Systematics and palaeoneurology of a new Pliocene raccoon dog (Canidae, Nyctereutes) from Jradzor (Armenia)
Authors/Creators
- 1. Universitat Autònoma de Barcelona, Barcelona, Spain|Università degli Studi di Firenze, Firenze, Italy
- 2. Ecole Nationale Vétérinaire de Toulouse, Toulouse, France
- 3. Università degli Studi di Firenze, Firenze, Italy
- 4. Università degli Studi di Firenze, Firenze, Italy|Department of Geology, Universitat Autònoma de Barcelona, Barcelona, Spain
- 5. National Museum of China, Beijing, China
- 6. Naturhistorisches Museum Basel, Basel, Switzerland
- 7. JURASSICA Museum, Porrentruy, Switzerland|University of Fribourg, Fribourg, Switzerland
Description
We describe a new fossil raccoon dog from the Late Pliocene site of Jradzor, Armenia, a key site to understand biogeographic connections and dispersal between Asia, Europe, and Africa. The specimens analysed here for the first time exhibit a mosaic of primitive and derived cranial and dental traits, distinguishing them from known taxa such as Nyctereutes tingi (e.g. elongated cranium, reduced subangular lobe), N. sinensis (e.g. molar proportions, sulcal pattern), and N. megamastoides (e.g. absence of squared M1 morphology). Virtual reconstruction of the endocranium reveals a unique sulcal and gyral pattern, including a parenthesis-shaped gyrus sigmoideus and a deeply developed sulcus pseudosylvius, placing it morphologically closer to Canini than extant Vulpini. Considering these features, we propose to ascribe it to a new Nyctereutes species. Morphometric and geometric morphometric analyses support its taxonomic distinctiveness and suggest ecological and behavioural affinities with jackal-like canids. Notably, the relative development of the rostral portion of the cerebrum implies a social structure more complex than that of extant Nyctereutes, suggesting the possibility of seasonal pack formation. Additionally, we reassess several neuroanatomical traits previously considered diagnostic, such as the cerebellar vermis and sulcus endomarginalis, revealing greater variability within Nyctereutes and challenging previous phylogenetic assumptions. These findings underscore the Pliocene diversification of the genus and highlight the Caucasus as a pivotal region in canid evolution. The erection of a new Nyctereutes species enriches our understanding of canid paleobiology and provides new perspectives on the evolutionary trajectories of hypocarnivorous canids.
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References
- Adams DC, Collyer M, Kaliontzopoulou A, Sherratt E (2016) geomorph: Software for geometric morphometric analyses. [Available from:] https://rune.une.edu.au/web/handle/1959.11/21330 [February 9, 2025]
- Alberdi MT, Cerdeño E, López-Martínez N, Morales J, Soria MD (1997) La fauna villafranquiense de El Rincon-1 Albacete. Estudios Geológicos 53: 69–93. https://doi.org/10.3989/egeol.97531-2248
- Alcalá L (1994) Macromamíferos Neógenos de la fosa de Alfambra-Teruel. Instituto de Estudios Turolenses y Museo Nacional de Ciencias Naturales, Teruel, Spain, 554 pp.
- Allman J (1990) Evolution of Neocortex. In: Jones EG, Peters A (Eds) Comparative Structure and Evolution of Cerebral Cortex, Part I. Springer US, Boston, MA, 269–283. https://doi.org/10.1007/978-1-4757-9622-3_7
- Argant A (2004) The Carnivores of the late Pliocene site of Saint-Vallier (Drome, France). Geobios 37: S133–S182. https://doi.org/10.1016/S0016-6995(04)80013-5
- Argant A (2024) Chapter 9 - The Carnivores of Senèze (Haute-Loire). In: Delson E, Faure M, Guérin C (Eds) Senèze: Life in Central France Around Two Million Years Ago: Paleontology, Geochronology, Stratigraphy and Taphonomy. Vertebrate Paleobiology and Paleoanthropology. Springer Nature Switzerland, Cham. https://doi.org/10.1007/978-3-031-64415-3
- Bardeleben C, Moore RL, Wayne RK (2005) A molecular phylogeny of the Canidae based on six nuclear loci. Molecular Phylogenetics and Evolution 37: 815–831. https://doi.org/10.1016/j.ympev.2005.07.019
- Bartolini-Lucenti S (2017) Nyctereutes megamastoides (Canidae, Mammalia) from the early and middle Villafranchian (late Pliocene and early Pleistocene) of the Lower Valdarno (Firenze and Pisa, Tuscany, Italy). Rivista Italiana di Paleontologia e Stratigrafia 123: 211–218.
- Bartolini-Lucenti S, Rook L, Morales J (2018) Nyctereutes (Mammalia, Carnivora, Canidae) from Layna and the Eurasian raccoon-dogs: An updated revision. Rivista Italiana di Paleontologia e Stratigrafia 124: 597–616. https://doi.org/10.13130/2039-4942/10739
- Barton RA, Harvey PH (2000) Mosaic evolution of brain structure in mammals. Nature 405: 1055–1058. https://doi.org/10.1038/35016580
- Bates DMA (1937) Palaeontology: The fossil fauna of the Wady el-Mughara caves. In: The Stone Age of Mount Carmel. Excavations at the Wady El-Mughara. Clarendon Press, Oxford, UK, 137–233.
- Batsch AJGC (1788) Versuch einer Anleitung zur Kenntniss und Geschichte der Thiere und Mineralien: für akademische Vorlesungen entworfen, und mit den nöthigsten Abbildungen versehen. Academischen Buchhandlung, Jena, Germany, 528 pp. [Available from:] https://ia800609.us.archive.org/32/items/versucheineranle11788bats/versucheineranle11788bats.pdf [April 22, 2024]
- Berta A (1988) Quaternary evolution and biogeography of the large South American Canidae (Mammalia, Carnivora). Univ. of California Press, Berkeley, Calif., 149 pp.
- Boch M, Karadachka K, Loh KK, Benn RA, Roumazeilles L, Bertelsen MF, Manger PR, Wriggelsworth E, Spiro S, Spocter MA, Johnson PJ, Avelino-de-Souza K, Patzke N, Lamm C, Miller KL, Sallet J, Khrapitchev AA, Tendler BC, Mars RB (2024) Comparative neuroimaging of the carnivoran brain: Neocortical sulcal anatomy, 2024.06.03.597118. https://doi.org/10.1101/2024.06.03.597118
- Bowdich ET (1821) An analysis of the natural classifications of Mammalia, for the use of students and travellers. J. Smith, Paris, France, 115 pp.
- Broom R, Schepers GWH (1946) 2 The South African Fossil Ape-Men, The Australopithecinae. Transvaal Museum, Pretoria, South Africa.
- Butler AB, Hodos W (2005) Comparative Vertebrate Neuroanatomy: Evolution and Adaptation. 1st ed. Wiley. https://doi.org/10.1002/0471733849
- Chen L, Zhang H (2012) The complete mitochondrial genome and phylogenetic analysis of Nyctereutes procyonoides. Acta Ecologica Sinica 32: 232–239. https://doi.org/10.1016/j.chnaes.2012.07.003
- Czeibert K, Nagy G, Csörgő T, Donkó T, Petneházy Ö, Csóka Á, Garamszegi LZ, Kolm N, Kubinyi E (2024) High-resolution computed tomographic (HRCT) image series from 413 canid and 18 felid skulls. Scientific Data 11: 753. https://doi.org/10.1038/s41597-024-03572-x
- Czyżewska T (1969) Nyctereutes sinensis Schlosser (Canidae, Mammalia) from the Pliocene breccia in Węże, Poland. Acta Zoologica Cracoviensia 14: 441–450.
- Czyżewska T (1981) Natural Endocranial Casts of the Canidae from Węże I near Działoszyn (Poland). Acta Zoologica Cracoviensia 25: 251–260.
- Daguenet T, Sen S (2019) Phylogenetic relationships of Nyctereutes Temminck, 1838 (Canidae, Carnivora, Mammalia) from early Pliocene of Çalta, Turkey. Geodiversitas 41: 663. https://doi.org/10.5252/geodiversitas2019v41a18
- Datta R, Lee J, Duda J, Avants BB, Vite CH, Tseng B, Gee JC, Aguirre GD, Aguirre GK (2012) A Digital Atlas of the Dog Brain. Paxinos G (Ed.). PLoS ONE 7: e52140. https://doi.org/10.1371/journal.pone.0052140
- De Villalta-Comella JF (1952) Contribución al conocimiento de la fauna de mamíferos fósiles del Plioceno de Villaroya. Boletin del Instituto Geologico y Minero de España 64: 1–204.
- Del Campana D (1917) Sulla presenza di (Canis) Cerdocyon megamastoides Pomel, nel Pliocene del Valdarno inferiore. Rivista Italiana di Paleontologia e Stratigrafia 23: 29–36.
- Depéret MC (1890) Les Animaux pliocènes du Roussillon. Mémoires de la Société Géologique de France (Paleontologie) 3: 4–194.
- Dermitzakis MD, Van der Geer AE, Lyras G (2004) The phylogenetic position of raccoon dogs: implications of their neuroanatomy. Proceedings of 5th International Symposium on Eastern Mediterranean Geology.
- Domingo L, Koch PL, Hernández Fernández M, Fox DL, Domingo MS, Alberdi MT (2013) Late Neogene and Early Quaternary Paleoenvironmental and Paleoclimatic Conditions in Southwestern Europe: Isotopic Analyses on Mammalian Taxa. PLoS ONE 8: e63739. https://doi.org/10.1371/journal.pone.0063739
- Dong W, Liu J, Fang Y (2013) The large mammals from Tuozidong (eastern China) and the Early Pleistocene environmental availability for early human settlements. Quaternary International 295: 73–82. https://doi.org/10.1016/j.quaint.2012.01.014
- Dong W, Zhang L-M, Liu W-H (2017) New material of the Early Pleistocene mammalian fauna from Chutoulang, Chifeng, eastern Nei Mongol, China and binary faunal similarity analyses. Vertebrata PalAsiatica 7: 257–275.
- Dozo MT, Paulina-Carabajal A, Macrini TE, Walsh S [Eds] (2023) Paleoneurology of Amniotes: New Directions in the Study of Fossil Endocasts. Springer International Publishing, Cham. https://doi.org/10.1007/978-3-031-13983-3
- Ekdale EG (2013) Comparative Anatomy of the Bony Labyrinth (Inner Ear) of Placental Mammals. Soares D (Ed.). PLoS ONE 8: e66624. https://doi.org/10.1371/journal.pone.0066624
- Engelman RK (2022) Occipital condyle width (OCW) is a highly accurate predictor of body mass in therian mammals. BMC Biology 20: 37. https://doi.org/10.1186/s12915-021-01224-9
- Fedorov A, Beichel R, Kalpathy-Cramer J, Finet J, Fillion-Robin J-C, Pujol S, Bauer C, Jennings D, Fennessy F, Sonka M, Buatti J, Aylward S, Miller JV, Pieper S, Kikinis R (2012) 3D Slicer as an image computing platform for the Quantitative Imaging Network. Magnetic Resonance Imaging 30: 1323–1341. https://doi.org/10.1016/j.mri.2012.05.001
- Ficcarelli G, Torre D, Turner A (1984) First evidence for a species of raccoon dog, Nyctereutes Temminck 1838, in South African Plio-Pleistocene deposits. Bollettino della Società Paleontologica Italiana 23: 125–130.
- Gabunia LK, Mchedlidze GA (1986) Basic features of the brain evolution in some groups of ancient mammals. Metsniereba, Tbilisi, Georgia.
- Gaspard M (1964) La region de l'angle mandibulaire chez les canidae. Mammalia 28: 249–329. https://doi.org/10.1515/mamm.1964.28.2.249
- Geraads D (1997) Carnivores Du Pliocene Terminal de Ahl Al Oughlam (Casablanca Maroc). Geobios 30: 127–164. https://doi.org/10.1016/S0016-6995(97)80263-X
- Geraads D (2011) A revision of the fossil Canidae (Mammalia) of north-western Africa. Palaeontology 54: 429–446. https://doi.org/10.1111/j.1475-4983.2011.01039.x
- Geraads D, Alemseged Z, Bobe R, Reed D (2010) Nyctereutes lockwoodi, n. sp., a new canid (Carnivora: Mammalia) from the middle Pliocene of Dikika, Lower Awash, Ethiopia. Journal of Vertebrate Paleontology 30: 981–987. https://doi.org/10.1080/02724631003758326
- Ginsburg L (1998) Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 5. Carnivores. Sen S (Ed.). Geodiversitas 20: 379–396.
- Gittleman JL (1986) Carnivore brain size, behavioral ecology, and phylogeny. Journal of Mammalogy 67: 23–36. https://doi.org/10.2307/1380998
- Gliozzi E, Abbazzi L, Argenti P, Azzaroli A, Caloi L, Barbato LC, Stefano GD, Esu D, Ficcarelli G, Girotti O, Kotsakis T, Masini F, Mazza P, Mezzabotta C, Palombo MR, Petronio C, Rook L, Sala B, Sardella R, Zanalda E, Torre D (1997) Biochronology of selected mammals, molluscs and ostracods from the Middle Pliocene to the Late Pleistocene in Italy. the State of the Art. Rivista Italiana di Paleontologia e Stratigrafia 103: 369–388. https://doi.org/10.13130/2039-4942/5299
- Gray J (1834) 2 Illustrations of Indian Zoology, chiefly selected from the collection of Major-General Hardwicke. Volume 2. Treuttel, Wurz, Treuttel Jun. and Richter, and Parbury, Allen and Co. Publishers, London, UK, 216 pp.
- Grewal JS, Gloe T, Hegedus J, Bitterman K, Billings BK, Chengetanai S, Bentil S, Wang VX, Ng JC, Tang CY, Geletta S, Wicinski B, Bertelson M, Tendler BC, Mars RB, Aguirre GK, Rusbridge C, Hof PR, Sherwood CC, Manger PR, Spocter MA (2020) Brain gyrification in wild and domestic canids: Has domestication changed the gyrification index in domestic dogs? Journal of Comparative Neurology 528: 3209–3228. https://doi.org/10.1002/cne.24972
- Grohé C, Tseng ZJ, Lebrun R, Boistel R, Flynn JJ (2016) Bony labyrinth shape variation in extant Carnivora: a case study of Musteloidea. Journal of Anatomy 228: 366–383. https://doi.org/10.1111/joa.12421
- Hecht EE, Smaers JB, Dunn WD, Kent M, Preuss TM, Gutman DA (2019) Significant neuroanatomical variation among domestic dog breeds. Journal of Neuroscience 39: 7748–7758. https://doi.org/10.1523/JNEUROSCI.0303-19.2019
- Heffner HE (1983) Hearing in large and small dogs: Absolute thresholds and size of the tympanic membrane. Behavioral Neuroscience 97: 310–318. https://doi.org/10.1037/0735-7044.97.2.310
- Hemprich WF, Ehrenberg CG (1833) Symbolae physicae, seu icones et descriptiones corporum naturalium novorum aut minus cognitorum quae ex itineribus per Libyam, Aegyptium, Nubiam, Dongalam, Syriam, Arabiam et Habessiniam. Pars zoologica II. Berolini, Ex Officina Academica, Berlin, 81 pp. [Available from:] https://www.biodiversitylibrary.org/item/189238
- Hendelman WJ (2006) Atlas of Functional Neuroanatomy. Second Edition. Taylor & Francis Group, Boca Raton, Florida, USA, 253 pp. https://doi.org/10.1201/9780849330841
- Hennig W (1966) Phylogenetic Systematics. Illinois University Press, Chicago, 263 pp.
- Holekamp KE, Benson-Amram S (2017) The evolution of intelligence in mammalian carnivores. Interface Focus 7: 20160108. https://doi.org/10.1098/rsfs.2016.0108
- Horowitz A (1979) The Quaternary of Israel. The Quaternary of Israel 1979: 257–290. https://doi.org/10.1016/B978-0-12-356170-1.50013-1
- Ivanoff DV, Wolsan M, Marciszak A (2014) Brainy stuff of long-gone dogs: A reappraisal of the supposed Canis endocranial cast from the Pliocene of Poland. Naturwissenschaften 101: 645–651. https://doi.org/10.1007/s00114-014-1200-4
- Jiang H, Liu J, Jiangzuo Q, Ning J (2024) The latest ancestor of extant raccoon-dog from Zhoukoudian, Beijing, China, highlights a complex or convoluted transition on the dietary habits. Historical Biology 36: 2739–2761. https://doi.org/10.1080/08912963.2023.2276147
- Jin C, Liu J [Eds] (2009) Paleolithic Site - The Renzidong Cave, Fanchang, Anhui Province. Science Press, Beijing, China.
- Kelava I, Lewitus E, Huttner WB (2013) The secondary loss of gyrencephaly as an example of evolutionary phenotypical reversal. Frontiers in Neuroanatomy 7: 16. https://doi.org/10.3389/fnana.2013.00016
- Kim SI, Oshida T, Lee H, Min MS, Kimura J (2015) Evolutionary and biogeographical implications of variation in skull morphology of raccoon dogs (Nyctereutes procyonoides, Mammalia: Carnivora). Biological Journal of the Linnean Society 116: 856–872. https://doi.org/10.1111/bij.12629
- Kim SI, Park SK, Lee H, Oshida T, Kimura J, Kim YJ, Nguyen ST, Sashika M, Min MS (2013) Phylogeography of Korean raccoon dogs: Implications of peripheral isolation of a forest mammal in East Asia. Journal of Zoology 290: 225–235. https://doi.org/10.1111/jzo.12031
- Kingdon J, Hoffmann M [Eds] (2013) Volumes I–VI Mammals of Africa. Bloomsbury Publishing Plc, London, UK, 3763 pp.
- Koufos GD (1993) Late Pliocene carnivores from western Macedonia (Greece). Paläontologische Zeitschrift 67: 357–376. https://doi.org/10.1007/BF02990288
- Koufos GD (1997) The canids Eucyon and Nyctereutes from the Ruscinian of Macedonia, Greece. Paleontologia i Evolucio 30–31: 39–48.
- Koufos GD (2014) The Villafranchian carnivoran guild of Greece: implications for the fauna, biochronology and paleoecology. Integrative Zoology 9: 444–460. https://doi.org/10.1111/1749-4877.12061
- Kretzoi M (1945) Bemerkungen über das Raubtiersystem. Annales Historico-Naturales Musei Nationalis Hungarici 38: 59–83.
- Kurtén B (1965) The Carnivora of the Palestine Caves. Acta Zoologica Fennica 107: 3–74.
- Kurtén B, Crusafont-Pairó M (1977) Villafranchian carnivores (Mammalia) from La Puebla de Valverde (Teruel, Spain). Commentationes Biologicae 85: 1–39.
- Lazarev S, Maul LC, Kuiper K, Becker D, Bukhsianidze M, Hovakimyan H, Sahakyan L, Vasilyan D (2023) Pliocene volcanic terrain of the South Caucasus as a missing link in Eurasian palaeobiogeography: Dating and palaeoenvironments of vertebrate assemblages of the Jradzor locality, Armenia. Palaeogeography, Palaeoclimatology, Palaeoecology 625: 111685. https://doi.org/10.1016/j.palaeo.2023.111685
- Li Q, Wang X, Qiu Z (2003) Pliocene mammalian fauna of Gaotege in Nei Mongol (Inner Mongolia), China. Vertebrata PalAsiatica 41: 104–114.
- Lindblad-Toh K, Wade CM, Mikkelsen TS, Karlsson EK, Jaffe DB, Kamal M, Clamp M, Chang JL, Kulbokas EJ, Zody MC, Mauceli E, Xie X, Breen M, Wayne RK, Ostrander EA, Ponting CP, Galibert F, Smith DR, DeJong PJ, Kirkness E, Alvarez P, Biagi T, Brockman W, Butler J, Chin C-W, Cook A, Cuff J, Daly MJ, DeCaprio D, Gnerre S, Grabherr M, Kellis M, Kleber M, Bardeleben C, Goodstadt L, Heger A, Hitte C, Kim L, Koepfli K-P, Parker HG, Pollinger JP, Searle SMJ, Sutter NB, Thomas R, Webber C, Baldwin J, Abebe A, Abouelleil A, Aftuck L, Ait-Zahra M, Aldredge T, Allen N, An P, Anderson S, Antoine C, Arachchi H, Aslam A, Ayotte L, Bachantsang P, Barry A, Bayul T, Benamara M, Berlin A, Bessette D, Blitshteyn B, Bloom T, Blye J, Boguslavskiy L, Bonnet C, Boukhgalter B, Brown A, Cahill P, Calixte N, Camarata J, Cheshatsang Y, Chu J, Citroen M, Collymore A, Cooke P, Dawoe T, Daza R, Decktor K, DeGray S, Dhargay N, Dooley KKK, Dooley KKK, Dorje P, Dorjee K, Dorris L, Duffey N, Dupes A, Egbiremolen O, Elong R, Falk J, Farina A, Faro S, Ferguson D, Ferreira P, Fisher S, FitzGerald M, Foley K, Foley C, Franke A, Friedrich D, Gage D, Garber M, Gearin G, Giannoukos G, Goode T, Goyette A, Graham J, Grandbois E, Gyaltsen K, Hafez N, Hagopian D, Hagos B, Hall J, Healy C, Hegarty R, Honan T, Horn A, Houde N, Hughes L, Hunnicutt L, Husby M, Jester B, Jones C, Kamat A, Kanga B, Kells C, Khazanovich D, Kieu AC, Kisner P, Kumar M, Lance K, Landers T, Lara M, Lee W, Leger J-P, Lennon N, Leuper L, LeVine S, Liu J, Liu X, Lokyitsang Y, Lokyitsang T, Lui A, Macdonald J, Major J, Marabella R, Maru K, Matthews C, McDonough S, Mehta T, Meldrim J, Melnikov A, Meneus L, Mihalev A, Mihova T, Miller K, Mittelman R, Mlenga V, Mulrain L, Munson G, Navidi A, Naylor J, Nguyen TT, Nguyen N, Nguyen C, Nguyen TT, Nicol R, Norbu N, Norbu C, Novod N, Nyima T, Olandt P, O'Neill B, O'Neill K, Osman S, Oyono L, Patti C, Perrin D, Phunkhang P, Pierre F, Priest M, Rachupka A, Raghuraman S, Rameau R, Ray V, Raymond C, Rege F, Rise C, Rogers J, Rogov P, Sahalie J, Settipalli S, Sharpe T, Shea T, Sheehan M, Sherpa N, Shi J, Shih D, Sloan J, Smith C, Sparrow T, Stalker J, Stange-Thomann N, Stavropoulos S, Stone C, Stone S, Sykes S, Tchuinga P, Tenzing P, Tesfaye S, Thoulutsang D, Thoulutsang Y, Topham K, Topping I, Tsamla T, Vassiliev H, Venkataraman V, Vo A, Wangchuk T, Wangdi T, Weiand M, Wilkinson J, Wilson A, Yadav S, Yang S, Yang X, Young G, Yu Q, Zainoun J, Zembek L, Zimmer A, Lander ES, Clubs B, Bac B, Resource B, Bac T, Biosciences A, Lindblad-Toh K, Wade CM, Mikkelsen TS, Karlsson EK, Jaffe DB, Kamal M, Clamp M, Chang JL, Kulbokas EJ, Zody MC, Mauceli E, Xie X, Breen M, Wayne RK, Ostrander EA, Ponting CP, Galibert F, Smith DR, DeJong PJ, Kirkness E, Alvarez P, Biagi T, Brockman W, Butler J, Chin C-W, Cook A, Cuff J, Daly MJ, DeCaprio D, Gnerre S, Grabherr M, Kellis M, Kleber M, Bardeleben C, Goodstadt L, Heger A, Hitte C, Kim L, Koepfli K-P, Parker HG, Pollinger JP, Searle SMJ, Sutter NB, Thomas R, Webber C, Baldwin J, Abebe A, Abouelleil A, Aftuck L, Ait-Zahra M, Aldredge T, Allen N, An P, Anderson S, Antoine C, Arachchi H, Aslam A, Ayotte L, Bachantsang P, Barry A, Bayul T, Benamara M, Berlin A, Bessette D, Blitshteyn B, Bloom T, Blye J, Boguslavskiy L, Bonnet C, Boukhgalter B, Brown A, Cahill P, Calixte N, Camarata J, Cheshatsang Y, Chu J, Citroen M, Collymore A, Cooke P, Dawoe T, Daza R, Decktor K, DeGray S, Dhargay N, Dooley KKK, Dooley KKK, Dorje P, Dorjee K, Dorris L, Duffey N, Dupes A, Egbiremolen O, Elong R, Falk J, Farina A, Faro S, Ferguson D, Ferreira P, Fisher S, FitzGerald M, Foley K, Foley C, Franke A, Friedrich D, Gage D, Garber M, Gearin G, Giannoukos G, Goode T, Goyette A, Graham J, Grandbois E, Gyaltsen K, Hafez N, Hagopian D, Hagos B, Hall J, Healy C, Hegarty R, Honan T, Horn A, Houde N, Hughes L, Hunnicutt L, Husby M, Jester B, Jones C, Kamat A, Kanga B, Kells C, Khazanovich D, Kieu AC, Kisner P, Kumar M, Lance K, Landers T, Lara M, Lee W, Leger J-P, Lennon N, Leuper L, LeVine S, Liu J, Liu X, Lokyitsang Y, Lokyitsang T, Lui A, Macdonald J, Major J, Marabella R, Maru K, Matthews C, McDonough S, Mehta T, Meldrim J, Melnikov A, Meneus L, Mihalev A, Mihova T, Miller K, Mittelman R, Mlenga V, Mulrain L, Munson G, Navidi A, Naylor J, Nguyen TT, Nguyen N, Nguyen C, Nguyen TT, Nicol R, Norbu N, Norbu C, Novod N, Nyima T, Olandt P, O'Neill B, O'Neill K, Osman S, Oyono L, Patti C, Perrin D, Phunkhang P, Pierre F, Priest M, Rachupka A, Raghuraman S, Rameau R, Ray V, Raymond C, Rege F, Rise C, Rogers J, Rogov P, Sahalie J, Settipalli S, Sharpe T, Shea T, Sheehan M, Sherpa N, Shi J, Shih D, Sloan J, Smith C, Sparrow T, Stalker J, Stange-Thomann N, Stavropoulos S, Stone C, Stone S, Sykes S, Tchuinga P, Tenzing P, Tesfaye S, Thoulutsang D, Thoulutsang Y, Topham K, Topping I, Tsamla T, Vassiliev H, Venkataraman V, Vo A, Wangchuk T, Wangdi T, Weiand M, Wilkinson J, Wilson A, Yadav S, Yang S, Yang X, Young G, Yu Q, Zainoun J, Zembek L, Zimmer A, Lander ES (2005) Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature 438: 803–819. https://doi.org/10.1038/nature04338
- Linnaeus C von (1758) Tomus I Systema Naturae per Regna Tria Naturae secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Editio Decima. Laurentii Salvii, Stockholm, Sweden, 823 pp. https://doi.org/10.5962/bhl.title.542
- Liu P, Qin H, Li S, Yuan B (2021) Magnetostratigraphic dating of the Danangou and Dongyaozitou mammalian faunas in the Nihewan Basin, North China. Quaternary Science Reviews 257: 106855. https://doi.org/10.1016/j.quascirev.2021.106855
- Liu W (2019) Nyctereutes from Hongya Yangshuizhan Locality at Nihewan Basin, and the Systematic Revision on the Genus Nyctereutes. Unpublished Ph.D. Thesis. Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Science.
- Lyras GA (2009) The evolution of the brain in Canidae (Mammalia: Carnivora). Scripta Geologica 139: 1–39.
- Lyras GA, Van Der Geer AAE (2003) External brain anatomy in relation to the phylogeny of Caninae (Carnivora: Canidae). Zoological Journal of the Linnean Society 138: 505–522. https://doi.org/10.1046/j.1096-3642.2003.00067.x
- Marciszak A, Kropczyk A, Gornig W, Kot M, Nadachowski A (2023) History of Polish Canidae (Carnivora, Mammalia) and Their Biochronological Implications on the Eurasian Background. Genes 14: 1–27. https://doi.org/10.3390/genes14030539
- Martin R (1971) Les affinités de Nyctereutes megamastoides (Pomel), Canidae du gisement villafranchien de Saint-Vallier (Drôme, France). Palaeovertebrata 4: 39–58. https://doi.org/10.18563/pv.4.2.39-58
- Matsumoto H (1930) Report of the mammalian remains obtained from the sited at Aoshima and Hibiku, Province fo Rikuzen. Science reports of the Tohoku Imperial University. 2nd series, Geology 13: 59–93.
- Michaud M, Toussaint SLD, Gilissen E (2022) The impact of environmental factors on the evolution of brain size in carnivorans. Communications Biology 5: 998. https://doi.org/10.1038/s42003-022-03748-4
- Moehlman PD, Hayssen V (2018) Canis aureus. Mammalian Species 50: 14–25. https://doi.org/10.1093/mspecies/sey002
- Monguillon A, Spassov N, Argant A, Kauhala K, Viranta S (2004) Nyctereutes vulpinus comb. et stat. nov. (Mammalia, Carnivora, Canidae) from the late Pliocene of Saint-Vallier (Drôme, France). Geobios 37: S183–S188. https://doi.org/10.1016/S0016-6995(04)80014-7
- Morales J (2016) 6. Los carnivoros de Villaroya. In: Alberdi MT, Azanza B, Cervantes E (Eds) Villaroya, yacimiento llave de la paleotologia Riojana. Instituto de Estudios Riojanos, Logroño, 119–142.
- Müller S (1836) Jahresbericht tiber die Fortschritte der anatomisch-physiologischen Wissenschaften im Jahre 1834. Archiv fur Anatomie und Physiologie 1835: 1–151, 225–243.
- Pei W (1934) On the Carnivora from Locality 1 of Choukoutien. Palaeontologia Sinica Series C 8: 1–216.
- Penrose F, Kemp GJ, Jeffery N (2016) Scaling and accommodation of jaw adductor muscles in Canidae. The Anatomical Record 299: 951–966. https://doi.org/10.1002/ar.23355
- Penrose F, Cox P, Kemp G, Jeffery N (2020) Functional morphology of the jaw adductor muscles in the Canidae. The Anatomical Record 303: 2878–2903. https://doi.org/10.1002/ar.24391
- Poe S, Wiens JJ (2000) Character selection and the methodology of morphological phylogenetics. Phylogenetic analysis of morphological data: 20–36.
- Pomel A (1842) Nouvelle espèce de Chien fossile dècouverte dans les alluvions volcaniques de l'Auvergne. Bulletin de la Société Géologique de France 14: 38–41.
- Qiu Z, Wang X, Li Q (2013) Neogene Faunal Succession and Biochronology of Central Nei Mongol (Inner Mongolia). In: Wang X, Flynn L, Fortelius M (Eds) Fossil Mammals of Asia. Columbia University Press, New York, USA, 155–186. https://doi.org/10.7312/columbia/9780231150125.003.0005
- R Core Team (2024) R: A language and environment for statistical computing, Vienna, Austria. [Available from:] https://www.R-project.org/
- Radinsky LB (1973) Evolution of the Canid Brain. Brain, Behavior and Evolution 7: 169–202. https://doi.org/10.1159/000124409
- Renda S, Roux A le (2017) The sensory ecology of prey detection in the bat-eared fox (Otocyon megalotis). https://doi.org/10.1163/1568539X-00003419
- Reynolds SC (2012) Nyctereutes terblanchei: The raccoon dog that never was. South African Journal of Science 108: 1–10. https://doi.org/10.4102/sajs.v108i1/2.589
- Rook L, Bartolini-Lucenti S, Bukhsianidze M, Lordkipanidze D (2017) The Kvabebi Canidae record revisited (late Pliocene, Sighnaghi, eastern Georgia). Journal of Paleontology 91: 1–14. https://doi.org/10.1017/jpa.2017.73
- RStudio Team (2024) RStudio: integrated development environment for R. Boston, MA. [Available from:] http://www.rstudio.com/
- Sakai ST, Arsznov BM (2017) Carnivoran Brains: Effects of Sociality on Inter- and Intraspecific Comparisons of Regional Brain Volumes. In: Evolution of Nervous Systems. Elsevier, 413–428. https://doi.org/10.1016/B978-0-12-804042-3.00054-3
- Savvidou A, Kostopoulos DS (2025) New and revised middle Villafranchian canid material from Dafnero-3 (Greece): Implications to Nyctereutes systematics and zoogeography. Palaeoworld: 201011. https://doi.org/10.1016/j.palwor.2025.201011
- Schlosser M (1903) Die fossilen Säugethiere Chinas nebst einer Odontographie der recenten Antilopen. Abhandlungen der Bayerischen Akademie der Wissenschaften 22: 1–222.
- Schweizer AV, Lebrun R, Wilson LAB, Costeur L, Schmelzle T, Sánchez-Villagra MR (2017) Size Variation under Domestication: Conservatism in the inner ear shape of wolves, dogs and dingoes. Scientific Reports 7: 13330. https://doi.org/10.1038/s41598-017-13523-9
- Shikama T (1949) The Kuzuü ossuaries: geological and palaeontological studies of the limestone fissure deposits in Kuzuü, Tochigi Prefecture. Scientific Report of the Tohoku University 23.
- Sillero-Zubiri C, Hoffmann M, Macdonald DW (2004) Canids: Foxes, Wolves, Jackals and Dogs Canids: Foxes, Wolves, Jackals and Dogs. Status Survey and Conservation Action Plan, 1–443.
- Simões TR, Vernygora O, Paparella I, Jimenez-Huidobro P, Caldwell MW (2017) Mosasauroid phylogeny under multiple phylogenetic methods provides new insights on the evolution of aquatic adaptations in the group. PLoS ONE 12: e0176773. https://doi.org/10.1371/journal.pone.0176773
- Soria D, Aguirre E (1976) El cánido de Layna: revisión de los Nyctereutes fósiles. Trabajos sobre Neogeno y Cuaternario 5: 83–116.
- Sotnikova M (2004) The Middle Pliocene assemblage of Carnivore from Shamar locality (Northern Mongolia). In: The International Conference on the Paleontology of the Central Asia. Paleontological Institute, Moscow, Russia, 54–56.
- Sotnikova MV, Baigusheva VS, Titov VV (2002) Carnivores of the Khapry faunal assemblage and their stratigraphic implications. Stratigraphy and Geological Correlation 10: 375–390.
- Spassov N (2000) Biochronology and zoogeographic affinities of the Villafranchian faunas of Bulgaria and south Europe. Historia naturalis Bulgarica 12: 89–128.
- Storr GCC (1780) Prodromvs methodi mammalivm. Litteris Reissianis, Tvbingæ, Germany, 43 pp.
- Tamvakis A, Savvidou A, Spassov N, Youlatos D, Merceron G, Kostopoulos DS (2023) New insights on Early Pleistocene Nyctereutes from the Balkans based on material from Dafnero-3 (Greece) and Varshets (Bulgaria). Palaeoworld 32: 555–572. https://doi.org/10.1016/j.palwor.2022.09.006
- Tedford RH, Qiu Z (1991) Pliocene Nyctereutes (Carnivora: Canidae) from Yushe, Shanxi, with comments on chinese fossil racoon-dogs. Vertebrata PalAsiatica 29: 176–189.
- Tedford RH, Taylor BE, Wang Xiaoming (1995) Phylogeny of the Caninae (Carnivora: Canidae): the living taxa. American Museum Novitates 3146: 1–40.
- Tedford RH, Wang X, Taylor BE (2009) Phylogenetic Systematics of the North American Fossil Caninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 2009: 1–218. https://doi.org/10.1206/574.1
- Teilhard de Chardin P, Piveteau J (1930) Les mammifères fossiles de Nihowan (Chine). Annales de Paleontologie 19: 1–134.
- Temminck C (1838) Over de kennis en de Verbreiding der Zoogdieren van Japan. Tijdschrift Natuurlijke Geschiedeins en Physiologie 5: 273–293.
- Torre D (1979) The Ruscinian and the Villafranchian dogs of Europe. Bollettino della Società Paleontologica Italiana 18: 162–165.
- Vekua A (1972) Kvabebi Fauna of Akchagylian Vertebrates. Moscow, Russia, 258 pp.
- Viret J (1954) Le lœss à bancs durcis de Saint-Vallier (Drôme), et sa faune de mammifères villafranchiens (avec une analyse granulométrique et une analyse pollinique). Nouvelle Archives du Musèum d'Histoide Naturelle de Lyon 4: 1–258. https://doi.org/10.3406/mhnly.1954.986
- Wada MY, Imai HT (1991) On the robertsonian polymorphism found in the Japanese raccoon dog (Nyctereutes procyonoides viverrinus). Idengaku Zasshi 66: 1–11. https://doi.org/10.1266/jjg.66.1
- Wada MY, Lim Y, Wurster-Hill DH (1991) Banded karyotype of a wild-caught male Korean raccoon dog, Nyctereutes procyonoides koreensis. Genome 34: 302–306. https://doi.org/10.1139/g91-049
- Wang X, Flynn LJ, Fortelius M [Eds] (2013) Fossil Mammals of Asia Fossil Mammals of Asia. Columbia University Press, New York, 759 pp. https://doi.org/10.7312/columbia/9780231150125.001.0001
- Ward OG, Wurster-Hill DH (1990) Nyctereutes procyonoides. Mammalian Species 358: 1–5. https://doi.org/10.2307/3504213
- Ward OG, Wurster-Hill DH, Ratty FJ, Song Y (1987) Comparative cytogenetics of Chinese and Japanese raccoon dogs, Nyctereutes procyonoides. Cytogenetics and Cell Genetics 45: 177–186. https://doi.org/10.1159/000132451
- Wayne RK, Ostrander EA (2007) Lessons learned from the dog genome. Trends in Genetics 23: 557–567. https://doi.org/10.1016/j.tig.2007.08.013
- Wayne RK, Lau LM, Geffen E, Girman DJ, Koepfli KP, Marshall CR (1997) Molecular systematics of the Canidae. Systematic Biology 46: 622–653. https://doi.org/10.1093/sysbio/46.4.622
- Werdelin L, Peigné S (2010) Carnivora. In: Werdelin L, Sanders W (Eds) Cenozoic Mammals of Africa. University of California Press, Berkeley, California, USA, 609–633. https://doi.org/10.1525/california/9780520257214.003.0032
- Werdelin L, Dehghani R (2011) Carnivora. In: Harrison T (Ed.) Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 2: Fossil Hominins and the Associated Fauna. Vertebrate Paleobiology and Paleoanthropology. Springer International Publishing, New York, 189–232. https://doi.org/10.1007/978-90-481-9962-4
- Westbury M, Dalerum F, Norén K, Hofreiter M (2017) Complete mitochondrial genome of a bat-eared fox (Otocyon megalotis), along with phylogenetic considerations. Mitochondrial DNA, Part B, Resources 2: 298–299. https://doi.org/10.1080/23802359.2017.1331325
- Wickham H, Chang W, Wickham MH (2016) Package 'ggplot2'. Create elegant data visualisations using the grammar of graphics. Version 2: 1–189. https://doi.org/10.1007/978-3-319-24277-4_9
- Wiley EO, Siegel-Causey D, Brooks DR, Funk VA (1991) The Compleat Cladist - A Primer of Phylogenetic Procedures. University of Kansas Museum of Natural History Special Publication 19: 1–12. https://doi.org/10.5962/bhl.title.4069
- Zhao C, Zhang H, Liu G, Yang X, Zhang J (2016) The complete mitochondrial genome of the Tibetan fox (Vulpes ferrilata) and implications for the phylogeny of Canidae. Comptes Rendus - Biologies 339: 68–77. https://doi.org/10.1016/j.crvi.2015.11.005
- Zimmermann EAW von (1780) Geographische Geschichte des Menschen und der allgemein verbreiteten vierfüßigen Tiere. Zweiter Band. Weygandschen Buchhandlung, Liepzig. [Available from:] https://cir.nii.ac.jp/crid/1130848327417958144 [January 8, 2024]
- Zrzavý J, Duda P, Robovský J, Okřinová I, Pavelková Řičánková V (2018) Phylogeny of the Caninae (Carnivora): Combining morphology, behaviour, genes and fossils. Zoologica Scripta 47: 373–389. https://doi.org/10.1111/zsc.12293