Turkish Journal of Earth Sciences Turkish J Earth Sci (2019) 28: 822-833 http://journals.tubitak.gov.tr/earth/ © TÜBİTAK Research Article doi:10.3906/yer-1905-7

Primary report of first documentation of mammal footprints from the late Oligocene in the Basin,

1, 2 3 Bekir Levent MESCİ *, Ahmet Cem ERKMAN , Şakir Önder ÖZKURT  1 Department of Geological Engineering, Faculty of Engineering, Sivas Cumhuriyet University, Sivas, Turkey 2 Department of Anthropology, Faculty of Science and Letters, Ahi Evran University, Kırşehir, Turkey 3 Department of Science Teaching, Faculty of Education, Ahi Evran University, Kırşehir, Turkey

Received: 12.05.2019 Accepted/Published Online: 06.09.2019 Final Version: 07.11.2019

Abstract: The footprints in this study, which were considered to have been made by ungulates, were discovered on mudstone that was approximately 5 cm thick, near an abundant mud-cracked plane located on the stratigraphic subsurface of the late Oligocene Karayün Formation, which crops out over wide areas in southern Sivas (Turkey) and has terrestrial, fluvial sediment characteristics. These ungulate footprints documented from the late Oligocene of the Karayün Formation in southern Sivas represent the first reported vertebrates in Anatolia. The footprints of three different species of ungulates were identified. The shapes, depth, and widths of the footprints provided some basic ichnotaxonomic and TrackMaker information, but based on the poor preservation of the footprints, ichnotaxa identification is difficult. This study aimed to use an ichnotaxonomic approach to contribute to the late Oligocene biochronology in Anatolia due to the small amount of footprint findings in the literature. Ungulate herds left mixed footprints in wetland areas along the banks of flooding rivers. The late Oligocene period was a time characterized by large climate changes in Anatolia; hence, it may have hosted different ecosystems and taxa.

Key words: Anatolia, ungulate footprints, paleoichnology, late Oligocene

1. Introduction the literature, as it is the first paleoichnology study from Although clastic deposits of continental environments the Oligocene in Anatolia. Fossil footprints provide limited were not as fully suited to the fossilization of organisms information for biostratigraphy and biochronology due to as other sedimentary rocks, such as carbonate, these types difficult genus or species level identifications. In addition of sediments may preserve the body fossils and footprints to paleobiogeography and paleoecological studies, they of terrestrial vertebrates. The majority of fossil footprints present very valuable information for researchers in have been found on mudstone or sandstone close to terms of behavior, habitat adaptation, and the anatomy of pond, lake, river, or coastal environments. The vertebrate organisms (Sarjeant, 1975; Demathieu et al., 1984; Kordos, mammal footprints from the Cenozoic appear to have 1985; Hunt and Lucas, 2007; Lucas, 2007a; McDonald et been studied less in the literature when compared with al., 2007). dinosaur footprints from the Mesozoic (e.g., Rivera-Sylva Found vertebrate footprints from the Oligocene et al., 2017). When the literature is investigated, generally, formations of the Sivas region in eastern Anatolia were these studies had ecological and ichnological taxonomic evaluated in this study. The study area is located at the features. Although some studies have been conducted Budaklı village crossroads, which is 17 km south of about bird and mammal footprints from the Cenozoic in Sivas and located on the Sivas I38-d4 topographical map Europe and Asia (Hernández-Pacheco, 1929; Vyalov, 1965, 1/25,000 in scale (Figure 1). According to WGS84 data, 1966; Casanova and Santafé, 1982; Santamaria et al., 1990; it is located geographically at the intersection point of López et al., 1992; Anton et al., 1993, 2004; Lockley and 39°35′46.23″N and 37°2′16.08″E. Meyer, 2000; Astibia et al., 2007; Lucas, 2007b; Abbassi et The coincidental preservation of footprints belonging al., 2015; Xing et al., 2017), few studies of invertebrate fossil to mammals provides information that enables one to traces are known from the Oligocene of Turkey (Demircan understand the faunal composition and habitat in the and Uchman, 2016). As a result, this study is important for late Oligocene. Thus, much of the paleoenvironment * Correspondence: [email protected] 822

This work is licensed under a Creative Commons Attribution 4.0 International License. MESCİ et al. / Turkish J Earth Sci of the study area could be explained. The main purpose Neotethyan Ocean in the middle Eocene. These ophiolitic of this study is the ichnotaxonomic report of ungulate mélange nappes and metamorphic rocks, which belong to footprints. With few literature examples, especially when the Kırşehir Massif, form the basement of the basin. Due compared to the rich records of the Cenozoic in Anatolia, to this primary relationship, it has been accepted by many this study may create a clear difference by contributing researchers who have conducted studies in this region to the Anatolian late Oligocene geochronology. Herein, (Cater et al., 1991; Guezou et al., 1996; Poisson et al., 1996). three different ungulate species footprints are presented. Formation of the Tertiary Sivas Basin began in the Eocene, This paper provides a synthetic view of the vertebrate as a foreland basin, with north-south direction in front of ichnological record from the late Oligocene in the Sivas the Pontides. Görür et al. (1998) believed that the Sivas Basin Basin of Turkey. is a peripheral foreland basin. Central Anatolian volcanics and volcanoclastics were formed in the middle-late Eocene 2. Geological setting in the basin (Yalçın et al., 2004), in relation to the collision The Tertiary Sivas Basin is bordered by a tectonic contact, mentioned above (Boztuğ, 2000), and important gypsum which evolved through the overthrust of ophiolitic mélange beds were formed in the shallowing basin throughout the onto basin sediments from the north to the south. The region in the Oligocene. The Sivas Basin is represented by basin belongs to the İzmir-Ankara-Erzincan Suture Zone, both marine and terrestrial sediments ranging in age from representing the closure of the northern branch of the the Eocene to early Miocene. Numerous studies have been

Figure 1. Location map of the study area.

Figure 2. Geological maps of the Sivas Basin and study area (simplified from Kergaravat et. al. (2016) (a) and Bilgiç (2002) (b)).

823 MESCİ et al. / Turkish J Earth Sci carried out on the Oligocene sediments in the basin, due to Karayün formation above a thrust fault. Due to intense displayed lateral and vertical facies changes and different deformation, the dip angle of the layers at the location, environmental conditions (Poisson et al., 1996; Poisson which was observed in detail, was measured as 65°, in a et al., 2015; Kergaravat et al., 2016). Early Miocene-late northwestern direction. From this tectonic border to the Pliocene fluvial and lake sediments have been noted as the north, the dip directions remain the same and the dip youngest rock units in the basin (Figure 2). angles decrease to 30° to 40°. In the study area, only 2 sedimentary rock units were observed: the Hafik and Karayün formations, which are 3. Formation and description of footprints described below in detail. The Hafik Formation comprises The outcrop where the footprints were located was studied massive gypsum beds that crop out over wide areas in the in five different sections according to the distribution and northern, northeastern, and southwestern sides of the walking directions of the footprints (Figure 4). There were study area. Stchepinsky (1939), Lahn (1950), Kurtman abundant mud cracks on mudstone of approximately 5–10 (1961, 1973), Artan and Sestini (1971), Sümengen et al. cm thick (Figure 4). It was thought that the footprints (1990), and Poisson et al. (2015) stated that the Hafik (Figures 5, 6a, 6b, 7a–7d, 8a, 8b, 9a, 9b, 10a, 10b, 11a, Formation is Oligocene in age. According to Poisson and 11b), which were observed in the upper levels of the et al. (1996), there were Eocene marine sediments mudstone layer, were formed when the water level receded. below the Hafik Formation dated as Oligocene. Massive They were preserved because the mudstone layer that gypsum masses showed that the conditions of a lagoon contained the footprints was covered with another layer environment dominated during the sedimentation of this of clastic sediment due to the water level rising once again. unit (Kurtman, 1973). Similar formations were observed in floodplain beds that The massive gypsums that formed the Hafik Formation are still in existence today (Figure 5). The presence of were dislocated towards different stratigraphic levels from these mud cracks indicates seasonal water level changes in their normal positions in parallel with diapiric ascents the paleostream, and when the water level decreased, the and/or due to tectonic compression forces. Therefore, mudstone was exposed to the atmospheric conditions that this formation has tectonic contact over younger late formed desiccation cracks (Figures 6a and 6b). Oligocene-Miocene rock units in different areas of the Ungulate mammals have very complicated walking basin. patterns because of their anatomy and tetrapedal The Karayün Formation, which stratigraphically sits locomotion, and they can change their walking speed and style according to the terrain and other circumstances. above the Hafik Formation, comprises alternations of The water saturation of the sediments on the ground, and fluvial, red-colored, medium-thick bedded conglomerate, thus increased softness, caused negative effects on the sandstone, siltstone, and mudstone. Channel fills, cross- formation and preservation of the footprints. This situation bedding, and current ripple marks (asymmetrical ripple creates difficulties in making precise ichnotaxonomic marks) in this unit indicate that the Karayün Formation identifications and separation of trackways.In these types was affected by active fluvial regimes. According to of situations, generally, trackways cannot be distinguished. Koşun and Çiner (2002), the Karayün Formation is 850 As a result, true measurements cannot be obtained due m in thickness. to increased softness. When measuring a track on the Cater et al. (1991) and Poisson et al. (2015) accepted ground, care should be taken to measure the true size of the Karayün Formation as late Oligocene in age, as it is the footprint. When animals step on heavy muddy ground, unconformably overlain by lower Miocene sediments the edges of the footprints are disrupted when they lift in some sections of the Sivas Basin. Similarly, according their feet. In this context, it is very difficult to say which to Çubuk and İnan (1998) and Kavak and İnan (2001), animal left the footprints observed at Location V (Figures the formation is late Oligocene in age based on its 7c and 7d). However, footprints described in a monograph stratigraphic position. of a study of Cenozoic vertebrate footprints by Oleg According to Kergaravat et al. (2016), the Karayün Stepanovich Vyalov, in 1966, from Dobrotovsky Svita in Formation formed in the playa lake, fluvial braided the western Ukraine region (16–20 Ma) were considered rivers, and saline lacustrine environments, which significant, as they showed similarities to the tracks found stratigraphically formed from bottom to top (Figure 3). in the current study (Lucas, 2007b). When the Vyalov There are many structures and thrust faults developing reproduction was examined (1966, pl. 37), the Hippipeda linked to salt tectonics within the Sivas Basin (Çubuk, aurelianus (holotype) described as possible anchithere 1998; Poisson et al., 2015; Kergaravat et al., 2016). As horse track traces appeared similar to those in Figures 7c, a result, immediately south of the studied location and 7d, 9b, and 10b, although it was difficult to make a clear north of Budaklı village, the Hafik formation overlies the deduction.

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Figure 3. Regional lithostratigraphic column showing depositional environments in the central Sivas Basin and stratigraphic level of footprints (simplified from Kergaravat et al., 2016).

Figure 4. Panoramic view of the footprint locations.

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Figure 5. Stages of the footprint and mud crack formations.

4. Systematic remarks to that of a small gazelle, nearly the same size as a small Bovid footprints are defined as inversely heart-shaped ruminant (like the modern Thomson’s gazelle: Eudorcas tracks, with 2 convex edges and the front tips of the thomsonii). However, it is not clear for ichnotaxonomic hooves stuck into the tracks. The morphology and size definition. of the footprint, which was identified by only 1 footprint, The monograph by Oleg Stepanovich Vyalov stated showed that it was probably made by a small artiodactyl that cervid tracks were ~60 mm long in early Miocene (Figure 7b). The footprint is 3.5 cm in length and 3 cm in fossil footprints (collected in the pre-Carpathian flexure width. Its morphology and size suggested that it is similar (foreland) of western Ukraine) (Lucas, 2007b). The

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Figure 6. View of the mud cracks and footprints in the study area (a), and recent mud cracks and footprints (b).

Figure 7. Footprints observed at Location V.

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Figure 8. Diagonal trace at Location II. holotype of the cervid footprint was Pecoripeda (Cervipeda) incapable of providing an accurate measurement. However, dicroceroides (Vyalov, 1966). When Vyalov’s (1965, pl. 16; there was one example: the length of the footprint was 1966, pl. 3.53) reproduction was examined, the Pecoripeda 8.7 cm and the front hoof ends had stuck into the mud (Gazellipeda) gazella and Pecoripeda (Ovipeda) satyri (Figures 7c and 8b). The animal that stepped on the heavy samples and the Pecoripeda hamori (Szarvas, 2007) muddy ground damaged the edges of the track while it sample from the Ipolytarnoc fossil site (early Miocene) was lifting up its foot. The actual size of the track on the in northern Hungary, found in 1836, appeared similar to ground, not the whole track, was taken into consideration small ruminants in Figures 7b and 8b. when measuring. Artiodactyla and Perissodactyla footprints were The footprint could have been formed with a 2-beat reported from Oligocene Navarre deposits (Astiba et al., gait, where the front-right, rear-left and front-left, rear- 2007). According to Astiba et al. (2007), the morphology of right limbs moved forward diagonally at the same time, the Navarrese footprints looked like that of Entelodontipus and the tracks were left one after another at an approximate viai from the Oligocene. Although the examples shown in distance of 55 cm at Location II (Figure 8a). Since the Figures 7b and 8b appeared to have similar features, this ground was soft, the distortion of the actual footprint was far from a taxonomic determination. increased. The traces were superficial and the edge was Location II included 15 Artiodactyla footprints, probably disrupted when the bovids lifted their feet, which which were thought to have morphologic characteristics was likely due to the heavy sludge on the ground. The belonging to a medium-sized bovid. Due to the ground distance between the animal’s steps showed a diagonal being soft, the deformation of the real track was increased. walking style in the same direction. Generally, it is difficult for a real track to be distinguished There were several footprints that were intermingled, in such cases. The observed tracks were superficial and the length of which was approximately 10–12 cm (Figure

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Figure 9. Mammal (perissodactyl) trackways from Location III.

9b). Because certain tracks were left one on top of the In the general view of Location III, we could observe other, they were more complicated and wider. The fact views of different footprints. However, though there that the ground was soft increased the deformation of was no clear evidence of which group they belonged to, the real tracks. The observed tracks were superficial and they were considered to have been left by a group close most of them were incapable of providing an accurate to Perissodactyla (Figures 10a and 10b). When Vyalov’s measurement and a clear view of the hooves. The herd was reproduction was examined (1966, pl. 37), although observed to have stopped here (Figures 9a and 9b). The the holotype of Hippipeda aurelianus that was described depth of the track in the mud showed the direction of the as possible anchithere horse tracks was similar to that footprints. in Figures 7c, 9b, and 10b, it was overly ambitious to

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Figure 10. Mammal tracks from Location IV include possible perissodactyl (Rhinocerotidae?) footprints on the mudstone. state anything clearly. However, anchithere horse tracks also remarked on the occurrence of Anchitherium sp. (at would probably have to be early Miocene rather than late Bayraktepe 1, Sarptepe Formation, Çanakkale), which Oligocene in age. Hence, the footprints may belong to a dates back to the late Astaracian (also equivalent to the medium-sized Perissodactyla. middle Miocene) (Ozansoy, 1973). Anchitherium is commonly known from the early Footprints at Locations III and IV were partly well and middle Miocene faunas from Turkey, such as in preserved and measurements could be taken (Figures 9 Bursa-Paşalar, Ankara-Çandır, Inönü-1, Muğla-Sarıçay, and 10). The footprints had a mean length of 10–9.3 cm and Kultak-Muğla (Becker-Platen et al., 1975; Atalay, and width of 10–12 cm. The oval morphology and size of 1981; Köhler, 1987; Kaya, 1989; Forsten, 1990). Although the footprints could be compared to the Rhinoceripeda Anchitherium does not appear in the fossil record of the from the Lower Miocene in Hungary, based on the analysis Balkan Peninsula during this period, which may suggest by Vyalov (1965). However, it was not possible to fully that this region was isolated from Anatolia (Koufos et al., identify these footprints with an oval hoof shape. Although 2005), the lack of Anchitherium in the Balkan Peninsula the footprints were roughly similar to Rhinoceripeda was probably due to the lack of early and middle Miocene in terms of the oval main outline, they appeared to be localities with a reliable record. Ozansoy (1969) mentioned relatively smaller. The footprints were observed to go in the occurrence of Anchitherium sp. (large size) in the Uşak 4 different directions and step length measurements were Akçalköy locality and stated that this situation constituted recorded at Location I (Figure 11a). Although it was very an exception for Turkey. He related this situation to the difficult to conclude which species it was close to, it could continuity of the genus Anchitherium’s vertical spread in be considered more similar to a single-hoofed animal Anatolia, as was seen in Europe (Ozansoy, 1969). He (Figure 11b).

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Figure 11. View of the walking tracks at Location I.

5. Conclusions into subbasins, allowing geographic differentiation, and Though some of the ungulate tracks were well preserved as a result, each region experienced a unique geodynamic in the Karayün Formation, we were unable to make and faunal evolution. During the late Oligocene to early genus or species level descriptions. However, general Miocene, many mammal groups spread out and became measurements including shapes, depths, and widths more diverse (Kemp 2005). The depositional environment of the footprints provided some information about the of the footprint site indicates that the area was seasonally sizes and order of the ungulates. These findings indicate flooded. The present habitat in the locality was the shore of that large and medium-sized ungulates with substantial a wetland area and could also be defined as an ecosystem biodiversity were present in Central Anatolia during the in which half-open meadows provided nutrition for species late Oligocene. Additionally, the Karayün Formation from that are ecological shareholders. This situation showed the late Oligocene of Central Eastern Anatolia has great that the water level along the river bank was periodically potential to yield more fossil mammals footprints. reduced, and after organisms living in the region left their Anatolia is located in the transition zone between Europe, footprints in the mud, it experienced a period of drying. Asia, and Africa and has the features of a small continent The mudstone layer containing the footprints was preserved because of its climatic and geographic conditions. The after being covered by clastic sediments again, due to a later southern side of the European continent was an archipelago rise in the water level. Mud cracks observed on the rock in terms of geography, which did not allow mammals to formed due to the same mechanism. Although there was no migrate towards the end of the Oligocene period. Pollen and clear information about what animal group left the tracks, spore studies (Akgün et al., 2007), geodynamic evolution they suggest the idea that tracks may have been left by two (Husing et al., 2009), and paleoclimatological research artiodactyl track-makers, one small and the other medium (Fortelius et al., 2006) covering the late Oligocene and in size, and probably medium-sized perissodactyl groups. early Miocene period showed that the climate in Central Anatolia comprised subtropical environmental conditions Acknowledgment in terms of global climate. Additionally, continuing tectonic We would like to thank the reviewers for their detailed compression and uplifting in the region divided the basins comments and suggestions for the manuscript.

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