Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa

Abstract

Comparative morphometric study of recently recovered fossil elephant molars from Natodomeri, Kenya identifies them as belonging to Elephas jolensis and confirms the presence of this species in Members I and II of the Kibish Formation. Improved datation of these geological units constrains them between 205 and 130 ka. Elephas jolensis is also reported from localities in northern, northwestern, eastern, and southern Africa. Thus, including its Natodomeri occurrence, E. jolensis appears to have been pan-African in distribution. Despite the wide geographic distribution of the species, molars of E. jolensis are remarkably uniform morphometrically. They are characterized by their extreme hypsodonty, high amplitude of enamel folding, high lamellar frequency, and plates that are anteroposteriorly thick relative to transverse valley interval spacing. In addition, they exhibit only a modest number of plates (<20 in M3/m3). Elephas jolensis either evolved from or represents the last stage of Elephas recki, the dominant elephant species in East Africa during the late Pliocene-Pleistocene. The dental morphology and isotopic composition of E. jolensis indicates that, like E. recki, it was a dedicated grazer. In the Kibish Formation, E. jolensis is succeeded by Loxodonta africana at 130 ka, coincident with an intensely cool, dry interval marked by episodes of extreme drought. This marked the extirpation of Elephas on the continent. The intensity and increased rate of climate fluctuation may have played an important role in the demise of the specialist, grazing E. recki-E. jolensis lineage in favor of a generalist, mixed feeder such as L. africana.

This is a preview of subscription content, access via your institution.

We’re sorry, something doesn't seem to be working properly.

Please try refreshing the page. If that doesn't work, please contact support so we can address the problem.

Fig. 1
Fig. 2

References

  1. Arambourg C (1938) Mammifères fossils du Maroc. Mem Soc Sci Nat Maroc 46:1–74

  2. Arambourg C (1952) Note préliminaire sur quelques Éléphants fossils de Berbérie. Bull Mus Natl His Nat 2e sér 24:407–418

  3. Arambourg C (1960) Au sujet de Elephas iolensis Pomel. Bull Archéol Marocaine 3:93–104

  4. Arambourg C (1970) Les Vertébrés du Pléistocène de l’Afrique du Nord. Arch Mus Natl Hist Nat 10:1–126

  5. Assefa Z, Yirga S, Reed KE (2008) The large-mammal fauna from the Kibish Formation. J Hum Evol 55:501–512

  6. Beaumont PB, Vogel JC (2006) On a timescale for the past million years of human history in central South Africa. So Afr J Sci 102:217–228

  7. Beden M (1980) Elephas recki Dietrich, 1915 (Proboscidea, Elephantidae). Évolution au cours du Plio-Pléistocène en Afrique Orientale. Géobios 13:891–901

  8. Beden M (1983) Family Elephantidae. In: Harris JM (ed) Koobi Fora Research Project. Volume 2. The Fossil Ungulates: Proboscidea, Perissodactyla, and Suidae. Clarendon Press, Oxford, pp 40–129

  9. Beden M (1985) Les Proboscidiens des Grands Gisements à Hominidés Plio-Pléistocènes d’Afrique Orientale. In: L’Environnement des Hominidés au Plio-Pléistocène. Fondation Singer-Polignac, Masson, Paris, pp 21–44

  10. Beden M (1987) Les Faunes Plio-Pléistocène de la Vallée de l’Omo. Tome 2. Les Eléphantidés (Mammalia, Proboscidea). Éditions du Centre National de la Recherche Scientifique, Paris

  11. Behrensmeyer AK (1976) Lothagam Hill, Kanapoi, and Ekora: a general summary of stratigraphy and faunas. In: Coppens Y, Howell FC, Isaac GL, Leakey REF (eds) Earliest Man and Environments in the Lake Rudolf Basin. University of Chicago Press, Chicago, pp 163–170

  12. Behrensmeyer AK, Potts R, Plummer T, Tauxe L, Opdyke N, Jorstad T (1995) The Pleistocene locality of Kanjera, western Kenya: stratigraphy, chronology and paleoenvironments. J Hum Evol 29:247–274

  13. Biberson P (1952a) Découverte d’une molaire d’Eléphant a la carrière de Sidi-Abder-Rahmane. Bull Soc Sci Nat Maroc 2:34–35

  14. Biberson P (1952b) Présentation d’un fragment de mandibule d’Eléphant découverte à la carrière de la S.T.I.C. près de Casablanca. Bull Soc Sci Nat Maroc 4:81–83

  15. Biberson P, Ennouchi E (1952) Présence de Elephas recki Dietrich dans la carrière de Sidi-Abder-Rahmane, à Casablanca. C R Soc Geol Fr 6:90–92

  16. Bobe R (2011) Fossil mammals and paleoenvironments in the Omo-Turkana Basin. Evol Anthropol 20:254–263

  17. Boule M (1900) Étude paléontologique et archéologique sur la station paléolithique du lac Karar (Algérie). L’Anthropologie 11:1–21

  18. Brandt AL, Ishida Y, Georgiadis NJ, Roca AL (2012) Forest elephant mitochondrial genomes reveal that elephantid diversification in Africa tracked climate transitions. Mol Ecol 21:1175–1189

  19. Brown FH, Fuller CR (2008) Stratigraphy and tephra of the Kibish Formation, southwestern Ethiopia. J Hum Evol 55:366–403

  20. Brown FH, McDougall I (2011) Geochronology of the Turkana depression of northern Kenya and southern Ethiopia. Evol Anthropol 20:217–227

  21. Brown FH, McDougall I, Fleagle JG (2012) Correlation of the KHS tuff of the Kibish Formation to volcanic ash layers at other sites, and the age of early Homo sapiens (Omo I and Omo II). J Hum Evol 63:577–585

  22. Butzer KW, Brown FG, Thurber DL (1969) Horizontal sediments of the lower Omo Valley: the Kibish Formation. Quaternaria 11:15–29

  23. Butzer KW, Helgren DM, Fock GJ, Stuckenrath R (1973) Alluvial terraces of the lower Vaal River, South Africa: a reappraisal and reinvestigation. J Geol 81:341–362

  24. Butzer KW, Isaac GL, Richardson JL, Washbourn-Kamau C (1972) Radiocarbon dating of east African lake levels. Science 175:1069–1076

  25. Butzer KW, Thurber DL (1969) Some late Cenozoic sedimentary formations of the lower Omo Basin. Nature 222:1138–1143

  26. Cerling TE, Andanje SA, Blumenthal SA, Brown FH, Chritz KL, Harris JM, Hart JA, Kirera FM, Kaleme P, Leakey LN, Leakey MG, Levin NE, Manthi FK, Passey BH, Uno KT (2015) Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 million years ago. Proc Natl Acad Sci USA 112:11467–11472

  27. Cerling TE, Harris JM (1999) Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia 120:347–363

  28. Cerling TE, Harris JM, Leakey MG (1999) Browsing and grazing in elephants: the isotope record of fossil and modern proboscideans. Oecologia 120:364–374

  29. Cerling TE, Omondi P, Macharia AN (2007) Diets of Kenyan elephants from stable isotopes and the origin of confiscated ivory in Kenya. Afr J Ecol 45:614–623

  30. Cerling TE, Passey BH, Ayliffe LK, Cook CS, Ehleringer JR, Harris JM, Dhidha MB, Kasiki SM (2004) Orphans’ tales: seasonal dietary changes in elephants from Tsavo National Park, Kenya. Palaeogeogr Palaeoclimatol Palaeoecol 206:367–376

  31. Cerling TE, Wittemyer G, Ehleringer JR, Remien CH, Douglas-Hamilton I (2009) History of animals using isotope records (HAIR): a 6-year dietary history of one family of African elephants. Proc Natl Acad Sci USA 106:8093–8100

  32. Coffing K, Feibel CS, Leakey M, Walker A (1994) Four-million-year-old hominids from East Lake Turkana, Kenya. Am J Phys Anthropol 93:55–65

  33. Cohen AS, Stone JR, Beuning KRM, Park LE, Reinthal PN, Dettman D, Scholz CA, Johnson TC, King JW, Talbot MR, Brown ET, Ivory SJ (2007) Ecological consequences of early late Pleistocene megadroughts in tropical Africa. Proc Natl Acad Sci USA 104:16422–16427

  34. Cooke HBS (1939) Discussion on H.B. Maufe’s paper, “new sections in the Kalahari beds at the Victoria Falls, Rhodesia.” Proc Geol Soc So Afr 42:51–52

  35. Cooke HBS (1947) Variation in the molars of the living African elephant and a critical revision of the fossil Proboscidea of southern Africa. Am J Sci 245:434–457

  36. Cooke HBS (1949) Fossil mammals of the Vaal River deposits. Geological Survey (South Africa) Memoir 35 (III):1–169

  37. Cooke HBS (1960) Further revision of the fossil Elephantidae of southern Africa. Palaeontol Afr 7:46–58

  38. Cooke HBS, Clark JD (1939) New fossil elephant remains from the Victoria Falls, northern Rhodesia, and a preliminary note on the geology and archaeology of the deposit. Trans R Soc So Afr 27:287–319

  39. Cooke HBS, Coryndon SC (1970) Pleistocene mammals from the Kaiso Formation and other related deposits in Uganda. In: Leakey LSB, Savage RJG (eds) Fossil Vertebrates of Africa, Volume 2. Academic Press, London, pp 107–224

  40. Cooke HBS, Maglio VJ (1972) Plio-Pleistocene stratigraphy in East Africa in relation to proboscidean and suid evolution. In: Bishop WW, Miller JA (eds) Calibration of Hominoid Evolution: Recent Advances in Isotopic and Other Dating Methods Applicable to the Origin of Man. Wenner-Gren Foundation for Anthropological Research, Scottish Academic Press, New York, pp 303–329

  41. Coppens Y, Gaudant M (1976) Découverte d’Elephas iolensis Pomel dans le Tyrrhénien de Tunisie. Bull Soc Géol Fr 18:171–177

  42. Coppens Y, Maglio VJ, Madden CT, Beden M (1978) Proboscidea. In: Maglio VJ, Cooke HBS (eds) Evolution of African Mammals. Harvard University Press, Cambridge, pp 336–367

  43. Dart RA (1927) Mammoths and man in the Transvaal. Suppl Nature 3032:41–48

  44. Dart RA (1929) Mammoths and other fossil elephants of the Vaal and Limpopo watersheds. So Afr J Sci 26:698–731

  45. de Lamothe, L. (1904) Note sur les relations stratigraphiques qui paraissent exister entre les anciennes lignes de rivage de la côte Algérienne et celles signalées sur la côte niçoise. Bull Soc Géol Fr 4:14–38

  46. Deperet C, Mayet L (1923) Les rameaux phyletiques des Elephants. C R Acad Sci Paris 176:1278–1281

  47. Feibel CS (2003) Stratigraphy and depositional setting of the Pliocene Kanapoi Formation, lower Kerio Valley. In: Harris JM, Leakey, MG (eds) Geology and Vertebrate Paleontology of the Early Pliocene Site of Kanapoi, Northern Kenya. Nat Hist Mus LA County Contrib Sci 498:9–20

  48. Fleagle JG, Assefa Z, Brown FH, Shea JJ (2008) Paleoanthropology of the Kibish Formation, southern Ethiopia: introduction. J Hum Evol 55:360–365

  49. Geraads D (1980) Le faune des sites à “Homo erectus” des carrières Thomas (Casablanca, Maroc). Quaternaria 22:65–94

  50. Haile-Selassie Y (2001) Late Miocene mammalian fauna from the Middle Awash Valley, Ethiopia. Unpublished PhD dissertation, University of California, Berkeley, 425 pp

  51. Harris JM, Leakey MG, Cerling TE (2003) Early Pliocene tetrapod remains from Kanapoi, Lake Turkana Basin, Kenya. In: Harris JG, Leakey MG (eds) Geology and Vertebrate Paleontology of the Early Pliocene Site of Kanapoi, Northern Kenya. Nat Hist Mus LA County Contrib Sci 498:39–113

  52. Haughton SH (1932) On some South African Proboscidea. Trans R Soc So Afr 21:1–18

  53. Hayward MW, Hayward MD (2012) Waterhole use by African fauna. So Afr J Wildl Res 42:117–127

  54. Hendey QB (1967) A specimen of ‘Archidiskodon’ cf. transvaalensis from the south-western Cape Province. So Afr Archaeol Bull 22:53–56

  55. Hopwood AT, Hollyfield JP (1954) An Annotated Bibliography of the Fossil Mammals of Africa (1742–1950). Fossil Mammals of Africa No. 8. British Museum (Natural History), London

  56. Kalb JE, Mebrate A (1993) Fossil elephantoids from the hominid-bearing Awash group, Middle Awash Valley, Afar depression, Ethiopia. Trans Am Philos Soc 83:1–114

  57. Klein RG (1984) The large mammals of southern Africa: late Pliocene to recent. In: Klein, RG (ed) Southern African Prehistory and Paleoenvironments. AA Balkema, Rotterdam, pp 107–146

  58. Klein RG (1988) The archaeological significance of animal bones from Acheulean sites in southern Africa. Afr Archaeol Rev 6:3–25

  59. Klein RG (2000) The earlier Stone age of southern Africa. So Afr Archaeol Bull 55:107–122

  60. Kullmer O, Sandrock L, Viola TB, Hujer W, Said H, Seidler H (2008) Suids, elephantoids, paleochronology, and paleoecology of the Pliocene hominid site Galili, Somali region, Ethiopia. Palaios 23:452–464

  61. Lister AM (2004) Ecological interactions of elephantids in Pleistocene Eurasia: Palaeoloxodon and Mammuthus. In: Goren-Inbar N, Speth JD (eds) Human Paleoecology in the Levantine Corridor. Oxbow Books, Oxford, pp 53–60

  62. Lister AM (2013) The role of behavior in adaptive morphological evolution of African proboscideans. Nature 500:331–334

  63. Lister AM, Dirks W, Assaf A, Chazan M, Goldberg P, Applbaum YH, Greenbaum N, Horwitz LK (2013) New fossil remains of Elephas from the southern Levant: implications for the evolutionary history of the Asian elephant. Palaeogeogr Palaeoclimatol Palaeoecol 386:119–130

  64. Maglio VJ (1970a) Four new species of Elephantidae from the Plio-Pleistocene of northwestern Kenya. Breviora 341:1–43

  65. Maglio VJ (1970b) Early Elephantidae of Africa and a tentative correlation of African Plio-Pleistocene deposits. Nature 225:328–332

  66. Maglio VJ (1973) Origin and evolution of the Elephantidae. Trans Am Philos Soc 63:1–149

  67. Maglio VJ, Hendey QB (1970) New evidence relating to the supposed stegolophodont ancestry of the Elephantidae. So Afr Archaeol Bull 25:85–87

  68. Manthi FK, Brown FH, Plavcan MJ, Werdelin L (2017) Gigantic lion, Panthera leo, from the Pleistocene of Natodomeri, eastern Africa. J Paleontol. https://doi.org/10.1017/jpa.2017.68

  69. Marinheiro J, Mateus O, Alaoui A, Amani F, Nami M, Ribeiro C (2014) New Quaternary fossil sites from the middle Atlas of Morocco. Com Geol Portugal 101, Especial I:485–488

  70. McDougall I, Brown FH (2008) Geochronology of the pre-KBS tuff sequence, Omo group, Turkana Basin. J Geol Sci 165:549–562

  71. McDougall I, Brown FH, Fleagle JG (2005) Stratigraphic placement and age of modern humans from Kibish, Ethiopia. Nature 433:733–736

  72. McDougall I, Feibel CS (2003) Numerical age control for the Miocene-Pliocene succession at Lothagam, a hominoid-bearing sequence in the northern Kenya rift. In: Leakey MG, Harris JM (eds) Lothagam: The Dawn of Humanity in Eastern Africa. Columbia University Press, New York, pp 43–64

  73. Meyer M, Palkopoulou E, Baleka S, Stiller M, Penkman KEH, Alt KW, Ishida Y, Mania D, Mallick S, Meijer T, Meller H, Nagel S, Nickel B, Ostritz S, Rohland N, Schauer K, Schüler T, Roca AL, Reich D, Shapiro B, Hofreiter M (2017) Palaeogenomes of Eurasian straight-tusked elephants challenge the current view of elephant evolution. eLife. https://doi.org/10.7554/eLife.25413.001

  74. Murata Y, Yonezawa T, Kihara I, Kashiwamura T, Sugihara Y, Nikaido M, Okada N, Endo H, Hasegawa M (2009) Chronology of the extant African elephant species and case study of the species identification of the small African elephant with the molecular phylogenetic method. Gene 441:176–186

  75. O’Regan HJ, Bishop LC, Lamb A, Elton S, Turner A (2005) Large mammal turnover in Africa and the Levant between 1.0 and 0.5 ma. In: Head MJ, Gibbard PL (eds) Early-Middle Pleistocene Transitions: The Land-Ocean Evidence. Geological Society of London Special Publications 247, London, pp 231–249

  76. Osborn HF (1928) Mammoths and man in the Transvaal. Nature 121:672–673

  77. Osborn HF (1934) Primitive Archidiskodon and Palaeoloxodon of South Africa. Am Mus Novitates 741:1–15

  78. Osborn HF (1942) Proboscidea: A Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World: Vol. II. Stegodontoidea, Elephantoidea. American Museum Press, New York, 828 pp

  79. Passey BH, Perkins ME, Voorhies MR, Cerling TE, Harris JM, Tucker ST (2002) Timing of C4 biomass expansion and environmental change in the Great Plains: an isotopic record from fossil horses. J Geol 110:123–140

  80. Passey BH, Robinson TF, Ayliffe LK, Cerling TE, Sponheimer M, Dearing MD, Roeder BL, Ehleringer JR (2005) Carbon isotopic fractionation between diet, breath, and bioapatite in different mammals. J Archaeol Sci 32:1459–1470

  81. Plummer TW, Potts R (1989) Excavations and new findings at Kanjera, Kenya. J Hum Evol 18:269–276

  82. Pomel A (1895) Les Éléphants quaternaires. In: Monographies des Vertébrés fossiles de l’Algérie. Publ Serv Carte Géol Algérie 6:1–68

  83. Porat N, Chazan M, Grün R, Aubert M, Eisenmann V, Horwitz LK (2010) New radiometric ages for the Fauresmith industry from Kathu Pan, southern Africa: implications for the earlier to middle Stone Age transition. J Archaeol Sci 37:269–283

  84. Potts R, Behrensmeyer AK, Faith JT, Tryon CA, Brooks AS, Yellen JE, Deino AL, Kinyanjui R, Clark JB, Haradon CM, Levin NE, Meijer HJM, Veatch EG, Owen RB, Renaut RW (2018) Environmental dynamics during the onset of the middle Stone age in eastern Africa. Science 360:86–90

  85. Potts R, Deino A (1995) Mid-Pleistocene change in large mammal faunas of East Africa. Quaternary Res 43:106–113

  86. Rohland N, Malaspinas A, Pollack JL, Slatkin M, Matheus P, Hofreiter M (2007) Proboscidean mitogenomics: chronology and mode of elephant evolution using mastodon as outgroup. PLoS Biol 5:1663–1671

  87. Saarinen J, Karme A, Cerling T, Uno K, Säilä L, Kasiki S, Ngene S, Obari T, Mbua E, Manthi FK, Fortelius M (2015) A new tooth wear-based dietary analysis method for Proboscidea (Mammalia). J Vertebr Paleontol. https://doi.org/10.1080/02724634.2014.918546

  88. Saegusa H, Gilbert WH (2008) Elephantidae. In: Gilbert WH, Asfaw B (eds), Homo erectus. Pleistocene Evidence from the Middle Awash. University of California Press, Berkeley, pp 193–226

  89. Sanders WJ (2006) Comparative description and taxonomy of proboscidean fossils from Langebaanweg, South Africa. Afr Nat Hist 2:196–197

  90. Sanders WJ (2007) Taxonomic review of fossil Proboscidea (Mammalia) from Langebaanweg, South Africa. Trans R Soc So Afr 62:1–16

  91. Sanders WJ, Gheerbrant E, Harris JM, Saegusa H, Delmer C (2010) Proboscidea. In: Werdelin L, Sanders WJ (eds) Cenozoic Mammals of Africa. University of California Press, Berkeley, pp 161–251

  92. Sanders WJ, Haile-Selassie Y (2012) A new assemblage of mid-Pliocene proboscideans from the Woranso-mille area, Afar region, Ethiopia: taxonomic, evolutionary, and paleoecological considerations. J Mammal Evol 19:105–128

  93. Scholz CA, Johnson TC, Cohen AS, King JW, Peck JA, Overpeck JT, Talbot MR, Brown ET, Kalindekafe L, Amoako PYO, Lyons RP, Shanahan TM, Castañeda IS, Heil CW, Forman SL, McHargue LR, Beuning KR, Gomez J, Pierson J (2007) East African megadroughts between 135 and 75 thousand year ago and bearing on early-modern human origins. Proc Natl Acad Sci USA 104:16416–16421

  94. Sukumar R, Bhattacharya SK, Krishnamurthy RV (1987) Carbon isotope evidence for different feeding patterns in an Asian elephant population. Curr Sci 56:11–14

  95. Sukumar R, Ramesh R (1992) Stable carbon isotope ratios in Asian elephant collagen: implications for dietary studies. Oecologia 91:536–539

  96. Tamrat E, Thouveny N, Taieb M, Opdyke ND (1995) Revised magnetostratigraphy of the Plio-Pleistocene sedimentary sequence of the Olduvai Formation (Tanzania). Palaeogeogr Palaeoclimatol Palaeoecol 114:273–283

  97. Tassy P (2003) Elephantoidea from Lothagam. In: Leakey MG, Harris JM (eds) Lothagam: The Dawn of Humanity in Eastern Africa. Columbia University Press, New York, pp 331–358

  98. Tassy P, Debruyne R (2001) The timing of early Elephantinae differentiation: the palaeontological record, with a short comment on molecular data. In: Cavarretta G, Gioia P, Mussi M, Palombo MR (eds) Proceedings of the First International Congress of La Terra Degli Elefanti: The World of Elephants. Consiglio Nazionale Delle Richerche, Rome, pp 685–687

  99. Tejada-Lara JV, MacFadden BJ, Bermudez L, Gianmarco R, Salas-Gismondi R, Flynn JJ (2018) Body mass predicts isotope enrichment in herbivorous mammals. Proc R Soc B 285:20181020

  100. Tierney JE, deMenocal PB, Zander PD (2017) A climatic context for the out-of-Africa migration. Geology. https://doi.org/10.1130/G39457.1

  101. Todd NE (2001) African Elephas recki: time, space and taxonomy. In: Cavarretta G, Gioia P, Mussi M, Palombo MR (eds) Proceedings of the First International Congress of La Terra Degli Elefanti: The World of Elephants. Consiglio Nazionale Delle Richerche, Rome, pp 693–697

  102. Todd NE (2005) New phylogenetic analysis of the family Elephantidae based on cranial-dental morphology. Anat Rec 293:74–90

Download references

Acknowledgments

We thank the Government of Kenya for permission to work at Natodomeri and the National Museums of Kenya (NMK) for their support of this project. Special appreciation is extended to Rose Nyaboke for collections assistance at the NMK and hospitality, and to Carol Abraczinskas for expert production of figures. The Turkana County administration, the Turkana people in the Natodomeri area, and West Turkana Palaeo Project crew members provided invaluable support to the project. Funding for the project was provided by the L.S.B. Leakey Foundation and the Palaeontological Scientific Trust (PAST) of South Africa. Research travel of WJS was supported by National Science Foundation grant NSF SBR 124811. The stable isotope work for the project was supported by National Science Foundation grant NSF-1740383.

Data Availability Statement

All data generated and analyzed during the current study are included in this published article.

Author information

Affiliations

Author notes

  1. Francis H. Brown is deceased. This paper is dedicated to his memory.

    • Francis H. Brown
Authors

Corresponding author

Correspondence to William J. Sanders.

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Manthi, F.K., Sanders, W.J., Plavcan, J.M. et al. Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa. J Mammal Evol 27, 483–495 (2020). https://doi.org/10.1007/s10914-019-09474-9

Download citation

Keywords

  • Natodomeri, Kenya
  • Kibish Formation
  • Elephantidae
  • Elephas jolensis
  • Late middle Pleistocene