Middle and late Eocene fish otoliths from the eastern and southern USA

  • Chien-Hsiang Lin Biodiversity Research Center, Academia Sinica, 128 Academia Road, Sec. 2, Nankang, Taipei 11529, Taiwan https://orcid.org/0000-0002-9843-9729
  • Dirk Nolf Royal Belgian Institute of Natural Sciences, 29, Rue Vautier, 1000 Brussels, Belgium
Keywords: paleodiversity, Claiborne and Jackson groups, paleoecology, Sciaenidae, faunal comparison


The fossil otoliths of the southern USA have been known for more than 130 years and are among the richest assemblages worldwide. However, previous studies are often scattered and with outdated systematic scheme. A collection of over 25000 otoliths ranging in age from the Lutetian to the Priabonian from 47 sites in five states in the eastern and southern USA is analysed here. Combined with the earlier described material, at least 101 otolith-based taxa are documented, of which 83 are identified at species level. Fourteen of these are introduced as new species: Elopothrissus bernardlemorti sp. nov., “Muraenesoxbarrytownensis sp. nov., Pseudophichthys texanus sp. nov., Paraconger wechesensis sp. nov., Neoopisthopterus weltoni sp. nov., “aff. Glyptophidiumstringeri sp. nov., Symmetrosulcus dockeryi sp. nov., Mene garviei sp. nov., “Citharusvarians sp. nov., Waitakia beelzebub sp. nov., Astroscopus compactus sp. nov., Parascombrops yanceyi sp. nov., Anisotremus rambo sp. nov., and Pagellus pamunkeyensis sp. nov. The assemblages are distinct fom contemporary European faunas by the complete lack of mesopelagic fish otoliths, and by the presence of sciaenids. Dominant taxa in the American Eocene are the Ophidiidae, Sciaenidae, Lactariidae, and Congridae. They indicate shallow-water environments for all the sampled sites. The notable abundance of those taxa suggests that they could have a higher turnover rate, and provided fundamental nutrition in the local Paleogene marine ecosystem. Further analyses of the species in the stratigraphic succession revealed that a faunal turnover between the Claiborne and Jackson seas was evident in teleosts, and it might be more widespread in other marine organisms in the region.


Agnini C., Fornaciari E., Raffi I., Catanzariti R., Pälike H., Backman J. & Rio D. 2014. Biozonation and biochronology of Paleogene calcareous nannofossils from low and middle latitudes. Newsletters on Stratigraphy 47: 131–181. https://doi.org/10.1127/0078-0421/2014/0042

Aguilera O., Schwarzhans W., Moraes-Santos H. & Nepomuceno A. 2014. Before the flood: Miocene otoliths from eastern Amazon Pirabas Formation reveal a Caribbean-type fish fauna. Journal of South American Earth Sciences 56: 422–446. https://doi.org/10.1016/j.jsames.2014.09.021

Aguilera O., Schwarzhans W. & Béarex P. 2016. Otoliths of the Sciaenidae from the Neogene of tropical America. Palaeo Ichthyologica 14: 7–90.

Bandy O.L. 1949. Eocene and Oligocene foraminifera from Little Stave Creek, Clarke County, Alabama. Bulletins of American Paleontology 32: 5–209.

Bannikov A.F. & Carnevale G. 2016. †Calomonnius quasigobius gen et sp. nov: the first gobioid fish from the Eocene of Monte Bolca, Italy. Bulletin of Geosciences 91 (1): 13–22. https://doi.org/10.3140/bull.geosci.1577

Bannikov A.F., Carnevale G. & Landini W. 2009. A new Early Miocene genus of the family Sciaenidae (Teleostei, Perciformes) from the eastern Paratethys. Comptes Rendus Palevol 8: 535–544. https://doi.org/10.1016/j.crpv.2009.03.001

Bannikov A.F., Schwarzhans W. & Carnevale G. 2018. Neogene Paratethyan croakers (Teleostei, Sciaenidae). Rivista Italiana di Paleontologia e Stratigrafia 124 (3): 535–571. https://doi.org/10.13130/2039-4942/10696

Betancur-R. R., Broughton R.E., Wiley E.O., Carpenter K., López J.A., Li C., Holcroft N.I., Arcila D., Sanciangco M., Cureton J.C., Zhang F., Buser T., Campbell M.A., Ballesteros J.A., Roa-Varon A., Willis S., Borden W.C., Rowley T., Reneau P.C., Hough D.J., Lu Q., Grande T., Arratia G. & Ortí G. 2013. The tree of life and a new classification of bony fishes. PLoS Currents 5: ecurrents.tol.53ba26640df0ccaee75bb165c8c26288. https://doi.org/10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288

Brandl S.J., Goatley C.H.R., Bellwood D.R. & Tornabene L. 2018. The hidden half: ecology and evolution of cryptobenthic fishes on coral reefs. Biological Reviews of the Cambridge Philosophical Society 93: 1846–1873. https://doi.org/10.1111/brv.12423

Breard S.Q. & Stringer G.L. 1999. Integrated paleoecology and marine vertebrate fauna of the Stone City Formation (middle Eocene), Brazos River section, Texas. Transactions of the Gulf Coast Association of Geological Societies 49: 132–42.

Bybell L.M. & Gibson T.G. 1994. Paleogene stratigraphy of the Putneys Mill, New Kent County, Virginia, corehole. US Geological Survey Open-File Report 94: 1–38. https://doi.org/10.3133/ofr94217

Carnevale G., Johnson G.D., Marrama G. & Bannikov A.F. 2017. A reappraisal of the Eocene priacanthid fish Pristigenys substriata (Blainville, 1818) from Monte Bolca, Italy. Journal of Paleontology 91: 554–565. https://doi.org/10.1017/jpa.2017.19

Case G.R. 1994. Fossil fish remains from the late Paleocene Tuscahoma and early Eocene Bashi formations of Meridian, Lauderdale County, Mississippi. Part II. Teleosteans. Palaeontographica, Abteilung A 230: 139–153.

Cicimurri D.J. & Knight J.L. 2009. New record of an extinct fish, Fisherichthys folmeri Weems (Osteichthyes), from the lower Eocene of Berkeley County, South Carolina, USA. PaleoBios 29: 24–28. https://doi.org/10.5070/P9291021805

Cushing E.M., Boswell E.H. & Hosman R.L. 1964. General geology of the Mississippi embayment. US Geological Survey Professional Paper 448-B: 1–28. https://doi.org/10.3133/pp448B

Datovo A., de Pinna M.C.C. & Johnson G.D. 2014. The infrabranchial musculature and its bearing on the phylogeny of percomorph fishes (Osteichthyes: Teleostei). PLoS ONE 9: e110129. https://doi.org/10.1371/journal.pone.0110129

Davidoff A.J. & Yancey T.E. 1993. Eustatic cyclicity in the Paleocene and Eocene: data from the Brazos River Valley, Texas. Tectonophysics 222: 371–395. https://doi.org/10.1016/0040-1951(93)90360-V

Dockery D.T. III. 1977. Mollusca of the Moodys Branch Formation, Mississippi. Mississippi Geological, Economic and Topographical Survey, Bulletin 120: 1–211.

Dockery D.T. III. 1980. Invertebrate macropaleontology of Clarke County, Mississippi Area. Mississippi Bureau of Geology, Bulletin 122: 1–387.

Dockery D.T. III. 1986a. Dobys Bluff tongue of the Kosciusko Formation and the Archusa Marl Member of the Cook Mountain Formation at Dobys Bluff on the Chickasawhay River, Clarke County, Mississippi. In: Neathery T.L. (ed.) Southeastern Section of the Geological Society of America: Decade of North American Geology, Centennial Field Guide. Vol. 6: 379–382. Southeastern Section of the Geological Society of America. https://doi.org/10.1130/0-8137-5406-2.379

Dockery D.T. III. 1986b. The Cockfield (Claiborne Group), Moodys Branch and Yazoo (Jackson Group) Formations at the Riverside Park locality in Jackson, Mississippi. In: Neathery T.L. (ed.) Southeastern Section of the Geological Society of America: Decade of North American Geology, Centennial Field Guide vol. 6: 401–403. Southeastern Section of the Geological Society of America. https://doi.org/10.1130/0-8137-5406-2.401

Dockery D.T. III. 1986c. Toward a revision of the generalized stratigraphic column of Mississippi. Mississippi Geology 17 (1): 1–9.

Dockery D.T. III. & Thompson D.E. 2016. The Geology of Mississippi. University Press of Mississippi, Jackson. Mississippi Department of Environmental Quality, Jackson, MS.

Ebersole J.A., Cicimurri D.J. & Stringer G.L. 2019. Taxonomy and biostratigraphy of the elasmobranchs and bony fishes (Chondrichthyes and Osteichthyes) of the lower–to–middle Eocene (Ypresian to Bartonian) Claiborne Group in Alabama, USA, including an analysis of otoliths. European Journal of Taxonomy 585: 1–274. https://doi.org/10.5852/ejt.2019.585

Fisher W.L., Rodda P.U. & Dietrich J.W. 1964. Evolution of Athleta petrosa Stock (Eocene, Gastropoda) of Texas. University of Texas, Austin, TX.

Flis J.E., Yancey T.E. & Flis C.J. 2017. Middle Eocene storm deposition in the Northwestern Gulf of Mexico, Burleson County, Texas, U.S.A. Gulf Coast Association of Geological Societies 6: 201–225.

Fornaciari E., Agnini C., Catanzariti R., Rio D., Bolla E.M. & Valvasoni E. 2010. Mid-latitude calceraous nannofossil biostratigraphy and biochronology across the middle to late Eocene transition. Stratigraphy 7 (4): 229–264.

Friedman M. & Johnson G.D. 2005. A new species of Mene (Perciformes: Menidae) from the Paleocene of South America, with notes on paleoenvironment and a brief review of menid fishes. Journal of Vertebrate Paleontology 25 (4): 770–783. https://doi.org/10.1671/0272-4634(2005)025[0770:ANSOMP]2.0.CO;2

Frizzell D.L. 1965. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes, Albulidae). Senckenbergiana Lethaea 46a: 85–110.

Frizzell D.L. & Dante J.H. 1965. Otoliths of some early Cenozoic fishes of the Gulf Coast. Journal of Paleontology 39: 687–718.

Frizzell D.L. & Lamber C.K. 1961. New genera and species of myripristid fishes, in the Gulf Coast Cenozoic, known from otoliths (Pisces, Beryciformes). Bulletin of the University of Missouri School of Mines and Metallurgy, Technical Series 100: 1–25.

Frizzell D.L. & Lamber C.K. 1962. Distinctive “congrid type” fish otoliths from the lower Tertiary of the Gulf Coast (Pisces: Anguilliformes). Proceedings of the California Academy of Science, Series 4: 87–101.

Frost G.A. 1933. Otoliths of fishes from the lower Tertiary formations of southern England. I. Isospondyli, Apodes, Berycomorphi. Annals and Magazine of Natural History 12: 387–396. https://doi.org/10.1080/00222933308673702

Garvie C.L. 2013a. Additions to the molluscan macrofauna of the Reklaw Formation (Eocene: Lower Claibornian) and two new taxa from the Middle Claibornian in Texas. Bulletins of American Paleontology 384: 131–152.

Garvie C.L. 2013b. New Eocene Mollusca from the collections of the Texas Natural Science Center. Bulletins of American Paleontology 386: 163–183.

Gaudant J. & Quayle W.J. 1988. New palaeontological studies on the Chapel Corner Fish Bed (Upper Eocene, Isle of Wight). Bulletin of the British Museun (Natural History), Geology 44 (1): 15–39.

Girone A. & Nolf D. 2009. Fish otoliths from the Priabonian (Late Eocene) of North Italy and South-East France – their palaeobiogeographical significance. Revue de Micropaléontologie 52: 195–218. https://doi.org/10.1016/j.revmic.2007.10.006

Hart M.B., Yancey T.E., Leighton A.D., Miller B., Liu C., Smart C.W. & Twitchett R.J. 2012. The Cretaceous-Paleogene boundary on the Brazos River, Texas: New stratigraphic sections and revised interpretations. Gulf Coast Association of Geological Societies 1: 69–80.

Hosman R.L. 1996. Regional stratigraphy and subsurface geology of Cenozoic deposits, Gulf Coastal Plain, south-central United States. US Geological Survey Professional Paper 1416-G: 1–35. https://doi.org/10.3133/pp1416G

Ivany L.C. 1998. Sequence stratigraphy of the Middle Eocene Claiborne Stage, US Gulf Coastal Plain. Southeastern Geology 38: 1–20.

Ivany L.C., Patterson W.P. & Lohmann K.C. 2000. Cooler winters as a possible cause of mass extinctions at the Eocene/Oligocene boundary. Nature 407: 887–890. https://doi.org/10.1038/35038044

King C., Underwood C. & Steurbaut E. 2014. Eocene stratigraphy of the Wadi Al-Hitan World Heritage Site and adjacent areas (Fayum, Egypt). Stratigraphy 11 (3): 185–234.

Koken E. 1884. Ueber Fisch-Otolithen, insbesondere über diejenige der norddeutschen Oligocän-Ablagerungen. Zeitschrift der deutschen geologischen Gesellschaft 36: 500–565.

Koken E. 1888. Neue Untersuchungen an tertiären Fisch-Otolithen. Zeitschrift der deutschen geologischen Gesellschaft 40: 274–305.

Lin C.-H. & Chang C.-W. 2012. Otolith Atlas of Taiwan Fishes. National Museum of Marine Biology and Aquarium, Pingtung, Taiwan.

Lin C.-H. & Chien C.-W. 2022. Late Miocene otoliths from northern Taiwan: insights into the rarely known Neogene coastal fish community of the subtropical northwest Pacific. Historical Biology 34 (2): 361–382. https://doi.org/10.1080/08912963.2021.1916012

Lin C.-H., Girone A. & Nolf D. 2016. Fish otolith assemblages from Recent NE Atlantic sea bottoms: A comparative study of palaeoecology. Palaeogeography, Palaeoclimatology, Palaeoecology 446: 98–107. https://doi.org/10.1016/j.palaeo.2016.01.022

Lin C.-H., Brzobohatý R., Nolf D. & Girone A. 2017a. Tortonian teleost otoliths from northern Italy: taxonomic synthesis and stratigraphic significance. European Journal of Taxonomy 322: 1–44. https://doi.org/10.5852/ejt.2017.322

Lin C.-H., Nolf D., Steurbaut E. & Girone A. 2017b. Fish otoliths from the Lutetian of the Aquitaine Basin (SW France), a breakthrough in the knowledge of the European Eocene ichthyofauna. Journal of Systematic Paleontology 15 (11): 879–907. https://doi.org/10.1080/14772019.2016.1246112

Lin C.-H., Taviani M., Angeletti L. & Girone A. 2017c. Fish otoliths in superficial sediments of the Mediterranean Sea. Palaeogeography, Palaeoclimatology, Palaeoecology 471: 134–143. https://doi.org/10.1016/j.palaeo.2016.12.050

Lin C.-H., Chiang Y.-P., Tuset V.M., Lombarte A. & Girone A. 2018. Late Quaternary to Recent diversity of fish otoliths from the Red Sea, central Mediterranean, and NE Atlantic sea bottoms. Geobios 51: 335–358. https://doi.org/10.1016/j.geobios.2018.06.002

Lin C.-H., De Gracia B., Pierotti M.E.R., Andrews A.H., Griswold K. & O’Dea A. 2019. Reconstructing reef fish communities using fish otoliths in coal reef sediments. PLoS ONE 14: e0218413. https://doi.org/10.1371/journal.pone.0218413

Lin C.-H., Chien C.-W., Lee S.-W. & Chang C.-W. 2021. Fossil fishes of Taiwan: a review and prospection. Historical Biology 33 (9): 1362–1372. https://doi.org/10.1080/08912963.2019.1698563

Lo P.-C., Liu S.-H., Chao N.L., Nunoo F.K.E., Mok H.-K. & Chen W.-J. 2015. A multi-gene dataset reveals a tropical New World origin and Early Miocene diversification of croakers (Perciformes: Sciaenidae). Molecular Phylogenetics and Evolution 88: 132–143. https://doi.org/10.1016/j.ympev.2015.03.025

Maisch H.M., Becker M.A., Raines B.W. & Chamberlain J.A. 2014. Chondrichthyans from the Tallahatta-Lisbon Formation contact (Middle Eocene), Silas, Choctaw County, Alabama. Paludicola 9: 183–209.

Mancini E.A. & Tew B.H. 1991. Relationships of Paleogene stage and planktonic foraminiferal zone boundaries to lithostratigraphic and allostratigraphic contacts in the eastern Gulf Coastal Plain. Journal of Foraminiferal Research 21 (1): 48–66. https://doi.org/10.2113/gsjfr.21.1.48

Manning E.M. 2003. The Eocene/Oligocene transition in marine vertebrates of the Gulf Coastal Plain. In: Prothero D.R., Ivany L.C. & Nesbitt E.A. (eds) From Greenhouse to Icehouse, the Marine Eocene–Oligocene Transition: 366–385. Columbia University Press, New York.

Martini E. 1971. Standard Tertiary and Quaternary calcareous nannoplankton zonation. In: Proceedings 2nd Planktonic Conference (Roma, 1970) (ed.) Technoscienza 2: 739–785.

Meyer O. 1889. Fish otoliths of the southern old-Tertiary. American Naturalist 23: 42–43.

Müller A. 1999. Ichthyofaunen aus dem atlantischen Tertiär der USA. Leipziger Geowissenschaften 9/10: 1–360.

Murray G.E. 1947. Cenozoic deposits of Central Gulf Coastal Plain. American Association of Petroleum Geologists Bulletin 31: 1825–50.

Murray G.E. 1961. Geology of the Atlantic and Gulf coastal province of North America. Harper & Brothers, New York.

Nelson J.S., Grande T.C. & Wilson M.V.H. 2016. Fishes of the World. John Wiley & Sons, Inc., Hoboken, NJ. https://doi.org/10.1002/9781119174844

Nolf D. 1976. Les otolithes de téléostéens néogènes de Trinidad. Eclogae Geologicae Helvetiae 69 (3): 703–742.

Nolf D. 1980. Étude monographique des otolithes des Ophidiiformes actuels et révision des espèces fossiles (Pisces, Teleostei). Mededelingen van de Werkgroep voor Tertiaire en Kwartaire Geologie 17 (2): 71–195.

Nolf D. 1985. Otolithi Piscium. In: Schultze H.P. (ed.) Handbook of Palaeoichthyology, Volume 10: 1–145. Gustav Fischer Verlag, Stuttgart.

Nolf D. 1988. Les otolithes de téléostéens éocènes d’Aquitaine (sud-ouest de la France) et leur intérêt stratigraphique. Mémoire de l’Académie royale de Belgique, Classe des Sciences, 4°, 2e série, 19 (2): 1–147.

Nolf D. 1995. Studies on fossil otoliths – the state of the art. In: Secor D., Dean J. & Campana S. (eds) Recent Developments in Fish Otolith Research: 513–544. University of South Carolina Press, Columbia, SC.

Nolf D. 2003. Revision of the American otolith-based fish species described by Koken in 1888. Louisiana Geological Survey, Geological Pamphlet 12: 1–20.

Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels.

Nolf D. 2018. Otoliths of Fishes from the North Sea and the English Channel. Royal Belgian Institute of Natural Sciences, Brussels.

Nolf D. & Aguilera O. 1998. Fish otoliths from the Cantaure Formation (early Miocene of Venezuela). Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 68: 237–262.

Nolf D. & Bajpai S. 1992. Marine middle Eocene fish otoliths from India and Java. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 62: 195–221.

Nolf D. & Cappetta H. 1976. Observations nouvelles sur les otolithes des téléostéens du calcaire Grossier (Eocène du Bassin de Paris). Geobios 9: 251–277.

Nolf D. & Girone A. 2008. Early Oligocene fish otoliths from the Castellane area (SE France) and an overview of Mediterranean teleost faunas at the Eocene–Oligocene boundary. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 248: 139–157. https://doi.org/10.1127/0077-7749/2008/0248-0139

Nolf D. & Lapierre H. 1979. Otolithes de poissons nouveaux ou peu connus du Calcaire Grossier et de la Formation d’Auvers (Eocène du Bassin parisien). Bulletin du muséum national d’histoire naturelle, Section C, Sciences de la Terre, Paléontologie, Géologie, Minéralogie 2: 79–125.

Nolf D. & Rundle A. 2018. Fish otoliths from the Paleocene and basal Ypresian of the North Sea Basin and dependencies (London Basin, Belgium, Paris Basin). Bulletin d’Information des Géologues du Bassin de Paris 55: 12–46.

Nolf D. & Steurbaut E. 1989. Importance and restrictions of the otolith-based fossil record of gadiform and ophidiiform fishes. In: Cohen D.M. (ed.) Papers on the Systematics of Gadiform Fishes 32: 47–58. Natural History Museum, Los Angeles.

Nolf D. & Stringer G.L. 1992. Neogene paleontology in the northern Dominican Republic 14. Otoliths of teleostean fishes. Bulletins of American Paleontology 102 (340): 41–81.

Nolf D. & Stringer G.L. 2003. Late Eocene (Priabonian) fish otoliths from the Yazoo Clay at Copenhagen, Louisiana. Louisiana Geological Survey Geological Pamphlet 13: 1–23.

Nolf D., De Potter H. & Lafond-Grellety J. 2009. Homage to Joseph Chaine and Jean Duvergier: The Diversity and Variability of Fish Otoliths. Palaeo Publishing and Library vzw, Mortsel, Belgium.

Pomerol C. 1973. Stratigraphie et Paléogéographie. Ere cénozoique (Tertiaire et Quaternaire). Doin, Paris.

Priem F. 1913. Sur les otolithes de l’Eocene du Cotentin et de Bretagne. Bulletin de la Société géologique de France 13: 151–158.

Přikryl T., Brzobohatý R. & Gregorová R. 2016. Diversity and distribution of fossil codlets (Teleostei, Gadiformes, Bregmacerotidae): review and commentary. Palaeobiodiversity and Palaeoenvironments 96: 13–39. https://doi.org/10.1007/s12549-015-0222-z

Přikryl T., Brzobohatý R. & Carnevale G. 2021. Skeletal remains with otoliths in situ of the Miocene croaker Trewasciaena cf. kokeni (Teleostei, Sciaenidae) from the Pannonian of the Vienna Basin. Bulletin of Geosciences 96 (1): 19–28. https://doi.org/10.3140/bull.geosci.1813

Schiebout J.A. & van den Bold W.A. 1986. Montgomery Landing Site, Marine Eocene (Jackson) of Central Louisiana. Proceedings of a Symposium, Gulf Coast Association of Geological Societies Annual Meeting, Baton Rouge, LA.

Schulte P., Speijer R., Mai H. & Kontny A. 2006. The Cretaceous–Paleogene (K–P) boundary at Brazos, Texas: Sequence stratigraphy, depositional events and the Chicxulub impact. Sedimentary Geology 184: 77–109. https://doi.org/10.1016/j.sedgeo.2005.09.021

Schwarzhans W. 1981a. Die Entwicklung der Familie Pterothrissidae (Elopomorpha; Pisces), rekonstruiert nach Otolithen. Senckenbergiana lethaea 62 (2–6): 77–91.

Schwarzhans W. 1981b. Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner geowissenschaftliche Abhandlungen A 32: 63–122.

Schwarzhans W. 1993. A comparative morphological treatise of recent and fossil otoliths of the family Sciaenidae (Perciformes). In: Pfeil F. (ed.) Piscium Catalogus, Otolithi Piscium: 1–245. Verlag Dr. Freidrich Pfeil, Munich.

Schwarzhans W. 1999. A comparative morphological treatise of recent and fossil otoliths of the order Pleuronectiformes. In: Pfeil F. (ed.) Piscium Catalogus, Otolithi Piscium: 1–391. Verlag Dr. Freidrich Pfeil, Munich.

Schwarzhans W. 2010. Otolithen aus den Gerhartsreiterschichten (Oberkreide: Maastricht) des Gerhartsreiter Grabens (Oberbayern). Palaeo Ichthyologica 4: 1–100.

Schwarzhans W. 2013. Otoliths from dredges in the Gulf of Guinea and off the Azores – an actuo-paleontological case study. Palaeo Ichthyologica 13: 7–40.

Schwarzhans W. 2019a. Reconstruction of the fossil marine bony fish fauna (Teleostei) from the Eocene to Pleistocene of New Zealand by means of otoliths. Memorie della Società Italiana di Scienze naturali e del Museo di Storia naturale di Milano 46: 3–326.

Schwarzhans W. 2019b. A comparative morphological study of Recent otoliths of the Congridae, Muraenesocidae, Nettastomatidae and Colocongridae (Anguilliformes). Memorie della Società Italiana di Scienze naturali e del Museo di Storia naturale di Milano 46: 327–354.

Schwarzhans W. 2019c. A comparative morphological study of Recent otoliths of the so-called Trachinoidei. Memorie della Società Italiana di Scienze naturali e del Museo di Storia naturale di Milano 46: 371–388.

Schwarzhans W. & Prokofiev A.M. 2017. Reappraisal of Synagrops Günther, 1887 with rehabilitation and revision of Parascombrops Alcock, 1889 including description of seven new species and two new genera (Perciformes: Acropomatidae). Zootaxa 4260 (1): 1–74. https://doi.org/10.11646/zootaxa.4260.1.1

Schwarzhans W. & Stringer G.L. 2020. Fish otoliths from the late Maastrichtian Kemp Clay (Texas, USA) and the early Danian Clayton Formation (Arkansas, USA) and an assessment of extinction and survival of teleost lineages across the K–Pg boundary based on otoliths. Rivista italiana di Paleontologia e Stratigraphia 126: 395–446. https://doi.org/10.13130/2039-4942/13425

Siesser W.G. 1983. Paleogene calcareous nannoplankton biostratigraphy: Mississippi, Alabama, and Tennessee. Bulletin Mississippi Department of Natural Resources Bureau of Geology 125: 1–61.

Smith E.A. & Johnson L.C. 1887. Tertiary and Cretaceous strata of the Tuscaloosa, Tombigbee, and Alabama Rivers. US Geological Survey Bulletin 43: 1–189.

Speijer R.P., Pälike H., Hollis C.J., Hooker J.J. & Ogg J.G. 2020. The Paleogene Period. In: Gradstein F.M., Ogg J.G., Schmitz M.D. & Ogg G.M. (eds) The Geologic Time Scale 2020: 1087–1140. Elsevier, Amsterdam. https://doi.org/10.1016/B978-0-12-824360-2.00028-0

Stanton R.J. & Nelson P.C. 1980. Reconstruction of the trophic web in paleontology: community structure in the Stone City Formation (Middle Eocene, Texas). Journal of Paleontology 54 (1): 118–135.

Stefanini G. 1924. Relations between American and European Tertiary echinoid faunas. Geological Society of America, Bulletin 35: 827–846.

Stenzel H.B., Krause E.K. & Twining J.T. 1957. Pelecypoda from the type locality of the Stone City Beds (middle Eocene) of Texas. The University of Texas Publication 5704: 1–237.

Stiassny M.L.J. 1993. What are grey mullets? Bulletin of Marine Science 52 (1): 197–219.

Stinton F.C. 1975. Fish otoliths from the English Eocene (part 1). Palaeontographical Society Monographs 544: 1–56.

Stinton F.C. 1977. Fish otoliths from the English Eocene (part 2). Palaeontographical Society Monographs 548: 57–126.

Stinton F.C. 1978. Fish otoliths from the English Eocene (part 3). Palaeontographical Society Monographs 555: 127–189.

Stinton F.C. 1980. Fish otoliths from the English Eocene (part 4). Palaeontographical Society Monographs 558: 191–258.

Stinton F.C. 1984. Fish otoliths from the English Eocene (part 5). Palaeontographical Society Monographs 565: 259–320.

Stinton F.C. & Nolf D. 1970. A teleost otolith fauna from the Sands of Lede, Belgium. Bulletin de la Société belge de Géologie, de Paléontologie et d’Hydrologie 78: 219–234.

Strickland L. 1985. Pamunkey River Field Trip. George Washington University, Washington, DC.

Stringer G.L. 1977. A Study of the Upper Eocene Otoliths and Related Fauna of the Yazoo Clay in Caldwell Parish, Louisiana. Master’s thesis, University of Louisiana at Monroe, Monroe, LA.

Stringer G.L. 1979. A study of the upper Eocene otoliths of the Yazoo Clay in Caldwell Parish, Louisiana. Tulane Studies in Geology and Paleontology 15: 95–105.

Stringer G.L. 1986. Teleostean otoliths and their paleoecological implications at the Montgomery Landing site. In: Schiebout J.A. & van den Bold W.A. (eds) Montgomery Landing Site, Marine Eocene (Jackson) of Central Louisiana: 209–222. Proceedings of a Symposium, Gulf Coast Association of Geological Societies, Baton Rouge, LA.

Stringer G.L. 2016. Evidence and implications of marine invertebrate settlement on Eocene otoliths from the Moodys Branch Formation of Montgomery Landing (Louisiana, USA). Cainozoic Research 16: 3–12.

Stringer G.L. & Bell D. 2018. Teleostean otoliths reveal diverse Plio–Pleistocene fish assemblages in coastal Georgia (Glynn County). Bulletin of the Florida Museum of Natural History 56 (3): 83–108.

Stringer G.L. & Breard S.Q. 1997. Comparison of otolith-based paleoecology to other fossil groups: an example from the Cane River Formation (Eocene) of Louisiana. Transactions of the Gulf Coast Association of Geological Societies 47: 563–570.

Stringer G.L. & Shannon K. 2019. The Pliocene Elizabethtown otolith assemblage (Bladen County, North Carolina, USA) with indications of a primary fish nursery area. Historical Biology 32 (8): 1108–1119. https://doi.org/10.1080/08912963.2019.1566324

Stringer G.L. & Starnes J.E. 2020. Significance of late Miocene fish otoliths (Micropogonias undulatus) from a Rangia johnsoni bed in the Pascagoula Formation in the subsurface of Mississippi. Southeastern Geology 54 (1): 21–28.

Stringer G.L, Ebersole J. & Ebersole S. 2020. First description of the fossil otolith-based sciaenid, Equetulus silverdalensis n. comb., in the Gulf Coastal Plain, USA, with comments on the enigmatic distribution of the species. PaleoBios 37: 1–12. https://doi.org/10.5070/P9371049670

Stringer G.L, Ebersole J., Starnes J. & Ebersole S. 2021. First Pliocene otolith assemblage from the Gulf Coastal Plain, Dauphin Island, Mobile County, Alabama, USA. Historical Biology 33 (10): 2147–2170. https://doi.org/10.1080/08912963.2020.1773457

Stringer G.L., Parmley D. & Quinn A. 2022. Eocene teleostean otoliths, including a new taxon, from the Clinchfield Formation (Bartonian) in Georgia, USA, with biostratigraphic, biogeographic, and paleoecologic implications. Palaeovertebrata 45 (1): e1. https://doi.org/10.18563/pv.45.1.e1

Taverne L. & Nolf D. 2010. Les Priacanthidae (Teleostei, Perciformes) des Sables de Lede (Éocène moyen, Belgique): ostéologie et otolithes. Bulletin de l’Institut royal des Sciences naturelles de Belgique 80: 187–243.

Toulmin L.D. 1977. Stratigraphic distribution of Paleocene and Eocene fossils in the Eastern Gulf Coast region. Geological Survey of Alabama, Monograph 13: 1–602.

Van der Laan R., Eschmeyer W. & Fricke R. 2014. Family-group names of Recent fishes. Zootaxa 3882 (1): 1–230. https://doi.org/10.11646/zootaxa.3882.1.1

Ward L.W. 1985. Stratigraphy and characteristic mollusks of the Pamunkey Group (Lower Tertiary) and the Old Church Formation of the Chesapeake Group-Virginia coastal plain. US Geological Survey Professional Paper 1346: 1–78. https://doi.org/10.3133/pp1346

Weems R.E., Self-Trail J.M. & Edwards L.E. 2004. Supergroup stratigraphy of the Atlantic and Gulf Coastal Plains (Middle? Jurassic through Holocene, eastern North America). Southeastern Geology 42: 191–216.

Weiler W. 1958. Fisch-Otolithen aus dem Ober-Oligozän und dem Mittelmiozän der Niederrheinischen Bucht. Fortschritte in der Geologie von Rheinland und Westfalen 1: 323–361.

Westgate J.W. 2001. Paleoecology and biostratigraphy of marginal marine Gulf Coast Eocene vertebrate localities. In: Gunnell G.F. (ed.) Eocene Biodiversity: Unusual Occurrences and Rarely Sampled Habitats: 263–297. Kluwer Academic/Plenum Publishing, New York. https://doi.org/10.1007/978-1-4615-1271-4_11

Wigley R.L. & Stinton F.C. 1973. Distribution of macroscopic remains of recent animals from marine sediments off Massachusetts. Fishery Bulletin 71 (1): 1–40.

Xu T., Tang D., Cheng Y. & Wang R. 2014. Mitogenomic perspectives into sciaenid fishes’ phylogeny and evolution origin in the New World. Gene 539: 91–98. https://doi.org/10.1016/j.gene.2014.01.048

Yancey T.E. & Yancey E.S. 1988. The lower Tertiary of the Texas Gulf Coast. In: Hayward O.T. (ed.) Geological Society of America Centennial Field Guide – South Central Section: 377–382. Geological Society of America, Boulder, CO.

Yancey T.E., Heizler M.T., Miller B.V. & Guillemette R.N. 2018. Eocene–Oligocene chronostratigraphy of ignimbrite flareup volcanic ash beds on the Gulf of Mexico coastal plains. Geosphere 14 (3): 1232–1252. https://doi.org/10.1130/GES01621.1

Zuschin M. & Stanton R.J. 2002. Paleocommunity reconstruction from shell beds: a case study from the Main Glauconite Bed, Eocene, Texas. Palaios 17: 602–614. https://doi.org/cb8kj5

How to Cite
Lin, C.-H., & Nolf, D. (2022). Middle and late Eocene fish otoliths from the eastern and southern USA. European Journal of Taxonomy, 814(1), 1-122. https://doi.org/10.5852/ejt.2022.814.1745