Abstract
The UNESCO World Heritage Site of Wadi Al-Hitan near Fayum in the Western Desert of Egypt is well-known for its remarkable abundance and diversity of late Eocene marine fossils, especially well-preserved cetaceans and sirenians. However, the taphonomy of the early Priabonian whales and sea cows is still not fully understood. These marine mammals often occur on specific horizons in the Birket Qarun Formation, for example, on a distinctive firmground called the Camp White Layer (CWL). Vertical, rod-like structures in the CWL were previously interpreted as “mangrove pneumatophores,” which have influenced hypotheses about the habitat of early whales. Here, we reexamine the rod-like structures of the CWL and show that they do not represent mangrove rooting structures based on an analysis of their morphology and an extensive comparison with rooting structures of living mangroves. Instead, the morphology of the rod-like structures indicates that they are trace fossils representing five size classes of marine invertebrate burrows, including burrows of weakly umbonate clams and Ophiomorphidae. The complex overprinting of all burrows toward the top of the CWL indicates stratigraphic condensation caused by low overall rates of sedimentation on a marine flooding surface in the transgressive systems tract. The large number of whale carcasses, abundant invertebrate skeletal remains, logs, marine invertebrate-encrusted celestine crystals, and an extensive, diverse burrow network reflect a combination of environmental and temporal condensation. A peculiar feature of the CWL, celestine crystal aggregates and gastropod celestine pseudomorphs, indicate an evaporative phase that is difficult to reconcile with the current sequence stratigraphic framework.
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References
Abdel-Fattah, Z. A., Gingras, M. K., Caldwell, M. W., & Pemberton, S. G. (2010). Sedimentary environments and depositional characteristics of the Middle to Upper Eocene whale-bearing succession in the Fayum Depression, Egypt. Sedimentology, 57, 446–476.
Andrews, C. W. (1906). A descriptive catalogue of the Tertiary vertebrata of the Fayum, Egypt. London: British Museum (Natural History) 324 pp.
Arab Republic of Egypt. (2004). Nomination file for the inscription of Wadi Al-Hitan (Whale Valley), the western desert of Egypt on the Natural World Heritage list. http://whc.unesco.org/uploads/nominations/1186.pdf. Accessed 8 February 2017.
Beadnell, H. J. L. (1905). The topography and geology of the Fayum Province of Egypt. Cairo: Survey Department of Egypt 101 pp.
Bebej, R. M., Zalmout, I. S., Abed El-Aziz, A. A., Antar, M. S., & Gingerich, P. D. (2016). First remingtonocetid archaeocete (Mammalia, Cetacea) from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution. Journal of Paleontology, 89, 882–893. https://doi.org/10.1017/jpa.2015.57.
Blanckenhorn, M. (1903). Neue geologisch-stratigraphische Beobachtungen in Aegypten. Sitzungsberichte der Mathematisch-physikalischen Classe der königlichen bayerischen Akademie der Wissenschaften. München, 32, 353–433.
Bown, T. M., & Kraus, M. J. (1988). Geology and paleoenvironment of the Oligocene Jebel Qatrani Formation and adjacent rocks, Fayum Depression, Egypt. U.S. Geological Survey Professional Paper, 1452, 1–60.
Dames, W. B. (1894). Über Zeuglodonten aus Aegypten und die Beziehungen der Archaeoceten zu den übrigen Cetaceen. Geologische und Paläontologische Abhandlungen, 5, 189–222.
El-Saadawi, W. E. (2005). A fossil rhizome at the mangrove site of Wadi Hitan, Egypt. Taeckholmia, 25, 129–136.
Fraas, E. (1904). Neue Zeuglodonten aus dem unteren Mitteleocän vom Mokattam bei Cairo. Geologische und Paläontologische Abhandlungen, Neue Folge, 6, 197–220.
Fraas, E. (1906). Wüstenreise eines Geologen in Ägypten. Kosmos, Handweiser für Naturfreunde, Stuttgart, 3, 263–269.
Gastaldo, R. A., & Huc, A.-Y. (1992). Sediment facies, depositional environments, and distribution of phytoclasts in the Recent Mahakam River delta, Kalimantan, Indonesia. Palaios, 7, 574–590.
Gingerich, P. D. (1992). Marine mammals (Cetacea and Sirenia) from the Eocene of Gebel Mokattam and Fayum, Egypt: stratigraphy, age, and paleoenvironments. University of Michigan, Papers on Paleontology 30, 84 pp.
Gingerich, P. D. (2007). Stromerius nidensis, new archaeocete (Mammalia, Cetacea) from the upper Eocene Qasr El-Sagha Formation, Fayum, Egypt. Contributions from the Museum of Paleontology, University of Michigan, 31, 363–378.
Gingerich, P. D., Smith, B. H., & Simons, E. L. (1990). Hind limbs of Eocene Basilosaurus isis: evidence of feet in whales. Science, 249, 154–157.
Gingerich, P. D., Abd-Elshafy, E., Metwally, H. M., Zalmout, I. S., & Antar, M. S. M. (2011). Paleoenvironmental and taphonomic assessment of Eocene marine faunas of Wadi Al-Hitan and Siwa areas, Egypt. Egyptian Journal of Paleontology, 11, 171–190.
Gingerich, P. D., Zalmout, I. S., Antar, M. S. M., Williams, E. M., Carlson, A. E., Kelly, D. C., & Peters, S. E. (2012). Large-scale glaciation and deglaciation of Antarctica during the late Eocene: reply. Geology, 40, e255.
Haq, B. U., Hardenbol, J., & Vail, P. R. (1987). Chronology of fluctuating sea levels since the Triassic. Science, 235, 1156–1167.
Hardenbol, J., Jacques, T., Martin, F. B., Pierre-Charles, G., & Vail, P. R. (1998). Mesozoic and Cenozoic sequence chronostratigraphic framework of European basins. Society for Sedimentary Geology Special Publication, 60, 3–13.
Mosbrugger, V., Gee, C. T., Belz, G., & Ashraf, A. R. (1994). Three-dimensional reconstruction of an in situ Miocene peat forest from the Lower Rhine Embayment, NW Germany—new methods in palaeovegetation analysis. Palaegeography, Palaeoclimatology, Palaeoecology, 10, 295–317.
Owen, R. (1875). On fossil evidences of a sirenian mammal (Eorherium aegyptiacum) from the nummulitic Eocene of the Mokattam cliffs, near Cairo. Quarterly Journal of the Geological Society of London, 31, 100–105.
Peters, S. E., Antar, M. S. M., Zalmout, I. S., & Gingerich, P. D. (2009). Sequence stratigraphic control on preservation of late Eocene whales and other invertebrates at Wadi Al-Hitan, Egypt. Palaios, 24, 290–302.
Peters, S. E., Carlson, A. E., Kelly, D. C., & Gingerich, P. D. (2010). Large-scale glaciation and deglaciation of Antarctica during the Late Eocene. Geology, 38, 723–726.
Salem, R. (1976). Evolution of Eocene–Miocene sedimentation patterns in parts of northern Egypt. AAPG Bulletin, 60, 34–64.
Schweinfurt, G. A. (1886). Reise in das Depressionsgebiet im Umkreise des Fajum im Januar 1886. Zeitschrift der Gesellschafi fir Erdkurde zu Berlin, 21, 96–149.
Seilacher, A. (2007). Trace fossil analysis. Berlin: Springer Verlag 226 pp.
Stromer, E. (1903). Zeuglodon-Reste aus dem oberen Mitteleocän des Fajum. Beiträge zur Paläontologie und Geologie Österreich-Ungarns und des Orients, Wien, 15, 65–100.
Stromer, E. (1908). Die Archaeoceti des ägyptischen Eozäns. Beiträge zur Paläontologie und Geologie Österreich-Ungarns und des Orients, Wien, 21, 106–178.
Strougo, A. (2008). The Mokattamian stage: 125 years later. Middle East Research Center, Ain Shams University, Cairo, Earth Science Series, 22, 47–108.
Strougo, A., Faris, M., Haggag, M. A. Y., Abul-Nasr, R. A., & Gingerich, P. D. (2013). Planktonic foraminifera and calcareous nannofossil biostratigraphy through the middle to late Eocene transition at Wadi Hitan, Fayum Province, Egypt. Contributions from the Museum of Paleontology, University of Michigan, 32, 111–138.
Tomlinson, P. B. (2016). The botany of mangroves (2nd ed.). Cambridge: Cambridge University Press 454 pp.
Uhen, M. D. (2004). Form, function, and anatomy of Dorudon atrox (Mammalia, Cetacea): an archaeocete from the middle to late Eocene of Egypt. University of Michigan Papers on Paleontology, 34, 1–222.
Vondra, C. F. (1974). Upper Eocene transitional and nearshore marine Qasr el Sagha Formation, Fayum Depression, Egypt. Annals of the Geological Survey of Egypt, 4, 79–94.
Zalmout, I. S., & Gingerich, P. D. (2012). Late Eocene sea cows (Mammalia, Sirenia) from Wadi Al Hitan in the Western Desert of Fayum, Egypt. University of Michigan Papers on Paleontology, 37, 1–158.
Acknowledgements
We are indebted to the park rangers and staff of the Wadi El-Rayan Protected Area for their able and tireless assistance, to the Egyptian Environmental Affairs Agency for encouraging and supporting our field studies, and to Georg Oleschinski (University of Bonn) for his assistance with Figs. 3d–g and 7a. We also thank reviewers Robert A. Gastaldo (Colby College, Maine, USA) and Michael R.W. Amler (Universität zu Köln, Germany) for helpful reviews.
Funding
This research was conducted during the tenure of research funds from Deutsche Forschungsgemeinschaft FOR 533 to CTG and PMS. Field work was supported by the National Geographic Society grant 7226-04 and National Science Foundation grant EAR-0920972 to PDG.
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Gee, C.T., Sander, P.M., Peters, S.E. et al. Fossil burrow assemblage, not mangrove roots: reinterpretation of the main whale-bearing layer in the late Eocene of Wadi Al-Hitan, Egypt. Palaeobio Palaeoenv 99, 143–158 (2019). https://doi.org/10.1007/s12549-018-0337-0
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DOI: https://doi.org/10.1007/s12549-018-0337-0