Abstract
The rhinocerotids (Mammalia: Rhinocerotidae) represent a herbivorous perissodactyl family that includes the extant rhinoceroses and their phylogenetically related extinct species. The family appears in the fossil record during the Eocene, and today comprises five extant species, two of which survive in the Afrotropical realm of Sub-Saharan Africa and three in the Indomalayan realm of Southern and Southeastern Asia. In Greece, the occurrence of rhinocerotids has been noted in ca. 80 fossiliferous localities ranging from the Middle Miocene to the Late Pleistocene. The reassessment of the Greek fossil record has revealed the presence of 11 extinct rhinocerotid species, representing eight genera and two subfamilies. During the Middle Miocene the family Rhinocerotidae is represented in Greece by the species Brachypotherium brachypus and Aceratherium simorrense; during the Late Miocene by the species Acerorhinus neleus, Chilotherium schlosseri, Chilotherium samium, Dihoplus pikermiensis, and Miodiceros neumayri gen. nov.; and during the Pliocene and Pleistocene by the species Stephanorhinus jeanvireti, Stephanorhinus etruscus, Stephanorhinus hemitoechus, and Coelodonta antiquitatis. The peak in the taxonomic diversity of the family in Greece is achieved during the Late Miocene, when up to three different rhinocerotid species with distinctive anatomical and dietary adaptations have been documented as sympatric in the classical localities of Pikermi (Attica), Samos Island (Aegean Sea), and Axios Valley (Macedonia).
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Acknowledgments
For facilitating access to specimens under their care, as well as constructive discussions, I am greatly indebted to G. Theodorou, M. Dermitzakis, S. Roussiakis, A. Athanassiou (AMPG); J. Hooker, A. Currant, R. Sabin (BMNH); K. Heissig, G. Rössner (BSPG); M. Bertling (GPMM); O. Sandrock (HLMD); D. Nagel, G. Rabeder (IPUW); G. Lenglet (IRSNB); E. Gilissen (KMMA); G. Koufos, E. Tsoukala, D. Kostopoulos (LGPUT); L. Kordos, K. Palotás (MAFI); P. Tassy, C. Sagne, C. Lefevre, J. Cuisin, J.-G. Michard (MNHN); L. van de Hoek Ostende, J. de Vos, N. den Ouden, H.J. van Grouw, P. Kamminga (Naturalis); O. Hampe, N. Klein (NHMB); U. Göhlich, G. Höck, F. Zachos, F. Spitzenberger, B. Herzig, A. Bibl (NHMW); P. Benda (NMP); Berhane Asfaw (MNE); N. Solounias (NYIT); T. White, R. Bernor (RHOI); F. Schrenk, G. Plodowski, O. Kullmer, J.L. Franzen (SMF); E. Frey, H.W. Mittmann, W. Munk, D. Schreiber (SMNK); E. Heizmann (SMNS); W. von Koenigswald, M. Sander (STIB); H. Baagøe (ZMUC). Bibliographical research has been greatly facilitated by the Rhino Resource Center and the Biodiversity Heritage Library. Financial support for comparative studies was provided by the Deutscher Akademischer Austausch Dienst (DAAD), the European Commission’s Research Infrastructure Action (EU-SYNTHESYS: GB-TAF-574, FR-TAF-1226, NL-TAF-2513, FR-TAF-2545, BE-TAF-5664, CZ-TAF-4227, DK-TAF-5242, AU-TAF 5710), the European Science Foundation (ESF-EEDEN/2003/EX05), and the Revealing Hominid Origins Initiative (RHOI, under NSF award BCS-0321893). The present contribution is dedicated to memory of our beloved colleague and friend Katerina Vasileiadou.
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Appendices
Appendix 1
List of the Greek localities containing Pliocene and Pleistocene rhinocerotid fossils. Locality numbers refer to the collection numbers of the PaleoBiology Database (PBDB). Biochronological data (age) are primarily based on Koufos (2013, 2016b) and Koufos and Kostopoulos (2016), as well as on evaluation of the biostratigraphical and faunal context of each locality. A detailed taxonomic history of the rhinocerotid record of each locality is provided in chronological order.
Localities | Age | Taxonomic history | Key references |
|---|---|---|---|
Aggitis Cave (204391) | Late Pleistocene Aurelian MNQ 26 34.0–27.9 Ka | Coelodonta antiquitatis1,2 | 2Trantalidou (2013) 1Koufos (1981) |
Agia Triada (204745) | Pleistocene (imprecise) | Rhinocerotidae indet.2 = Dicerorhinus sp.1 | Symeonidis et al. (2006) 1Marinos (1965) |
Aivaliki (195881) | Early Pleistocene Villafranchian | Stephanorhinus etruscus2 = Stephanorhinus cf. etruscus1 | 2Present study 1Symeonidis et al. (2006) |
Aliakmon | Pleistocene (imprecise) | Stephanorhinus sp.3 = Rhinocerotidae indet.2 = Dicerorhinus etruscus1 | 3Tsoukala (2018) 2Symeonidis et al. (2006) 1Psarianos (1958) |
Alikes (34782) | Early Pleistocene L. Villafranchian MNQ 19 | Rhinocerotidae indet.1,2,3 | 3Symeonidis et al. (2006) 2Athanassiou (1998) 1Athanassiou (1996) |
Allatini (182683) | Early Pliocene Ruscinian | Rhinocerotidae indet.2 = Rhinoceros sp.1 | 2Symeonidis et al. (2006) 1Marinos (1965) |
Angelochori (211694) | Late Pliocene E. Villafranchian MN16 | Stephanorhinus jeanvireti 1 | 1Tsoukala (2018) |
Apolakkia, Rhodes (34607) | Early Pliocene Ruscinian MN15 | Rhinocerotidae indet.1,2,3 | 3Symeonidis et al. (2006) 2Koufos (2006a) 1Van de Weerd et al. (1982) |
Apollonia (34784) | Early Pleistocene Epivillafranchian MNQ 19 1.3–1.0 Ma | Rhinocerotidae indet.1,2,3,4 | 4Symeonidis et al. (2006) 3Koufos (2001) 2Koufos and Kostopoulos (1997) 1Koufos et al. (1992) |
Asprochaliko (208564) | Late Pleistocene Aurelian MNQ 26 | Rhinocerotidae indet.2 = Dicerorhinus kirchbergensis1 | 2Present study 1Bailey et al. (1983) |
Gephyra | ?Pleistocene | Rhinocerotidae indet.3 = Coelodonta antiquitatis praecursor1,2 | 3Present study 2Tsoukala (1992) 1Tsoukala (1991a) |
Kalamakia (184245) | Late Pleistocene Aurelian MNQ 26 100–39 Ka | Stephanorhinus sp.4 = Rhinoceros sp.1,2,3 | 4Harvati et al. (2013) 3Gardeisen et al. (2001) 2Darlas and Lumley (2000) 1Gardeisen and Trantalidou (1998) |
Kalamoto 1 (200083) | Early Pleistocene Epivillafranchian MNQ 19 | Stephanorhinus etruscus2 = Dicerorhinus etruscus1 | 2Tsoukala (2018) 1Tsoukala and Chatzopoulou (2005) |
Kalamoto 2 (200084) | Early Pleistocene Epivillafranchian MNQ 19 | Stephanorhinus etruscus2 = Dicerorhinus etruscus1 | 2Tsoukala (2018) 1Tsoukala and Chatzopoulou (2005) |
Karnezeika (202122) | Early Pleistocene Villafranchian | Stephanorhinus sp.1 | 1Kokotini et al. (2019) |
Krimni (34762) | Early Pleistocene L. Villafranchian MNQ 18 | Stephanorhinus etruscus2,3,4 = Stephanorhinus cf. etruscus5 = Dicerorhinus etruscus1 | 5Symeonidis et al. (2006) 4Koufos (2001) 3Koufos and Kostopoulos (1997) 2Tsoukala (1992) 1Sakellariou-Mane et al. (1979) |
Kyparissia (194477) | Mid. Pleistocene Galerian MNQ 20-21 | Stephanorhinus sp.2 = Rhinocerotidae indet.1 | 2Athanassiou (2018b) 1Athanassiou (2016) |
Kythera Island | ?Pleistocene (imprecise) | Rhinocerotidae indet.1,2 | 2Symeonidis et al. (2006) 1Bartsiokas (1998) |
Lakonis I (207778) | Late Pleistocene Aurelian MNQ 26 100–40 Ka | Rhinocerotidae indet.1 | 1Panagopoulou et al. (2004) |
Livakos (34764) | Early Pleistocene L. Villafranchian MNQ 18 1.8–1.6 Ma | Stephanorhinus etruscus2,4 = Stephanorhinus cf. etruscus3 = Dicerorhinus etruscus1 = Dicerorhinus sp.1 = Dicerorhinus? sp.1 | 4Present study 3Symeonidis et al. (2006) 2Tsoukala (1992) 1Steensma (1988) |
Megalopolis, Marathousa (34809) | Mid. Pleistocene Galerian MNQ 20–21 | Rhinocerotidae indet.4 = Stephanorhinus etruscus1,2,3 | 4Symeonidis et al. (2006) 3Koufos and Kostopoulos (1997) 2Tsoukala (1992) 1Sickenberg (1975) |
Megalopolis (old col.) | Middle–Late Pleistocene (imprecise) | Stephanorhinus sp.6,10 = Stephanorhinus hemitoechus5,9 = Stephanorhinus kirchbergensis5,9 = Dicerorhinus hemitoechus3,4 = Dicerorhinus merckii 3,4 = Rhinoceros sp.2 = Rhinocerotidae indet.1 Non Coelodonta antiquitatis3–10 | 10Present study 9Tsoukala (2018) 8Iliopoulos et al. (2010) 7Lyras (2007) 6Symeonidis et al. (2006) 5Tsoukala (1992) 4Melentis (1965) 3Melentis (1964) 2Skouphos (1905) 1Bürchner (1903) |
Milia, Grevena (185859–62, 195290–99) | Late Pliocene E. Villafranchian MN16 | Stephanorhinus jeanvireti 4,5,6 = Dicerorhinus jeanvireti 3 = Dicerorhinus etruscus2 = Rhinocerotidae indet.1 | 6Present study 5Tsoukala (2018) 4Vlachos et al. (2018) 3Guérin and Tsoukala (2013) 2Tsoukala (2005) 1Tsoukala (2000) |
Molikrio | Early Pleistocene Villafranchian | Stephanorhinus cf. etruscus2,3,4 = Dicerorhinus cf. etruscus1 | 4Symeonidis et al. (2006) 3Koufos and Kostopoulos (1997) 2Tsoukala (1992) 1Symeonidis et al. (1986) |
Neapolis | Pleistocene (imprecise) | Rhinocerotidae indet.2 = Rhinoceros sp.1 | 2Symeonidis et al. (2006) 1Brunn (1956) |
Nigrita (202533) | Plio-Pleistocene Villafranchian (imprecise) | Rhinocerotidae indet.1 | 1Athanassiou (2017) |
Perdikas | Middle–Late Pleistocene | Rhinocerotidae indet.3 = Coelodonta antiquitatis1,2 | 3Present study 2Koufos and Pavlides (1988) 1Pavlides (1985) |
Penios (old col.) | Late Pleistocene Aurelian MNQ 26 | Stephanorhinus cf. hemitoechus2,4 = Dicerorhinus cf. hemitoechus1,3 | 4Symeonidis et al. (2006) 3Athanassiou (2002) 2Tsoukala (1992) 1Boessneck (1965) |
Penios riverbank (204397) | Late Pleistocene Aurelian MNQ 26 45–35 Ka | Stephanorhinus hemitoechus1 | 1Athanassiou (2011) |
Petralona Cave (183123) | Mid. Pleistocene Galerian110 | Stephanorhinus hemitoechus9,10,11,13,14 = Dicerorhinus hemitoechus7,8,12 = Dicerorhinus cf. hemitoechus5,6 = Stephanorhinus sp.4 = Didermoceros sp.3 = Didermoceros kirchbergensis2 = Rhinoceros mercki1 | 14Present study 13Tsoukala (2018) 12Tsoukala and Guérin (2016) 11Symeonidis et al. (2006) 10Fortelius et al. (1993) 9Tsoukala (1992) 8Tsoukala (1991a) 7Tsoukala (1989) 6Poulianos (1981) 5Fortelius and Poulianos (1979) 4Kretzoi (1977) 3Sickenberg (1971) 2Sickenberg (1964) 1Kanellis (1962) |
Platanochori (204658) | Early Pleistocene Epivillafranchian MNQ 19 | Stephanorhinus sp.2 = Stephanorhinus hundsheimensis1 | 2Present study 1Konidaris et al. (2015) |
Psychiko | Early Pleistocene (imprecise) | Stephanorhinus cf. etruscus2 = Rhinoceros cf. etruscus1 | 2Symeonidis et al. (2006) 1Paraskevaidis (1953) |
Richea (195310) | Early Pleistocene Villafranchian | Stephanorhinus etruscus2 = Stephanorhinus cf. etruscus1 | 2Present study 1Kampouridis et al. (2018) |
Saint George Priporos | Late Pliocene E. Villafranchian MN16 | Stephanorhinus jeanvireti 3 = Dicerorhinus jeanvireti 2 = Dicerorhinus etruscus1 | 3Present study 2Guérin and Tsoukala (2013) 1Tsoukala (2005) |
Serres Basin | Pleistocene (imprecise) | Rhinocerotidae indet.2 = Dicerorhinus cf. etruscus1 | 2Symeonidis et al. (2006) 1Marinos (1965) |
Sesklon (34614) | Early Pleistocene L. Villafranchian MN 17 | Stephanorhinus sp.2,3,4,6 = Stephanorhinus cf. etruscus5 = Dicerorhinus cf. etruscus1 | 6Athanassiou (2018a) 5Symeonidis et al. (2006) 4Koufos (2006a) 3Athanassiou (1998) 2Athanassiou (1996) 1Symeonidis (1992) |
Sesklon (lower) (205043) | Late Pliocene E. Villafranchian MN16 | Stephanorhinus sp.2 = Dicerorhinus cf. etruscus1 | 2Athanassiou (2018a) 1Symeonidis (1992) |
Tourkovounia 3-5 (34592) | Early Pleistocene M. Villafranchian MN 17 | Stephanorhinus cf. etruscus2-5 = Dicerorhinus cf. etruscus1 | 5Present study 4Symeonidis et al. (2006) 3Koufos (2006a) 2Tsoukala (1992) 1Symeonidis and de Vos (1976) |
Tsiotra Vrisi (197943) | Early Pleistocene L. Villafranchian | Stephanorhinus sp.1 | 1Konidaris et al. (2015) |
Vatera DS (183344) | Early Pleistocene M. Villafranchian MN 17 | Stephanorhinus cf. etruscus1,2,3 | 3Symeonidis et al. (2006) 2Koufos (2006a) 1de Vos et al. (2002) |
Volax (34593) | Early Pleistocene M. Villafranchian MN 17 | Rhinocerotidae indet.3,4,5 = Stephanorhinus cf. etruscus2 = Didermoceros sp.1 | 5Symeonidis et al. (2006) 4Koufos (2006a) 3Koufos and Kostopoulos (1997) 2Sickenberg (1975) 1Sickenberg (1968) |
Appendix 2
List of the Greek localities containing Miocene rhinocerotid fossils. Locality numbers refer to the collection numbers of the PaleoBiology Database (PBDB). Biochronological data (age) are primarily based on Koufos (2013, 2016b), as well as on evaluation of the biostratigraphical and faunal context of each locality. A detailed taxonomic history of the rhinocerotid record of each locality is provided in chronological order.
Localities | Age | Taxonomic history | Key references |
|---|---|---|---|
Asprogiannos 1 (205337) | Late Miocene Turolian MN 13 5.96–5.33 Ma | Rhinocerotidae indet.1,2 | 2Koufos and Vasileiadou (2015) 1Sylvestrou (2002) |
Atalanti | Middle Miocene Astaracian MN 7+8 | Aceratherium simorrense 3 = Aceratheriini indet.2 = Aceratherium sp.1 | 3Present study 2Giaourtsakis (2003) 1Symeonidis (1974) |
Chalkoutsi (205344) | Late Miocene Turolian | Rhinocerotidae indet.1,2 | 2Giaourtsakis (2003) 1Theodorou et al. (1995) |
Chrysavgi (205366) | Middle Miocene Astaracian MN 7 + 8 | Brachypotherium brachypus 6,7 = ?Brachypotherium sp.5 = Rhinocerotidae indet.3,4 = Dicerorhinus orientalis2 = Diceros pachygnathus1 | 7Present study 6Koufos and Kostopoulos (2013) 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Dimopoulos (1972) 1Psarianos (1958) |
Dytiko 1 (32374) | Late Miocene Turolian MN 13 6.81–5.96 Ma | Miodiceros neumayri 6 = Ceratotherium neumayri 3,4,5 = Dicerorhinus orientalis1,2 | 6Present study 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Bonis et al. (1988) 1Koufos (1980) |
Halmyropotamos (202213) | Late Miocene Turolian MN 12 | Dihoplus pikermiensis 5–8 = Dicerorhinus pikermiensis 3,4 = Dicerorhinus orientalis 1,2 | 8Present study 7Giaourtsakis et al. (2006) 6Koufos (2006a) 5Giaourtsakis (2003) 4Bonis and Koufos (1999) 3Geraads (1988) 2Melentis (1969) 1Melentis (1968b) |
Kerassia (Ke) (old collection) (195431) | Late Miocene Turolian MN 11–12 | Miodiceros neumayri 3 = “Diceros” neumayri 1,2 | 3Present study 2Giaourtsakis et al. (2020) 1Kampouridis et al. (2019) |
Kerassia 1 (K1) (195432) | Late Miocene Turolian MN 11–12 | Miodiceros neumayri 3 = “Diceros” neumayri 1,2 | 3Present study 2Giaourtsakis et al. (2020) 1Kampouridis et al. (2019) |
Kerassia 3 (K3) (195434) | Late Miocene Turolian MN 11–12 | Miodiceros neumayri 12 = “Diceros” neumayri 8,9,10,11 = Ceratotherium neumayri 3,5,6,7 = Rhinocerotidae indet.1,2,4 | 12Present study 11Giaourtsakis et al. (2020) 10Kampouridis et al. (2019) 9Athanassiou et al. (2014) 8Giaourtsakis (2009) 7Giaourtsakis et al. (2006) 6Koufos (2006a) 5Theodorou et al. (2003) 4Giaourtsakis (2003) 3Theodorou et al. (2001) 2Theodorou et al. (1998) 1Theodorou et al. (1995) |
Kerassia 4 (K4) (195435) | Late Miocene Turolian MN 11–12 | Miodiceros neumayri 12 = “Diceros” neumayri 8,9,10,11 = Ceratotherium neumayri 3,5,6,7 Dihoplus pikermiensis 7-12 = Dihoplus cf. pikermiensis 6 = “Dicerorhinus” cf. pikermiensis 5 = Dicerorhinus pikermiensis 3 = Rhinocerotidae indet.1,2 Acerorhinus neleus 9,10,11,12 = Rhinocerotidae n.sp.5,6 = Rhinocerotidae indet.2,3 | 12Present study 11Giaourtsakis et al. (2020) 10Kampouridis et al. (2019) 9Athanassiou et al. (2014) 8Giaourtsakis (2009) 7Giaourtsakis et al. (2006) 6Koufos (2006a) 5Theodorou et al. (2003) 4Giaourtsakis (2003) 3Theodorou et al. (2001) 2Theodorou et al. (1998) 1Theodorou et al. (1995) |
Kerassia 5 (K5) (195436) | Late Miocene Turolian MN 11–12 | Miodiceros neumayri 3,4,5 = Dihoplus cf. pikermiensis 2 = “Dicerorhinus” cf. pikermiensis1 | 5Present study 4Giaourtsakis et al. (2020) 3Kampouridis et al. (2019) 2Koufos (2006) 1Theodorou et al. (2003) |
Maramena (32189) | Mio-Pliocene boundary MN 13–14 5.40–5.23 Ma | Rhinocerotidae indet.3-7 = Diceros neumayri 2 = Diceros neymari (sic)1 | 7Present study 6Koufos (2006a) 5Giaourtsakis (2003) 4Bonis and Koufos (1999) 3Schmidt-Kittler et al. (1995) 2Meulen and Kolfschoten (1986) 1Karistineos (1984) |
Nea Silata (191612) | Mio-Pliocene boundary MN 13–14 5.40–5.23 Ma | Rhinocerotidae indet.1,2,3,4 | 4Koufos and Vasileiadou (2015) 3Koufos (2006c) 2Giaourtsakis (2003) 1Syrides (1990) |
Nikiti 1 (202729) | Late Miocene Vallesian MN 10 9.3–8.7 Ma | Miodiceros neumayri 6 = “Diceros” neumayri 5 = Rhinocerotidae indet.1,2,3,4 Dihoplus pikermiensis 5,6 | 6Present study 5Koufos et al. (2016) 4Koufos (2006a) 3Giaourtsakis (2003) 2Bonis and Koufos (1999) 1Kostopoulos (1994) |
Nikiti 2 (73869) | Late Miocene Turolian MN 11 8.7–8.3 Ma | Miodiceros neumayri 3 = “Diceros” neumayri 2 = Rhinocerotidae indet.1 | 3Present study 2Koufos (2016a) 1Kostopoulos et al. (1996) |
Pentalophos 1 (202119) | Late Miocene Vallesian MN 9–10 | Miodiceros neumayri 10 = “Diceros” neumayri 7,9 = Ceratotherium neumayri 1–6,8 Acerorhinus sp.7,8,9,10 = Acerorhinus cf. zernowi 6 = Acerorhinus zernowi 4 =p. Aceratherium kiliasi 1,2,3 Chilotherium samium 10 = Chilotherium cf. samium 9 = Chilotherium (Eochilotherium) kiliasi 8 = Chilotherium sp.7 = Chilotherium kiliasi 2,4,6 =p. Aceratherium kiliasi 1,3,5 | 10Present study 9Athanassiou et al. (2014) 8Geraads and Spassov (2009) 7Giaourtsakis (2009) 6Koufos (2006a) 5Giaourtsakis (2003) 4Heissig (1999) 3Bonis and Koufos (1999) 2Heissig (1996) 1Geraads and Koufos (1990) |
Perivolaki (194879) | Late Miocene Turolian MN12 7.3–7.1 Ma | Miodiceros neumayri 3 = Ceratotherium neumayri 2 = Ceratotherium sp.1 | 3Present study 2Koufos (2006d) 1Koufos (2006a) |
Pikermi (182754) Classic collections: MNHN, BSPG, AMPG, BMNH, NHMW, IPUW, SMNS, MNHB, STIBP, NRM, USNM, YPM, UCMP | Late Miocene Turolian MN 12 7.33–7.29 Ma | Miodiceros neumayri 33 = “Diceros” neumayri 29–32 = Ceratotherium neumayri 23–28 = Diceros neumayri 20 = Diceros pachygnathus pachygnathus 19 = Pliodiceros pachygnathus 17 = Diceros pachygnathus 15,16,21,22 = Atelodus pachygnathus 11 = Rhinoceros (Atelodus) pachygnathus 10 = Atelodus (Colodus) pachygnathus 9 =p. Rhinoceros schleiermacheri 7 =p. Rhinoceros pachygnathus 7 = Rhinoceros pachygnathus 6,8,12 Non Chalicotherium (Colodus) pachygnathus 5 non Rhinoceros pachygnathus 4 = Rhinoceros schleiermacheri 2 Non Rhinoceros pachygnathus 2 | 33Present study 32Giaourtsakis et al. (2018) 31Athanassiou et al. (2014) 30Giaourtsakis (2009) 29Giaourtsakis et al. (2009) 28Koufos (2006a) 27Giaourtsakis et al. (2006) 26Giaourtsakis (2003) 25Heissig (1999) 24Heissig (1996) 23Geraads (1988) 22Solounias (1981) 21Guérin (1980) 20Heissig (1975) 19Thenius (1955) 18Kretzoi (1942b) 17Kretzoi (1942a) 16Ringström (1924) (continued below) |
Pikermi (continued) | Dihoplus pikermiensis 26–33 = Stephanorhinus pikermiensis24,25 = Dicerorhinus schleiermacheri 22 = Stephanorhinus pachygnathus 20 = Dicerorhinus pikermiensis17,18,23 = Dicerorhinus orientalis 16,21 = Ceratorhinus aff. schleiermacheri15 = Rhinoceros (Ceratorhinus) schleiermacheri orientalis 14 = Rhinoceros schleiermacheri pikermiensis 13 = Ceratorhinus schleiermacheri11 = Rhinoceros (Dihoplus) schleiermacheri 10 = Dihoplus schleiermacheri 9 =p. Rhinoceros pachygnathus 7 =p. Rhinoceros schleiermacheri 7 = Rhinoceros schleiermacheri 3,4,6,8,12 = Rhinoceros pachygnathus 2 = Rhinoceros sp.1 Acerorhinus neleus 31,32,33 = Acerorhinus sp.30 = Aceratherium sp.28 = Aceratheriini indet.26 = Aceratherium cf. incisivum 22 = Aceratherium incisivum 10 = Acerotherium sp.7 | 15Abel (1922) 14Schlosser (1921) 13Toula (1906) 12Weber (1904) 11Osborn (1900) 10Zittel (1893) 9Brandt (1878) 8Weithofer (1888) 7Gaudry (1862–1867) 6Gaudry (1864) 5Wagner (1861) 4Wagner (1857) 3Roth and Wagner (1854) 2Wagner (1848) 1Goldfuss (1841) | |
Pikermi Chomateri (195562) | Late Miocene Turolian MN 12 7.17–7.11 Ma | Dihoplus pikermiensis 7 Acerorhinus neleus 6,7 = Aceratheriini indet.4 = Diceros aff. pachygnathus 2 = Aceratherium sp.1,2,3,5 | 7Present study 6Athanassiou et al. (2014) 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Marinos and Symeonidis (1974) 1Marinos and Symeonidis (1973) |
Pikermi Valley-1 (202630) | Late Miocene Turolian MN 12 ~7.27 Ma | Miodiceros neumayri 3 = “Diceros” neumayri 2 Dihoplus pikermiensis 2,3 = Rhinocerotidae indet.1 Acerorhinus neleus 3 | 3Present study 2Filis et al. (2019) 1Roussiakis et al. (2014) |
Pikermi Valley-3 (202631) | Late Miocene Turolian MN 12 7.33–7.29 Ma | Miodiceros neumayri 2 Dihoplus pikermiensis 1,2 Acerorhinus neleus 2 | 2Present study 1Roussiakis et al. (2014) |
Platania (182682) | Late Miocene Turolian MN12 | Miodiceros neumayri 3 = Ceratotherium neumayri 2 = Rhinocerotidae indet.1 | 3Present study 2Tsoukala (2018) 1Vlachos and Tsoukala (2014) |
Prokopi (=Achmet Aga) Euboea (214079) | Late Miocene Turolian | Rhinocerotidae indet.1,2,3 | 3Koufos (2006a) 2Giaourtsakis (2003) 1Woodward (1901) |
Pyrgos Vassilissis (=Tour la Reine) (195555) | Late Miocene Turolian MN 12 7.18–7.17 Ma | Miodiceros neumayri 7 = ?Ceratotherium neumayri 5,6 = Rhinocerotidae indet.4 = Stephanorhinus pachygnathus2 = Dicerorhinus ?orientalis 1,3 | 7Present study 6Böhme et al. (2017) 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Paraskevaidis (1977) 1Freyberg (1951) |
Ravin de la Pluie (191070) | Late Miocene Vallesian MN10 ~9.3 Ma | Rhinocerotidae indet.1,2,3,4 | 4Koufos (2006a) 3Giaourtsakis (2003) 2Bonis and Koufos (1999) 1Bonis et al. (1988) |
Ravin des Zouaves 5 (195489) | Late Miocene Turolian MN 11 ~8.2 Ma | Miodiceros neumayri 6 = Ceratotherium neumayri 3,4,5 = Dicerorhinus orientalis 1,2 | 6Present study 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Bonis et al. (1988) 1Koufos (1980) |
Ravin X (182745) | Late Miocene Turolian MN 12 | Miodiceros neumayri 5 = Ceratotherium neumayri 2,3,4 = Rhinoceros pachygnathus1 | 5Present study 4Koufos (2006a) 3Giaourtsakis (2003) 2Bonis and Koufos (1999) 1Arambourg and Piveteau (1929) |
Samos (various localities and collections) | Late Miocene Turolian MN11–13 | Miodiceros neumayri 1 Dihoplus pikermiensis1 Chilotherium schlosseri 1 Chilotherium samium1 | 1Present study (see separate Appendix 3) |
Servia (202257) | Late Miocene | Rhinocerotidae indet.2,3 = Rhinoceros sp.1 | 3Koufos (2006a) 2Giaourtsakis (2003) 1Paraskevaidis (1977) |
Tanagra | Late Miocene Turolian | Rhinocerotidae indet.3,4,5 = Rhinocerotidarum gen. indet.1,2 | 5Koufos (2006a) 4Giaourtsakis (2003) 3Bonis and Koufos (1999) 2Mitzopoulos (1961) 1Mitzopoulos (1960) |
Thermopigi (73553) | Late Miocene Turolian MN 12 | Dihoplus pikermiensis 1,2 | 2Tsoukala (2018) 1Geraads et al. (2007) |
Vathylakkos 2 (202703) | Late Miocene Turolian MN 12 ~7.3 Ma | Rhinocerotidae indet.1,2,3 | 3Koufos (2006a) 2Giaourtsakis (2003) 1Bonis and Koufos (1999) |
Vathylakkos 3 (182750) | Late Miocene Turolian MN 12 ~7.3 Ma | Miodiceros neumayri 10 = Stephanorhinus pikermiensis 7 = Ceratotherium neumayri 5,6,8,9 = Dicerorhinus orientalis 3,4 = Diceros pachygnathus pachygnathus 2 = Rhinoceros pachygnathus 1 Aceratheriini indet.8,10 = Rhinocerotidae indet.6 = Aceratherium sp.1 | 10Present study 9Koufos (2006a) 8Giaourtsakis (2003) 7Heissig (1999) 6Bonis and Koufos (1999) 5Geraads (1988) 4Bonis et al. (1988) 3Koufos (1980) 2Thenius (1955) 1Arambourg and Piveteau (1929) |
Xirochori (195490) | Late Miocene Vallesian MN 10 9.64–9.58 Ma | Rhinocerotidae indet.1,2,3 | 3Koufos (2006a) 2Giaourtsakis (2003) 1Bonis and Koufos (1999) |
Appendix 3
List of the various collections of rhinocerotid fossils on Samos Island. Some records correspond to precise and biochronologically well-calibrated points of collections (localities). Other records refer to old fossil collections housed in various institutions, whose exact point of excavation may have not been documented. Locality numbers refer to the collection numbers of the PaleoBiology Database (PBDB). Biochronological data (age) are based on Koufos et al. (2009b) and Koufos (2013, 2016b). A detailed taxonomic history of the rhinocerotid record of each collection is provided in chronological order.
Localities | Age | Taxonomic history | Key references |
|---|---|---|---|
Samos BSPG col. (207137) (horned taxa) | Late Miocene Turolian | Miodiceros neumayri 23 = “Diceros” neumayri 21,22 = Ceratotherium neumayri 14–20 = Diceros neumayri 11 = Diceros pachygnathus pachygnathus10 = Pliodiceros pachygnathus8 = Diceros pachygnathus7,12,13 = Rhinoceros pachygnathus3,5,6 =p. Rhinoceros pachygnathus1 Dihoplus pikermiensis19–23 = Stephanorhinus pikermiensis17,18 = “Dicerorhinus” schleiermacheri16 = Dicerorhinus schleiermacheri 13 = Stephanorhinus pachygnathus11 = Dicerorhinus pikermiensis8,9,14,15 = Dicerorhinus orientalis7,12 = Rhinoceros (Ceratorhinus) schleiermacheri orientalis5 = Rhinoceros schleiermacheri samius4 = Rhinoceros aff schleiermacheri3,6 =p. Rhinoceros pachygnathus1 =p. Rhinoceros schleiermacheri 1 | 23Present study 22Giaourtsakis (2009) 21Giaourtsakis et al. (2009) 20Giaourtsakis et al. (2006) 19Giaourtsakis (2003) 18Heissig (1999) 17Heissig (1996) 16Bernor et al. (1996) 15Geraads and Koufos (1990) 14Geraads (1988) 13Solounias (1981) 12Guérin (1980) 11Heissig (1975) 10Thenius (1955) 9Kretzoi (1942b) 8Kretzoi (1942a) 7Ringström (1924) 6Andrée (1921) 5Schlosser (1921) 4Toula (1906) 3Weber (1907) 2Weber (1905) 1Weber (1904) |
Samos BSPG col. (207137) (hornless taxa) | Late Miocene Turolian | Chilotherium schlosseri 6,7,9,12–14,16,17 = Chilotherium (Chilotherium) schlosseri 8.11,15 = Aceratherium schlosseri 1–5 Chilotherium samium6,7,9,12–14,16,17 = Chilotherium (Eochilotherium) samium15 = Chilotherium (Chilotherium) samium8,11 = Aceratherium sp.? 2,3 = Aceratherium samium1,2,3,5,10 | 17Present study 16Giaourtsakis (2009) 15Geraads and Spassov (2009) 14Giaourtsakis (2003) 13Heissig (1999) 12Bernor et al. (1996) 11Heissig (1996) 10Geraads and Koufos (1990) 9Solounias (1981) 8Heissig (1975) 7Borsuk-Bialynicka (1969) 6Ringström (1924) 5Andrée (1921) 4Krokos (1917) 3Kiernik (1914) 2Weber (1907) 1Weber (1905) |
Samos GPMM col. (207193) | Late Miocene Turolian | Miodiceros neumayri 12 = "Diceros" neumayri 11 = Ceratotherium neumayri 6,9,10 = Diceros neumayri 6 = Diceros pachygnathus4,7 = Rhinoceros pachygnathus1 Dihoplus pikermiensis10,11,12 = Stephanorhinus pikermiensis9 = Dicerorhinus pikermiensis8 = Stephanorhinus pachygnathus6 = Dicerorhinus orientalis4 = Rhinoceros schleiermacheri 1 Chilotherium schlosseri 4,7,9,10,11,12 = Chilotherium kowalevskii 9,10 = Chilotherium(Chilotherium) schlosseri 6 = Chilotherium (Chilotherium) kowalevskii 6 = Chilotherium angustifrons4 = Chilotherium wegneri 4 = Aceratherium habereri2,3 = Aceratherium schlosseri 1 = Aceratherium angustifrons1 = Aceratherium wegneri 1,5 | 12Present study 11Giaourtsakis (2009) 10Giaourtsakis (2003) 9Heissig (1999) 8Geraads (1988) 7Solounias (1981) 6Heissig (1975) 5Leonardi (1947) 4Ringström (1924) 3Killgus (1922b) 2Killgus (1922a) 1Andrée (1921) |
Samos Mytilinii-1A NHMA col. (202215) | Late Miocene Turolian MN 12 ~7.1 Ma | Miodiceros neumayri 4 = “Diceros” neumayri 3 Dihoplus pikermiensis2,3,4 = Dicerorhinus pikermiensis1 | 4Present study 3Giaourtsakis (2009) 2Koufos (2006a) 1Koufos et al. (1997) |
Samos Mytilinii-1B NHMA col. (202216) | Late Miocene Turolian MN 12 ~7.1 Ma | Miodiceros neumayri 4 = “Diceros” neumayri 3 = Dihoplus cf. pikermiensis2 = Dicerorhinus cf. pikermiensis1 | 4Present study 3Giaourtsakis (2009) 2Koufos (2006a) 1Koufos et al. (1997) |
Samos Mytilinii-1C NHMA col. (202217) | Late Miocene Turolian MN 12 ~7.1 Ma | ?Rhinocerotidae indet.3 = Rhinocerotidae indet2 = Dicerorhinus sp.1 | 3Present study 2Koufos (2006a) 1Koufos et al. (1997) |
Samos Mytilinii-1D NHMA col. (202220) | Late Miocene Turolian MN 12 ~7.1 Ma | Miodiceros neumayri 2 = “Diceros” neumayri 1 | 2Present study 1Giaourtsakis (2009) |
Samos Mytilinii-3 NHMA col. (202218) | Late Miocene Turolian MN 12 ~7.3 Ma | Miodiceros neumayri 3 = “Diceros” neumayri 2 Dihoplus pikermiensis2,3 = Rhinocerotidae indet.1 | 3Present study 2Giaourtsakis (2009) 1Koufos (2006a) |
Samos Mytilinii-4 NHMA col. (202219) | Late Miocene Turolian MN 12 ~7.5 Ma | Miodiceros neumayri 3 = “Diceros” neumayri 2 = Rhinocerotidae indet.1 | 3Present study 2Giaourtsakis (2009) 1Koufos (2006a) |
Samos (PMMS) NHMA col. | Late Miocene Turolian MN 12 ~7.1 Ma | Miodiceros neumayri 3 = “Diceros” neumayri 2 = Dicerorhinus orientalis1 | 3Present study 2Giaourtsakis (2009) 1Melentis (1968a) |
Samos Q1 AMNH col. (95691) | Late Miocene Turolian MN 12 ~7.1 Ma | Miodiceros neumayri 2 = Diceros pachygnathus1 ?Dihoplus pikermiensis2 = Dicerorhinus schleiermacheri1 Chilotherium sp.2 = Chilotherium samium1 | 2Present study 1Solounias (1981) |
Samos Q2 AMNH col. (206460) | Late Miocene Turolian MN 12 ~7.5 Ma | Chilotherium sp.2 = Chilotherium samium1 | 2Present study 1Solounias (1981) |
Samos Q4 AMNH col. (95689) | Late Miocene Turolian MN 11 8.0–7.6 Ma | Miodiceros neumayri 2 = Diceros pachygnathus1 Chilotherium sp.2 = Chilotherium samium1 | 2Present study 1Solounias (1981) |
Samos Q5 AMNH col. (95690) | Late Miocene Turolian MN 13 6.9–6.7 Ma | Miodiceros neumayri 2 = Diceros pachygnathus1 ?Dihoplus pikermiensis2 = Dicerorhinus schleiermacheri 1 | 3Present study 2Koufos (2006a) 1Solounias (1981) |
Samos S4 CM col. (205353) | Late Miocene Turolian MN 12 ~7.1 Ma | Chilotherium sp.3 = Chilotherium samium1,2 | 3Present study 2Solounias (1981) 1Black et al. (1980) |
Samos CM col. | Late Miocene Turolian | Miodiceros neumayri 1 Chilotherium sp.1 | 1Present study |
Samos UCMP col. | Late Miocene Turolian | Rhinocerotidae indet.1 | 1Present study |
Samos SMF col. (202725) | Late Miocene Turolian | Miodiceros neumayri 6 = “Diceros” neumayri 5 = Ceratotherium neumayri 3 = Atelodus pachygnathus1 Dihoplus pikermiensis6 Chilotherium sp.5,6 = Chilotherium (Eochilotherium) samium4 = Chilotherium schlosseri 1,2 | 6Present study 5Giaourtsakis (2009) 4Geraads and Spassov (2009) 3Giaourtsakis et al. (2006) 2Heissig (1999) 1Drevermann (1930) |
Samos HLMD col. (202724) | Late Miocene Turolian | Miodiceros neumayri 4 = “Diceros” neumayri 3 = Ceratotherium neumayri 2 Dihoplus pikermiensis4 Chilotherium schlosseri 1,4 | 4Present study 3Giaourtsakis (2009) 2Giaourtsakis et al. (2006) 1Heissig (1999) |
Samos SMNS col. | Late Miocene Turolian | Miodiceros neumayri 3 = “Diceros” neumayri 2 = Ceratotherium neumayri 1 Chilotherium sp.2,3 | 3Present study 2Giaourtsakis (2009) 1Geraads (1988) |
Samos GPIH col. (43007) | Late Miocene Turolian | Miodiceros neumayri 6 = “Diceros” neumayri 5 = Ceratotherium neumayri 3,4 = Diceros neumayri 2 Chilotherium schlosseri 2–6 = Aceratherium sp.1 | 6Present study 5Giaourtsakis (2009) 4Giaourtsakis (2003) 3Heissig (1999) 2Lehmann (1984) 1Gürich (1912) |
Samos NHMW col. (182751) | Late Miocene Turolian | Miodiceros neumayri 3 = “Diceros” neumayri 2 = Ceratotherium neumayri 1 Dihoplus pikermiensis1,2,3 Chilotherium schlosseri 3 = Chilotherium sp.2 | 3Present study 2Giaourtsakis (2009) 1Giaourtsakis et al. (2006) |
Samos IPUW col. | Late Miocene Turolian | Miodiceros neumayri 1 | 1Present study |
Samos MCGL col. (202120) | Late Miocene Turolian | Miodiceros neumayri 7 = “Diceros” neumayri 6 = Ceratotherium neumayri 5 = Diceros pachygnathus4 = Rhinoceros pachygnathus1,2,3 Dihoplus pikermiensis6,7 = Dicerorhinus schleiermacheri 4 = Rhinoceros schleiermacheri 3 =?Rhinoceros schleiermacheri 2 | 7Present study 6Giaourtsakis (2009) 5Heissig (1999) 4Solounias (1981) 3Weber (1904) 2Forsyth-Major (1894) 1Forsyth-Major (1888) |
Samos NHMB col. | Late Miocene Turolian | Miodiceros neumayri 3 = Ceratotherium neumayri 2 = Diceros pachygnathus1 Chilotherium sp.3 | 3Present study 2Geraads (1988) 1Viret (1958) |
Samos NMBE col. | Late Miocene Turolian | Miodiceros neumayri 2 = Rhinoceros (Atelodus) pachygnathus1 Dihoplus pikermiensis2 = Rhinoceros (Ceratorhinus) schleiermacheri 1 | 2Present study 1Studer (1911) |
Samos AMPG col. (208570) | Late Miocene Turolian | Miodiceros neumayri 3 = “Diceros” neumayri 1,2 Dihoplus pikermiensis1,2,3 Chilotherium sp.1,2,3 | 3Present study 2Svorligkou et al. (2019) 1Giaourtsakis (2009) |
Samos MNHN col. | Late Miocene Turolian | Miodiceros neumayri2 = “Diceros” neumayri1 | 2Present study 1Giaourtsakis (2009) |
Samos BMNH col. | Late Miocene Turolian | Miodiceros neumayri 2 = “Diceros” neumayri 1 Chilotherium sp.1,2 | 2Present study 1Giaourtsakis (2009) |
Samos MAFI col. | Late Miocene Turolian | Miodiceros neumayri 2 =p. Rhinoceros sp.1 Dihoplus pikermiensis2 =p. Rhinoceros sp.1 | 2Present study 1Kormos (1913) |
Samos MGPP col. | Late Miocene Turolian | Chilotherium schlosseri 3,4 = Chilotherium wegneri 1,2 | 4Present study 3Giaourtsakis (2009) 2Piccoli et al. (1975) 1Leonardi (1947) |
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Giaourtsakis, I.X. (2022). The Fossil Record of Rhinocerotids (Mammalia: Perissodactyla: Rhinocerotidae) in Greece. In: Vlachos, E. (eds) Fossil Vertebrates of Greece Vol. 2. Springer, Cham. https://doi.org/10.1007/978-3-030-68442-6_14
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