The Late Miocene Maragheh Fauna, Northwest Iran

Recent Advances in Paleobiological Research of the Late Miocene Maragheh Fauna, Northwest Iran

Majid Mirzaie Ataabadi1, Raymond L. Bernor 2,3, Dimitris S. Kostopoulos4, Dominik Wolf 3, Zahra Orak5, Gholamreza Zare5, Hideo Nakaya6, Mahito Watabe7 and Mikael Fortelius1,8

1 Department of Geology, University of Helsinki, P.O. Box 64, FIN-00014, Finland

2 National Science Foundation, GEO:EAR Sedimentary Geology and Paleobiology Program

3 College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, Howard University,

520 W St. N.W., Washington D.C. 20059, USA

4University of Thessaloniki, Geological Department, Museum of Geology and Paleontology, 54124 Thessaloniki, Greece

5Department of Environment, MMTT, Tehran, Iran

6Department of Earth and Environmental Sciences, Faculty of Science, Kagoshima University, Japan

7 Hayashibara Museum of Natural Sciences, Okayama, Japan

8 Institute of Biotechnology, P.O. Box 56, FIN-00014 University of Helsinki, Finland

I. Historical Background

The fossil localities of Maragheh are located in the latter half of the 19th century (Abich 1858, the eastern Azarbaijan province, northwest Iran Brandt 1870, Grewingk 1881). These early at 37° 21' 08'' N latitude and 46° 24' 40'' E works provided data on Maragheh’s similarity longitude. The Maragheh fauna has long been to Pikermi. The Austrian paleontologist Pohlig considered one of the three most preeminent was invited by a merchant from the nearby city western Eurasian late Miocene Pikermian of Tabriz to visit the locality in 1884 and it was faunas, along with Samos and Pikermi in Pohlig that made the first comprehensive Greece. As with Pikermi and Samos, collection and geological study of Maragheh Maragheh is a true “Lagerstätte” because of (Pohlig, 1886). Pohlig explored extensively the shear abundance and diversity of its fauna. across the Maragheh basin and would appear It is unique amongst the three classical to have sampled fossils from nearly all, if not Pikermian faunas in its clear stratigraphic all of the Maragheh section. The Pohlig display and layer-cake stratigraphy with collection in the Naturhistorisches Museum, several, laterally continuous volcanic ashes Wien is extraordinary as an early collection that are readily amenable to radioisotopic because much of it preserves locality dating. information which facilitates understanding of A Russian explorer, Khanikoff has been its stratigraphic provenance. Two other credited with first finding the Maragheh site in Austrian paleontologists, Rodler and Kittl, 1840 and sending a small collection to Dorpat visited Maragheh and made an extensive University (now University of Tartu, Estonia). collection of fossils which were later published The Maragheh fauna was initially studied in by Kittl (1887), Rodler (1890), Rodler and MIRZAIE ATAABADI ET AL. / RECENT ADVANCES IN MARAGHEH RESEARCH

Weithofer (1890) and Schlesinger (1917). There are three important outcomes Damon, from the British Museum of Natural from the field work undertaken in the 1970’s History, London made a small collection including: a) collection of fossils with close briefly communicated by Lydekker in 1886. In regard to stratigraphic provenance which has 1897, the French paleontologist Marcellin led to a biostratigraphy of the Maragheh fauna, Boule secured permission to conduct a b) study of all collections to better understand paleontological expedition to Maragheh in the taxonomy and diversity of the mammalian 1904. The 1904 French expedition to fauna, and c) application of a variety of Maragheh was organized at a very grand scale geochronologic tools to secure well resolved for this time in paleontology. A group of ages for the Maragheh section and its faunas. French paleontologists assisted by 12 local After 30 years cessation of excavation laborers excavated a large sample of Maragheh activities in the Maragheh basin, Iran’s fossils from Kingir, Kopran, Shollevend and Department of Environment (DOE) and Kermedjawa (Mecquenem 1905, 1906, 1908, National Museum of Natural History (MMTT) 1911, 1924-25). started a new initiative and sponsored new More than 50 years elapsed before other excavations in the area which resulted in reported expeditions occurred at Maragheh. nomination of 10 km2 of Maragheh Takai of Tokyo University collected Maragheh fossiliferous area as a national protected zone fossils from Kerjabad (Takai 1958). Robert and establishment of a field museum and Savage of Bristol University also visited research station in the Dare Gorg (Gort Daresi) Maragheh in 1958 and collected fossils. area. The MMTT-University of Helsinki Tobien from the Johannes-Gutenberg initiative known as the International Sahand University, Mainz made important excavations Paleobiology Expedition (INSPE) has also of the middle portion of the Maragheh been recently started and is currently in sequence in the 1960’s (Tobien, 1968). During progress. This program has undertaken three the 1970’s three scientific groups conducted field seasons between 2007 and 2009, research at Maragheh: a combined Dutch- discovering several new localities and German group led by Erdbrink (Erdbrink et al. numerous fossils. The program has further 1976, Erdbrink 1976 (a,b), 1977, 1978, 1982, reinitiated study of the mammalian fauna with 1988), a joint University of Kyoto-Geological the intention of bringing them into a Survey of Iran led by Kamei (Kamei et al. contemporary taxonomic context for 1977, Watabe 1990, Watabe and Nakaya comparative paleoecological and 1991a,b) and the Lake Rezaiyeh Expedition paleobiogeographic studies. (LRE) led by Campbell (Campbell et al. 1980). Bernor was a student charged with the study of Abbreviations vertebrate fauna for the LRE which resulted in his PhD (1978) and manuscripts on the fauna, INSPE - International Sahand Paleobiology biostratigraphy and zoogeographic Expedition relationships of the fauna (1986) and the MMTT - Muze Meli Tarikh Tabiei (National systematic, biostratigraphy and zoogeography Museum of Natural History), Tehran of the hipparionine horses (1985). An MNHN - Muséum National d’Histoire extensive review of the fauna with systematic, Naturelle, Paris chronologic and biogeographic comparisons to M2 – Upper second molar Pikermi and Samos was published by Bernor et NMW - Naturhistorisches Museum Wien al. (1996). UCR - University of California, Riverside

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II. Geology and Stratigraphy (Urmiyeh-Dokhtar) volcanic belt in the northeast and the Sanandaj-Sirjan metamorphic The Maragheh basin is bounded to the north by belt (Mendelassar transform) in the southwest the NW-SE trending Anatolian transform fault, (Fig.1). The Urmiyeh-Bazman volcanic belt also known as the Tabriz fault, on the west by with its northwest-southeast trend is believed the N-S trending Urmiyeh fault and on the to have resulted from the collision of the south by the NW-SE Mendelasar transform Arabian and Iranian plates. The Sanandaj- fault. Regionally the Maragheh Basin and its Sirjan metamorphic belt with a similar trend associated transform faults are dominated by lies between the main Zagros thrust (crush the Zagros crush zone to the south and west. zone) and the Urmiyeh-Bazman belt Also, there are the Urmiyeh-Bazman (Davoudzadeh et al. 1997).

Figure 1 – Geographic position and relationships of the Maragheh area to the major tectonic features (after Dewey et al. 1973 and Huber 1976) in N. W. Iran.

During the Paleogene, northwest Iran Miocene, the last Tethyan seaway incursion experienced a wide range of post collisional regressed from this area ending the local arc volcanic activities. After this magmatism carbonate deposition cycle (Aghanabati 2004). event, clastic, evaporate, and carbonate Consequently, at the beginning of the sediments were deposited during the late Neogene, this domain emerged above sea level Paleogene and early Neogene (Lower Red and and developed as incipient mountain ranges, Qom Formations). By the end of the early basin troughs, and a topography resembling

3 MIRZAIE ATAABADI ET AL. / RECENT ADVANCES IN MARAGHEH RESEARCH present conditions (Davoudzadeh et al. 1997). soils. These uppermost horizons are more than The most significant deposits of this time are 350 m thick south of Sahand, but can be as terrestrial sediments and evaporites known much as 1000 m in thickness in areas near the collectively as the Upper Red Formation. The Anatolian transform (Bernor et al. 1980). remains of these deposits are not found in the Maragheh area but mostly occur north of the The Maragheh Group Tabriz Fault and south of Urmiyeh fault We describe herein the sedimentary horizons (Fig.1). It seems that these major faults in the of the Maragheh Group as they are expressed area, which have been active since the in the Maragheh Basin. Paleozoic (Aghanabati 2004), structurally The Basal Tuff Fm. represents a single controlled and prevented deposition of these air-fall unit of rhyolite tuff with local thickness units in the Maragheh Basin. Volcanic activity of over 80 m. This unit is a uniform, unbedded, was reinitiated in the late Miocene to middle structureless deposit of white, devitrified ash Pliocene interval in the Maragheh Basin and with randomly oriented crystals of mica and adjacent areas (Moin-Vaziri and Amin- fresh fragments of feldspar and quartz. The Sobhani 1977). unit represents a tremendous pyroclastic event The Late Miocene Maragheh with substantial outcrops south and northeast stratigraphic sequence accumulated on the of the central fossiliferous area which has also southern flank of the Mt. Sahand volcanic proven to be useful for long-range intra-basin massif. Mt. Sahand is a large volcanic complex correlations (Campbell et al. 1980, Bernor et covering an area of about 10,000 km2 (Moin- al. 1980, Bernor 1986). Vaziri and Amin-Sobhani 1977) and regardless The Maragheh Fm. rests unconformably of its circular outline, is not a single volcano. on the Basal Tuff Fm. It is eroded to a A series of distinct volcanic cones are arranged thickness of 300 m and consists of strata made along an east-west trend collectively forming up exclusively of detrital fragments of this enormous volcanic massif. hornblende andesite and dacitic pumice, and is The late Miocene deposits of the interbedded at widely spaced intervals with Maragheh Basin consist of a thick sequence of layers of pumice-lapilli tuff. In general, the volcaniclastic continental strata with a basal volcaniclastic beds are unlaminated poorly pyroclastic unit. Kamei et al. (1977) named the sorted silty grits with lenses of andesite and entire 500-600 m thick late Miocene sequence pumice cobbles, in depositional units ranging the Maragheh Formation. They differentiated in thickness from 1-3 m. The top of each this formation into a lower fossiliferous depositional unit is generally marked by a member (160 m) and an upper non darker, hardly weathered zone with root-casts fossiliferous unit forming the upland hills of (Bernor et al. 1980). Mt. Sahand. Campbell et al. (1980) restricted Maragheh Fm. deposits are bound to the the Maragheh Fm. to the lower 300m thick north by the Anatolian transform (Tabriz fault) volcaniclastic and fossiliferous series. They (Fig.1). Between the northwest of Sahand also referred to the basal pyroclastic unit as the massif and the city of Tabriz possible Basal Tuff Formation. Hence, the fossil- Miocene-Pliocene deposits are unlike the bearing sequence of Maragheh Basin is Maragheh Fm. These beds are composed of confined to the lower 150 m of 300 m thick diatomites containing fish and mollusks and Maragheh Fm. The upper surface of the some lignites with plant remains. Hipparionine Maragheh Fm. is erosional with local Pliocene- teeth and scarce mammalian bones are Quaternary capping of heavily oxidized terrace recorded from these deposits (Rieben 1934), gravels, pumice breccias, and boulder-ridden whose nature is quite different from those of

4 MIRZAIE ATAABADI ET AL. / RECENT ADVANCES IN MARAGHEH RESEARCH the Maragheh Fm. Recently, abundant also known as the Sanandaj–Sirjan mammalian fossils have been discovered in the metamorphicbelt. areas north and northeast of Tabriz (Mirzaie Campbell et al. (1980) reported that Ataabadi et al. submitted). Although this fossil some lithological horizons in the Maragheh material resembles that of the Maragheh Fm., Fm. can be traced over wide areas allowing their geology, sedimentology, and taphonomy intra-basin correlations. The most distinctive differs. To the west, the Maragheh Fm. is unit for correlation is a diamictitic breccia bounded by Lake Urmiyeh (or Urmiah), which named “Loose Chippings”. This marker bed is a shallow, hypersaline body of water formed best outcrops in the central portion of the study in the Pleistocene by the activities of the area and has been used for stratigraphic Tabriz and Urmiyeh faults (Aghanabati 2004). correlation of vertebrate localities in this area To the southwest, the limit of Maragheh Fm. is (Fig. 2). This bed is recorded as the “scoria the Mendelassar ridge (Fig.1). This is an bed” by Kamei et al. (1977) and as the uplifted belt of lightly metamorphosed rocks “trachytic breccia” by Erdbrink et al. (1976).

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Figure 2 – Lithology and stratigraphy of the Maragheh Formation, N. W. Iran. (See Fig. 3 for location of sections A-H). Sites A and H are correlated based on the Basal Tuff. Sites B and D-G are correlated by the “Loose Chippings” marker bed. Site C is correlated to nearby sections by “pumice bed 2” and corresponds to the recent excavations (MMTT II, DRG1, and AZM1) in the Maragheh area. The position of section C above the basal tuff and below the “Loose Chippings” is certain. However, the details of the base and top of the section are not recorded. The basal tuff is shown at the base of most sections, with topographic elevation. Numbers and letters to the left of each column are fossil localities. R1-R12 and MULB-5 are sites from which radiometric age determinations were obtained (see also Fig. 3 and Table 1). 1A, 1B and 2 in site C indicate pumice beds.

Section C (Fig. 2) represents the recent part (Ilkhchi) which are the youngest (Bernor excavations in the Maragheh area by MMTT et al. 1980, Bernor 1986). and INSPE teams. It is correlated to the Major sedimentary facies that have been adjacent sections by a major pumice layer distinguished in Maragheh are pebble and known as the “Pumice Bed 2”. “Pumice Bed cobble conglomerate which make less than 5% 2” is 5-7 m thick and has been widely traced in of the studied sections, grey sandstone and the study area. This bed was likely breccia facies which makes up about 25% of accumulated from a single large-scale flow the sections, poorly sorted massive siltstones event (Sakai, pers. commun.). The sections in which constitute about 70% of Maragheh the extreme northeast and southwest (A and H) strata, and air-fall tuff deposits consisting are correlated based on the Basal Tuff Fm. almost entirely of pumice fragments. It seems Maragheh Fm. seems to rest with a low that the following sedimentary events are angle regional unconformity on the Basal Tuff responsible for deposition of the Maragheh Fm. Based on the studies in the central Fm.: a) erosion by small streams which made a fossiliferous area the regional dip of the small disconformity at the base, b) deposition Maragheh Fm. is west-southwest with about 5 of coarse clastics by lateral accretion in point m/km inclination. Triangulation from the bar deposits and fine clastics by vertical presently known exposures of the Basal Tuff accretion in overbank deposits, c) soil Fm. by the American team in the 1970’s also formation at the top of former units, and d) indicates a consistent dip to the west-southwest random airfall deposition. These processes with a general inclination of about 15 m/km. built the extensive Maragheh Fm. as a product The differences between the dips of these units of alluviation rather than volcanic activity or suggest that the Basal Tuff draped over a west lacustrine sedimentation (Campbell et al. 1980, sloping paleoslope/basin that was gradually Bernor et al. 1980, Bernor 1986). filled by the Maragheh Fm. These successive Fossil bones in Maragheh Fm. occur as beds prograded eastward as the base level rose localized concentrations within the (Campbell et al. 1980). This interpretation has unlaminated beds, floating in the sediments been generally supported by radiometric rather than lying on bedding-planes. A single (Campbell et al. 1980) and biostratigraphic complete articulated skeleton of the mustelid evidence (Bernor 1978, Bernor et al. 1979, Promeles palaeattica has been found from the Bernor et al. 1980, Bernor 1986). These data MMTT 13 quarry (Bernor et al. 1996). suggest that the beds and associated fossils Taphonomic studies of these fossil farthest to the west (Kopran) are the oldest accumulations indicate autochthonous bone compared to the localities in the easternmost assemblages accumulated on overbank or

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Figure 3 – Geographic sites and fossil localities (UCR–MMTT, MMTT, and INSPE) of the Maragheh Basin, N. W. Iran. 1- Kopran localities , 2-Varjoy localities, 3- Aliabad localities, 4- Mordagh (Mirduq, Mordaq) localities, 5- Dare Gorg (Gort Daresi) localities (including new MMTT and INSPE localities), 6- Karajabad (Kherjabad) localities, 7- Sumu Daresi locality, 8- E. Mordagh localities, 9- Shalilvand (Shol’avand) localities, 10- Ghartavol localities, 11- N. E. Shalilvand (Shol’avand) localities, 12- Khermejavand locality, 13- Ilkhchi localities and 14-Ahagh (Ahaga), W. Maragheh localities. |---A---| correspond to the stratigraphic columns of figure 2. Pyroclastic events such as mudflows or ash falls were not directly responsible for floodplain deposits of fluvial systems. mortality. On the other hand, biologic agents A large number of the bones are were the probable cause of death, as the bones preserved with articulation of distal limb were buried almost immediately or subaerially elements and early weathering stages. exposed only long enough to allow removal of some elements by scavengers (Morris 1997).

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III. Paleontology and Chronology

Bernor (1986) and Bernor et al. (1996) specifically the importance of the Maragheh provided an account of the mammalian species sequence. Figure 3 is a satellite image reported from Maragheh. Since 1996, there indicating the principal vertebrate paleontology have been a number of taxonomic revisions collecting areas in the Maragheh Basin. that affected the documentation of fossils at Vertebrate fossil localities crop out across the Maragheh and their comparisons with other Maragheh Basin and often are expressed as penecontemporaneous Eurasian and African dense concentrations of fossils up to a Meter in mammal faunas. Moreover, there have been a thickness and extending 10s to 100s of Meters number of studies of Eurasian (Bernor et al. laterally. This is particularly true for the Upper 2003, 2005, Eronen et al. 2009) and Eurasian- Maragheh locality MMTT 13 near the village African (Bernor and Rook 2008, Bernor et al. of Shol’avand. Since the latter part of the 19th 2009) biogeographic relationships for the late century coincident with the Austrian Miocene interval that have brought new exploration of the Maragheh Basin there has significance to understanding Old World been a growing archive for the stratigraphic (Pikermian) chronofaunas in general, and provenance of the Maragheh fauna (Bernor 1986).

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Figure 4 – Mammalian biostratigraphy of the Maragheh Formation, N. W. Iran. The Stratigraphic provenance of vertebrate localities is given above/below the “Loose Chippings” marker bed. Taxa with * are also recorded from Upper Maragheh Loc. 37

The work by Iranians with Japanese, includes two distinct skull morphologies and Dutch, German and American groups in the two distinct postcranial morphologies. The 1970’s brought marked improvements to this skulls with a large, single, preorbital fossa stratigraphic record. Figure 2 provides a (POF) placed close to the orbit are believed to summary of 8 stratigraphic sections, arrayed be associated with elongate slender from west to east, of these principal collecting metapodials and likely referable to areas with the UCR -MMTT localities (Bernor Cremohipparion aff. moldavicum, whereas the 1986) and newly discovered INSPE localities skull with a single, highly reduced POF is indicated. likely associated with short, stout metapodials Bernor (1978, 1986) integrated all referable to Hippotherium brachypus. In that known stratigraphic records of fossil mammals the MNHN assemblage is believed to be to develop the first Maragheh mammalian stratigraphically derived mostly from the biostratigraphy. He originally subdivided the middle portion of the section we now need to Maragheh Fm. into three units based on the recognize that the biostratigraphic subdivisions stage-of-evolution of the Hipparion s.s. are not based on the evolution of a continuous lineage: Lower Maragheh whose base was lineage, but simply as biozones. defined by the first occurrence of Hipparion Current evidence suggests the following gettyi at Kopran; Middle Maragheh whose base horse sequence: Hipparion gettyi occurs at was defined by the first occurrence of Kopran, the oldest set of localities in the Hipparion prostylum; and Upper Maragheh Maragheh Basin with its stratigraphic range whose base was defined by the first occurrence being from the -150 to -75 Meter interval of of Hipparion campbelli. Recent studies of the the section; Hippotherium brachypus and Maragheh hipparion samples housed by the Cremohipparion moldavicum occur in the -52 MNHN and Howard University to -25 Meter interval of the section; Hipparion Laboratory of Evolutionary Biology suggest campbelli is first known to occur at the -20 that Hipparion prostylum, originally defined Meter interval and is believed to be present at based on skull morphology alone, does not the + 7 Meter interval of the section. Small occur at Maragheh. The Paris sample horses which we believe to be best referred to (MNHN) originally believed to be referable to Cremohipparion?matthewi occur from about - Hipparion prostylum (Woodburne and Bernor 115 to + 7 Meter interval of the section. There 1980, Bernor et al. 1980, Bernor 1985) yields are likely multiple species of small no postcrania characteristic of Hipparion Cremohipparion known from Western Eurasia prostylum s.s (Mt. Luberon sample) or (i.e. Cremohipparion matthewi, Cremohipparion Hipparion campbelli (Howard University nikosi and Cremohipparion minus and Maragheh sample). However, there are potentially others) and there is simply too little postcrania in the Paris collection and Tobien’s cranial and complete metapodial data to collection that are referable to Hippotherium determine which of these occur at Maragheh. brachypus, and we believe that this is a more The Maragheh hipparion assemblage is likely referral for the Paris and Tobien numerically abundant and species diverse and collections of hipparions with a reduced fossa is undergoing extensive new systematic placed high on the skull. We find therefore that analysis by Mirzaie Ataabadi, Bernor, and the MNHN Maragheh Hipparion assemblage Wolf (in progress). Figure 4 updates Bernor’s

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(1986) and Bernor et al.’s (1996) initial report on zircon fission track ages and biostratigraphy of the Maragheh mammal K/Ar40 ages. These are summarized, by fauna. stratigraphic interval, in table 1 here which Swisher (1996) provided new single shows that the ages are internally coherent crystal Argon ages for the Maragheh fauna that with the oldest ages being stratigraphically significantly revised Campbell et al.’s (1980) succeeded by progressively younger ages.

Table 1 – Summary of the isotopic age determinations from the Maragheh Formation and Basal Tuff (after Bernor 1986 and Swisher III 1996).

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Table 2- Locality ages of UCR-MMTT fossil localities of Maragheh Formation, N. W. Iran, inferred from the estimated sedimentation rates. Ages with * are isotopic age determinations of zero level which corresponds to “Loose Chippings” and -110 level which corresponds to Mordaq (Murdag) tuff (see also Fig. 5 and Table 1). Localities with * have an estimated level with respect to “Loose Chippings”.

Bernor et al. (1996) suggested that based on estimated sedimentation rates the Maragheh fauna ranges from about 9 m. y. to 7.4 m. y. Table 2 represents a calculation of ages for localities in the western and central portion of the sequence based on the estimated sedimentation rates (0.008 m. y. per Meter). Thus, the oldest locality is Kopran I (8.96 m. y. estimated age) and the youngest is MMTT 26 (7.68 m. y. estimated age), located 7 Meters above the “Loose Chips” marker bed. Figure 5 summarizes the geochronology of the Maragheh Basin and shows that the Ilkhchi fauna is located in an eastern section calibrated as being between 7.58 +/- .11 m. y. (R8) and 7.42 +/- 0.11 m. y. (R10).

Table 3 provides a summary of commun.); within the Proboscidea we now Maragheh mammalian taxa by biostratigraphic recognize the deinothere as being interval as originally defined by Bernor (1986). Deinotherium gigantissimum following the Here we update a number of the mammalian specimens discovered at MMTT31 (Erdbrink groups. Taxonomic changes over Bernor et al. locality K1) by Schmidt-Kittler; the (1986) include: within the Hyaenidae we equids are as described above; amongst the recognize Adcrocuta (not Percrocuta) eximia; Rhinocerotidae we recognize Iranotherium the large machairodont cat is now recognized morgani and a Rhinocerotidae new gen. and as being Amphimachairodus (not sp. for Maragheh (Giaourtsakis, pers. Machairodus) aphanistus (Lars Werdelin, pers. commun.); the single suid occurring at

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Maragheh corresponds to a population of Oioceros atropatenes, Oioceros rothii, small-medium sized Microstonyx major Nisidorcas sp., Prostrepsiceros (=erymanthius) (Kostopoulos pers. data); for houtumschindleri, Prostrepsiceros cf. vinayaki, the giraffids, we currently recognize Prostrepsiceros fraasi, Prostrepsiceros cf. Helladotherium duvernoyi, Samotherium rotundicornis, Protragelaphus skouzesi, neumayri, Palaeotragus coelophrys and ?Palaeoreas sp., Skoufotragus laticeps, Bohlinia attica (not the specimen mentioned Palaeoryx sp., Urmiatherium polaki, by de Mequenem 1925: Pl. II, Fig. 3), but a ?Criotherium sp., Mirabilocerus cf. maius, revision of the material is certainly needed in Miotragocerus cf. vallenciennesi, Tragoportax order to clarify chronogeographic relationships cf. amalthea (not Miotragocerus rugosifrons), (Kostopoulos, pers. data); the Bovidae are and Samokeros minotaurus. The additional undergoing extensive revision (Kostopoulos presence of Skoufotragus schlosseri and Bernor, in progress) and we recognize the (=Pachytragus crassicornis) is possible but not following taxa for Maragheh: Gazella substantiated by the current revision. We also capricornis (not cf. deperdita), Gazella cf. recognize the presence of Hystrix in Maragheh ancyrensis, “Gazella” (not Gazella) rodleri, based on material in MNHN.

Figure 5 – Stratigraphic array of isotopic age determinations from the Maragheh Formation and Basal Tuff (after Bernor 1986 and Swisher III 1996) best-known Middle Maragheh localites (1A, In summary, the Maragheh fauna has a 1B, and 1C), where several groups have chronologic range of nearly 9 m.y. to less than collected fossils, the current oldest age is for 7.6 m.y. but the bulk of fossil material is from locality 1A, 8.16 m. y. (Table 2, estimated), the middle and upper parts of the section. The and the youngest localities in the Shol’avand Lower Maragheh fauna is mostly comprised of area (MMTT 26) date 7.68 m. y. (Table 2, taxa with very long time distributions that estimated). To the east, the youngest localities cannot be used in fine time- of Ilkhchi (MMTT 37) would appear to be ca resolution interregional comparisons. For the 7.4 m. y.

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Table 3- Mammalian species, biostratigraphic intervals with their estimated ages, and UCR-MMTT fossil localities of Maragheh Formation, N. W. Iran (modified after Bernor 1986).

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Table 3- cont.

The correlation of Maragheh with Miocene, is still open to discussion. Samos and Pikermi, the classical mammal Although this problem has been previously fossil localities of the Eurasian late addressed (Bernor et al. 1996) and local

14 MIRZAIE ATAABADI ET AL. / RECENT ADVANCES IN MARAGHEH RESEARCH biostratigraphy and geochronology of (Kostopoulos et al. 2003, Koufos et al. Maragheh and Samos has been greatly 2009a). This chronology (Fig. 6) would improved in the past years (Bernor et al. make Middle-Upper Maragheh generally 1996, Swisher III 1996, Kostopoulos et al. correlative with the lower fossil horizons 2003), Pikermi remains badly resolved, (PMAS) at Samos (7.8-7.4 m.y.), but the being dated only indirectly. Maragheh localities, as currently According to the latest understood, appear somewhat older than magnetostratigraphic correlation the Samos those producing the Samos Intermediary fauna ranges from ca 7.8 to 6.7 m.y., but (IMAS) and Dominant Mammal the core of the Samos fauna referred to as Assemblages (7.4-6.9 m.y., Koufos et al. the Samos Dominant Faunal Assemblage 2009a). (DMAS) is dated between 7.2 and 6.9 m.y.

Figure 6 – Chronostratigraphic position of Maragheh, Samos and Pikermi fossiliferous sites and faunal assemblages according to data presented in this paper and Koufos et al. (2009a).

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However, this conclusion is not always locality of Cr. moldavicum is Taraklija of the consistent with the evolutionary stage of late Turolian. several mammalian taxa and their combined Clearly a conclusive correlation between presence in both Samos and Maragheh (i.e. Maragheh, Samos, and Pikermi will have to Melodon (=Parataxidea) maraghanus, await resolution of the apparent mismatch Hyaenictitherium (=Thalassictis) wongii, between the available geochronological and Adcrocuta eximia, Hippotherium brachypus, biochronological evidence. Without “Cremohipparion” cf. matthewi, Diceros anticipating we note that the main explanations neumayri, Skoufotragus laticeps, available at this time are: a) mistakes in the Miotragocerus valenciennesi, Gazella radiometric or magnetostratigraphic dating, b) capricornis, Prostrepsiceros fraasi, mismatches in the correlation of old localities Tragoportax, Palaeoryx, Protragelaphus, with new stratigraphic evidence, c) major Palaeotragus, Samotherium; Kostopoulos, diachrony in the occurrence of species between pers. data), which dates to the 7.4-6.9 m.y. these localities, and d) a mistaken attribution of interval of Samos but appears to be older in ecomorphs to chronospecies. Maragheh. The Maragheh mammal association as it IV. Paleobiogeography and is shown in Fig. 4 is in part constructed from Paleoecology the old provenance data of the Vienna (NMW) and the Paris (MNHN) collections developed We follow recent investigations on in Bernor (1986). However, the Tobien paleobiogeographic analysis by Bernor et al. collection does have stratigraphic provenance (2001) and Fortelius et al. (1996) by and was collected from the same part of the undertaking genus-level faunal resemblance stratigraphic sequence where MMTT localities index (FRI) studies using both the Simpson’s 1A-1C were collected (Tobien, pers. commun.) (1943) and the Dice (Sokal and Sneath 1963) One of the present authors (RLB) indicies. Dice FRI is highly recommended by correlates Middle Maragheh, 8.1-7.9 m.y. Archer and Maples (1987) and Maples and (estimated) with Pikermi based principally on Archer (1988) and is calculated as 2C/2(A+B), the occurrence of Hippotherium brachypus where C is the number of shared taxa between which is very similar to the Pikermi form and two faunas, and A and B are the total number Cremohipparion moldavicum which is the of taxa present in fauna 1 and fauna 2. sister taxon to Cremohipparion mediterraneum, Simpson’s FRI also has a long tradition of use also known from Pikermi (Koufos, 1987). (Bernor 1978, Flynn 1986, Bernor and Another (DSK) holds that the apparent faunal Pavlakis 1987) and is calculated as C/smaller relationships between Pikermi and Maragheh, of (A and B). also evidenced by several artiodactyl taxa do We also used the ungulate tooth crown not confirm such ages, as the mammal height (Bernor et al. 2001) to show the contrast association of Pikermi appears to be more between the localities under consideration. The advanced than those from Vathylakkos-2, three part subdivision of crown height Prochoma and Perivolaki (Greece), includes: brachydont whereby M2 crown magnetostratigraphically dated between 7.4- length is greater than its crown height; 7.2 m. y. (Koufos et al. 2006). In addition, mesodont whereby M2 crown length is roughly Hippotherium brachypus and Cremohipparion the same as crown height; hypsodont whereby moldavicum are also present in Akkaşdaği M2 crown height is more than 2 times that of (Turkey, dated 7.1 m. y.) while the type crown length.

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All data have been downloaded from the Maragheh, the Western and Central European NOW (Neogene of Old World) database on localities, all plot with an index value of less September 30, 2009 (http://www.helsinki.fi than 20% for both Dice and Simpson’s FRI. /science/now/). Our analyses here consider However, Sinap loc.12 (hominid zone), only large mammals because small mammal Turkey, has 9 genera in common with records vary greatly across the faunas as a Maragheh and a Dice FRI of 30% and result of taphonomic and sampling bias. Simpson’s FRI of 47% which is the highest Figure 7 illustrates the plot of genus among these early Vallesian localities. The level (G) FRI in pair-wise comparisons shared genera between Sinap loc. 12 and between Maragheh and 10 other localities Maragheh are: Chilotherium, Choerolophodon, under consideration. Among the early Criotherium, Gazella, Hipparion, Orycteropus, Vallesian (MN 09) localities compared to Palaeoreas, Palaeotragus and Tragoportax.

Figure 7 – Genus level Faunal Resemblance Index (GFRI); pair wise comparison of localities under consideration with Maragheh: Can Llobateres, Spain; Eppelsheim, Germany; Rudabanya, Hungary; Sinap loc.12 , Turkey (MN09); Samos and Pikermi, Greece; Baode loc. 49, China (MN12); Baode loc.30, China; Sahabi, Libya and Middle Awash, Ethiopia (MN13 equivalent).

Analysis of crown height (Fig.8) also hypsodonty in these faunas is only due to demonstrate that Western and Central occurrence of the invasive species of European (MN09) localities of Rudabanya, hipparionine horses (Hippotherium) which Eppelsheim and Can Llobateres have low show adaptation to closed environments with a incidence of mesodont and hypsodont forms significant amount of browse in their diets (5-10% of mesodont and 5-10% hypsodont (Bernor et al. 2003). In remarkable contrast is taxa) compared to more than 85% of the Sinap loc. 12, showing stronger mesodonty brachydonts. This suggests that Central and and hypsodonty signals. Sinap loc. 12 exhibits Western Europe during Vallesian was forested 35 % brachydonts, 28% mesodonts and 35% with warm climates and low seasonality. The hypsodont forms. As demonstrated before

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Figure 8 – Crown height diagrams for Vallesian and Turolian localities under consideration: Can Llobateres, Spain; Eppelsheim, Germany; Rudabanya, Hungary; Sinap loc.12 , Turkey (MN09); Samos and Pikermi, Greece; Maragheh, Iran; Baode loc. 49, China (MN12); Baode loc.30, China; Sahabi, Libya and Middle Awash, Ethiopia (MN13 equivalent).

(Fortelius et al. 2003), later Vallesian Western temperate (more) wooded settings at this Eurasian faunas are very similar in their locality (Solounias 1999). community structure to regional Turolian Recently, Kostopoulos (2009) showed faunas. They might be called “proto- that the paleoecological profile in Pikermi Pikermian” showing the origin of “Pikermian includes the tree-dwelling semi-terrestrial Chronofauna” which is characterized best by primate Mesopithecus, diversified felids and the Maragheh-Pikermi-Samos triad (Eronen et mustelids, and browse-dependent proboscideans, al. 2009). rhinos, giraffes and bovids. In contrast, Samos Maragheh’s closest resemblance is to the has no primate and browse-dependent taxa and Greek locality of Samos. Maragheh has 28 has diversified hipparionine horses and genera (22 species) in common with Samos gazelles with grazing rhinos, giraffes, and and its GFRI is 68% for Dice and 74% for bovids. Maragheh paleoecological profile is Simpson’s. The next closest relationship of remarkable in this context by having a mixture Maragheh is clearly to Pikermi, Greece. of Samos and Pikermi characters (see Koufos Maragheh shares 28 genera (20 species) with et al. 2009b: Fig.7). Not only primates are this locality and its GFRI is 61% for Dice and present in Maragheh, diversified felids, 63% for Simpson’s. These classical mustelids and hipparionine horses also exist. “Pikermian Chronofaunas” show more stable Browsing and grazing proboscideans, rhinos, community structures. Maragheh and Samos giraffes and bovids also occur in Maragheh. have about 45-35% of brachydont taxa, 30% of Therefore, although Maragheh is more similar mesodonts and 25-35% of hypsodonts (Fig. 8). to Samos than it is to Pikermi (Fig. 7), However, Pikermi has a lower percentage of environmentally it has more wooded settings in hypsodont forms which reflects the warm its dominant grass/bushy vegetation. For this

18 MIRZAIE ATAABADI ET AL. / RECENT ADVANCES IN MARAGHEH RESEARCH reason, as mentioned by Strömberg et al. morgani, also occurs in early Late Miocene of (2007), presence of a climatic and vegetational Linxia basin, northwestern China (Deng 2005). gradient across the Greco-Iranian province This taxon has apparently first appeared in (east-to-west pattern) is not clearly evident. northwestern China during Vallesian and In the Middle Turolian (Kostopoulos immigrated to Maragheh later in the Turolian. 2009) evidently great intercontinental The terminal Miocene Libyan locality of dispersion of large carnivores and ungulates of Sahabi is also similar in community structure “Pikermian chronofauna” occurred and some to Maragheh. High percentage of hypsodont of the core Pikermian genera extended their taxa (32%) occurs with 23% mesodont and geographic range so that a sizable number of 45% brachydont forms in this locality which is genera are shared among Western Eurasian as similar to Maragheh as are the Chinese “Pikermian chronofaunas” (Eronen et al. Baode localities (Fig. 7). Taxa shared between 2009), Chinese MN 13-equivalant faunas this locality and Maragheh are: Adcrocuta, (Mirzaie Ataabadi et al. submitted), and the Amphimachairodus, Ceratotherium (Diceros), terminal Miocene north and east African Cremohipparion, Gazella, Indarctos, faunas (Bernor and Rook 2008, 2009). Prostrepsiceros, Samotherium, and Baode loc. 30 (MN13 equivalent) has 13 Tragoportax. genera in common with Maragheh and a Dice The late Miocene Middle Awash fauna, FRI of 40% and Simpson’s FRI of 56%. Baode Ethiopia is less than 20% similar to Maragheh loc. 49 (MN12 equivalent) has 11 genera in in terms of GFRI. However, it has almost common with Maragheh and its Dice FRI is similar community structure but with less 32% and Simpson’s FRI 41%. North African hypsodont and more brachydont taxa. These (Libyan) locality of Sahabi shows a similar African faunas clearly had “Pikermian” level of similarity. Sahabi has 9 genera in elements that were vicariant and evolved common with Maragheh and its Dice FRI is independently subsequent to an early-middle 27% and Simpson’s FRI 37%. Turolian extension (Bernor and Rook 2008, The Chinese localities of Baode 49 and 2009). 30, being the eastern extensions of the “Pikermian Chronofauna” (Mirzaie Ataabadi et Conclusions al. submitted) exhibited Pikermian-type community structures. The MN 12-equivalent Apart from widely distributed Pikermian taxa locality of Baode 49 is similar to Pikermi with (e.g. Adcrocuta, Amphimachairodus, Felis, about 12% hypsodont taxa, 30% mesodonts Deinotherium, Hipparion, Hyaenictitherium, and 59 % brachydonts. On the other hand, the Hysterix, Ictitherium, Indarctos, Metailurus, MN13-equivalent locality of Baode 30 is very Palaeotragus, Samotherium ), the giraffid similar to Maragheh and Samos with about Bohlinia and bovids such as “Gazella” rodleri, 33% hypsodont taxa (Fig.8). This locality is the large Palaeoryx, and Mirabilocerus cf. also more similar to Maragheh than locality 49 maius and hipparionine horses like in terms of GFRI (Fig. 7). Taxa shared Cremohipparion moldavicum indicate between Baode localities and Maragheh are: affinities of the Maragheh fauna to the Adcrocuta, Amphimachairodus, Chilotherium, Northern Black Sea region. Some taxa like Cremohipparion, Felis, Gazella, Hipparion, Prostrepsiceros aff. vinayaki show Ictitherium, Indarctos, Metailurus, Palaeoryx, relationships with the western Asian (Arabia, Palaeotragus, Parataxidea, Samotherium, Afghanistan and Indian sub-continent) region, Thalassictis and Urmiatherium. The large while Gazella cf. ancyrensis, Prostrepsiceros elasmothere rhinoceros, Iranotherium fraasi/houtumschindleri, Samokeros and

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Skoufotragus suggest affiliations with the area. We also thank the Governor, the Mayor Anatolia and Samos. Urmiatherium and and the local DOE office managers of Iranotherium are certainly common elements Maragheh and Tabriz for their hospitality and with China. Regardless of the absence of real assistance during INSPE fieldworks. MMA, time resolution and mismatch between some MF and field work in Maragheh were partially geochronological and biochronological data, it supported by the Academy of Finland, RHOI- is evident that the Maragheh area was affected NSF project and the Sasakawa Foundation. Z. by biogeographically distinct late Miocene Davoudifard extensively helped in preparation areas, representing the crossroads of several of several illustrations for this paper. RLB provinces. wishes to acknowledge the National Science Foundation, including EAR-0125009 (grant to Acknowledgements R.L. Bernor and M.O. Woodburne), BCS- 0321893 (grant to F.C. Howell and T.D. We appreciate the efforts of Dr D. Najafi White) and the Sedimentary Geology and Hajipour and Dr S. Montazami and their Paleobiology Program (GEO: EAR: SEP) for predecessors in Iran’s Department of supporting his research on this project. Environment for reinitiating and supporting new excavations in the Maragheh fossiliferous

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