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Oligocene ruminants from the Kızılırmak Formation, Çankırı-Çorum Basin, Central Anatolia, Turkey

Grégoire Métais, Ebru Albayrak, Pierre-Olivier Antoine, Ozan Erdal, Levent Karadenizli, Neşe Oyal, Gerçek Saraç, Yeşim Büyükmeriç, and Sevket Sen

Plain Language Abstract

The Oligocene is key interval for the origin and early diversification of modern groups of Ruminantia. This paper provides new data from five localities of the Oligocene Kızılırmak Formation in Central Anatolia. The fauna shows taxonomic affinities with both European and Asian ruminants faunas. The tragulids are represented by Iberomeryx parvus, a species so far exclusively known from the Late Oligocene of Georgia, and another large and unnamed species of Iberomeryx sp. The deciduous dentition and postcranial elements of Iberomeryx are described for the first time. A large lophiomerycid have been recognized as well as the stem pecoran Dremotherium guthi, cf. Palaeohypsodontus, and a large indeterminate Pecora. The occurrence of Dremotherium guthi predates its first occurrence in Europe, and the occurrence of Palaeohypsodontus in Central Anatolia significantly expands its geographical range, and suggest biogeographical affinities with Central Asia. In the five localities, the majority of the ruminant material is referred to Iberomeryx parvus, but the samples from the most recent localities of the Kızılırmak Formation display Pecoran species that are either absent or poorly represented in the oldest locality of the Kızılırmak Formation. Although incomplete, these data may reflect a drastic change in the ruminant faunas perhaps related to the Late Oligocene Warming event that strongly affected European faunas during the MP28 interval.

Resumen en Español

Rumiantes oligocenos de la Formación Kızılırmak, Cuenca Çankırı-Çorum, Anatolia Central, Turquía

Se describe una nueva asociación de rumiantes de 5 localidades oligocenas de la Formación Kızılırmak, Anatolia Central, Turquía. Se han reconocido los tragúlidos Iberomeryx parvus e Iberomeryx sp. (grande), así como también un probable lofiomerícido grande. El pecoro basal Dremotherium guthi, cf. Palaeohypsodontus y un Pecora grande indeterminado también han sido identificados. En las 5 localidades, la mayoría de los materiales de rumiantes se refieren a Iberomeryx parvus, pero la muestra de la localidad Tepe 641 (miembro superior de la Formación Kızılırmak) muestra algunos caracteres distintivos sugiriendo una especie/forma más derivada que probablemente vivía en ambientes más abiertos. Los taxones de rumiantes registrados en la Formación Kızılırmak son congruentes con una edad oligocena tardía, probablemente cercana en edad a la fauna Benara del sur de Georgia. La posible ocurrencia de Palaeohypsodontus en Anatolia Central expandiría significativamente su rango geográfico y sugeriría afinidades biogeográficas con Asia Central. La fauna de rumiantes de la Formación Kızılırmak sugiere la existencia de bosques de tierras bajas con paisajes más abiertos en Anatolia Central durante el Oligoceno tardío.
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Palabras clave: Ruminantia; Oligoceno; traguloideos; Pecora; Formación Kızılırmak; Turquía.

Traducción: Diana Elizabeth Fernández

Résumé en Français

LES RUMINANTS OLIGOCENE DE LA KIZILIRMAK FORMATION, BASSIN DE ÇANKIRI ÇORUM, ANATOLIE CENTRALE, TURQUIE

Les ruminants provenant de cinq localités fossilifères de la formation Kızılırmak en Anatolie Centrale (Turquie) sont décrits. Les tragulidés Iberomeryx parvus et Iberomeryx sp. (grande forme), un grand lophiomerycidé, le pecora basal Dremotherium guthi, Palaeohypsodontus, et un grand pecora indéterminé ont été identifiés. Dans les cinq localités, la majorité du matériel appartient à Iberomeryx parvus, mais le materiel provenant de la localité Tepe 641 (membre supérieur de la formation Kızılırmak) porte des caractères distinctifs qui suggèrent l'existance d'une autre espèce/forme plus dérivée qui vivait probablement dans des environnements plus ouverts. Les ruminants de la formation Kızılırmak confirment un âge Oligocène supérieur probablement proche de celui de la faune de Benara, située au sud de la Géorgie. La présence de Palaeohypsodontus en Anatolie centrale étend sa répartition géographique et suggère des affinités biogéographiques avec l'Asie centrale. La faune de ruminants de la formation Kızılırmak suggère la présence de forêts de plaines avec des paysages ouverts en Anatolie centrale durant l'Oligocène supérieur.

mots clés: Ruminantia; Oligocène; traguloids; Pecora; Formation Kızılırmak ; Turquie

Translator: Author

Deutsche Zusammenfassung

Oligozäne Ruminanten aus der Kızılırmak Formation, Çankırı-Çorum Becken, Zentral-Anatolien, Türkei

Es wird eine neue Ruminanten-Assemblage aus fünf verschiedenen oligozänen Fundstellen der Kızılırmak Formation aus Zentral-Anatolien, Türkei, beschrieben. Es wurden die Traguliden Iberomeryx parvus und Iberomeryx sp. (groß), ebenso wie eine möglicherweise große Lophiomerycide erkannt. Der Stamm-Stirnwaffenträger Dremotherium guthi, cf. Palaeohypsodontus und ein großer unbestimmter Pecora wurden ebenso identifiziert. An allen fünf Lokalitäten kann der Großteil des Ruminanten-Materials Iberomeryx parvus zugerechnet werden. Die Probe aus der Lokalität Tepe 641 (oberes Mitglied der Kızılırmak Formation) zeigt bestimmte Merkmale, die auf eine abgeleitete Art/Form hinweisen, die möglicherweise in einer offeneren Umgebung lebte. Die Ruminanten-Taxa aus der Kızılırmak Formation stimmen mit einem spätoligozänen Alter überein, möglicherweise nahe mit der Bengara Fauna aus dem südlichen Georgien. Das mögliche Auftreten von Palaeohypsodontus in Zentral-Anatolien würde seine geografische Reichweite signifikant erweitern und legt eine biogeografische Nähe mit Zentralasien nahe. Die Ruminanten-Fauna der Kızılırmak Formation weist auf Tieflandwälder mit mehreren offenen Landschaften in Zentralasien während des späten Oligozäns hin.

Schlüsselwörter: Ruminantia; Oligozän; Traguloide; Pecora; Kızılırmak Formation; Türkei,

Translator: Eva Gebauer

Arabic

in progress

Translator: Ashraf M.T. Elewa

 

 

TABLE 1. Measurements (in mm) of the dental material of taxa discussed in the text from the different fossil localities of the Kızılırmak. The values in brackets designate the width of the trigonid of lower molars, the non-bracketed values the maximal width. The asterisk designates estimated values when material is slightly broken or heavily worn/digested.

Specimen Taxon Tooth Length Width
GK2-11 Iberomeryx parvus d3 4.8 1.3
GK2-11 I. parvus d4 4.7 2.3
GK3-6 I. parvus i2-3? 3.2 2.5
GK3-7 I. parvus i1 2.4 2.9
GK3-11 I. parvus p3 3 1.2
GK3-36 I. parvus p4 4.4 1.9
GK3-8 I. parvus m1 5.3 2,8 (2,2)
641-5 I. parvus m1 5.5 2,9 (2,6)
641-4 I. parvus m1 5.4 3,2 (2,5)
GK3-3 I. parvus m1 5.6 3,1 (2,6)
GK3-42 I. parvus m2 6.1 3,4 (2,9)
641-7 I. parvus m2 6.3 3,4 (2,6)
641-3 I. parvus m2 6.8 3,7 (3)
GK3-34 I. parvus m2 7.2 4,5 (3,4)
GK3-33 I. parvus m2 6 3,5 (2,9)
641-6 I. parvus m2 7.1 4 (3,1)
641-2 I. parvus m3 7.8 3,4 (2,9)
GK3-2 I. parvus m3 7.9 3,1 (2,7)
GK2-4 I. parvus D2 5.1 2.1
GK2-4 I. parvus D3 5.2 2.8
GK2-4 I. parvus D4 4.5 4.4
GK2-4 I. parvus M1 5.3 5.2
GK3-4 I. parvus P2 6.3 3.7
GK3-5 I. parvus P4 3.8 4.4
KZ-7 I. parvus M1 5.4 5.1
641-13 I. parvus M1 5.3 5.5
GK3-31 I. parvus M2 6.4 6.8
641-12 I. parvus M2 6.3 6.2
641-14 I. parvus M2? 6.7 6.4
GK3-35 I. sp. p4 ? 3.2
641-9 lophiomerycid i ndet. p1? 7.3 2.9
BA-3 Dremotherium guthi m1 10.1 7.9
BA-6 D. guthi M2? 10.4 ?
641-10 cf. Palaeohypsodontus sp. M2? 9,5* ?

  

TABLE 2. Measurements (in mm) of the postcranial material of taxa discussed in the text from the different fossil localities of the Kızılırmak Formation. The asterisk designates estimated values when material is slightly broken or heavily worn/digested. Abbreviations: prox for proximal.

humerus (distal)
    width lateral depth medial depth
GK3-21 Iberomeryx parvus 10.3 4.2 6.7
GK3-22 I. parvus 10 4.7 7.1
         
metacarpal III+IV (distal)
    width depth  
GK3-54 I. parvus 8.9 ?  
         
astragali
    length prox width distal width
GK3-18 I.sp. 16.1 9,5* 8.7
641-23 I.sp. 15.9 9,0* 8,5*
641-24 I.sp. 16.6 9,6* 9*
641-25 I.sp. 15,5* 9,0* 8.8
GK3-25 I. parvus 12* 7,5* 6,7*
GK3-37 I. parvus 13.1 7,0* 6.5
GK3-38 I. parvus ? 5.7 ?
GK3-39 I. parvus ? 6* ?
641-21 I. parvus ? ? 5,5*
641-22 I. parvus 13.1 7 6.4
BA-2 lophiomerycid indet. 27* ? 15*
KZ-9 Pecora i ndet. 30.7 19.6 19.1
         
cubonavicular
    width depth max height
GK2-5 I. parvus 9.2 8,0* 8,5*
641-15 I. parvus 10.8 9.8 10
641-16 I. parvus 7.6 6.2 8.9
GK2-3 lophiomerycid indet. 17.7 ? 16.7
         
calcanei
    length prox width distal width
GK2-7 lophiomerycid indet. ? 15* ?
BA-3 lophiomerycid indet. ? 14* 11*
KZ-10 lophiomerycid indet. ? 15* ?
GK3-19 I. parvus ? 7* ?
GK3-23 I. parvus 21* 7,5* 5.5
GK3-24 I. parvus 18* ? 4*
641-17 I. parvus 17,5* ? 4.4
641-19 I. parvus 18* 7* 4.5
KZ-11 I. sp. ? 11* ?
         
phalanx prox
    length prox width distal width
GK3-13 I. parvus 8.7 3.2 2.4
GK3-46 I. parvus ? ? 3.3
KZ-4 I. parvus ? ? 3.1
KZ-5 I. parvus ? ? 2.8
KZ-6 I. parvus 13.4 4.4 2.7
641-29 I. sp. 13.2 5.2 4
KZ-3 I.sp. ? ? 5
BA-8 I. sp. ? ? 6.5

 

TABLE 3. Comparison of body mass estimations of taxa documented by an astragalus using the astragalus size (W-astragalus) from the method developed in Martinez and Sudre (1995), and comparisons with the mass range of the extant Tragulus kanchil (from Meijaard and Groves, 2004). The estimated body mass of I. parvus from Benara was obtained by using the m1 area (Wm1) of the type specimen (Gabunia, 1964, p. 182) and the method of Legendre (1989).

  Taxon Estimated body mass
(Kg)
TP641-21 Iberomeryx parvus 1.4
GK3-25 I. parvus 2
TP641-22 I. parvus 2.1
GK3-37 I. parvus 2.2
TP641-24 I.sp. 6.1
GK3-18 I. sp. 5.9
type (p4-m2) I. parvus 2.6
  Tragulus kanchil 1500 to 2500
KZ-9 large Pecora indet. 38

 

 

TABLE 4. Occurrence of identified taxa in the different localities of the Kızılırmak Formation.

  GK2 GK3 Bagdatli TP641 Kz
Iberomeryx parvus x x x x x
Iberomeryx sp. (large)   x x x x
Lophiomerycidae indet. x   x x x
Dremotherium guthi     x    
cf Palaeohypsodontus sp.       x  
large Pecora indet.     x   x
 

metaisGrégoire Métais. Sorbonne Universités, CR2P, MNHN, CNRS, UPMC, Muséum National d’Histoire Naturelle, 57 rue Cuvier, CP 38, 75005 Paris, France. This email address is being protected from spambots. You need JavaScript enabled to view it..

Grégoire MÉTAIS is a vertebrate paleontologist interested in how complex interactions among climate, tectonics, and other physical components of the Earth's environment impact the course of biotic evolution. He completed his PhD at the University of Montpellier, and he is, since 2008, CNRS researcher at the National Museum of Natural History, Paris. His work is primarily dedicated to ungulate mammals' systematic, phylogeny and historical biogeography, but also to the biostratigraphy as well as the sedimentary environments of terrestrial deposits. His fieldwork focuses on the Indo-Pakistan Subcontinent, and the Northern margin of the former Neothethys Ocean, from the Levant to Southeast Asia.

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albayrakEbru Albayrak. Natural History Museum, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Ebru ALBAYRAK is a vertebrate paleontologist primarily devoted to Proboscideans. She completed her undergraduate training and her PhD at Hacettepe University Department of Biology in Ankara. She was a research assistant at the same department. Her work primarily focuses on: 1) the evolution of elephantids, especially the primitive groups, 2) migration of elephantids out of Africa, 3) past distribution of Asian elephant.

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antoinePierre-Olivier Antoine. ISEM, Université de Montpellier, CNRS, IRD, EPHE, CC64, Place Eugène Bataillon, F-34095 Montpellier, France, This email address is being protected from spambots. You need JavaScript enabled to view it.

Pierre-Olivier ANTOINE is a vertebrate paleontologist primarily devoted to Cenozoic mammal evolution, phylogeny, and spatiotemporal distribution. He completed his undergraduate training at the University of Toulouse and his PhD at the Muséum National d'Histoire Naturelle in Paris. He was Post-doctorate fellow at the University of Montpellier and Assistant Professor in the University of Toulouse. His research combines phylogenetic analyses and fieldwork in remote areas such as Balochistan, Peruvian Amazonia, and the Andean Altiplano. Major focuses of his research are: i) Cenozoic stratigraphy, biochronology, and palaeogeography of Eurasia; ii) evolution of megaherbivores (anatomy, systematics, phylogeny, and biochronology), with respect to palaeoenvironmental changes and Cenozoic geodynamics; iii) Cenozoic Evolution of western Amazonian ecosystems and biodiversity, with respect to Andean geodynamics. Outside interests include running (marathon) and drumming (in punk rock and hardcore bands).

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erdalOzan Erdal. Istanbul Technical University, Eurasia Institute of Earth Sciences, 34469 Maslak-Istanbul, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Ozan ERDAL is currently a PhD student who focused on systematics, phylogeny and biostratigraphy of small mammals (mainly rodents) from the late Neogene and Quaternary of some basins formed along the North Anatolian Fault Zone. Formerly, he completed his undergraduate studies on biology at Uludag University (Bursa, Turkey) and he had his master degree on vertebrate palaeontology at University of Montpellier (France). He studied systematics, phylogeny and paleobiogeography of Palaeoamasia (Embrithopoda, Mammalia) from the Eocene of Turkey.

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karadenizliLevent Karadenizli. Geology Department, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Levent KARADENIZLI is Associated Professor and geologist at the General Directorate of Mineral Research and Exploration in Ankara, Turkey. He studied basin formation, stratigraphy and sedimentary environments of terrestrial and marine deposits in several Cenozoic basins of Turkey. He mapped several sedimentary basins in Turkey and provided data on the evolution of their sedimentary environments, lastly in the Çankiri and Sivas basins.

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oyalNeşe Oyal. Natural History Museum, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Ozan ERDAL is currently a PhD student who focused on systematics, phylogeny and biostratigraphy of small mammals (mainly rodents) from the late Neogene and Quaternary of some basins formed along the North Anatolian Fault Zone. Formerly, he completed his undergraduate studies on biology at Uludag University (Bursa, Turkey) and he had his master degree on vertebrate palaeontology at University of Montpellier (France). He studied systematics, phylogeny and paleobiogeography of Palaeoamasia (Embrithopoda, Mammalia) from the Eocene of Turkey.

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saracGerçek Saraç. Natural History Museum, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Gerçek SARAÇ is emeritus paleontologist at the Natural History Museum (MTA) of Ankara, Turkey. He is renowned specialist of Cenozoic rhinos from Turkey, and also biostratigraphy of Cenozoic terrestrial deposits. His catalog of fossil mammalian faunas of Turkey is frequently used by geologists. He published or co-authored many articles on Cenozoic mammals and biostratigraphy.

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yesimYeşim Büyükmeriç. Geological Engineering Department, Bülent Ecevit Üniversitesi, İncivez-Zonguldak, 67100 Zonguldak, Turkey, This email address is being protected from spambots. You need JavaScript enabled to view it.

Yesim BÜYÜKMERIÇ is professor of paleontology at the Bülent Ecevit University in Zonguldak, NW Turkey. She was formerly at the General Directorate of Mineral Research and Exploration in Ankara, where she was the head of the Natural History Museum. She is a worldwide renowned expert of Cenozoic marine mollusks. She took part in many national and international projects in collaboration with Turkish and foreign geologists-paleontologists and published numerous research papers and one monograph dealing with systematics of mollusks, biostratigraphy and paleobiogeography.

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sevketSevket SEN. Sorbonne Universités, CR2P, MNHN, CNRS, UPMC, Muséum National d’Histoire Naturelle, 57 rue Cuvier, CP 38, 75005 Paris, France, This email address is being protected from spambots. You need JavaScript enabled to view it.

Sevket Sen is emeritus Research Director at the National Research Center and works at the Natural History Museum of Paris, France. He investigated Oligocene and Tertiary mammal faunas around the Mediterranean realm and magnetostratigraphy of terrestrial deposits. He edited several volumes on mammalian faunas and published hundreds of academic and non-academic papers. 

 

FIGURE 1. Location map of Late Oligocene mammal localities and sections from the Kızılırmak Formation (Çankırı-Çorum Basin, north central Anatolia, Turkey). 1, geotectonic map of Central Anatolia showing the basins including the Çankırı-Çorum Basin, and the main structural features of the area; 2, Geological map of the studied area with the four fossil localities of the Kızılırmak Formation discussed in the text.

figure1 

FIGURE 2. The four sampled sections showing the stratigraphic position of fossil localities in the Kızılırmak Formation (modified from Karadenizli et al., 2004).

figure2 

FIGURE 3. Dental remains of Iberomeryx parvus and Iberomeryx sp. from the Kızılırmak Formation (early late Oligocene, Central Anatolia). Iberomeryx parvus : 1, Right maxilla with D2-M1 in occlusal view, GK2-4; 2, Left M2 in occlusal view, GK3-31; 3, Left M1 in occlusal view, KZ-7; 4-5, Left M2 in occlusal (4) and labial (5) views, TP641-12; 6-7, Right lower jaw with d3-d4 and the trigonid of m1 in occlusal (6, with line drawing below) and lingual (7) views, GK2-11; 8, Right P4 in occlusal view, GK3-5; 9-10, Right P2 in occlusal (9) and labial (10) views, GK3-4; 11, Left m1 in occlusal view, TP641-4; 12-13, Right m1 in occlusal (12) and labial (13) views, GK3-3; 14-15, Left lower jaw with the talonid of m1 and the trigonid of m2 in occlusal (14) and labial (15) views, TP641-1; 16-17, Left lower jaw with m3 in occlusal (16) and lingual (17) views, TP641-2; 18, Left i1 in lingual view, GK3-7; 19, Left i2 or i3 in lingual view GK3-6; 20, Right p3 in occlusal view, GK3-11; 21, Right p4 in occlusal view, GK3-36. Iberomeryx sp.: 22, Talonid of a left p4 in occlusal view, GK3-35. The scale bar 3 mm goes exclusively with the views 20 to 22.

figure3 

FIGURE 4. Postcranial remains of Iberomeryx parvus and Iberomeryx sp. from the Kızılırmak Formation (early late Oligocene, Central Anatolia). Iberomeryx parvus : 1, Right cubonavicular in anterior view, TP641-16; 2-3, Right astragalus in anterior (2) and posterior (3) views, GK3-39; 4, Right cubonavicular in anterior view, GK2-5; 5-6, Right astragalus in anterior (5) and posterior (6) views, GK3-37; 7, Left calcaneum in medial view, GK3-24. 8 , Right calcaneum in medial view, KZ-11; 9-11, Left distal humerus in anterior-cranial (9), lateral (10), and medial (11) views, GK3-22; 12-13, Proximal part of coalescent metatarsals III and IV in anterior (12), and medial (13) views, GK3-54; 14-16, phalanx proximalis in anterior (14), lateral (15), and proximal views, GK3-13; Iberomeryx sp. 17, Left cubonavicular in anterior view, KZ-2; 18-21, Right astragalus in anterior (18), posterior (19), lateral (20), medial (21) views, TP641-24. Anatomical abbreviations: acf, astragalo-calcaneal facet; ca, capitulum; daf, distal astragalar facet; fbf, fibular facet; imf, internal malleous facet; me, medial epicondyle; of, olecranon fossa; smt2, contact surface of the Mt2; stf, sustentacular facet.

 figure4

FIGURE 5. Dremotherium guthi : 1-3, Left m1 in occlusal (1), labial (2), and lingual (3) views, BA-1; 4, M2? In occlusal view, BA-6; Cf. Palaeohypsodontus sp.: 5-6, Right M2? in occlusal (5) and lingual (6) views, TP641-10; Lophiomerycidae indet.: 7, Right calcaneum in medial view, GK2-7; 8, Right p1 in labial view, TP641-9; 9, Right cubonavicular in anterior view, GK2-3. Large Pecora indet.: 10-11, Right astragalus in anterior (10) and posterior (11) views, KZ-9. Anatomical abbreviations: cf, calcaneal facet; fbf, fibular facet; stf, sustentacular facet. The scale bar 10 mm goes exclusively with the views 7 to 11.

figure5 

FIGURE 6. Stratigraphic ranges of Late Eocene to Oligocene ruminant species in Eurasia (adapted from Métais and Vislobokova, 2007); the Land Mammal Ages are based on Tong et al. (1995), and Woodburne (2006). The colored area designates the duration of the Late Oligocene Warming according to the chart of Zachos et al. (2001).

 figure6

FIGURE 7. Oligocene chronostratigraphic chart indicating the preliminary ages of the different fossil localities based on the unpublished magnetostratigraphic study (Sevket Sen, personal commun., 2016), and the Paleoclimatic context during the Oligocene. Oi1 and Oi2 designate main cooling and glaciation events (Adapted from Berggren et al. 1995, and Zachos et al., 2001 for the paleoclimatic chart)

figure7