The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla)

doi:10.1038/s42003-020-01205-8

. 2020 Sep 14;3(1):509.

doi: 10.1038/s42003-020-01205-8.

The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla)

Bin Bai^{1

2}, Jin Meng^{3

4

5}, Chi Zhang^{3

6}, Yan-Xin Gong^{3

6

7}, Yuan-Qing Wang^{8

9

10}

Affiliations

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, 100044, China. baibin@ivpp.ac.cn.
² CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China. baibin@ivpp.ac.cn.
³ Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China.
⁴ Division of Paleontology, American Museum of Natural History, New York, NY, 10024, USA.
⁵ Earth and Environmental Sciences, Graduate Center, City University of New York, New York, NY, 10016, USA.
⁶ CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China.
⁷ College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing, 100049, China.
⁸ Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China. wangyuanqing@ivpp.ac.cn.
⁹ CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China. wangyuanqing@ivpp.ac.cn.
¹⁰ College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing, 100049, China. wangyuanqing@ivpp.ac.cn.

PMID: 32929169
PMCID: PMC7490376
DOI: 10.1038/s42003-020-01205-8

The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla)

Bin Bai et al. Commun Biol. 2020.

. 2020 Sep 14;3(1):509.

doi: 10.1038/s42003-020-01205-8.

Authors

Bin Bai^{1

2}, Jin Meng^{3

4

5}, Chi Zhang^{3

6}, Yan-Xin Gong^{3

6

7}, Yuan-Qing Wang^{8

9

10}

Affiliations

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, 100044, China. baibin@ivpp.ac.cn.
² CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China. baibin@ivpp.ac.cn.
³ Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China.
⁴ Division of Paleontology, American Museum of Natural History, New York, NY, 10024, USA.
⁵ Earth and Environmental Sciences, Graduate Center, City University of New York, New York, NY, 10016, USA.
⁶ CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China.
⁷ College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing, 100049, China.
⁸ Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China. wangyuanqing@ivpp.ac.cn.
⁹ CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, China. wangyuanqing@ivpp.ac.cn.
¹⁰ College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing, 100049, China. wangyuanqing@ivpp.ac.cn.

PMID: 32929169
PMCID: PMC7490376
DOI: 10.1038/s42003-020-01205-8

Erratum in

Author Correction: The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla).
Bai B, Meng J, Zhang C, Gong YX, Wang YQ. Bai B, et al. Commun Biol. 2021 Jan 20;4(1):111. doi: 10.1038/s42003-021-01660-x. Commun Biol. 2021. PMID: 33473158 Free PMC article. No abstract available.

Abstract

Rhinoceroses have been considered to have originated from tapiroids in the middle Eocene; however, the transition remains controversial, and the first unequivocal rhinocerotoids appeared about 4 Ma later than the earliest tapiroids of the Early Eocene. Here we describe 5 genera and 6 new species of rhinoceroses recently discovered from the early Eocene to the early middle Eocene deposits of the Erlian Basin of Inner Mongolia, China. These new materials represent the earliest members of rhinocerotoids, forstercooperiids, and/or hyrachyids, and bridge the evolutionary gap between the early Eocene ceratomorphs and middle Eocene rhinocerotoids. The phylogenetic analyses using parsimony and Bayesian inference methods support their affinities with rhinocerotoids, and also illuminate the phylogenetic relationships and biogeography of Ceratomorpha, although some discrepancies are present between the two criteria. The nearly contemporary occurrence of various rhinocerotoids indicates that the divergence of different rhinocerotoid groups occurred no later than the late early Eocene, which is soon after the split between the rhinocerotoids and the tapiroids in the early early Eocene. However, the Bayesian tip-dating estimate suggests that the divergence of different ceratomorph groups occurred in the middle Paleocene.

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Conflict of interest statement

The authors declare no competing interests.

Figures

**Fig. 1. Fossil localities and distributions of early rhinocerotoids.**
a The location of the Erlian Basin of Inner Mongolia, China; b Paleogene fossil localities in the Erlian Basin. 1, Houldjin; 2, Arshanto; 3, Irdin Manha; 4, Daoteyin Obo; 5, Duheminboerhe; 6, Nuhetingboerhe; 7, Wulanboerhe; 8, Huheboerhe; 9, Chaganboerhe; 10, Bayan Ulan; 11, Nom Khong. The red dots refer to the localities where new materials were found. c Distributions of early controversial and unequivocal rhinocerotoids and new material from the early Eocene and early–middle Eocene in the Erlian Basin. The dark blue bars and stars show the distributions of new rhinocerotoids from the Erlian Basin. The light blue, gray, and green bars represent previously described early rhinocerotoids (or controversial rhinocerotoids) known from Asia and North America. Abbreviations: A. Amynodontidae, Br. Bridgerian, Forst. Forstercooperiidae, Rhino. Rhinocerotoidea.

**Fig. 2. Specimens of *Yimengia magna* sp. nov. from the upper part of the Nomogen Formation of the Erlian Basin.**
a, b Right mandible with dp4-m1 (a), m3 (b) (IVPP V 26234, holotype) in occlusal (a1, b1), buccal (a2, b2), and lingual (a3, c3) views; c right mandible with dp3–m1 (IVPP V 26235) in occlusal (c1), buccal (c2), and lingual (c3) views; d partial left P4 (IVPP V 26238.1) in occlusal (d1) and lingual (d2) views; e right M3 (IVPP V 26238.2) in occlusal (e1), buccal (e2), and lingual (e3) views; f right maxilla with M1–2 (IVPP V 26241) in occlusal (f1), buccal (f2), and lingual (f3) views.

**Fig. 3. Specimens of *Yimengia chaganense* sp. nov. from the lower and middle parts of the Arshanto Formation of the Erlian Basin.**
a Right (a1) and left (a2) maxillae with P4–M3 (IVPP V 26242.1, holotype) in occlusal view; b right maxilla with DP2-DP4 and M1 (IVPP V 26242.2) in occlusal (b1), buccal (b2), and lingual (b3) views; c left mandible with p3–4 (IVPP V 26243) in buccal (c1), lingual (c2), and occlusal (c3) views; d right m1/2 (IVPP V 26245.3) in occlusal (d1), buccal (d2), and lingual (d3) views; e right mandible with dp4 and m1 in the alveolus (IVPP V 26247.1) in occlusal view.

**Fig. 4. Scatter plots and box plot of dental proportions and length.**
*Yimengia*, *Rhodopagus*, and other early ceratomorphs (a, b), *Gobioceras*, *Pappaceras*, *Forstercooperia*, and *Juxia* (c, d). a Scatter plots of M1/2 proportion with the regression line for width as a function of length in *Yimengia*. b Scatter plots of m1 proportions with the regression line for width as a function of length in *Yimengia* and *Rhodopagus*. c Scatter plot of M3 proportion with the regression line for width as a function of length. d Box plot of m1–3 length. Box represents 25% and 75% quartiles, and the dotted line has a length of 1.5 times the interquartile range. n = 3, 1, 2, and 8 biologically independent samples, respectively.

**Fig. 5. Specimens of *Triplopus*? *youjingensis* sp. nov. and *Gobioceras wangi* gen. et sp. nov. from the base of the Arshanto Formation in the Erlian Basin.**
a T.? *youjingensis*, right mandible with p2–m3 (IVPP V 26248, holotype) in occlusal (a1), buccal (a2), and lingual (a3) views; b–d *G. wangi*, b right mandible with m1–3 (IVPP V 26249, holotype) in occlusal (b1), buccal (b2), and lingual (b3) views; c right M3 (IVPP V 26250.1) in occlusal (c1), buccal (c2), and lingual (c3) views; d fragmentary M2 ectoloph (IVPP V 26250.2) in occlusal (d1) and buccal (d2) views.

**Fig. 6. Specimens of *Ephyrachyus woodi* sp. nov. and *Hyrachyus*? *tumidus* from the Arshanto Formation of the Erlian Basin.**
a–f *E. woodi* (IVPP V 26252, holotype), a right maxilla with P2–M3 in occlusal (a1), buccal (a2), and lingual (a3) views; b fragmentary symphyseal region with the roots of incisors and canine; c right mandible with p2–3 in occlusal (c1), lingual (c2), and buccal (c3) views; d left mandible with p2–3 in occlusal view; e p4 fragment in occlusal (e1) and buccal (e2) views; f m1/2 fragment in occlusal (f1) and buccal (f2) views. g–i H.? *tumidus*; g right maxilla with P3–M2 (IVPP V 26253.1, holotype) in occlusal (g1) and buccal (g2) views; h m1 fragment (IVPP V 26253.2) in occlusal view; i m2 fragment (IVPP V 26253.3) in occlusal (i1), buccal (i2), and lingual (i3) views.

**Fig. 7. The strict consensus tree of two most parsimonious trees, showing the phylogeny of Ceratomorpha with paraphyletic ‘Isectolophidae’ as a sister group.**
All new taxa reported here are marked in red, and placed in Rhinocerotoidea. The geographic distribution was reconstructed using the parsimony criterion in Mesquite. The taxa marked in gray with asterisks in different clades are reconstructed on the right side with simplified phylogenetic relationships (scale bar equals 10 cm). The numbers and letters at the nodes show Bremer Support >1, and the clades discussed in the text with synapomorphies are listed in Supplementary Table 1.

**Fig. 8. Majority-rule (50%) consensus trees of Ceratomorpha using Bayesian phylogenetic tip-dating analyses.**
The node ages (divergence times) are the median estimates and node bars represent the 95% highest posterior density (HPD) intervals of the divergence times. The numbers at the internal nodes are the posterior probabilities of the corresponding clades. Eo. Eocene, Mio. Miocene, Oli. Oligocene, Pa. Paleocene.

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References

1. Holbrook LT. The phylogeny and classification of Tapiromorph Perissodactyls (Mammalia) Cladistics. 1999;15:331–350. - PubMed
1. Steiner CC, Ryder OA. Molecular phylogeny and evolution of the Perissodactyla. Zool. J. Linn. Soc. 2011;163:1289–1303.
1. Kosintsev P, et al. Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions. Nat. Ecol. Evol. 2019;3:31–38. - PubMed
1. Hooker, J. J. Character polarities in early Eocene perissodactyls and their significance for Hyracotherium and infraordinal relationships. In: The Evolution of Perissodactyls (eds Prothero, D. R. & Schoch R. M.). 79–101 (Oxford University Press, 1989).
1. Colbert MW. The facial skeleton of the early Oligocene Colodon (Perissodactyla, Tapiroidea) Palaeontol. Electron. 2005;8:1–27.

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[1] Holbrook LT. The phylogeny and classification of Tapiromorph Perissodactyls (Mammalia) Cladistics. 1999;15:331–350. - PubMed

[2] Holbrook LT. The phylogeny and classification of Tapiromorph Perissodactyls (Mammalia) Cladistics. 1999;15:331–350. - PubMed

[3] Steiner CC, Ryder OA. Molecular phylogeny and evolution of the Perissodactyla. Zool. J. Linn. Soc. 2011;163:1289–1303.

[4] Steiner CC, Ryder OA. Molecular phylogeny and evolution of the Perissodactyla. Zool. J. Linn. Soc. 2011;163:1289–1303.

[5] Kosintsev P, et al. Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions. Nat. Ecol. Evol. 2019;3:31–38. - PubMed

[6] Kosintsev P, et al. Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions. Nat. Ecol. Evol. 2019;3:31–38. - PubMed

[7] Hooker, J. J. Character polarities in early Eocene perissodactyls and their significance for Hyracotherium and infraordinal relationships. In: The Evolution of Perissodactyls (eds Prothero, D. R. & Schoch R. M.). 79–101 (Oxford University Press, 1989).

[8] Hooker, J. J. Character polarities in early Eocene perissodactyls and their significance for Hyracotherium and infraordinal relationships. In: The Evolution of Perissodactyls (eds Prothero, D. R. & Schoch R. M.). 79–101 (Oxford University Press, 1989).

[9] Colbert MW. The facial skeleton of the early Oligocene Colodon (Perissodactyla, Tapiroidea) Palaeontol. Electron. 2005;8:1–27.

[10] Colbert MW. The facial skeleton of the early Oligocene Colodon (Perissodactyla, Tapiroidea) Palaeontol. Electron. 2005;8:1–27.

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The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla)

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The origin of Rhinocerotoidea and phylogeny of Ceratomorpha (Mammalia, Perissodactyla)

Authors

Affiliations

Erratum in

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Conflict of interest statement

Figures

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