lesserThe amountsfossiliferous of yellow horizon and at SASorange. is approximately Within this ho 15- like gars, trionychid turtles, and crocodilians rizon, vertebrate fossils are common and haphazardly distributed and oriented, small white clay clasts are com- of sediment via a river system. Minerals like kyanite mon, and larger quartz clasts (>1 cm) are uncommon. andindicates amphibole very close suggest proximity local igneous to the shoreline and metamorphic and influx Numerous small phosphatic steinkerns representing at rocks were sediment sources for the fossiliferous bed least ten species of bivalves, ten species of gastropods, a at the SAS. scaphopod, a barnacle, possible branching bryozoa, and If extant elasmobranchs can be utilized as proxy chelipeds of at least three species of decapod crustaceans environmental indicators for fossil species, the taxa we are associated. The fossil horizon is located approxi- mately two meters below the contact with the overlying above hypotheses. Tawny nurse sharks (Nebrius) and se- Tobacco Road Formation. The unit directly underlies at vengillidentified sharks from (Notorynchus the Dry Branch) are Formationpredominantly support inshore the CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA 23

similar to features of bioerosion reported on Cretaceous 1999 (inhabitantsIsogomphodon where) prefer water turbid depth waters is between near river five tomouths 30 m we also observed the markings on steinkerns, phosphate toand depths one to of 50 40 m, m; respectively, the latter whereastwo species daggernose are apparently sharks pebblesfish teeth and by coprolites,Underwood and et ital. is ( unclear:fig. if 2e). these However, are the intolerant of hyposaline conditions (Compagno et al. result of pre-fossilization bioerosion or post-fossilization ). The discovery of a hexanchid in the Dry Branch chemical alteration. Formation, occurring at both NAS and SAS, is particularly notable2005 because these sharks have not previously been Connection to the Tethys Sea reported from the late Eocene of the Gulf and Atlantic The Dry Branch Formation elasmobranch assemblages coastal plains. Stingrays currently inhabiting coastal from NAS and SAS share generic similarities to the Eo- waters of South Carolina, including Hypanus sabinus, Hy. cene Tethyan region, with Squatina, Alopias, Galeocerdo, say ( ), and Hy. americanus (Hildebrand and Abdounia, Physogaleus, Negaprion, Rhizoprionodon, and Snel- Rhinoptera having been reported from Europe, Africa, Lesueur, 1817, McEachran and Carvalo 2002, Farmer and Asia ( , Adnet et al. 2010, Under- Schroeder,2004, Aguiar 1928 et), al. occur 2009 at depths of nineRhynchobatus to 25 m ( ) wood et al. 2011, Zalmout et al. 2012, Malyshkina et al. son et al. 1988 2013). HemipristisArambourg curvatus 1952 (Compagno and Last 1999a). ),Wedgefish and cow-nosed ( rays (Rhi- (Mustafa and Zalmout 2002) and northern Africa (Dames nopteracommonly) inhabit occur a invariety coastal of nearshore waters to habitatsdepths ofin water25 m , , has been, Caseidentified and Cappettain Jordan depths up to 26 m (Compagno and Last 1999b). Although 1990), and Pseudaetobatus was originally described extant angel sharks (Squatina) inhabit a wide range of 1883from thePriem Eocene 1905 of MoroccoStromer (1905 ). Dasyatis habitats, primarily remaining buried in the substrate tricuspidatus is known from Eocene strata of Belgium and waiting to ambush prey ( , Compagno et England (Casier 1946, Kemp etCappetta al. 1990), 1986 and Rhinobatos ), Case ( ) considered Squatina to be indica- steurbauti has been reported from the Paris Basin (Cap- tive of an estuarine depositionalCompagno environment, 1984 whereas ) and Uzbekistan (Case et al. 1996). Manningal. 2005 and Standhardt1981 ( ) believed the absence Notorynchus kempi occurs in the Eocene of England of the genus was evidence for fully marine conditions. (petta and Nolf, Kemp1981 et al. 1990). 1986 Sand have been used as an indication of a subtidal to in- ComparisonWard 1979a to Barnwell Group elasmobranch nerBarnacle shelf environment, faunas reported and the from SAS the Dry Griffins Branch Landing sample assemblages from Georgia contains at least one taxon (cf. Hesperibalanus Pilsbury, At the generic level, the elasmobranch assemblages re- 1916). We recovered several specimens of the coral En- dopachys and Dry Branch Formations) are similar in faunal com- species of this genus can be found in subtropical inner position,ported from and Barnwell the South Group Carolina strata Dry of GeorgiaBranch assemblage(Clinchfield to middle Milneshelf environments Edwards and (20–100Haime, 1848, m) where and extantwater contains most of these genera, including Galeocerdo, temperatures are between 21° and 22° C (Brook 1999, Squatina, Nebrius, Carcharias, Premontreia, Hemipristis, Abdounia, Negaprion, Physogaleus, Rhinoptera, Aetomy- coral, represented by several small and poorly preserved laeus, and Dasyatis ( , Case and Borodin 2000, conicalKeller and calyces, Os’Kina occurs 2008 at ).the An SAS. additional, unidentified Parmley and Cicimurri 2003, DJC and JLK unpublished Many specimens recovered from both sites exhibit data). The Georgia assemblagesCase 1981 also contain taxa not yet unusual circular depressions, including teeth of Hemipris- known in the South Carolina Dry Branch Formation, in- tis, Carcharias, Negaprion, Myliobatidae, Sphyraena, and cluding Otodus (Carcharocles) Jordan and Hannibal, 1923, crocodile, as well as mollusk steinkerns, phosphate peb- Macrorhizodus Glikman, 1964, Heterodontus Blainville, bles, and coprolites. These features are usually isolated , Pristis, and Propristis (see references and penetrate various depths into an object, and some- cited above). Conversely, the South Carolina Dry Branch times completely through. The surfaces of the markings paleofaunas1816 contain taxa notDames, known 1883 from Georgia, in- are white. The marks are found primarily on the roots of cluding Notorynchus, Isogomphodon, Rhynchobatus, and the selachian teeth (one was observed at the crown base Pseudaetobatus. These differences could be related to of a Carcharias tooth, where the enameloid covering is collecting biases, but the samples from the Dry Branch thinnest), directly on the crown of Sphyraena, and on the - crown and root of Myliobatidae teeth. These markings are tion of Georgia that we examined were obtained during Formation of South Carolina and the Clinchfield Forma 24 PALEOBIOS, VOLUME 36, JUNE 2019 multiple site visits over the course of several years, There are tantalizing clues that additional vertebrate through prospecting surfaces for exposed macrofossils faunas are preserved within other Eocene deposits in the and processing bulk matrix samples for microfossils. region. We examined two small collections of vertebrate These units are not time-equivalent, and the differences fossil that were made by Zullo and Kite ( ) during a could be related to variations in depositional environ- study of barnacles from the Dry Branch Formation. Both ment, with, for example, absence of Otodus (Carcharocles) samples were recovered from deposits attributed1985 to the and Macrorhizodus from the South Carolina assemblage and/or course sandy substrate. However, Zalmout et al. Griffins Landing Sand, and the various elasmobranch and (reflecting2012) reported the very both shallow, Otodus probably (Carcharocles turbid) andconditions Macro- thebony identity fish species of some we speciesidentified that are we listed collected in Appendix from the 1. rhizodus IrwintonThe shark Sand. teeth Kite are (well preserved) collected and an helped assortment confirm of Egypt, and perhaps these larger sharks were attracted elasmobranch remains from the middle Eocene Huber to the area, from where river-influened marine mammals bay/estuarine were apparentlydeposits in Formation, which underlies1982a the Dry Branch Formation plentiful. Marine mammals are thus far unknown from the Dry Branch Formation. her illustrated material are listed in Appendix 2. She also notedto the thateast her of Aiken, material and was the poorly species preserved, we identified fragile, from and CONCLUSIONS had been altered to clay. Not surprisingly, comparison of the vertebrate as- Finding additional vertebrate faunas within the Dry semblages from the NAS and SAS revealed a similar Branch Formation and other Barnwell Group strata in . Both assemblages are derived from the upper part of to note that, of eight stratigraphic sections of the Dry thetaxonomic Dry Branch composition Formation within and arethe separated fish component by only BranchSouth Carolina Formation may prepared be a difficult by Kite task. ( It is), interesting none were reported to contain vertebrate fossils. Attempts to revisit are thought to have been very close to the late Eocene these sites have been thwarted because1982b of landscape shoreline,13 km (SAS as crocodilianis nearly 7 andkm trionychidfurther inland). turtle Bothfragments sites alteration or they no longer exist. For example, a borrow were found at both sites . The SAS is thus far the furthest inland within the South Carolina Coastal Plain from which Eocene marine vertebrates have been reported. The SAS likepit in Kite’s northeastern ( ) HuberHollow Formation Creek Quadrangle site and (stopstop 47 of ZulloKite 1982b et al. )( is now; locality used as SCGS-004, an inert landfill. 33° Other sites, elasmobranch species, including Pseudaetobatus undu- ° 1982a latusis also ( significant because it has now) and yielded Isogomphodon two new subdivisions1982 (as are the NAS and SAS). Additionally, 29’ 27” N lat.,the aikenensis described herein. The Dry Branch Formation 81 49’ 5” W long.), are overgrown and part of housing sedimentsCicimurri occurring and Ebersole at both sites,2015 especially SAS, may of the region. have been derived from metamorphic rocks and/or peg- GriffinsEven Landingwith abundant Sand largely exposures, occurs discoveringin the subsurface elas- matites located in Aiken County. The two elasmobranch mobranch teeth at other sites may not occur unless assemblages we recovered are similar to, but less diverse lithostratigraphic beds contain large fossils and/or Parmley and concentrations of remains. Teeth are easily eroded from Cicimurri 2003) and Dry Branch ( , Case and the fossiliferous horizon at the SAS, and larger specimens than,Borodin those 2000 reported) Formations from ofthe central Clinchfield Georgia. ( Close prox- - imity to the shoreline could accountCase for lower 1981 taxonomic posure. As control samples, we collected matrix randomly diversity in the South Carolina Dry Branch assemblages, occurringfrom 110 cm,as float 100 are cm, common 60 cm, and around 30 cm the below base theof the fossil ex and large pelagic sharks like Macrorhizodus and Otodus (Carcharocles) may have preferred deeper water. How- two additional samples from the Tobacco Road Forma- ever, diagenetic alteration/obliteration of remains (i.e., horizon, as well as 10 cm and 165 cm above. We obtained current action, bioerosion, chemical leaching) may also be a phenomenon affecting species diversity. The variety eachtion, oneof these from horizons, within the only lowermost two teeth 5 cm,were and recovered one 30 cm in of elasmobranch and teleost tooth morphologies oc- theabove sample the base. from Of 60 the cm 4.5 below kg of the matrix fossiliferous processed horizon, from curring in the Dry Branch Formation indicates greater and a few from the sample 10 cm above (thin-bedded red prey species richness (i.e., invertebrates and small ver- sand and white sandy clay). In contrast, the fossiliferous tebrates) than is currently known. horizon at the SAS yielded approximately 30 specimens CICIMURRI & KNIGHT—ELASMOBRANCHS FROM THE DRY BRANCH FORMATION, SOUTH CAROLINA, USA

25 per kg of matrix sampled. (crocodiliens, poissons) du miocène marin de l’alagarve oc- cidentale. Ciências da Terra ACKNOWLEDGEMENTS - boids. Journal of the Marine 6:9–38. Biological Association of India We would like to express our appreciation to Andrew Applegate, S.P. 1972. A revision of the higher taxa of Orectolo - hours they spent sorting through matrix with a binocular 14(2):743–751. Notes et Mémoires du Service microscopeScheffield, Katie and countingWhite, and specimens. David Briedis Karin, for Ryan, the longand Arambourg,Géologique C. du1952. Maroc Les vertébrés fossils des gisements de phos phates (Maroc-Algérie-Tunisie). Stacy Knight helped JLK recover fossils during many Proceedings of the California 92:1–372. Academy of Natural Sciences visits to the Nouth Aiken site (NAS), and Christian and Ayres, W.O. 1855. Description of new species of California fishes. Morgan Cicimurri accompanied DJC and JLK to the North in Memoir of the Life and South Aiken sites (SAS). Mike Bruggeman assisted 1(1):23–77. Bennett, E.T. 1830. Class Pices. Pp. 686–694 and Public Services of Sir Thomas Stamford Raffles, particularly strata at the NAS belonged to the Dry Branch Formation. andin the resources government of the of Eastern Java, 1811–1816, Archipelago and and of selectionsBencoolen from and BobbyJLK at the and NAS, Susan and Congdon Ralph Willoughby donated a confirmed large collection that the of its dependencies, 1817–1824: with details of the commerce fossils from the SAS to SC. Scott Harris determined that greenish-black grains occurring in matrix from the SAS undhis correspondence. Fische. Hochschulbücher Lady Stamford für Biologie, Raffles, Berlin. London, 310 851pp. pp. were an amphibole mineral. Mark Smith and Christian Berg, L.S. 1958. System der Rezenten und Fossilen Fischartigen skates and rays; chimaeroids. Fishes of the western North At- Cicimurri arranged for the loan of McKissick Museum Bigelow,lantic, H.B., part andtwo. W.C. Memoirs Schroeder. of the 1953. Sears Sawfishes, Foundation , for Marine (University of South Carolina, Columbia) specimens to DJC. Editorial comments provided by Peter Kloess and - Diane M. Erwin, as well as input from Will Doar (South butionResearch, systematique Yale University. du 514règne pp. . Bulletin du Société Carolina Geological Survey, Columbia, SC) and Charlie Blainville,Philomathique H.M.D. du de. Paris 1816. Prodrome d’une nouvelle distri Geologist Underwood (Birkbeck, University of London), improved 8:105–112. upon our original manuscript. Blake,classi C.C. degli 1862. animali Shark’s vertebrati. teeth at Panama. Tomo III. Pesci. 5:316. Bonaparte, C.L. 1835. Icnografia della fauna italic perNuovi le Quattro Annali LITERATURE CITED delle Scienze Naturali Bonaparte, C.L. 1838. Selachorum tabula analytica. Adnet, S., P.-O. Antoine, S.R. Hassan Baqri, J.-Y. Crochet, L. Marivaux, Dongen Formation (Eocene) 1(2):195–214. in the . 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South Carolina Geology

Oligocene or early Miocene 34(1-2):57–67. age for the Dry Branch Formation Zullo,and V.A., Tobacco R.H. RoadWilloughby, Sand in and Aiken P.G. County, Nystrom, South Jr., Carolina?1982. A late Pp. in P.G. Nystrom, Jr. and R.H. Willoughby (eds.). Geological investigations related to the stratigraphy in the kaolin mining 34–45district, Aiken County, South Carolina. Carolina Geological Society, Field Trip Guidebook ______1982. Appendix 1. -

Fossil elasmobranch and bony fish spe- eredcies identified by Zullo and from Kite McKissick ( ). Museum samples of the Griffins______Landing Sand, Dry Branch Formation, recov McKissick Museum locality1985 03, Griffins Landing (Z –622). Chondrichthyes Carcharias sp. Hemipristis curvatus Abdounia enniskilleni Rhizoprionodon sp. cf. Rhiz. ganntourensis Rhinoptera sp. Dasyatis sp. cf. D. tricuspidata Osteichthyes cf. Sphyraenodus sp. Seriola sp. McKissick Museum locality 06 (=South Carolina Geological Survey locality 2-49), Aiken County auger hole (Z- 723). Chondrichthyes Carcharias sp. Negaprion gilmorei Physogaleus sp. Abdounia enniskilleni Osteichthyes cf. Sphyraenodus sp. ______Appendix 2. Elasmobranch taxa from the Huber Formation exposed to the east of Aiken, Aiken County,

( ): South______Carolina, based on material figured by Kite 1982aChondrichthyes Carcharias sp. ( Carcharhiniformes indet. ( RhizoprionodonKite sp. (1982a:fig. 2A-B) Myliobatinae indet. ( Kite 1982a:fig. 2C, left) Rhinoptera sp. ( Kite 1982a:fig. 2C, right) indeterminate batoidKite caudal 1982a spines:fig. ( 2D) 2F) Kite 1982a:fig. 2E) Kite 1982a:fig.