Journal Pre-proof Qurliqnoria (Bovidae, Mammalia) from the Upper Miocene of Çorakyerler (Central Anatolia, Turkey) and its biogeographic implications Dimitris S. Kostopoulos, Ayla Sevim Erol, Serdar Mayda, Alper Yener Yavuz, Erhan Tarhan PII: S1871-174X(19)30105-2 DOI: https://doi.org/10.1016/j.palwor.2019.10.003 Reference: PALWOR 539 To appear in: Palaeoworld Received Date: 11 July 2019 Revised Date: 15 October 2019 Accepted Date: 23 October 2019 Please cite this article as: Kostopoulos DS, Erol AS, Mayda S, Yavuz AY, Tarhan E, Qurliqnoria (Bovidae, Mammalia) from the Upper Miocene of Çorakyerler (Central Anatolia, Turkey) and its biogeographic implications, Palaeoworld (2019), doi: https://doi.org/10.1016/j.palwor.2019.10.003 This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier. Qurliqnoria (Bovidae, Mammalia) from the Upper Miocene of Çorakyerler (Central Anatolia, Turkey) and its biogeographic implications. Dimitris S. Kostopoulos a *, Ayla Sevim Erol b, Serdar Mayda c, Alper Yener Yavuz d, Erhan Tarhan e a School of Geology, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece; dkostop@geo.auth.gr b Ankara University, Faculty of Languages, History and Geography, Department of oo c Ege University, Faculty of Science, Department of Biology, Izmir, Turkey; serdarmayda2@yahoo.com Mehmet Akif Ersoy University, Faculty of Art & Sciences, Department of Anthropology, Burdur, Turkey; alpyenyav@gmail.com Hitit University, Faculty of Art & Sciences, Department of Anthropology, Çorum, Turkey; e- e pr d f Anthropology, Ankara, Turkey; ayla_sevim@yahoo.com * Corresponding author. Abstract Pr erhantarhan@hitit.edu.tr na l New bovid material from the Upper Miocene site of Çorakyerler (Çankırı basin, Anatolia, Turkey) is described and compared here. The described taxon is identified as a representative of the stem caprine genus Qurliqnoria, previously known from the peri-Tibetan area ur exclusively. The stronger horn-core divergence, weaker anterior keel, smoother horn-core surface, stronger lateral horn-core curvature, stronger and thicker interfrontal suture, less flexed Jo and less pneumatized frontals, and smaller supraorbital foramina differentiate the Çorakyerler Qurliqnoria from the type and only known species of the genus, Q. cheni from China, and demand the erection of a new species, Qurliqnoria chorakensis n. sp. A review of other late Miocene bovid records allows the recognition of Qurliqnoria in Sinap Tepe (Turkey) and Platania (Greece), suggesting a westward propagation of the genus during the Vallesian. Keywords: Caprini; Artiodactyla; Miocene; Anatolia; systematics Keywords: Carpini, Artiodactyla, Miocene, Anatolia, systematics, biogeography 1. Introduction Discovered in the early 1970s during a joint German-Turkish mineral exploration project (Sickenberg, 1975), the Çorakyerler fossil site is located in the Çankırı basin of north-central Anatolia (Turkey). The fossil site belongs to the fluvial flood-plain deposits of the Tüglu Formation and the local geological and stratigraphical setting have been provided by Kaymakçi et al. (2001) (see also Güleç et al., 2007; Kaya et al., 2016). During the 2000’s the site attracted f worldwide interest due to the discovery of important late Miocene hominoid material (e.g., oo Sevim et al., 2001; Begun et al., 2003; Güleç et al., 2007). Based mainly on biochronological evidence the fauna of Çorakyerler was originally regarded as Vallesian (Becker-Platen et al., 1975) but later tentatively correlated to the early Turolian European Land Mammal Age pr (MN11; e.g., Köhler, 1987; Ünay et al., 2006). Geraads (2013) further suggested an age slightly earlier than Pikermi, Greece (MN12, recently dated at ~7.3 Ma by Böhme et al., 2017) leaving, e- however, open the possibility to be even older than Kemiklitepe D, Turkey (7.9–7.6 Ma; Sen et al., 1994). Kaya et al. (2016) magnetostratigraphically correlated the normal interval Pr including the Çorakyerler fossil site with C4n, framing the age of the fauna between 8.11 and 7.64 Ma. Nevertheless, an alternative correlation option of the magnetostratigraphic results of Kaya et al. (2016) would be around the C4An–C4r boundary, showing a better about 8.9 Ma. na l biochronological match with Sinap Tepe, Turkey data and dating the Çorakyerler fauna at Systematic palaeontological excavations at the Çorakyerler fossil site under the direction of one of us (ASE) started in 2001 and are still ongoing. Apart from primates, nearly 4000 ur identifiable fossil mammal remains have been unearthed the last 18 years, representing mainly hipparionin horses, bovids, giraffids, rhinos, and carnivores. The Çorakyerler bovids have been Jo originally studied by Köhler (1987) based on the initial sparse collection by Sickenberg’s team in the early 70’s. Güleç et al. (2007), Bibi and Güleç (2008) and Kaya et al. (2016) presented successively updated faunal lists taking into account material from the new expeditions, whereas Geraads (2013) provided a brief description of the bovid material collected around 2000 (collection numbers up to ÇO 450). According to the last author, the bovid fauna includes: Miotragocerus (Pikermicerus) sp., Tragoportax? sp., cf. Prostrepsiceros sp., Criotherium? sp., Majoreas cf. woodwardi, Gazella sp.?, Oioceros rothi, Protoryx sp. and “Plesiaddax” inundatus. Bibi and Güleç (2008) and Kaya et al. (2016) also report Nisidorcas and Protragelaphus. Although correct in its core, this faunal list remains incomplete, and somewhat misleading in terms of relative composition of the bovid association (e.g., Bibi and Güleç, 2008, table 6). A study of the Çorakyerler bovid material collected between 2001 and 2018 reveals the presence of an additional new taxon, described and compared here. An exhaustive systematic study of the whole bovid assemblage is in progress. 2. Methodology oo f The studied material is housed in the Çankırı Museum, Turkey and bears the locality prefix ÇO. The material was studied during summer 2019 in the Dil ve Tarih Coğrafya Fakültesi of Ankara University, Turkey. Standard cranial terminology has been used for descriptions (e.g., pr Gentry, 1971; Kostopoulos, 2009). Horn-cores have been considered as heteronymously twisted when the right horn-core follows an anticlockwise torsion. All linear measurements are e- in mm; angles are in degrees (º). The term ‘oblique angle’ (sensu Wang et al., 2019) is used here to describe the angle of the greater basal axis of the horn-core compared to the sagittal 3. Systematic palaeontology Family Bovidae Gray, 1821 Pr plane. na l Subfamily Antilopinae Gray, 1821 (sensu Kingdon, 1982) Tribe Caprini Gray, 1821 ur Genus Qurliqnoria Bohlin, 1937 Emended diagnosis: Small caprine-like bovids with moderately long, slender, anteriorly Jo keeled and weakly twisted heteronymously horn-cores set on short pedicles; horn-core compression moderate to strong; backward curvature weak; lateral curvature (outward flaring) moderate to strong; greater basal axis of the horn-core at a large angle compared to the sagittal plane; supraorbital foramina small associated by shallow furrows; postcornual grooves large. Type species: Qurliqnoria cheni Bohlin, 1937. Qurliqnoria chorakensis n. sp. (Figs. 1, 2) Etymology: from the type locality of Çorakyerler. Holotype: ÇO 3226, part of cranium with right horn-core and palate; Fig. 1. Paratypes: ÇO 3534, frontlet; ÇO 895 frontlet; Fig. 2. Repository: Çankırı Museum, Turkey. Type locality: Çorakyerler, Turkey. Differential diagnosis: A species similar in size to the type species Q. cheni, from which it differs in the stronger horn-core divergence, weaker anterior keel, less intensively grooved oo f horn-core surface, horn-cores curved laterally more than caudally, stronger horn-core compression, stronger and thicker interfrontal suture, less flexed and pneumatized frontals, and smaller supraorbital foramina. pr Occurrence: Late Vallesian–Early Turolian of Anatolia. Description: e- A single partial cranium and two frontlets discovered during the 2004 to 2017 Çorakyerler excavations represent the only evidence of this taxon in the site (Figs. 1, 2). The holotype (Fig. are given in Table 1. Pr 1) belongs to a mature individual (M1 in mid wear, M3 in initial wear stage). Measurements The frontals are slightly raised between the horn-cores; they form a 110–130º angle along the sagittal plane. The interfrontal suture is strong, complicated and constricted all along its na l length as far as the supraorbital foramina (Fig. 2A, D, E). The fronto-parietal suture, is narrower, simpler and ‘T’ shaped (Fig. 2E). The frontal area outlined by these two sutures is moderately to strongly depressed (Fig. 2E). The fronto-nasal dorsal profile is smoothly concave ur (Fig. 1A). The postcornual groove is large, oval shaped and deep (height: 11.7–15.5 mm; width: 6.4–8.0 mm; deepness: 3.3–6.0 mm; Fig. 2B). The supraorbital foramina are small (height: Jo 5.0–6.0 mm; width: 2.0–3.0 mm), not sunken into depressions, moderately wide apart (see Table 1), and associated rostrally by short weak furrows (Fig. 2A, D). The orbits are rather small, rounded and not projecting laterally; their anterior margin is placed above the M3 (Fig. 1A). A large, rounded and fairly deep lachrymal depression occupies the posterodorsal side of the face in lateral view, extending rostrally to above the P4 (Fig. 1A). The horn-cores are inserted directly above the orbits on very short pedicles (Figs. 1A, 2A–C). Their major basal axis is obliquely set compared to the sagittal plane (oblique angle ≥ 45º; maximum measured oblique angle: 67º but possibly exaggerated by taphonomic deformation; Table 1, Figs. 1B, 2B, E). The horn-cores appear slightly heteronymously twisted and strongly divergent from the base up, curved mostly laterally and to a lesser degree caudally (Figs. 1A, 2C, E). They are moderately spaced near the base (Table 1) but 150 mm above, the distance between their medial sides already reaches 230 mm (Fig. 2A, B, D). No one of the available specimens is preserved above the ~150 mm in length (measured along the antero-medial surface) but we estimate that the total length may well have exceeded 200 mm. A weak anterior keel descending rostromedially is present, associated by a series of distinct striations on the mid horn-core surface, both rostrally and caudally (Figs. 1B, 2A, E). The posterior surface of the horn-cores is oo f flattened, the anterior one is more convex. The basal horn-core compression is strong, ranging from 52.3% to 66.2% (mean: 58.5%). The upper toothrow P2-M3 is 71.8 mm long (P2-P4 length: 30.7–31.7 mm; M1-M3 length: pr 42.7 mm). The upper premolars are not shortened compared to the molars (premolar to molar length ratio: 71.9%). The P3 has strong parastyle and paracone rib, and it appears wider distally e- in occlusal view (Fig. 1C). The P4 is rather simple with a strong parastyle (Fig. 1C). The upper molars have strong parastyle and paracone rib and a thinner mesostyle. On the M3 the metastyle Pr is equal to the mesostyle. The lingual crescents are angular and there are no basal pillars (Fig. 1C). Comparison: The saber-like, anteriorly keeled and compressed horn-cores and the slightly raised frontals easily affiliate the Çorakyerler taxon to the ‘protoryxoid’ bovids, a group of Late na l Miocene genera cluster around Protoryx Major (Gentry, 1971; Kostopoulos, 2009). Nevertheless, the particular horn-core morphology (moderately long, slightly heteronymously twisted, strongly divergent and with large oblique angle) in association with their relatively ur smaller size sharply differentiate them from any known genus of the Greco-Irano-Afghan province. The very same features bring the Çorakyerler taxon closer to a set of Chinese taxa Jo from ‘Qaidam (= Tsaidam) fauna’ originally described by Bohlin (1937) and recently revised by Wang et al. (2019). Although similar in overall size and horn-core compression (55–59%; data from Wang et al., 2019), Olonbulukia Bohlin, 1937 differs from the Çorakyerler taxon in the parallel trending and more widely spaced horn-cores on the frontals with a clear posterior curvature and no lateral projection (= outward flaring). In Olonbulukia, the supraorbital foramina are placed into large tear-shaped supraorbital pits, the postcornual grooves take a lower position and the interfrontal suture is not thickened. Tossunnoria Bohlin, 1937 also has similarly compressed horn-cores (53–57%; data from Wang et al., 2019) but it differs from the Çorakyerler taxon in the much more anteroposteriorly extended, relatively shorter and stronger keeled horn-cores, the stronger raised and more pneumatized frontals, and the absence of postcornual grooves. The Çorakyerler taxon matches Qurliqnoria Bohlin, 1937 in the short pedicles, the large postcornual grooves close to the horn-core bases, the slightly raised but thick and pneumatized frontals, the small supraorbital foramina extending into shallow furrows and the rather slender, well striated horn-cores, keeled anteriorly and weakly twisted heteronymously, significantly diverging in frontal view and curving laterally, and set on the frontals at a wide angle to the sagittal plane. The single known species Qurliqnoria cheni oo f Bohlin, 1937 appears more archaic than the Çorakyerler taxon in the less diverging and less compressed horn-cores with weaker outward flaring but is possibly more advanced in the likely more strongly flexed and more pneumatized frontals, the more intense surface grooving, and e- species of Qurliqnoria: Qurliqnoria chorakensis n. sp. pr the larger supraorbital foramina. Hence, we suggest referring the Çorakyerler taxon to a new 4. Discussion Pr According to Wang et al. (2011) and Wang et al. (2019) Qurliqnoria from Qaidam in northeastern Tibet comes from Tuosu Fauna dated at about 12–10 Ma, though Wang et al. (2013a) restrict this range to 11.15–9.98 Ma. The genus is also known from the neighboring Wuzhong Fauna of Ganhegou Formation dated at around the end of the same time interval (~10 na l Ma). Dmitrieva and Serdyuk (2011) also refer Qurliqnoria sp. (mistakenly spelled Quirliqnoria) from Kholu fauna, Tuva, Russia, considered as late Vallesian–early Turolian. Deng et al. (2011) and Wang et al. (2013b) report Qurliqnoria sp. from sites ZD0604, ZD0745, ur and ZD1202 of Zanda Formation, in southwestern Tibet, ranging between 5.29 and 3.31 Ma, and thus expanding significantly the chronological range of the genus up to the end of Early Jo Pliocene. Deng et al. (2011) and Wang et al. (2019) suggest that Qurliqnoria emerged from an endemic Middle Miocene stem caprine assemblage shaped during the Middle Miocene along the rising north Tibetan Plateau. According to the same authors, Qurliqnoria remained restricted to this geographic area (extended up to South Siberia) until its replacement by its alleged descendant the chiru (Pantholops Hodgson; see also Gentry, 1968) during the early Pleistocene (Ruan et al., 2005). The discovery of Qurliqnoria at Çorakyerler highlights the first — but not the only — evidence of the genus far outside its presumed place of origin, and together with available chronological data, leads to a more complex zoogeographic scenario. In an unpublished MTA report and a series of following papers, Ozansoy (1956, 1957, 1965) listed but did not describe the new species of both Qurliqnoria and Olonbulukia from the marl horizons of Middle Sinap, Turkey. The material on which they are based is unknown and the proposed species obviously represent nomina nuda (see also Sen, 2003). Nevertheless, Ozansoy (1956, 1957) also reports from the Lower and Middle Sinap horizons ‘Antilope gen. et sp. indet. II’, by reference to Bohlin’s, 1937 taxon, material on which it appears to establish later Capra bohlini Ozansoy, 1965. ‘Capra’ bohlini is also suggested by Gentry (2003) as a oo f likely match for some new material from Locality 49 of Middle Sinap, described under ‘Pseudotragus aff. capricornis’ (Table 2). The generic affiliations of this taxon have already been called into question by Bouvrain et al. (1994) and a re-examination of the holotype frontlet pr in MNHN, Paris (a not registered specimen from Middle Sinap partly illustrated by Ozansoy, 1965, pl. 8, fig. 1, and marked by the prefix TRQ, as most material of the Ozansoy’s collection e- in Paris, and the label 1955-15-31; DSK pers. obs. and additional illustrations provided by D. Geraads) indicates strong similarities with Q. chorakensis and Qurliqnoria in general (Fig. 3, Pr Table 2): small size (especially compared to other ‘protoryxoid’ bovids), small supraorbital foramina, large postcornual grooves, short pedicles, weakly flexed frontals, and well-divergent, weakly curved backward and gently flaring outward horn-cores with large oblique angle (≥ 45º), weak heteronymous torsion, fairly strong anterior keel, and moderate to strong surface na l grooving (especially anteriorly and posteriorly). We propose therefore Qurliqnoria as a most reliable affiliation for the Ozansoy’s taxon. Several unpublished horn-core specimens from Middle Sinap in MNHN, Paris, Maden Tetlik ve Arama (MTA) Museum in Ankara and the ur Natural History Museum of Ege University in Izmir, likely belong to the same species and a complete revision is needed. Jo Q. bohlini differs from Q. chorakensis in the slightly smaller size (Table 2), less divergent, less outward flaring, less compressed and stronger keeled horn-cores. In most respects — apart from the smaller supraorbital foramina and the less acute anterior keel — Q. bohlini stands morphologically closer to the type species Q. cheni from Qaidam. Ozansoy’s type series of Q. bohlini includes specimens from both the Lower and Middle Sinap (Ozansoy, 1957, 1965) suggesting a large chronological range covering most of the Vallesian but likely older than 9.5 Ma (e.g., Kappelman et al., 2003), and thus fairly contemporaneous with the Qaidam and Wuzhong records. Gentry’s Locality 49 taxon of Middle Sinap, probably represents the same genus; an attribution to the species level is not feasible at the moment, though horn-core compression (~63%) appears closer to the Çorakyerler taxon (Table 2). Locality 49 of Middle Sinap is magnetochronologically dated between 9.2 and 9.0 Ma (Kappelman et al., 2003), an age that might be proved not much older than that of Çorakyerler. Tekkaya (1974) also referred to as a new genus and species, ‘Sinapocerus ozansoyi’, some horn-cores from Middle Sinap, but did not illustrate nor describe the taxon, which apparently becomes another nomen nudum. According to Tekkaya (1974) the horn-cores of ‘Sinapocerus ozansoyi’ are fully comparable to those of Antilope gen. et sp. indet. of Ozansoy from Middle oo f Sinap and therefore good candidates for Qurliqnoria as well (D. Geraads, pers. com. to DSK, October 2019). A far more westward and fairly isochronous record comes from the newly discovered pr Platania fossil site in Northern Greece, yielding a late Vallesian–early Turolian vertebrate fauna (Vasileiadis et al., in press). Tragoreas? aff. oryxoides frontlets from this site can be attributed e- to Qurliqnoria with certain confidence based on the small size, small supraorbital foramina associated by narrow weak furrows, short pedicles, large postcornual grooves, weak frontal Pr flexion, weak heteronymous horn-core torsion, clear outward horn-core flaring, strong compression, and large oblique angle (Table 2). The Platania Qurliqnoria appears morphologically closer to the Çorakyerler taxon as concerns horn-core oblique angle, torsion and divergence but differs in the less deep postcornual grooves, weaker horn-core compression na l (similar to Q. cheni), and smoother grooved horn-core surface; more material is certainly needed for appreciating its species status. ur 5. Conclusions In contrast with the opinion of Wang et al. (2019), Qurliqnoria does not seem to fit the Jo hypothesis of a strictly endemic peri-Tibetan radiation. The similarities between the Tibetan Q. cheni and the Anatolian Q. bohlini suggest close phylogenetic relationships and allow assuming a mid Vallesian westward propagation of the genus. The south pan-Asian geographic range of Qurliqnoria, matches that of the rhino Chilotherium Ringström, the giraffid Samotherium Major, the ovibovine-like bovids Plesiaddax Bohlin, Urmiatherium Bohlin, and Sinotragus Bohlin, as well as that of some hyaenid taxa, which altogether attest the existence of some kind of biogeographic relations between China and the Balkans during the Late Miocene. In this respect, Qurliqnoria could be added to the list of taxa involved in the ‘Late Miocene Middle Asiatic Province’ proposed by Geraads et al. (2002). This idea of a common biogeographic province of open-like environments, previously put forward by Kurtén (1952), partly contradicts the results of Deng’s (2006) analysis (restricted, however, on the Greek and Chinese records), which suggests that the gradually increased diversity and similarity between the Hipparion faunas at the edges of this area do not necessarily imply similar environments and proposes that from the late Vallesian onwards the East Asian fauna extended into Europe. Furthermore, the East and West chronological ranges of the Late Miocene taxa of wide Asiatic distribution, indicate that their expansion does not correspond to a single event but to oo f diachronous ones. As the vast majority of Late Miocene mammal associations from the West and East remains distinct at the species level at least (Deng, 2006), a more selective dispersal between China and Anatolia may be presumed, involving taxa of higher ecological tolerance pr (Samartín, 2012). Within Anatolia, and based on available but not adequate chronological data, Q. bohlini e- and Q. chorakensis seem to be succeeding in time. From the morphological point of view, the former may stand as a possible antecedent of the later, whereas the relationships of the Locality Pr 49 of Middle Sinap and Platania Qurliqnoria remain uncertain. If the Tibetan stem caprines represent ideal candidates for the ancestry of some living Caprini (e.g., Gentry, 1968; Deng et al., 2011; Wang et al., 2019, fig. 6), the Anatolian Qurliqnoria might also meet the morphological and geographic requirements of the na l hypothetical ancestral stock of the Ammotragus-Arabitragus lineage, by the fairly short, obliquely set on the frontals, anteriorly keeled and flaring outward horn-cores, the short pedicles and the small supraorbital foramens associated by narrow-shallow furrows. Molecular ur evidence dates the split of the Ammotragus-Arabitragus clade between 5 and 9 Ma (Ropiquet and Hassanin, 2005) in accordance with Anatolian records of Qurliqnoria. This hypothesis Jo implies, however, a long ghost lineage, and requires a thorough in-depth study, which is beyond the scope of this paper. Acknowledgements Çorakyerler excavations are supported by the Turkish Ministry of Culture and Tourism, the General Directorate of Cultural Heritage and Museums, Ankara University, and the Turkish Historical Society; we are thankful to all of them. Thanks are also due to Wei Dong for fruitful comments and Denis Geraads for valuable suggestions, and information concerning Turkish specimens in his knowledge. References Becker-Platen, J.D., Sickenberg, O., Tobien, H., 1975. 3. Vertebraten-Lokalfaunen der Türkei und ihre Altersstellung. In: Sickenberg, O. 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Mio-Pleistocene Zanda Basin biostratigraphy and geochronology, pre-Ice Age fauna, and mammalian evolution in Jo ur na l western Himalaya. Palaeogeography, Palaeoclimatology, Palaeoecology 374, 81–95. Figure captions Fig. 1. Qurliqnoria chorakensis n. sp. from Çorakyerler, (Late Miocene, Turkey). Holotype ur na l Pr e- pr oo f partial cranium ÇO 3226 in lateral (A), dorsal (B), and palatal (C) view. Scale bar = 5 cm. Fig. 2. Qurliqnoria chorakensis n. sp. from Çorakyerler, (Late Miocene, Turkey). Paratype Jo frontlet ÇO 3534 in rostral (A), caudal (B), and lateral (C) view; and ÇO 895 in rostral (D) and dorsal (E) view. Scale bar = 5 cm. f oo pr ePr na l Fig. 3. Qurliqnoria bohlini (Ozansoy, 1965) from Middle Sinap, (Late Miocene, Turkey). ur Holotype frontlet in MNHN, Paris (suffix TRQ/1955-15-31 – no catalogue number) originally illustrated by Ozansoy (1965, pl. 8, fig. 1) in rostral (A), lateral (B), and caudal (C) view. Note Jo that the distal part of the left horn-core is restored to a greater length compared to the illustration by Ozansoy (courtesy of D. Geraads). Scale bar = 5 cm. na l ur Jo f oo pr e- Pr Table 1. Cranial and horn-core measurements of Qurliqnoria chorakensis n. sp. from Corakyerler, Turkey. All measurements apart from the oblique angle are in mm. Values in brackets are probably affected by taphonomic deformation. 72.5 72.9 30.2 34.0 40.5 42.1 f 122.8 oo 115.1 left 50.5 right 50.2 left 48.8 right 46.5 29.1 26.4 28.8 27.8 30.8 36 29 (67º) pr right 48.1 na l ur Jo ÇO 3226 81.0 40 26.9 60º (65º) Pr Max horn-core diameter at the base Min horn-core diameter at the base Max horn-core diameter at 7 cm above the base Min horn-core diameter at 7 cm above the base Oblique angle ÇO 895 e- Bi-orbital width Length of frontals Width of skull at the lateral edges of the horn-core bases Width of braincase just behind the horn-cores Internal horn-core distance (base) Distance of supraorbital foramens ÇO 3534 113.4 60º 39 25.7 45º Table 2. Morphometrical comparison between the type species and the west records of Qurliqnoria. * according to data provided by Gentry (2003). Length measurements in mm; angle measurements in degrees (º). Q. cheni Bohlin, 1937 Jo 63% Qurliqnoria Platania 36.2–41.9 (mean: 39.7) 52–66% 79% 44–56.5º (mean 51.5º) weakmoderate strong 45–60º (mean ~55º) ≥ 45º strong weak moderate strong intensively grooved large-deep moderately grooved large-deep moderate small small - moderately wideshallow 110–130º moderately wideshallow ~130º - very narrowshallow - ~137º - - oo - 45–60º (mean 54.5º) - moderate moderateweak - only distally pr moderately grooved large-deep 70–73% f 62–73% ur Frontal flexion 41.5 Qurliqnoria * Locality 49, Middle Sinap 42.4 e- Horn-core surface Postcornual grooves Supraorbital foramens Supraorbital furrows Q. bohlini (Ozansoy, 1965) type Pr Outward flaring Anterior keel 39.1–48.4 46.5–50.5 (mean: 43.1) (mean: 48.8) na l Largest basal horn-core diameter Basal horncore compression Oblique angle Q. chorakensis n. sp. - weakly grooved largeshallower small