diversity Review A Giant Ostrich from the Lower Pleistocene Nihewan Formation of North China, with a Review of the Fossil Ostriches of China Eric Buffetaut 1,2,* and Delphine Angst 3 1 Centre National de la Recherche Scientifique—CNRS (UMR 8538), Laboratoire de Géologie de l’Ecole Normale Supérieure, PSL Research University, 24 rue Lhomond, CEDEX 05, 75231 Paris, France 2 Palaeontological Research and Education Centre, Maha Sarakham University, Maha Sarakham 44150, Thailand 3 School of Earth Sciences, University of Bristol, Life Sciences Building, 24 Tyndall Avenue, Bristol BS8 1TQ, UK; [email protected] * Correspondence: [email protected] Abstract: A large incomplete ostrich femur from the Lower Pleistocene of North China, kept at the Muséum National d’Histoire Naturelle (Paris), is described. It was found by Father Emile Licent in 1925 in the Nihewan Formation (dated at about 1.8 Ma) of Hebei Province. On the basis of the minimum circumference of the shaft, a mass of 300 kg, twice that of a modern ostrich, was obtained. The bone is remarkably robust, more so than the femur of the more recent, Late Pleistocene, Struthio anderssoni from China, and resembles in that regard Pachystruthio Kretzoi, 1954, a genus known from the Lower Pleistocene of Hungary, Georgia and the Crimea, to which the Nihewan specimen is referred, as Pachystruthio indet. This find testifies to the wide geographical distribution Citation: Buffetaut, E.; Angst, D. A of very massive ostriches in the Early Pleistocene of Eurasia. The giant ostrich from Nihewan was Giant Ostrich from the Lower contemporaneous with the early hominins who inhabited that region in the Early Pleistocene. Pleistocene Nihewan Formation of North China, with a Review of the Keywords: ostrich; China; Nihewan; Pleistocene; femur Fossil Ostriches of China. Diversity 2021, 13, 47. https://doi.org/ 10.3390/d13020047 1. Introduction Academic Editor: Michael Wink The Lower Pleistocene fossiliferous beds of the Nihewan Basin (Figure1) in northern Received: 30 December 2020 Hebei Province (North China) have been known for their vertebrate remains since the 1920s. Accepted: 22 January 2021 More recently, abundant evidence of early human occupation has also come to light ([1], Published: 26 January 2021 and references therein). The fossil mammals from the various formations of the Nihewan Basin have attracted considerable attention, starting with the pioneering paper by Teilhard Publisher’s Note: MDPI stays neutral de Chardin and Piveteau [2]. However, although bird bones have been mentioned, few of with regard to jurisdictional claims in them have been described in detail, with the notable exception of a metatarsus belonging published maps and institutional affil- to a crow (Corvus)[3]. iations. Here we describe an ostrich femur, collected in the 1920s and kept in the paleontology collection of the Muséum National d’Histoire Naturelle (Paris, France). Although this bone is poorly preserved, a body mass estimate based on its circumference shows that it belonged to a giant ostrich, significantly larger than the living Struthio camelus. It provides new Copyright: © 2021 by the authors. evidence of the wide geographical distribution of giant ostriches in the Early Pleistocene Licensee MDPI, Basel, Switzerland. of Eurasia. This article is an open access article A note on spelling: in this paper we have used the modern p¯ıny¯ın spelling for place distributed under the terms and names. In the 1920s, a different transliteration was used by paleontologists working in conditions of the Creative Commons China: “Nihowan” instead of Nihewan, “Sangkan Ho” instead of Sanggan He, etc. Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ Institutional abbreviations: MNHN: Muséum National d’Histoire Naturelle, Paris, 4.0/). France. IVPP: Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China. Diversity 2021, 13, 47. https://doi.org/10.3390/d13020047 https://www.mdpi.com/journal/diversity Diversity 2021, 13, 47 2 of 12 Diversity 2021, 13, x FOR PEER REVIEW 2 of 13 Figure 1. MapFigure of part 1. Map of northern of part Chinaof northern showing China location showing of the locati Nihewanon of the Basin, Nihewan WNW Basin, of Beijing, WNW with of Beijing, general location in China (smallwith map general at lower location right corner). in China Modified (small map after at Teilhard lower right de Chardin corner). andModified Piveteau after [2 ].Teilhard de Char- din and Piveteau [2]. 2. Discovery and Geological Setting of the Specimen 2. Discovery and Geological Setting of the Specimen Following discoveries of fossil bones by Father Vincent, a missionary based in Ni- Followinghewan discoveries village of (some fossil 150bones km by NW Father of Beijing; Vincent, Figure a missionary1), in 1920 based [ 4–6 in], theNihe- area was visited wan village (someindependently 150 km NW but of almost Beijing; simultaneously Figure 1), in by1920 the [4–6], British the geomorphologist area was visited George B. Bar- independentlybour but almost [7] and simultaneously the French Jesuit by and the naturalist British geomorphologist Emile Licent in 1924 George [4]. B. Abundant Bar- vertebrate bour [7] and the(mainly French mammal) Jesuit and remains naturalist were Emile subsequently Licent in 1924 collected [4]. Abundant from the vertebrate Nihewan Basin in the (mainly mammal)course remains of field were trips subsequently led by Licent incollected 1925 and from Licent the andNihewan Teilhard Basin de Chardinin the in 1926 [6]. course of field tripsThe led firstby Licent mention in 1925 of an and ostrich Licent bone and fromTeilhard the Nihewande Chardin Basin in 1926 is in [6]. a section on the The first mentionantiquity of of an the ostrich ostrich inbone eastern from Asia the inNihewan the monograph Basin is by inBoule a section et al. on [8] the on the Paleolithic antiquity of thein ostrich China. in The eastern authors Asia note in th (p.e monograph 92) that Licent by Boule has et found al. [8] in on the the “Sanmenian” Paleo- [Lower lithic in China.Pleistocene] The authors beds note of(p. the 92) Sanggan that Licent He has (the found river whichin the “Sanmenian” flows through [Lower the Nihewan Basin) Pleistocene] bedsan ostrichof the Sanggan femur more He (the than river 340 which mm in flows length, through indicating the Nihewan a bird larger Basin) than the living an ostrich femurostrich. more Thisthan brief 340 mm mention in length, seems indicating to have attracted a bird larger little attention,than the living although os- it was noted trich. This briefby mention Lowe [9 seems] and Lambrechtto have attracted [10]. Later, little inattention, their study although of the it Nihewan was noted mammals, by Teilhard Lowe [9] and Lambrechtde Chardin [10]. and Later, Piveteau in thei [2]r briefly study mentionedof the Nihewan in a footnote mammals, (p. Teilhard 126) the fewde bird remains Chardin and Piveteauin their collection,[2] briefly mentioned viz. a humerus in a footnote of a large (p. vulture126) the andfew anbird ostrich remains femur. in These two their collection,bones viz. a are humerus kept together of a large at the vulture MNHN, and the an ostrich ostrich bone femur. bearing These number two bones NIH008. A second are kept togethernumber, at the 1927–13,MNHN, the refers ostrich to a bone catalog bearing entry number briefly listing NIH008. a collection A second ofnum- vertebrate fossils ber, 1927–13, refersfrom Nihewanto a catalog brought entry briefly back from listing his a secondcollection mission of vertebrate to China fossils by Teilhard from de Chardin Nihewan broughton 20 back November from his 1927. second The mission words to “Struthio China ”by and Teilhard “Femur” de Chardin are written on 20 in pencil on the November 1927.bone. The There words is “ thereforeStruthio” noand doubt “Femur” that theare femurwritten described in pencil below on the is bone. thatmentioned by There is thereforeTeilhard no doubt de Chardin that the and femur Piveteau described in 1930. below Whether is that it mentioned is the same boneby Teilhard as that mentioned by de Chardin andBoule Piveteau et al. in [8 ]1930. is not Whether so clear, it because is the same specimen bone as MNHN–NIH008, that mentioned inby its Boule present condition, et al. [8] is not isso 247clear, mm because in length, specimen while MNHN–NIH008, the length provided in its by pres Bouleent etcondition, al. [8] is is more 247 than 340 mm. mm in length, Thiswhile may the suggestlength provided that two by distinct Boule ostrich et al. [8] femora is more were than found 340 mm. in the This Nihewan may beds in the suggest that two1920s distinct and thatostrich one femora of them were may found have remainedin the Nihewan in China beds while in the the 1920s other and was sent to Paris. that one of themHowever, may have no remained ostrich femur in China is currently while the kept other at thewas Hoang sent to HoParis. Pai However, Ho Museum in Tianjin, no ostrich femurwhere is currently Licent’s collections kept at the are Hoang kept, andHo Pai there Ho is Museum no evidence in Tianjin, that such where a bone was part of Diversity 2021, 13, x FOR PEER REVIEW 3 of 13 Diversity 2021, 13, 47 3 of 12 Licent’s collections are kept, and there is no evidence that such a bone was part of the fossils that were transferred from the Hoang Ho Pai Ho Museum to Beijing in 1940 by theTeilhard fossils de that Chardin were transferred and Leroy from(see theLeroy Hoang [11] Hoabout Pai this Ho transfer) Museum and to Beijing are now in 1940kept byat Teilhard de Chardin and Leroy (see Leroy [11] about this transfer) and are now kept at the the IVPP in Beijing. We therefore suppose that MNHN–NIH008 is indeed the bone men- IVPP in Beijing.