Naoto HANDA, Ph.D.
Lake Biwa Museum, curator
e-mail: hndnaoto (at) gmail.com
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Research Interest
Paleobiogeography of Miocene Rhinocerotidae from Africa
Taxonomic revision of Cenozoic mammals in Asia
Elucidation of faunal composition of Mammoth fauna in Mongolia
Keywords
Vertebrate paleontology, Biostratigraphy, Paleobiogeography, Systematics
Publications
Please contact me if you are interested in reading PDF files of my publications.
1) Handa, N., Watabe, M., Tsogtbaatar, Kh. (2012)
New specimens of Protoceratops (Dinosauria: Neoceratopsia) from the Upper Cretaceous in Udyn Sayr, southern Gobi area, Mongolia. Paleontological Research. vol. 16, no. 3, 179-198. link
2) Handa, N., Nakada, K., Anso, J., Matsuoka, A. (2014)
Bathonian/Callovian (Middle Jurassic) ammonite biostratigraphy of the Kaizara Formation of the Tetori Group in central Japan. Newsletters on Stratigraphy. vol. 47, 283-297. link
3) Handa, N. (2014)
Review of the Neogene vertebrate fossils from Shinshushinmachi, Nagano Prefecture, central Japan. Bulletin of the Nagano City Museum, vol. 15, 108-118. (in Japanese)
4) Furumi, Y., Handa, N. Mizoguchi, M., Ito, T., Sato, T., Yoshino, K., Abe, Y., Murasawa, S., Takenouchi, K. (2014)
Elasmobranchs from the Upper Miocene Otogawa Formation in the Yatsuo Area, Toyama Prefecture, central Japan. Bulletin of the Itoigawa City Museums, no. 3, 33-42. (in Japanese)
5) Goto, M., Handa, N. (2014)
Ammonoid from the Lower Formation of the Kuzuryu Subgroup, Tetori Group in the Itoshiro area, Ono City, Fukui Prefecture, Central Japan. Memoir of the Fukui Prefectural Dinosaur Museum, vol. 13, 9-15. (in Japanese) link
6) Handa, N. (2015)
A Pleistocene rhinocerotid (Mammalia, Perissodactyla) from Yage, Shizuoka Prefecture, central Japan. Paleontological Research. vol. 19, 139-142. link
7) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Tsubamoto, T., Nakaya, H. (2015)
New specimens of Chilotheridium (Perissodactyla, Rhinocerotidae) from the Upper Miocene Namurungule and Nakali Formations, northern Kenya. Paleontological Research. vol. 19, 181-194. link
8) Handa, N., Pandolfi, L. (2016)
Reassessment of the middle Pleistocene Japanese rhinoceroses (Mammalia, Rhinocerotidae) and paleobiogeographic implications. Paleontological Research. vol. 20, 247-260. link
9) Handa, N., Kawabe, S. (2016)
Femur of Schizotheriinae (Perissodactyla, Chalicotheriidae) from the Lower Miocene Hiramaki Formation of the Mizunami Group in Gifu Prefecture, central Japan. Journal of Vertebrate Paleontology, vol. 36, e1131163 link
10) Handa, N., Takechi, Y. (2017)
A Pleistocene rhinocerotid (Mammalia, Perissodactyla) from Bisan-Seto area, western Japan. The Journal of the Geological Society of Japan, vol. 123, no. 6, 433-441. link
11) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2017a)
A new Elasmotheriini (Perissodactyla, Rhinocerotidae) from the upper Miocene of Samburu Hills and Nakali, northern Kenya. Geobios, vol. 50, 197-209. link
12) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2017b)
Additional specimens of Diceros (Perissodactyla, Rhinocerotidae) from the Upper Miocene Nakali Formation in Nakali, central Kenya. Historical Biology, 1-12. link
13) Handa, N., Izuho, M., Takahashi, K., Iizuka, F., Tsogtbaatar, B., Gunchinsuren, B., Odsuren, D., Ishtseren, L. (2017)
Discovery of a Late Pleistocene rhinocerotid fossil from Ondorkhaan, eastern Mongolia. Journal of the Geological Society of Japan, vol. 123, no. 12, v-vi (in Japanese)
14) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2018)
Brachypotherium (Perissodactyla, Rhinocerotidae) from the late Miocene of Samburu Hills, Kenya. Geobios. vol. 51, 391-399. link
15) Handa, N. (2019)
Reassessment of a Pleistocene rhinocerotid (Mammalia, Perissodactyla) from Aira, Kagoshima, southwestern Japan. Paleontological Research. vol. 53, 55-65. link
16) Handa, N., Kohno, N., Kudo, Y. (2019) Reappraisal of a middle Pleistocene rhinocerotid (Mammalia, Perissodactyla) from the Matsugae Cave, Fukuoka Prefecture, southwestern Japan. Historical Biology. link
17) Tsubamoto, T., Kunimatsu, Y., Sakai, T., Saneyoshi, M., Shimizu, D., Morimoto, N., Nakaya, H., Handa, N., Tanabe, Y., Manthi, F. K., Nakatsukasa, M. (2020)
A new species of Nyanzachoerus (Mammalia, Artiodactyla, Suidae, Tetraconodontinae) from the upper Miocene Nakali Formation, Kenya. Paleontological Research. vol. 24, 41-63. link
18) Handa, N. (2020)
Reappraisal of a rhinocerotid (Mammalia, Perissodactyla) from the lower Miocene Yotsuyaku Formation, Northeast Japan, with an overview of the early Miocene Japanese rhinocerotids. Paleontological Research. vol. 23, 183-191. link
19) Handa, N., Nishioka, Y., Duangkrayom, J., Jintasakul, P. (2020)
Brachypotherium perimense (Perissodactyla, Rhinocerotidae) from the Miocene of Nakhon Ratchasima, Northeastern Thailand, with comments on fossil records of Brachypotherium. Historical Biology. link
20) Handa, N., Kato, T. (2020)
A Pliocene rhinocerotid (Mammalia, Perissodactyla) from Ajimu, Oita Prefecture, southwestern Japan, with comments on the Japanese Pliocene rhinocerotid fossil records. Paläontologische Zeitschrift. link
21) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Tsubamoto T. and Nakaya, H. (2021)
The Chalicotheriidae (Mammalia, Perissodactyla) from the upper Miocene Nakali Formation, Kenya. Historical Biology. link
22) Handa, N., Tsubamoto, T. (2021)
Reappraisal of a supposed chalicotheriid perissodactyl femur from the Pliocene of Yenangyoung, central Myanmar. Journal of Fossil Research, vol. 54 (1), 11-14.
23) Handa, N., Izuho, M., Takahashi,K. Iizuka, F., Tsogtbaatar, B., Gunchinsuren, B., Odouren, D., Ishitseren, L. (2022)
The woolly rhinoceros (Coelodonta antiquitatis) from Ondorkhaan, eastern Mongolia. Boreas.vol. 51 (3), 584-605. https://doi.org/10.1111/bor.12582
24) Handa, N., Kato, T., Takahashi, K., Baba, R., Kitabayashi, E. (2023)
An additional remain of Pliocene Rhinocerotidae from Ajimu, western Japan. Historical Biology, https://doi.org/10.1080/08912963.2023.2217197
25) Handa, N. (2023)
The Rhinocerotid Fossils of Naora Collection, Bulletin of the National Museum of Japanese History, vol. 243, 71-79. https://rekihaku.repo.nii.ac.jp/records/2000039
26) Naksri, W., Nishioka, Y., Duangkrayom, J., Métais, G., Handa, N., Jintasakul, P., Martin, J.E., Sila, S., Sukdi, W., Suasamong, K., Tong, H., Claude, J. (2023)
A new Miocene and Pleistocene continental locality from Nakhon Ratchasima in Northeastern Thailand and its importance for vertebrate biogeography, Annales de Paléontologie, vol. 109, Issue 4, https://doi.org/10.1016/j.annpal.2023.102659
27) Longuet, M., Handa, N., Zin-Maung- Maung-Thein, Thaung- Htike, Man-Thit- Nyein, Takai, M. (2024)
Post-cranial remains of Rhinocerotidae from the Neogene of central Myanmar: morphological descriptions and comparisons with ratios, Historical Biology, https://10.1080/08912963.2024.2408617
28) Handa, N., Takahashi, K. (2024)
A review of the taxonomy, biostratigraphy and paleobiogeography of Plio-Pleistocene rhinoceroses in Japan. Journal of Asian Earth Science, https://doi.org/10.1016/j.jseaes.2024.106357
29) Handa, N., Taru, (2025)
Taxonomic revision of a late Miocene rhinoceros from Japan with an overview of Brachypotherium from East Asia. Historical Biology, https://doi.org/10.1080/08912963.2025.2456950
Abstracts
1) Handa, N., Ito T., Sato, T., Mizoguchi, M., Yoshino, K., Murasawa S., Hurumi, A. (2007)
Fossil teeth of Elasmobranch from the Upper Miocene Otogawa Formation in the Yatsuo Area, Toyama Prefecture, central Japan. The 2007 Annual Meeting of the Paleontological Society of Japan. Osaka, Japan
2) Handa, N., Anso, J., Yin, J., Matsuoka, A. (2008)
Molluscan biostratigraphy of the Kaizara Formation of the Tetori Group in the Kuzuryugawa area, Fukui Prefecture. The 2008 Annual Meeting The Paleontological Society of Japan, Miyagi, Japan
3) Matsuoka, A., Handa, N., Nakada, K., Terabe, K., Anso, J., Sato, T. (2008)
Stratigraphic levels of Pseudoneuqueniceras-bearing beds in the Kaizara Formation of the Kuzuryu Subgroup and biometrical analysis of Pseudoneuqueniceras. The 115th Annual Meeting of the Geological Society of Japan. Akita, Japan
4) Matsuoka, A., Nakada, K., Nikaido, T., Kokubo, S., Ito, T., Yoshioka, T., Ishida, N., Kurihara, T., Terabe, K., Handa, N., Yoshino, K., Sato, T. (2009)
Report of the events of Asahimachi Museum of Niigata University. The 158th Regular Meeting of the Paleontological Society of Japan. Okinawa, Japan
5) Handa, N., Matsumoto, Y., Watabe, M., Suzuki, S., Nakaya, H. (2009)
Morphology of protoceratopsid from Upper Cretaceous in Udyn Sayr, Mongolia.The 116th Annual Meeting of the Geological Society of Japan. Okayama, Japan
6) Handa, N., Matsumoto, Y., Watabe, M., Suzuki, S., Nakaya, H., Tsogtbaater, Kh. (2009)
Morphology of Protoceratops from the Upper Cretaceous in Udyn Sayr, Mongolia. 4th International Symposium of the IGCP507. Kumamoto, Japan
7) Handa, N., Watabe, M., Tsogtbaatar, Kh., Nakaya, H. (2011)
New protoceratopsid specimen from the Upper Cretaceous in Udyn Sayr, Mongolia. The 160th Regular Meeting Paleontological Society of Japan. Kochi, Japan
8) Handa, N., Watabe, M., Tsogtbaatar, Kh., Nakaya, H. (2011)
Protoceratopsid fossils from the Upper Cretaceous in Udyn Sayr, Mongolia. World Conference on Paleontology and Stratigraphy 2011. Nachon Ratchashima, Thailand
9) Miyata, K., Saegusa, H., Nakaya, H., Ugai, H., Kato, T., Nakata, A., Hattori, H., Akashi, N., Handa, N., Nakatani, D., Ode, S., Yamada, E., Akahoshi, M., Sekiya, H., Murakami, T., Otsuka, K., Onodera, M., Iwai, J., Watanabe, H., Kitagawa, K., Sakado, M. (2012)
Early Miocene rhinocerotid specimen from the "Takashima Formation", Takashima island (Taka Island), Matsuura City, Nagasaki Prefecture. The 161th Regular Meeting Paleontological Society of Japan. Gunma, Japan
10) Handa, N., Nakaya, H., Nakatsukasa, M., Kunimatsu, Y. (2012)
The Late Miocene Rhinocerotidae (Perissodactyla, Mammalia) fossils from the Namurungule and Nakali Formations of northern Kenya. The 49th Annual Meeting of Japan Association for African Studies. Osaka, Japan
11) Handa, N., Nakaya, H., Nakatsukasa, M., Kunimatsu, Y. (2012)
The Late Miocene Elasmotheriini (Rhinocerotidae, Mammalia) from the Namurungule and Nakali Formations of northern Kenya. Japan Geoscience Union Meeting 2012. Chiba, Japan. link
12) Handa, N., Nakaya, H., Nakatsukasa, M., Kunimatsu, Y. (2012)
New Specimens of Elasmotheriini (Rhinocerotidae, Perissodactyla) from the Namurungule and Nakali Formations (early Late Miocene) of northern Kenya. 72th Annual Meeting of the Societ of Vertebrate Paleontology, USA.
13) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2013)
Chilotheridium (Perissodactyla, Rhinocerotidae) from the Upper Miocene Namurungule and Nakali Formations, northen Kenya, The 162th Regular meeting of the Paleontological Society of Japan, Yokohama, Japan.
14) Ito, M., Handa, N., Ode, S., Katsumura, T., Tanabe, Y., Iwai, J., Watanabe, H., Inuzuka, N., Nakaya, H. (2013)
Mammalian fossils from the Miocene Kukinaga Group of Tanegashima Island in Kagoshima Prefecture, The 162th Regular meeting of the Paleontological Society of Japan, Yokohama, Japan.
15) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2013)
The Late Miocene Chilotheridium (Rhinoceortidae, Perissodactyla) from Samburu Hills and Nakali, Northern Kenya, The 50th Annual Meeting of Japan Association for African Studies Tokyo, Japan.
16) Handa, N. (2014)
A rhinocerotid incisor fossil of the Nishizawa collection, The 163th Regular meeting of the Paleontological Society of Japan, Hyogo, Japan.
17) Handa, N., Nakada, K., Anso, J., Matsuoka, A. (2014)
Ammonite biostratigarphy of the Middle Jurassic Kaizara Formation of the Tetori Group in central Japan. International Symposium on Asian Dinosaurs in Fukui, Fukui, Japan.
18) Handa, N., Nakatsukasa, M., Kunimatsu, Y. (2014)
Late Miocene rhinocerotid assemblage from Samburu Hills and Nakali, northern Kenya. The 2014 Annual Meeting of the Palaeontological Society of Japan, Fukuoka, Japan.
19) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Hideo Nakaya (2015)
The Miocene rhinocerotid faunal change in East Africa. The 2015 Annual Meeting of the Palaeontological Society of Japan, Tsukuba, Japan.
20) Handa, N. and Pandolfi, L. (2016)
Taxonomic revision of Dicerorhinus nipponicus (Perissodactyla, Rhinocerotidae) from the middle Pleistocene in Mine City, Yamaguchi Prefecture. The 165th Regular meeting of the Paleontological Society of Japan, Kyoto, Japan.
21) Shikazawa, Y., Sakai, Y., Handa., N., Matsuoka, A. (2016)
Jurassic marine bivalve fossils from the Tetori Group in the Itoshiro and Nagano areas, Ono City, Fkukui Prefecture, central Japan. The 165th Regular meeting of the Paleontological Society of Japan, Kyoto, Japan.
22) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Hideo Nakaya (2016)
Diceros (Mammalia, Perissodactyla) from the Upper Miocene Nakali Formation in northern Kenya and its paleobiogeographic significance. The 2016 Annual Meeting of the Palaeontological Society of Japan, Fukui, Japan.
23) Pandolfi, L., Handa, N., Lu, X. (2016)
Systematics and phylogeny of Old world aceratheres (Mammalia, Rhinocerotidae).
XIV Annual Meeting of the European Association of Vertebrate Paleontologists, Haarlem, the Netherlands. p. 158.
24) Shikazawa, Handa., N., Y., Sakai, Y.,Matsuoka, A. (2016)
Bivalve fossils and taphonomy in the Kaizara Fromation, Kuzuryu Subgroup, Tetori Group, distributed in Izumi Region, Ono City, Fkukui Prefecture, Central Japan.
The 123th Annual Meeting of the Geological Society of Japan. Tokyo, Japan
25) Handa, N., Izuho, M., Takahashi, K., Batmunkh Tsogtbaatar, Byambaa Gunchinsuren, Davaakhuu Odsuren (2017)
Preriminary report of the Pleistocene rhinoceros fossil from Ondorkhaan, eastern Mongolia.
The 166th Regular meeting of the Paleontological Society of Japan, Tokyo, Japan
26) Handa, N. (2017)
A review of Japanese Pleistocene Rhinocerotidae (Mammalia, Perissodactyla) and paleobiogeographical significance. 77th Annual Meeting of the Societ of Vertebrate Paleontology, Calgary, Canada.
27) Handa, N. (2017)
Revision of the Plio-Pleistocene rhinocerotid from Japan and its stratigraphy. The 124th Annual Meeting of the Geological Society of Japan.Ehime, Japan
28) Handa, N., Nishioka, Y. (2018)
Brachypotherium (Perissodactyla, Rhinocerotidae) from the Miocene of Nakhon Ratchashima, northeastern Thailand. The 167th Regular meeting of the Palaeolontological Society of Japan, Ehime, Japan.
29) Handa, N. (2018)
A review of the Miocene Rhinocerotidae (Mammalia, Perissodactyla) of Japan. The 2018 Annual Meeting of the Palaeontological Society of Japan, Sendai, Japan.
30) Handa, N., Nakatsukasa, M., Kunimatsu, Y., Nakaya, H. (2018)
Late Miocene rhinocerotid assemblage from Samburu Hills and Nakali in Kenya. 5th International Palaeontological Congress, Paris, France
31) Handa, N., Nakaya, N., Kunimatsu, Y., Saegusa, H. (2019)
Miocene Rhinocerotidae and Chalicotheriidae (Perissodactyla) from Chiang Muan, Northern Thialand. The 168th Regular meeting of the Palaeolontological Society of Japan, Odawara, Japan.
32) Handa, N. (2019)
Taxonomic revision of "Rhinoceros sp." from the Pleistocene in Kuzuu area, Tochigi Prefecture. The 2019 Annual Meeting of the Palaeolontological Society of Japan, Shizuoka, Japan.
33) Handa, N., Kato, T. (2020)
A review of Japanese Pliocene rhinocerotids (Perissodactyla). The 168th Regular meeting of the Palaeolontological Society of Japan, Tokyo, Japan.
34) Ito, M., Handa, N., Nakaya, H., Tsubamoto, T. (2021)
An rhinocerotid incisor from the Miocene Kukinaga Group in Tanegashima Island, Kagoshima, Japan, The 2021 Annual Meeting of the Palaeontological Society of Japan, Okayama, Japan
35) Tsubamoto,T., Handa, N. (2021)
Reappraisal of a supposed chalicotheriid (Mammalia,Perissodactyla) femur from the Pliocene of Central Myanmar. 81st Annual Meeting of the Society of Vertebrate Paleontology, virtual meeting.
36) Handa, N., Izuho, M., Takahashi, K., Iizuka, F., Tsogtbaatar, B., Gunchinsuren, B., Odsuren, D., Ishitseren, L. (2022)
The woolly rhinoceros (Coelodonta antiquitatis) from Ondorkhaan, eastern Mongolia. The 171st Regular meeting of the Palaeolontological Society of Japan, Nagoya, Japan.
37) Nishioka, Y., Kunimatsu, Y., Kusaka, S., Handa, N. (2022)
Miocene mammalian fossil assemblages from Thailand: a preliminary report on field research in 2017-2018. The 171st Regular meeting of the Palaeolontological Society of Japan, Nagoya, Japan.
38) Handa, N., Kato, T., Takahashi, K., Kitabayashi, E. (2022)
An additional remain of the Pliocene Rhinocerotidae from Oita Prefecture. The 2022 Annual Meeting of the Palaeontological Society of Japan, Kanazawa, Japan
39) Handa, N. (2023)
A taxonomic revision of the Late Miocene Rhinocerotidae from Tunisia. The 172nd Regular meeting of the Palaeolontological Society of Japan, Hakata, Japan
40) Handa, N. (2023)
A review of late Cenozoic fossil Rhinocerotidae from Japan. 2nd Asian Palaeontological Congress, Tokyo, Japan
41) Handa, N. (2023)
An overview of the study of Plio-Pleistocene rhinocerotids in Japan, Annual meeting of the Mammal Society of Japan 2023, Okinawa, Japan
42) Handa, N. Taru, H. (2023)
An taxonomic revision of late Miocene rhinocerotid from the Oiso Formation, Kanagawa Prefecture, The 130th Annual Meeting of the Geological Society of Japan. Kyoto, Japan
43) Handa, N., Kunimatsu, Y., Nishioka, Y., Kusaka, S. (2024)
Miocene rhinocerotid assemblage from Phra Phut sandpit, Khorat area in Northeastern Thailand. The 173th Regular meeting of the Palaeolontological Society of Japan, Miyagi, Japan
44) Handa, N., Tomiya, S. (2024)
A preliminary report of a Pliocene carnivoran from the Ueno Formation, Kobiwako Group. The 2024 Annual Meeting of the Palaeontological Society of Japan, Kochi, Japan
45) Handa, N. (2025)
An overview on the Plio-Pleistocene Japanese rhinocerotids. The 2025 Annual Meeting of Fossil Research Society of Japan, Japan
46) Handa, N., Nishioka, Y. (2025)
Taxonomic revision of a Middle Pleistocene Bovidae from the Kobiwako Group. The 2025 Annual Meeting of the Palaeontological Society of Japan, Hokkaido, Japan
Grants
1) Fujiwara Natural History Foundation Grant (2012)
2) Sasakawa Scientific Research Grant (2012)
3) Grant of Nagano Society for the Promotion of Science (2013)
4) Grant of Nagano Society for the Promotion of Science (2015)
5) Fujiwara Natural History Foundation Grant (2016)
6) Grants-in-Aid for Scinetific Research (Early-Career Scientists) (2018)
7) Grants-in-Aid for Scinetific Research (C) (2021)
8) Grants-in-Aid for Scinetific Research (B) (2022)
9) Grants-in-Aid for Scinetific Research (B) (2024)
Degree
B. Sc. (March 2008), Department of Geology, Faculty of Science, Niigata University.
Thesis: Lithostratigraphy and molluskan biostratigraphy of the Middle Jurassic to Lower Cretaceous Tetori Group in Fukui Prefecture, central Japan
M.Sc. (March 2010), Department of Earth and Environment of Science, Kagoshima University.
Thesis: Ceratopsian dinosaur from the Upper Cretaceous in southern Gobi area, Mongolia
Ph.D. (March 2016), Graduate school of Science and Engineering, Kagoshima University.
Thesis: Fossil rhinoceroses from the Late Miocene of Kenya - Miocene faunal changes of the Rhinocerotidae (Mammalia, Perissodactyla) in sub-Saharan East Africa -