RESEARCH ARTICLE Comparisons of Schansitherium tafeli with Samotherium boissieri (Giraffidae, Mammalia) from the Late Miocene of Gansu Province, China 1,2,3 4 5 4,6 Sukuan HouID *, Michael Cydylo , Melinda Danowitz , Nikos SolouniasID 1 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China, 2 CAS Center for Excellence in Life and Paleoenvironment, Beijing, China, 3 College of Earth and Planetary a1111111111 Sciences, University of Chinese Academy of Sciences, Beijing, China, 4 Department of Anatomy, New York a1111111111 Institute of Technology College of Osteopathic Medicine, Old Westbury, NY, United States of America, a1111111111 5 Department of Pediatrics, Alfred I. duPont Hospital for Children, Wilmington, DE, United States of America, a1111111111 6 Department of Paleontology, American Museum of Natural History, New York, NY, United States of a1111111111 America * [email protected] OPEN ACCESS Abstract Citation: Hou S, Cydylo M, Danowitz M, Solounias We are describing and figuring for the first time skulls of Schansitherium tafeli, which are N (2019) Comparisons of Schansitherium tafeli with Samotherium boissieri (Giraffidae, Mammalia) abundant in the Gansu area of China from the Late Miocene. They were animals about the from the Late Miocene of Gansu Province, China. size of Samotherium with shorter necks that had two pairs of ossicones that merge at the PLoS ONE 14(2): e0211797. https://doi.org/ base, which is unlike Samotherium. The anterior ossicones consist of anterior lineations, 10.1371/journal.pone.0211797 which may represent growth lines. They were likely mixed feeders similar to Samotherium. Editor: Suzannah Rutherford, Fred Hutchinson Schansitherium is tentatively placed in a very close position to Samotherium. Samotherium Cancer Research Center, UNITED STATES and Schansitherium represent a pair of morphologically very similar species that likely coex- Received: June 26, 2018 isted similarly to pairs of modern species, where the main difference is in the ossicones. Accepted: January 21, 2019 Pairs of ruminants in Africa, for example, exist today that differ mostly in their horn shape Published: February 12, 2019 but otherwise are similar in size, shape, and diet. The absence of Schansitherium from Europe is interesting, however, as Samotherium is found in both locations. While is it chal- Copyright: © 2019 Hou et al. This is an open access article distributed under the terms of the lenging to interpret neck length and ossicone shape in terms of function in combat, we offer Creative Commons Attribution License, which our hypothesis as to how the two species differed in their fighting techniques. permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper. Introduction Funding: This work was supported by the Strategic Priority Research Program of Chinese Academy of Giraffidae are Pecora ruminants [1]. There are approximately twenty-five species of Giraffidae, Sciences (XDA20070203, XDB26000000), the two extant and the rest are extinct [2±3]. The modern giraffe possesses exceptionally elongated National Natural Science Foundation of China cervical vertebrae and metapodials; the extinct taxa exhibit varying degrees of neck and limb (41872005, 41202002, 41430102), the Key Frontier Science Research Program of the Chinese elongation [4±6]. Giraffidae possess specialized cranial appendages termed ossicones, which Academy of Sciences (QYZDY-SSW-DQC002), and start as separate ossifications that subsequently fuse to the skull [2, 7±9]. Among the Giraffidae, the China Scholarship Council. several taxa exhibit atypical ossicone structure, including the sivatheres Sivatherium and PLOS ONE | https://doi.org/10.1371/journal.pone.0211797 February 12, 2019 1 / 29 Schansitherium versus Samotherium Competing interests: The authors have declared Bramatherium and the samothere Schansitherium. The first two have been reported and fig- that no competing interests exist. ured [10±12]. Unlike these, Schansitherium is poorly described and studied. Schansitherium tafeli is a giraffid from the late Miocene (Beodian age) of North China [2, 13±14]. Although it is a fairly abundant species in North China and is represented by complete cranial and postcranial material, it has not yet been figured nor studied in detail. It is a species similar to Samotherium, which is also found at the same deposits [14]. For example, some of the key similarities are: absence of sinuses in the calvaria, the position of the main ossicone (posterior ossicones of Schansitherium and the only pair of ossicones of Samotherium), the masseteric fossa size, the basicranium and area surrounding the foramen magnum, the occipi- tal shape, the dentition. Schansitherium tafeli was named on an isolated skull by Kilgus in 1922 from a collection of fossils from Shanxi, which is housed in the University Museum of TuÈbingen in Germany. Samotherium boissieri was named on a complete, crushed skull from Samos by Major in 1888 [3, 15±16]. The cranial and postcranial material of Samotherium bois- sieri is housed at the Natural History Museum NHMUK in London. Since 1922, several skulls and numerous postcranial specimens have been attributed to Schansitherium tafeli and Samotherium boissieri that have been found in North China. The most recent and statistically supported cladogram of the Giraffidae places Schansither- ium adjacent and primitive to Samotherium [17]. Of the various Samotherium species, we chose Samotherium boissieri to compare with Schansitherium tafeli, based on similarities between the skulls, dentitions and postcranial elements of the two taxa [18±19]. In addition, Samotherium boissieri is the most abundant Samotherium species in Gansu, where it coexisted with Schansitherium tafeli in the Miocene. We provide the first detailed description of Schan- sitherium tafeli, and we compare this taxon to Samotherium boissieri. The studied Schansitherium tafeli material was found in the Linxia Basin. The Linxia Basin is located in the transitional zone between the Tibetan and Loess plateaus, which is filled with 700±2000 m of Cenozoic deposits [20±22]. The Schansitherium tafeli specimens were exca- vated from the Upper Miocene Liushu Formation, which consists of mudstones and marls. Four representative Late Miocene faunas are recognized, from older to younger: the Guonigou, Dashengou, Yangjiashan, and the Qingbushan faunas. Schansitherium tafeli are abundant in the Dashengou Fauna and Yangjiashan Fauna, dated at about 8±10 Ma. Material and methods We studied Schansitherium and Samotherium material in Beijing IVPP and Hezheng HPM. We also studied in person the Samotherium material in AMNH, NHMUK, and other muse- ums. We describe the skull, third cervical vertebra, metacarpal and metatarsal of Schansither- ium tafeli from Gansu and of Samotherium boissieri from AMNH, NHMUK, and Gansu. The material from AMNH and NHMUK were previously studied by Kostopoulos [23], Hamilton [2], and Bohlin [14]. The material from Gansu are described for the first time in this paper. The Samotherium boissieri skull used for the comparisons is of great value as it is complete and not crushed. This skull enables us to study the details of the ossicone and its apex, the occipital and the basicranium. We compare the cranial and post-cranial material of these two taxa. We use cervical vertebral terminology and characteristics established by Danowitz and Solounias [24] and Danowitz et al. [25]. We use metapodial terminology established by Rios et al. [6]. Schansitherium tafeli: Skulls: AMNH 30502 (plaster cast holotype skull from TuÈbingen); HMV 1740, 0945, 1943, 1931, 1416, 1321, 1934, 1932, 1572 Vertebra: HMV 1988 Metacarpal: HMV 1951 PLOS ONE | https://doi.org/10.1371/journal.pone.0211797 February 12, 2019 2 / 29 Schansitherium versus Samotherium Metatarsal: HMV 1986 Samotherium boissieri: Skulls: IVPP V20167, V20271 Vertebra: C3 ±NHMUK 4250 Metacarpal and metatarsal: AMNH 15876 Institutional abbreviations AMNH, American Museum of Natural History, New York, USA. NHMUK, Natural His- tory Museum United Kingdom, London, UK. HPM Hezheng Paleozoological Museum, Gansu, China. IVPP, Institute of Vertebrate Paleotology and Palaeoanthropology, Chinese Academy of Sciences, Beijing, China. Description of Schansitherium tafeli (Tables 1 and 2 summarize the morphology and size of the specimens) Cranial Four ossicones are present. The two pairs of ossicones share a common base on each side, which is positioned above the orbit. The base of the anterior pair is merged with the posterior pair, making the common base long and oval in cross section (Figs 1±4). The posterior ossicone is large and long, and is positioned above the orbit. The ossicone gently curves posteriorly, most notably at the distal end. The ossicone base is straight, com- pressed medio-laterally, and set slightly posterior to the orbit. The base laterally merges with the superior orbital rim. Thus, the rim is no longer distinct (Figs 1±4). There is considerable variation of the surface of these ossicones. The surficial grooving of the ossicone is irregular and forms long streaks separated by fine grooves. Certain specimens have this morphology (Fig 2). There are also secondary bone growth streaks descending from the apex forming four distinct ridges circumferentially in one specimen.