Published March 6, 2025 | Version v1
Journal article Open

A taxonomic review of the genus Rhinoceros with emphasis on the distinction of Eurhinoceros (Perissodactyla, Rhinocerotidae)

  • 1. IUCN SSC Asian Rhino Specialist Group and Conservation Planning Specialist Group, Gland, Switzerland
  • 2. Bayerische Staatssammlung für Paläontologie und Geologie, München, Germany

Description

This study examines the ecomorphological characteristics of two Asian rhinoceros species: the critically endangered Sundaic rhinoceros and the vulnerable Indian rhinoceros. Among the five living rhinoceros taxa, the three Asian species are notable for their tusked incisors. Fossil evidence highlights the divergence between Rhinoceros and Eurhinoceros in cheek tooth morphology, linked to different dietary specialisations. The Sundaic rhinoceros, a generalist browser restricted to the Ujung Kulon peninsula of Java, exhibits distinctive features such as a grey hide with polygonal patterns, a typical 'saddle' on the nape, a slender head shape and a protrusion instead of a horn in females. The latter is a unique trait among Rhinocerotini species. In contrast, the Indian rhinoceros, a variable grazer, inhabits riverine grasslands in northern India and southern Nepal, displaying deep skin folds and tubercles. Ecological behaviours differ significantly, with the Sundaic rhinoceros being solitary wanderers and Indian rhinoceros forming temporary crashes. Both species possess unique adaptations for survival, emphasising the importance of understanding their systematics for effective conservation. The study further examines the interrelationships among the one-horned Asian species of the Rhinocerotidae family, highlighting their distinct features. The revision delves into skull morphology, dentition, and ecological dynamics, revealing evolutionary patterns and ancestral traits. Both single horned rhinoceroses went a separate and diverging way of evolution that was not triggered by geographical separation but by niche partitioning. Comparative analyses shed light on the evolutionary trajectory and ecological adaptations of each species. The fossils, the ecological and morphological adaptations of both species, suggest designating 'Rhinoceros' sondaicus as distinct from Rhinoceros unicornis, under the one-horned rhinoceros Eurhinoceros, as proposed by Gray (1868). Eurhinoceros sondaicus emerges as a persistently more primitive form.

Files

ZK_article_127858.pdf

Files (5.3 MB)

Name Size Download all
md5:71de30634e374a11fd75d7651be734d0
5.3 MB Preview Download

System files (241.2 kB)

Name Size Download all
md5:85896a2440ebf8a5c0cc48a0bbc88a25
241.2 kB Download

Linked records

Additional details

References

  • Abdul AM, Cerdeño E, Akhtar M, Khan MA, Siddiq Kh (2014) An account of the Upper Siwalik rhinocerotids of Pakistan. Abstract volume of the 4th International Palaeontological Congress, Mendoza, Argentina, 549 pp.
  • Amman H (1986) Contributions to the ecology and sociology of the Javan rhinoceros (Rhinoceros sondaicus, Desmarest). Basel University, Inaugural Dissertation.
  • Antoine P-O (2012) Pleistocene and Holocene rhinocerotids (Mammalia, Perissodactyla) from the Indochinese Peninsula. Comptes Rendus Palevol, Elsevier 11: 159–168. https://doi.org/10.1016/j.crpv.2011.03.002
  • Bacon AM, Demeter F, Schuster M, Vu TL, Nguyen KR, Antoine P-O, Sevket S, Ha HN, Nguyen MH (2004) The Pleistocene Ma U'Oi cave, northern Vietnam: palaeontology, sedimentology and palaeoenvironments. Geobios 37: 305–314. https://doi.org/10.1016/j.geobios.2003.03.010
  • Bales GS (1996) Skull evolution in the Rhinocerotidae (Mammalia, Perissodactyla): Cartesian transformations and functional interpretations. Journal of Mammalian Evolution 3(3): 261–279. https://doi.org/10.1007/BF01458183
  • Barry JC, Morgan ME, Flynn LJ, Pilbeam D, Behrensmeyer AK, Raza SM, Khan AI, Badgley C, Hicks J, Kelley J. (2002) Faunal and environmental change in the late Miocene Siwaliks of northern Pakistan. Paleobiology 28 (S2): 1–71. https://doi.org/10.1666/0094-8373(2002)28[1:FAECIT]2.0.CO;2
  • Beden M, Guerin C (1973) Le gisement des vertébrés du Phnom Loang (Province de Kampot, Cambodge). Travaux et Documents de l'ORSTOM 27: 3–97.
  • Benoit J, Legendre LJ, Farke AA, Neenan JM, Mennecart B, Costeur L, Merigraud S, Manger PR (2020) A test of the lateral semicircular canal correlation to head posture, diet and other biological traits in "ungulate" mammals. Scientific Reports 10: 1–21. https://doi.org/10.1038/s41598-020-76757-0
  • Bhattacharya A (2020a) Ecology, behaviour and management practices of Great Indian one Horned Rhinoceros at Gorumara, Jaldapara and Kaziranga national parks, India. Thesis presented to Raiganj University.
  • Bhattacharya A (2020b) Communication types in Indian Rhinoceros (R. unicornis, Linnaeus). International Journal of Trend in Research and Development 7(4): 188–194.
  • Bhattacharya A, Chakraborty K (2016) Defecation behaviour of great Indian one horned rhinoceros (R. unicornis, Linnaeus). International Journal of Science and Research 5(7): 923–978.
  • Blumenbach JF (1799) Handbuch der Naturgeschichte. Sechste Ausgabe (6th ed). Johann Christian Dieterich, Goettingen, 703 pp.
  • Burchell WJ (1817) Ueber eine neue Gattung Nashorn. Isis (Oken) 1: 1318–1319. [figs 1, 2]
  • Cave AJE (1969) Hairs and vibrissae in the Rhinocerotidae. Journal of Zoology, London 157(2): 247–257. https://doi.org/10.1111/j.1469-7998.1969.tb01700.x
  • Cave AJE (1985) An unrecorded specimen of the Javan rhinoceros (Rhinoceros sondaicus). Journal of Zoology, London 207: 527–535. https://doi.org/10.1111/j.1469-7998.1985.tb04949.x
  • Cerling T, Harris J, MacFadden B, Leakey MG, Quade J, Eisenmann V, Ehleringer J.R (1977) Global vegetation change through the Miocene/Pliocene boundary. Nature 389: 153–158. https://doi.org/10.1038/38229
  • Choudhury A (1985) Distribution of Indian one-horned rhinoceros (Rhinoceros unicornis). Tiger Paper 12(2): 25–30.
  • Colbert EH (1934) A new rhinoceros from the Siwalik beds of India. American Museum Novitates 749: 1–13. https://doi.org/10.70249/9798893981964-002
  • Colbert EH (1942) Notes on the Lesser one-horned rhinoceros, Rhinoceros sondaicus, 2. The position of Rhinoceros sondaicus in the phylogeny of the genus Rhinoceros. American Museum Novitates 1207: 1–5.
  • Cranbrook Earl of (1986) A review of fossil and prehistoric remains of rhinoceroses of Borneo. Sabah Museum and Archives Journal 1(1): 50–110.
  • Cranbrook Earl, Piper PJ (2007) The Javan rhinoceros Rhinoceros sondaicus in Borneo. Raffles Bulletin of Zoology 55(1): 217–220.
  • Desmarest AG (1822) Mammalogie, ou description des espèces des Mammifères. Veuve Agasse, Paris, 277–555.
  • Dinerstein E (2003) The return of the unicorns: natural history and conservation of the Greater one-horned rhinoceros. Columbia University Press, New York, 316 pp. https://doi.org/10.7312/dine08450
  • Dinerstein E (2011) Family Rhinocerotidae (Rhinoceroses). In: Handbook of the mammals of the world, vol. 2: Hoofed mammals. Lynx, New York, 144–181.
  • Dubois E (1908) Das geologische Alter der Kendeng-oder Trinilfauna. Tijdschrift van het Koninklijk Nederlandsch Aardrijkskundig Genootschap (2)25: 1235–1270.
  • Endo H, Kobayashi H, Koyabu D, Hayashida A, Jogahara T, Taru H, Oishi M, Itou T, Koie H, Sakai T (2009) The morphological basis of the armor-like folded skin of the Greater Indian Rhinoceros as a thermoregulator. Mammal Study 34(4): 195–200. https://doi.org/10.3106/041.034.0403
  • Erxleben JCP (1777) Systema regni animalis per classes, ordines, genera, species, varietates, cum synonymia et historia animalium. Weygandianis, 636 pp. https://doi.org/10.5962/bhl.title.15933
  • Falconer H, Cautley PT (1847) Fauna antiqua sivalensis, being the fossil zoology of the Sewalik Hills, in the North of India Illustrations, part VIII: Suidae and Rhinocerotidae. Smith, Elder and Company, London, pls 69–80. https://doi.org/10.5962/bhl.title.61447
  • Falconer H (1859) [In Gaudin CT (Ed.)] Modifications apportés par Mr. Falconer a la faune du Val d'Arno. Bulletin de la Société Vaudoise des Sciences Naturelles 6(44): 130–131.
  • Falconer H, Cautley PT (1836) Note on the fossil hippopotamus of the Siwalik Hills. Asiatic Researches 19(3): 39–53.
  • Flower WH (1876) On some cranial and dental characters of the existing species of rhinoceroses. Proceedings of the Zoological Society of London 1876 May 16: 443–457. https://doi.org/10.1111/j.1096-3642.1876.tb02583.x
  • Forster-Cooper C (1934) The extinct Rhinoceroses of Baluchistan. Philosophical Transactions of the Royal Society of London (B) 223: 569–616. https://doi.org/10.1098/rstb.1934.0013
  • Ghosh T, Kumar S, Sharma K, Kakati P, Sharma A, Mondol S (2022) Consideration of genetic variation and evolutionary history in future conservation of Indian one-horned rhinoceros (Rhinoceros unicornis). BMC Ecology and Evolution 22(92): 1–13. https://doi.org/10.1186/s12862-022-02045-2
  • Ginsburg L (1974) Les Rhinocérotidés du Miocène de Sansan (Gers). Compte Rendu des Séances de l'Académie des Sciences, Paris 278 (5) D: 597–600.
  • Gippoliti S, Cotterill FPD, Groves CP (2013) Mammal taxonomy without taxonomists: a reply to Zachos and Lovari. Hystrix Italian Journal Mammalogy 24: 145–147.
  • Gray JE (1868) Observations on the preserved specimens and skeletons of the Rhinocerotidae in the collection of the British Museum and Royal College of Surgeons, including the description of three new species. Proceedings of the Zoological Society of London: 1003–1032.
  • Griffiths M (1993) The Javan rhino of Ujung Kulon: an investigation of its population and ecology through camera trapping. Directorate General of Forest Protection and Nature Conservation and the World Wide Fund for Nature Indonesia Program, Jakarta, 1–92.
  • Groves CP (1967) On the rhinoceroses of South-East Asia. Säugetierkundliche Mitteilungen 15(3): 221–237.
  • Groves CP (1971) Species characters in rhinoceros horns. Zeitschrift fur Saugetierkunde 36(4): 238–252. [figs 1–22]
  • Groves CP, Guérin C (1980) Le Rhinoceros sondaicus annamiticus d'Indochine: distinction taxinomique et anatomique; relations phylogénétiques. Geobios 13(2): 199–208. https://doi.org/10.1016/S0016-6995(80)80028-3
  • Groves CP, Leslie Jr DM (2011) Rhinoceros sondaicus (Perissodactyla: Rhinocerotidae). Mammalian Species 43(887): 190–208. https://doi.org/10.1644/887.1
  • Guérin C (1973) Rhinocerotidae: 19–50, pls. 1–4. In Beden M and Guérin C (1973) Le gisement des vertébrés du Phnom Loang (Province de Kampot, Cambodge). Travaux et Documents de l'ORSTOM 27: 3–97.
  • Guérin C (1980) Les rhinocéros (Mammalia, Perissodactyla) du Miocène terminal au Pléistocène supérieur en Europe occidentale; comparaison avec les espèces actuelles. Documents du Laboratoire de Géologie de la Faculté des Sciences de Lyon 79: 3–1183.
  • Hariyadi ARS, Sajuthi D, Astuti DA, Alikodra HS, Maheshwari H (2016) Analysis of nutrition quality and food digestibility in male Javan rhinoceros (Rhinoceros sondaicus) in Ujung Kulon National Park. Pachyderm 57: 86–96. https://doi.org/10.69649/pachyderm.v57i.408
  • Harper F (1945) Extinct and vanishing mammals of the old world. New York, American Committee for International Wild Life protection, Special Publication no. 12: 1–850. https://doi.org/10.5962/bhl.title.19520
  • Haryono M, Rahmat UM, Daryan M, Raharja AS, Muhtarom A, Firdaus AY, Rohaeti A, Subchiyatin I, Nugraheni A, Khairani KO, Kartina (2015) Monitoring of Javan Rhino population in Ujung Kulon National Park. Pachyderm 56: 82–86. https://doi.org/10.69649/pachyderm.v56i.374
  • Hazarika BC, Saikia PK (2010) A study on the behaviour of great Indian one-horned rhinoceros (Rhinoceros unicornis) in the Rajiv Gandhi National Park, Assam, India. NeBio 1(2): 62–74. https://doi.org/10.5402/2012/259695
  • Heißig K (1969) Die Rhinocerotidae (Mammalia) aus der oberoligozänen Spaltenfüllung von Gaimersheim bei Ingolstadt in Bayern und ihre phylogenetische Stellung. Verlag der Bayerischen Akademie der Wissenschaften; in Kommission bei der Beck'schen CH Verlagsbuchhandlung, München: 1–133.
  • Heißig K (1972) Paläontologische und geologische Untersuchungen im Tertiär von Pakistan, 5. Rhinocerotidae (Mammalia.) aus den unteren und mittleren Siwalik-Schichten. Bayerische Akademie der Wissenschaften Mathematisch−Naturwissenschaftliche Klasse, Abhandlungen, München. Neue Folge 152: 1–112.
  • Heißig K (1973) Die Unterfamilien und Tribus der rezenten und fossilen Rhinocerotidae (Mammalia). Säugetierkundliche Mitteilungen 21: 25–30.
  • Heißig K (1981) Probleme bei der cladistischen Analyse einer Gruppe mit wenigen eindeutigen Apomorphien: Rhinocerotidae. Palaeontologisches Zeitschrift 55(1): 117–123. https://doi.org/10.1007/BF02986041
  • Heißig K (1989) 21 The Rhinocerotidae. In: Prothero DR, Schoch RM (Eds) The Evolution of Perissodactyls. Oxford University Press, New York, 399–417.
  • Hernesniemi E, Blomstedt K, Fortelius M (2011) Multi-view stereo three-dimensional reconstruction of lower molars of Recent and Pleistocene rhinoceroses for mesowear analysis. Palaeontologia Electronica: 1–15.
  • Heude PM (1892) Études odontologiques. 1. Herbivores trizygodontes et dizygodontes. Mémoires concernant l'histoire naturelle de l'Empire Chinois 2: 65–84.
  • Hitchins PM (1986) Earlessness in the black rhinoceros–a warning. Pachyderm 7: 8–10. https://doi.org/10.69649/pachyderm.v7i1.634
  • Hoogerwerf A (1970) Udjung Kulon, the land of the last Javan rhinoceros. With local and general data on the most important faunal species and their preservation in Indonesia. Brill RJ, Leiden, 512 pp. https://doi.org/10.1163/9789004646698
  • Hooijer DA (1946) The evolution of the skeleton of Rhinoceros sondaicus Desmarest. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Amsterdam 49(6): 671–676.
  • Hooijer DA (1964) New records of mammals from the middle pleistocene of Sangiran, central Java. Zoologische Mededelingen 40(10): 73–88.
  • Hullot M, Antoine P-O, Ballatore M, Merceron G (2019) Dental microwear textures and dietary preferences of extant rhinoceroses (Perissodactyla, Mammalia). Mammal Research 64: 397–409. https://doi.org/10.1007/s13364-019-00427-4
  • Jäger GF (1839) Über die fossilen Säugethiere welche in Württemberg in verschiedenen Formationen aufgefunden worden sind, nebst geognostischen Bemerkungen über diese Formationen. Verlag Carl Erhard, Stuttgart. Abtheilung 2: 1–214.
  • Khan AM (2009) Taxonomy and distribution of rhinoceroses from the Siwalik Hills of Pakistan. PhD Thesis, University of the Punjab, Lahore.
  • Kim HL, Li T, Kalsi N, Nguyen HTT, Shaw TA, Ang KC, Cheng KC, Ratan A, Peltier WR, Samanta D, Pratapneni M, Schuster SC, Horton BP (2023) Prehistoric human migration between Sundaland and South Asia was driven by sea-level rise. Commununications Biology 6: 1–10. https://doi.org/10.1038/s42003-023-04510-0
  • Koenigswald GHR van (1935) Die fossilen Säugetierfaunen Javas. Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen 38: 88–98.
  • Laurie WA (1978) The ecology and behaviour of the greater one-horned rhinoceros. Ph.D. dissertation. University of Cambridge, Cambridge.
  • Laurie WA (1982) Behavioural ecology of the Greater one-horned rhinoceros (Rhinoceros unicornis). Journal of Zoology, London 196(3): 307–341. https://doi.org/10.1111/j.1469-7998.1982.tb03506.x
  • Laurie WA, Lang RM, Groves CP (1983) Rhinoceros unicornis. Mammalian Species 211, The American Society of Mammalogists: 1–6. https://doi.org/10.2307/3504002
  • Lesson RP (1836) Complement des œuvres de Buffon ou histoire naturelle des animaux rares découverts par les naturalistes et les voyageurs depuis la mort de Buffon, vol. 10: Oiseaux et Mammifères. Histoire naturelle générale et particulière des mammifères et oiseaux découverts depuis la mort de Buffon: Oiseaux et mammifères. Pourrat Frères et Roret, Paris, 414 pp.
  • Linnaeus C (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis Editio decima, reformata [edn 10]. Holmiae, Laurentii Salvii: i–iv, 1–824. https://doi.org/10.5962/bhl.title.542
  • Liu S, Dalen L, Gilbert T, Rookmaaker LC and 35 others (2021) Ancient and modern genomes unravel the evolutionary history of the rhinoceros family. Cell 184: 4874–4885. https://doi.org/10.1016/j.cell.2021.07.032
  • Loch CW (1937) Rhinoceros sondaicus: the Javan or Lesser one-horned rhinoceros and its geographical distribution. Journal of the Malayan Branch of the Royal Asiatic Society 15(2): 130–149.
  • Longuet M, Zin-Maung-Maung-Thein , Thaung-Htike , Man-Thit-Nyein , Masanaru (2023) New fossil remains of Rhinocerotidae (Perissodactyla) from the early Late Miocene Tebingan area, central Myanmar, Historical Biology: 1–14. https://doi.org/10.1080/08912963.2024.2408617
  • Longuet M, Handa N, Maung-Thein ZM, Htike T, Nyein MT, Takai M (2024) Post-cranial remains of Rhinocerotidae from the Neogene of central Myanmar: morphological descriptions and comparisons with ratios. Historical Biology: 1–15. https://doi.org/10.1080/08912963.2024.2408617
  • Lydekker R (1907) The game animals of India, Burma and Tibet. Being a new and revised edition of "The great and small game of India, Burma and Tibet". Rowland Ward, London: 1–409. https://doi.org/10.5962/bhl.title.16137
  • MacFadden BJ (1998) Tale of two rhinos: isotopic ecology, paleodiet, and niche differentiation of Aphelops and Teleoceras from the Florida Neogene. Paleobiology 24(2): 274–286. https://doi.org/10.1666/0094-8373(1998)024[0274:TOTRIE]2.3.CO;2
  • Margaryan A, Sinding MS, Liu S, Vieira FG, Chan YL, Nathan S, Moodley Y, Bruford MW, Gilbert MTP (2020) Recent mitochondrial lineage extinction in the critically endangered Javan rhinoceros. Zoological Journal of the Linnean Society 190(1): 372–383. https://doi.org/10.1093/zoolinnean/zlaa004
  • Matschiner M (2021) Museum specimens tell the history of rhinoceroses. Cell 184(19): 4841–4842. https://doi.org/10.1016/j.cell.2021.08.026
  • Matthew WD (1929) Critical observations upon Siwalik Mammals. Bulletin of the American Museum of Natural History, New York 56(7): 437–560.
  • Matthew WD (1931) Critical observations on the phylogeny of the rhinoceroses. University of California Publications, in Geological Science 20: 1–8.
  • McNeely JA, Cronin EW (1972) Rhinos in Thailand. Oryx 11(6): 457–460. https://doi.org/10.1017/S0030605300010735
  • Medway L (1969) The wild mammals of Malaya and offshore islands, including Singapore. Kuala Lumpur, Oxford University Press: 1–127.
  • Morley RJ (2018) Assembly and division of the South and South-East Asian flora in relation to tectonics and climate change. Journal of Tropical Ecology 34(4): 209–234. https://doi.org/10.1017/S0266467418000202
  • Nardelli F (1988) The Rhinoceros: a Monograph. Basilisk Press, London, 133 pp.
  • Nardelli F (2013) The mega-folivorous mammals of the rainforest: feeding ecology in nature and in controlled environment: A contribution to their conservation. International Zoo News 60(5): 323–339.
  • Nardelli F (2016) Current status and conservation prospects for the Javan rhinoceros Rhinoceros sondaicus Desmarest 1822. International Zoo News 63(3): 180–202.
  • Nardelli F, Robovský J (2023) New data on the ecology and conservation of the Javan rhinoceros Rhinoceros sondaicus Desmarest, 1822 (Perissodactyla, Rhinocerotidae). Gazella (Praha) 49(2022): 183–205.
  • Neese HC (1975) Survival of the Javan rhinoceros in Laos. Report: 1–27.
  • Owen R (1870) On fossil remains of mammals found in China. Quarterly Journal of the Geological Society London 26: 417–434. https://doi.org/10.1144/GSL.JGS.1870.026.01-02.40
  • Owen-Smith RN (1988) Megaherbivores: The influence of very large body size on ecology. Cambridge University Press, Cambridge, 369 pp. https://doi.org/10.1017/CBO9780511565441
  • Pandolfi L (2015) Sistematica e filogenesi dei Rhinocerotini (Mammalia, Rhinocerotidae). Dissertation presented to the University of Rome, Italy.
  • Pandolfi L, Maiorino L (2016) Reassessment of the largest Pleistocene rhinocerotine Rhinoceros platyrhinus (Mammalia, Rhinocerotidae) from the Upper Siwaliks (Siwalik Hills, India). Journal of Vertebrate Paleontology 36(2): 1–12. https://doi.org/10.1080/02724634.2015.1071266
  • Pandolfi L, Bartolini-Lucenti S, Cirilli O, Bukhsianidze M, Lordkipanidze D, Rook L (2021) Paleoecology, biochronology, and paleobiogeography of Eurasian Rhinocerotidae during the Early Pleistocene: The contribution of the fossil material from Dmanisi (Georgia, Southern Caucasus). Journal of Human Evolution 156: 1–13. https://doi.org/10.1016/j.jhevol.2021.103013
  • Peacock EH (1933) A game book for Burma and adjoining territories. H&F, Witherby G, London, 292 pp.
  • Piras P, Maiorino L, Raia P, Marcolini F, Salvi D, Vignoli L, Kotsakis T (2010) Functional and phylogenetic constraints in Rhinocerotinae craniodental morphology. Evolutionary Ecology Research 12: 897–928.
  • Poole CM, Duckworth JW (2005) A documented 20th century record of Javan Rhinoceros Rhinoceros sondaicus from Cambodia. Mammalia 69(3–4): 443–444. https://doi.org/10.1515/mamm.2005.039
  • Reynolds EAP (1954) Burma rhino. Burmese Forester 4(2): 104–108.
  • Rookmaaker LC (1977) The rhinoceros of Borneo: a 19th century puzzle. Journal of the Malayan Branch of the Royal Asiatic Society 50(1): 52–62.
  • Rookmaaker LC (1980) The distribution of the rhinoceros in Eastern India, Bangladesh, China and the Indo-Chinese region. Zoologische Anzeiger 205(3/4): 253–268.
  • Rookmaaker LC (1997) Records of the Sundarbans rhinoceros (Rhinoceros sondaicus inermis) in India and Bangladesh. Pachyderm 24: 37–45. https://doi.org/10.69649/pachyderm.v24i1.903
  • Rookmaaker LC (2000) Records of the rhinoceros in Pakistan and Afghanistan. Pakistan Journal of Zoology 32(1): 65–74.
  • Rookmaaker LC (2002) Historical records of the Javan rhinoceros in North-East India. Newsletter of the Rhino Foundation of Nature in North-East India 4: 11–12.
  • Rookmaaker LC (2006) Distribution and extinction of the rhinoceros in China: review of recent Chinese publications. Pachyderm 102: 102–106. https://doi.org/10.69649/pachyderm.v40i1.1267
  • Rookmaaker LC (2016) On the alleged presence of the two-horned Sumatran rhinoceros and one-horned Javan rhinoceros in the Himalayan Kingdom of Bhutan. Pachyderm 57: 116–117. https://doi.org/10.69649/pachyderm.v57i.399
  • Rookmaaker LC, Carpentier H (2007) Early references to the rhinoceros on the Chinese island of Hainan. Journal of the Royal Asiatic Society Hong Kong Branch 45(2005): 235–236.
  • Schaller GB, Dang NX, Thuy LD, Son VT (1990) Javan rhinoceros in Vietnam. Oryx 24(2): 77–80. https://doi.org/10.1017/S0030605300034712
  • Schellhorn R (2018) A potential link between lateral semicircular canal orientation, head posture, and dietary habits in extant rhinos (Perissodactyla, Rhinocerotidae). Journal of Morphology 279: 50–61. https://doi.org/10.1002/jmor.20753
  • Setiawan R, Gerber B, Ujang MR, Daryang D, Firdaus AY, Haryono M, Khairani KO, Kurniawan Y, Long B, Lyet A, Muhiban M, Mahmud R, Muhtarom A, Purastuti E, Ramono W, Subrata D, Sunarto S (2017) Preventing global extinction of the Javan Rhino: Tsunami risk and future conservation direction. Conservation Letters 11(1): 1–9. https://doi.org/10.1111/conl.12366
  • Siddiq MKh, Akhtar M, Khan MA, Ghaffar A, Sarwar Kh, Khan AM (2016) New fossils of rhinoceros (Rhinocerotidae) from the Soan Formation (Plio-Pleistocene) of Northern Pakistan. Pakistan Journal of Zoology 48(3): 657–664.
  • Singh S, Parkash B, Awasthi AK, Singh T (2012) Palaeoprecipitation record using O-isotope studies of the Himalayan Foreland Basin sediments, NW India. Palaeogeography, Palaeoclimatology, Palaeoecology 331–332: 39–49. https://doi.org/10.1016/j.palaeo.2012.02.031
  • Sody HJV (1959) Das Javanische Nashorn, Rhinoceros sondaicus, historisch und biologisch. Zeitschrift fur Saugetierkunde 24(3/4): 109–240.
  • Strien NJ van (1985) The Sumatran rhinoceros in the Gunung Leuser National Park, its distribution, ecology and conservation. Doorn, Van Strien: 1–207.
  • Suraprasit K, Jaeger JJ, Chaimanee Y, Chavasseau O, Yamee C, Tian P, Panha S (2016) The Middle Pleistocene vertebrate fauna from Khok Sung (Nakhon Ratchasima, Thailand): biochronological and paleobiogeographical implications. ZooKeys 613: 1–157. https://doi.org/10.3897/zookeys.613.8309
  • Tougard C, Delefosse T, Hoenni C, Montgelard C (2001) Phylogenetic relationships of the five extant rhinoceros species (Rhinocerotidae, Perissodactyla) based on mitochondrial cyctochrome b and 12s rRNA genes. Molecular Phylogenetics and Evolution 19(1): 34–44. https://doi.org/10.1006/mpev.2000.0903
  • Toula F (1902) Das Nashorn von Hundsheim Rhinoceros (Ceratorhinus Osborn) hundsheimensis nov. form. mit Ausführungen über die Verhältnisse von elf Schädeln von Rhinoceros (Ceratorhinus) sumatrensis. Abhandlungen der Kaiserlich-Königlichen Geologischen Reichsanstalt 19: 1–92.
  • Willerslev E, Gilbert M, Binladen J, Ho S, Campos P, Ratan A, Tomsho L, Fonseca R da, Sher A, Kuznetsova T, Nowak-Kemp M, Roth T, Miller W, Schuster S (2009) Analysis of complete mitochondrial genomes from extinct and extant rhinoceroses reveals lack of phylogenetic resolution. BMC Evolutionary Biology 9: 1–30. https://doi.org/10.1186/1471-2148-9-95
  • Wilson SG (2021) Factors shaping the conservation of the critically endangered Javan Rhinoceros–Rhinoceros sondaicus. Ph.D. thesis, School of Biological Sciences, University of Queensland, Brisbane.
  • Yan Y, Wang Y, Zhu M, Zhang Y, Qin D, Jin C (2023) New rhino remains from Middle to Late Pleistocene of Chongzuo, Guangxi with discussion on Quaternary Rhinoceros evolution in Southern China. Quaternary Sciences 43(3): 777–792.
  • Zeuner FE (1934) Die Beziehungen zwischen Schädelform und Lebensweise bei den rezenten und fossilen Nashornern. Berichte der Naturforschenden Gesellschaft zu Freiburg im Breisgau 34: 21–80.
  • Zin-Maung-Maung-Thein , Thaung-Htike , Tsubamoto T, Takai M, Egi N, Maung-Maung (2006) Early Pleistocene Javan rhinoceros from the Irawaddy Formation, Myanmar. Asian Paleoprimatology, Primate Research Institute, Kioto University 4: 197–204.
  • Zin-Maung-Maung-Thein , Takai M, Tsubamoto T, Egi N, Thaung-Htike , Nishimura T, Maung-Maung , Zaw-Win (2010) A review of fossil rhinoceroses from the Neogene of Myanmar with description of new specimens from the Irrawaddy Sediments. Journal of Asian Earth Sciences 37(2): 154–165. https://doi.org/10.1016/j.jseaes.2009.08.009
  • Zschokke S (2016) Genetic structure of the wild populations of the Indian rhinoceros (Rhinoceros unicornis). Indian Journal of History of Science 51(2.2): 380–389. https://doi.org/10.16943/ijhs/2016/v51i2.2/48451
  • Zschokke S, Armbruster GFJ, Ursenbacher S, Baur B (2011) Genetic differences between the two remaining wild populations of the endangered Indian rhinoceros (Rhinoceros unicornis). Biological Conservation 144: 2702–2709. https://doi.org/10.1016/j.biocon.2011.07.031