Fossil Mammals of Asia

Chapter 25

Recent Advances in Paleobiological Research of the Late Miocene Maragheh Fauna, Northwest Iran MAJID MIRZAIE ATAABADI, RAYMOND L. BERNOR, DIMITRIS S. KOSTOPOULOS, DOMINIK WOLF, ZAHRA ORAK, GHOLAMREZA ZARE, HIDEO NAKAYA, MAHITO WATABE, AND MIKAEL FORTELIUS

The fossil localities of Maragheh are located in the eastern Azarbaijan province, northwest Iran, between 37°20'– 37°30'N latitude and 46°10'–46°35' E longitude. The Maragheh fauna has long been considered one of the three most preeminent western Eurasian Late Miocene Pikermian faunas, along with those of Samos and Pikermi in Greece. As with Pikermi and Samos, Maragheh is a true “Lagerstätte” because of the shear abundance and diversity of its fauna. It is unique among the three classical Pikermian faunas in its clear layer-cake stratigraphy with several, laterally continuous volcanic ashes that are readily amenable to radioisotopic dating. A Russian explorer, M. Khanikoff, has been credited with fi rst fi nding the Maragheh site in 1840 and sending a small collection to Dorpat University (now University of Tartu, Estonia). The Maragheh fauna was initially studied in the latter half of the nineteenth century (Abich 1858; Brandt 1870; Grewingk 1881). These early works provided data on Maragheh’s similarity to Pikermi. The Austrian paleontologist H. Pohlig was invited by a merchant from the nearby city of Tabriz to visit the locality in 1884, and it was Pohlig (1886) who made the fi rst comprehensive collection and geological study of Maragheh. He explored extensively across the Maragheh Basin and would appear to have sampled fossils from nearly all, if not all, of the Maragheh sections. The Pohlig collection in the Naturhistorisches Museum, Vienna, is extraordinary as an early collection because much of it preserves locality information that facilitates an understanding of its stratigraphic provenance. Two other Austrian paleon-

tologists, A. Rodler and E. Kittl, visited Maragheh and made an extensive collection of fossils, which were later studied (Kittl 1887; Rodler 1890; Rodler and Weithofer, 1890; Schlesinger 1917). R. Damon, from the British Museum of Natural History, London, purchased a small collection of Maragheh fossils, which was briefly communicated by R. Lydekker in 1886. In 1897, the French paleontologist M. Boule secured permission to conduct a paleontological expedition to Maragheh in 1904. The 1904 French expedition to Maragheh was orga nized at a very grand scale for this time in paleontology. A group of French paleontologists assisted by 12 local laborers excavated a large sample of Maragheh fossils from Kingir, Kopran, Shol’avand, and Kermedjawa (de Mecquenem 1905, 1906, 1908, 1911, 1924–1925). More than 50 years elapsed before other reported expeditions occurred at Maragheh. F. Takai (1958) of Tokyo University collected Maragheh fossils from Kerjabad. R. Savage of Bristol University also visited Maragheh in 1958 and collected fossils. H. Tobien (1968) from the Johannes-Gutenberg University, Mainz, made important excavations of the middle portion of the Maragheh sequence in the 1960s. During the 1970s, three scientific groups conducted research at Maragheh: a combined Dutch– German group led by B. Erdbrink (Erdbrink 1976a, 1976b, 1977, 1978, 1982, 1988; Erdbrink et al. 1976), a joint University of Kyoto– Geological Survey of Iran team led by T. Kamei (Kamei et al. 1977; Watabe 1990; Watabe and Nakaya 1991a, 1991b), and the Lake Rezaiyeh Expedition (LRE) led by B. Campbell

RECENT ADVANCES IN PALEOBIOLOGICAL RESEARCH OF THE LATE MIOCENE MARAGHEH FAUNA, NORTHWEST IRAN

(Campbell et al. 1980). R. Bernor was a student charged with the study of vertebrate fauna for the LRE, which resulted in his Ph.D. (1978) and manuscripts on the fauna, biostratigraphy, and zoogeographic relationships of the fauna (Bernor 1986) as well as the systematics, biostratigraphy, and zoogeography of the hipparionine horses (Bernor, Woodburne, and Van Couvering 1980; Bernor 1985). An extensive review of the fauna with systematic, chronologic, and biogeographic comparisons to Pikermi and Samos was published by Bernor et al. (1996). There are three important outcomes from the field work undertaken in the 1970s, including (1) collection of fossils with close regard to stratigraphic provenance, which has led to a biostratigraphy of the Maragheh fauna; (2) study of all collections to better understand the taxonomy and diversity of the mammalian fauna; and (3) application of a variety of geochronologic tools to secure well-resolved ages for the Maragheh section and its faunas. After a 30-year cessation of excavation activities in the Maragheh Basin, Iran’s Department of the Environment (DOE) and National Museum of Natural History (MMTT) started a new initiative and sponsored new excavations in the area, which resulted in the nomination of 10 km2 of the Maragheh fossiliferous area as a national protected zone and the establishment of a field museum and research station in this area. The recent MMTT– University of Helsinki initiative, known as the International Sahand Paleobiology Expedition (INSPE), is currently in progress. Th is program has undertaken three field seasons between 2007 and 2009, discovering several new localities and numerous fossils. The program has further reinitiated studies of the mammalian fauna with the intention of bringing them into a contemporary taxonomic context for comparative paleoecological and paleobiogeographic studies. GEOLOGY AND STRATIGRAPHY

The Maragheh Basin is bounded to the north by the northwest– southeast-trending Anatolian transform fault, also known as the Tabriz fault, to the west by the north–south-trending Urmiyeh fault and to the south by the northwest–southeast Mendelasar transform fault. Regionally, the Maragheh Basin and its associated transform faults are dominated by the Zagros crush zone to the south and west. Also, there are the Urmiyeh– Bazman (Urmiyeh–Dokhtar) volcanic belt in the northeast and the Sanandaj– Sirjan metamorphic belt (Mendelassar transform) in the southwest (figure 25.1). The

547

Figure 25.1 Geographic position and relationships of the Maragheh area to the major tectonic features. After Dewey et al. (1973) and Huber (1976) in northwest Iran.

Urmiyeh–Bazman volcanic belt with its northwest– southeast trend is believed to have resulted from the collision of the Arabian and Ira nian plates (Davoudzadeh, Lammerer, and Weber-Diefenbach 1997). The Sanandaj– Sirjan metamorphic belt with a similar trend lies between the main Zagros thrust (crush zone) and the Urmiyeh–Bazman belt. During the Paleogene, northwest Iran experienced a wide range of postcollisional arc volcanic activities. After this magmatism event, clastic, evaporite, and carbonate sediments were deposited during the late Paleogene and early Neogene (Lower Red and Qom formations). By the end of the Early Miocene, the last Tethyan seaway incursion regressed from this area resulting in the local carbonate deposition cycle (Aghanabati 2004). Consequently, at the beginning of the Neogene, this domain emerged above sea level and developed as incipient mountain ranges, basin troughs, and a topography resembling present conditions (Davoudzadeh, Lammerer, and WeberDiefenbach 1997). The most significant deposits of this time are terrestrial sediments and evaporites known collectively as the Upper Red Formation. The remains of these deposits are not found in the Maragheh area but mostly occur north of the Tabriz fault and south of the Urmiyeh fault (see figure 25.1). It seems that these major faults in the area, which have been active since the Paleozoic (Aghanabati 2004), structurally controlled and prevented deposition of these units in the Maragheh Basin. Volcanic activity was reinitiated in

548

WEST ASIA AND ADJACENT REGIONS

the Late Miocene to Middle Pliocene interval in the Maragheh Basin and adjacent areas (Moin-Vaziri and Amin-Sobhani 1977). The Late Miocene Maragheh stratigraphic sequence accumulated on the southern fl ank of the Mount Sahand volcanic massif. Mount Sahand is a large volcanic complex which covers an area of about 10,000 km 2 (Moin-Vaziri and Amin-Sobhani 1977) and, despite its circular outline, is not a single volcano. A series of distinct volcanic cones are arranged along an east– west trend collectively forming this enormous volcanic massif. The Late Miocene deposits of the Maragheh Basin consist of a thick sequence of volcaniclastic continental strata with a basal pyroclastic unit. Kamei et al. (1977) named the entire 500–600-m-thick Late Miocene sequence the Maragheh Formation. They differentiated this formation into a lower fossiliferous member (160 m) and an upper nonfossiliferous unit forming the upland hills of Mt. Sahand. Campbell et al. (1980) restricted the Maragheh Formation to the lower 300 m volcaniclastic and fossiliferous series. They also referred to the basal pyroclastic unit as the Basal Tuff Formation. Hence, the fossil-bearing sequence of Maragheh Basin is confi ned to the lower 150 m of a 300-m-thick Maragheh Formation. The upper surface of the Maragheh Formation is erosional with a local Pliocene– Quaternary capping of heavily oxidized terrace gravels, pumice breccias, and boulder-ridden soils. These uppermost horizons are more than 350 m thick south of Sahand, but they can be as much as 1000 m in thickness in areas near the Anatolian transform (Bernor, Woodburne, and Van Couvering 1980). THE MARAGHEH GROUP

We describe herein the sedimentary horizons of the Maragheh Group as they are expressed in the Maragheh Basin. The Basal Tuff Formation represents a single air-fall unit of rhyolite tuff with local thickness of over 80 m. Th is unit is a uniform, unbedded, and structureless deposit of white, devitrified ash with randomly oriented crystals of mica and fresh fragments of feldspar and quartz. The unit represents a tremendous pyroclastic event with substantial outcrops south and northeast of the central fossiliferous area and has proven to be useful for long-range intrabasin correlations (Campbell et al. 1980; Bernor, Woodburne, and Van Couvering 1980; Bernor 1986).

The Maragheh Formation rests unconformably on the Basal Tuff Formation. It is eroded to a thickness of 300 m and consists of strata made up exclusively of detrital fragments of hornblende andesite and dacitic pumice and is interbedded at widely spaced intervals with layers of pumice-lapilli tuff. In general, the volcaniclastic beds are unlaminated and poorly sorted silty grits with lenses of andesite and pumice cobbles, in depositional units ranging in thickness from 1m to 3 m. The top of each depositional unit is generally marked by a darker, weathered zone with root casts (Bernor, Woodburne, and Van Couvering 1980). Maragheh Formation deposits are bound to the north by the Anatolian transform (Tabriz fault) (see figure 25.1). Between the northwest of Sahand massif and the city of Tabriz possible Miocene-Pliocene deposits are unlike the Maragheh Formation. These beds are composed of diatomites containing fish and mollusks and some lignites with plant remains. Hipparionine teeth and scarce mammalian bones are recorded from these deposits (Rieben 1934), whose nature is quite different from those of the Maragheh Formation. Recently, abundant mammalian fossils have been discovered in the areas north and northeast of Tabriz (Mirzaie Ataabadi, Zaree, and Orak 2011; Mirzaie Ataabadi, Mohammadalizadeh, Zhang 2011). Although this fossil material resembles that of the Maragheh Formation, their geology, sedimentology, and taphonomy differ. To the west, the Maragheh Formation is bounded by Lake Urmiyeh (or Urmiah), which is a shallow, hypersaline body of water formed in the Pleistocene by the activities of the Tabriz and Urmiyeh faults (Aghanabati 2004). To the southwest, the limit of Maragheh Formation is the Mendelassar ridge (see figure 25.1). Th is is uplifted lightly metamorphosed rocks known as the Sanandaj–Sirjan metamorphic belt. Campbell et al. (1980) reported that some lithological horizons in the Maragheh Formation can be traced over wide areas allowing intrabasin correlations. The most distinctive unit for correlation is a diamictitic breccia named “Loose Chippings.” Th is marker bed crops out best in the central portion of the study area and has been used for stratigraphic correlation of vertebrate localities in this area (figure 25.2). Th is bed is recorded as the “scoria bed” by Kamei et al. (1977) and as the “trachytic breccia” by Erdbrink et al. (1976). Section C (see figure 25.2) represents the recent excavations in the Maragheh area by MMTT and INSPE teams. It is correlated to adjacent sections by a major pumice layer known as “Pumice Bed 2.” Pumice Bed 2 is 5–7 m thick and has been widely traced in the study area. Th is bed was likely accumulated from a single


549

Figure 25.2 Lithology and stratigraphy of the Maragheh Formation, northwest Iran. (For location of sections A– H, see figure 25.3.) 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 Maragheh. 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 MUL6-5 are sites from which radiometric age determinations were obtained (see also figure 25.3 and table 25.1). 1A, 1B, and 2 in site C refers to pumice beds.

large-scale flow event (T. Sakai, pers. comm.). The sections in the extreme northeast and southwest (A and H) are correlated based on the Basal Tuff Formation. The Maragheh Formation seems to rest with a low angle regional unconformity on the Basal Tuff Formation. Based on the studies in the central fossiliferous area the regional dip of the Maragheh Formation is westsouthwest with about 5 m/km inclination. Triangulation from the presently known exposures of the Basal Tuff Formation by the American team in the 1970s also indicates a consistent dip to the west-southwest with a general inclination of about 15 m/km. The differences

between the dips of these units suggest that the Basal Tuff draped over a west-sloping paleoslope/basin that was gradually fi lled by the Maragheh Formation. These successive beds prograded eastward as the base level rose (Campbell et al. 1980). Th is interpretation has been generally supported by radiometric (Campbell et al. 1980) and biostratigraphic evidence (Bernor 1978, 1986; Bernor, Tobien, and Van Couvering 1979; Bernor, Woodburne, and Van Couvering 1980). These data suggest that the beds and associated fossils farthest to the west (Kopran) are the oldest compared to the localities in the easternmost part (Ilkhchi), which are the youngest

550


(Bernor, Woodburne, and Van Couvering 1980; Bernor 1986). Major sedimentary facies that have been distinguished in Maragheh Formation are pebble and cobble conglomerate, which make up less than 5% of the studied sections, gray sandstone and breccia facies, which make up about 25% of the sections, poorly sorted massive siltstones, which constitute about 70% of Maragheh Formation, and occasional air-fall tuff deposits consisting almost entirely of pumice fragments. It seems that the following sedimentary events are responsible for deposition of the Maragheh Formation: • Erosion by small streams, which made a small disconformity at the base • Deposition of coarse clastics by lateral accretion in point bar deposits and fi ne clastics by vertical accretion in overbank deposits • Soil formation at the top of these units • Random airfall deposition These processes built the extensive Maragheh Formation as a product of alluviation rather than volcanic activity or lacustrine sedimentation (Bernor, Woodburne, and Van Couvering 1980; Campbell et al. 1980; Bernor 1986). Fossil bones in the Maragheh Formation occur as localized concentrations within the unlaminated beds, floating in the sediments rather than lying on bedding planes. A single complete articulated skeleton of the mustelid Promeles palaeattica has been found from the MMTT 13 quarry (Bernor et al. 1996). Taphonomic studies of these fossil accumulations indicate autochthonous bone assemblages accumulated on overbank or floodplain deposits of fluvial systems. A large number of the bones are preserved with articulation of distal limb elements and early weathering stages. Pyroclastic events such as mudflows or ash falls were not directly responsible for the mortality of animals. On the other hand, biologic agents were the probable cause of death, as the bones were buried almost immediately or subaerially exposed only long enough to allow removal of some elements by scavengers (Morris 1997). PALEONTOLOGY AND CHRONOLOGY

Bernor (1986) and Bernor et al. (1996) provided an account of the mammalian species reported from Maragheh. Since 1996, there have been a number of taxonomic revisions that affected the documentation of fossils at Maragheh and their comparisons with other penecon-

temporaneous Eurasian and African mammal faunas. Moreover, there have been a number of studies of Eurasian (Bernor, Kordos, and Rook 2003, 2005; Eronen et al. 2009) and Eurasian-African (Bernor and Rook 2008; Bernor, Rook, and Haile-Selassie 2009) biogeographic relationships for the Late Miocene interval that have brought new significance to our understanding of Old World (Pikermian) chronofaunas in general, and specifically the importance of the Maragheh sequence. Figure 25.3 is a satellite image indicating the principal vertebrate fossil producing areas in the Maragheh Basin. Vertebrate fossil localities crop out across the Maragheh Basin and often are expressed as dense concentrations of fossils up to a meter in thickness and extending tens to hundreds of meters laterally. Th is is particularly true for the Upper Maragheh locality MMTT 13 near the village of Shol’avand. Since the latter part of the nineteenth century, coincident with the Austrian exploration of the Maragheh Basin, there has been a growing archive for the stratigraphic provenance of the Maragheh fauna (Bernor 1986). The work by Japanese, Dutch, German, and American groups in the 1970s brought marked improvements to this stratigraphic record. Figure 25.2 provides a summary of eight stratigraphic sections, arrayed from west to east, of these principal collecting areas with the University of California, Riverside (UCR)–MMTT localities (Bernor 1986) and newly discovered INSPE localities indicated. Bernor (1978, 1986) integrated all known stratigraphic records of fossil mammals to develop the fi rst Maragheh mammalian biostratigraphy. He originally subdivided the Maragheh Formation into three units based on the stage of evolution of the Hipparion s.s. lineage: “Lower Maragheh” whose base was defi ned by the fi rst occurrence of Hipparion gettyi at Kopran; “Middle Maragheh” by the fi rst occurrence of Hipparion prostylum; and “Upper Maragheh” by the fi rst occurrence of Hipparion campbelli. Recent studies of the Maragheh Hipparion samples housed by the Muséum National d’Histoire Naturelle, Paris (MNHN), and Howard University Laboratory of Evolutionary Biology suggest that Hipparion prostylum, originally defi ned based on skull morphology alone, may not occur at Maragheh. The MNHN sample originally referred to Hipparion prostylum (Woodburne and Bernor 1980; Bernor, Woodburne, and Van Couvering 1980; Bernor 1985) is not supported by the postcranial material: there are no metapodials or phalanges of Hipparion prostylum s.s (type species from Mount Luberon, France) in the MNHN sample. On the other hand, Hipparion campbelli’s postcrania (Howard University Maragheh sample) are morphologically similar to


551

Figure 25.3 Geographic sites and fossil localities (UCR- MMTT, MMTT, and INSPE) of the Maragheh Basin, northwest 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. |—x—| correspond to the stratigraphic columns of figure 25.2.

Hipparion prostylum s.s. The postcrania in the collections of the MNHN and the Bayerische Staatssammlung für Paläontologie und Historische Geologie, Munich (BSP), are referable to Hippotherium brachypus; they co-occur with skulls that have a reduced preorbital fossa like

H. prostylum. Th is consistent co-occurrence in two distinct paleontologic collections suggests that it is possible that these skulls and postcrania may be of the same taxon, Hippotherium brachypus. We further fi nd that the MNHN Maragheh Hipparion assemblage includes two

Figure 25.4 Mammalian biostratigraphy of the Maragheh Formation, northwest Iran. The stratigraphic provenance of vertebrate localities is given above/below the “Loose Chippings” (LC) marker bed. Taxa with * are also recorded from “Upper Maragheh” Loc. 37.

554


distinct skull morphologies and two distinct postcranial morphologies. The skulls with a large and single preorbital fossa (POF) placed close to the orbit are believed to be associated with the elongate and slender metapodials and likely referable to Cremohipparion aff . C. moldavicum, whereas the skull with a single and highly reduced POF is likely associated with the short, stout metapodials referable to Hippotherium brachypus. In that the MNHN assemblage is believed to be stratigraphically derived mostly from the middle portion of the section (“Middle Maragheh”), we now need to recognize that the biostratigraphic subdivisions may not be attributable to the evolution of a continuous lineage, but simply as biozones. Current evidence suggests the following horse sequence: Hipparion gettyi occurs at Kopran, the oldest set of localities in the Maragheh Basin with its stratigraphic range being from the –150 m to –75 m interval of the section (“–” and “+” refer to below and above the level of the “Loose Chippings”); Hippotherium brachypus and Cremohipparion moldavicum occur in the –52 m to –25 m interval of the section; Hipparion campbelli is fi rst known to occur at the –20 m interval and is believed to be present at the +7 m interval of the section. Small horses that we believe to be best referred to Cremohipparion?matthewi occur from about the –115 m to + 7 m interval of the section. There are likely multiple species of small Cremohipparion from Western Eurasia (i.e., Cremohipparion matthewi, Cremohipparion nikosi, Cremohipparion minus, and potentially others), and there is simply too little cranial and complete metapodial data to determine which of these occur at Maragheh. The Maragheh hipparion assemblage is numerically abundant and diverse in species and is undergoing extensive new systematic analysis by Mirzaie Ataabadi, Bernor, and Wolf (in progress). Figure 25.4 updates Bernor’s (1986) and Bernor et al.’s (1996) biostratigraphy of the Maragheh mammal fauna. Swisher (1996) provided new single crystal Argon ages for the Maragheh fauna that significantly revised Campbell et al.’s (1980) initial report on zircon fi ssion track ages and K/Ar40 ages. These are summarized, by stratigraphic interval, in table 25.1, which shows that the ages are internally coherent with the oldest being stratigraphically succeeded by progressively younger ages. Bernor et al. (1996) suggested that based on estimated sedimentation rates the Maragheh fauna ranges from about 9 Ma to 7.4 Ma. Table 25.2 represents a calculation of ages for localities in the western and central portion of

the sequence based on the estimated sedimentation rates (0.008 million yr/m). Thus, the oldest locality is Kopran I (8.96 Ma) and the youngest is MMTT 26 (7.68 Ma), located 7 m above the “Loose Chippings” marker bed. Figure 25.5 summarizes the geochronology of the Maragheh Basin and shows that the Ilkhchi locality, which is located in an eastern section, calibrates between 7.58 ± 0.11 Ma (R8) and 7.42 ± 0.11 Ma (R10). Table 25.3 provides a summary of Maragheh mammalian taxa by biostratigraphic interval as originally defi ned by Bernor (1986). Here we update a number of the mammalian groups. Taxonomic changes include the following: within the Hyaenidae, we recognize Adcrocuta (not Percrocuta) eximia and Hyaenictitherium wongii; the large machairodont cat is now recognized as Amphimachairodus (not Machairodus) aphanistus (L. Werdelin, pers. comm.); within the Proboscidea, we now recognize the deinothere as Deinotherium gigantissimum following the specimens discovered at MMTT31 (Erdbrink et al. 1976; locality K1) by Schmidt-Kittler; the equids are as described earlier; among the Rhinocerotidae, we recognize Iranotherium morgani and Rhinocerotidae n. gen. and sp. for Maragheh (I. Giaourtsakis, pers. comm.); the single suid occurring at Maragheh corresponds to a population of small/medium-size Microstonyx major; and for the giraffids, we currently recognize Helladotherium duvernoyi, Samotherium neumayri, Palaeotragus coelophrys and Bohlinia attica (not the specimen mentioned by de Mecquenem [1924–1925:pl. II, fig. 3]), but a revision of the material is certainly needed in order to clarify chronogeographic relationships. Additionally, the Bovidae are extensively revised (Kostopoulos and Bernor, 2011), and we recognize the following taxa for Maragheh: Gazella capricornis (not cf. G. deperdita), Gazella cf. G. ancyrensis, Demecquenemia n. gen. (not Gazella) rodleri, Oioceros atropatenes, Oioceros rothii, Nisidorcas sp., Prostrepsiceros houtumschindleri, Prostrepsiceros cf. P. vinayaki, Prostrepsiceros fraasi, Prostrepsiceros cf. P. rotundicornis, Protragelaphus skouzesi, cf. Palaeoreas sp., Skoufotragus laticeps, Palaeoryx sp., Urmiatherium polaki, Miotragocerus cf. M. maius, Miotragocerus sp., Tragoportax cf. T. amalthea (not Miotragocerus rugosifrons), and Samokeros minotaurus. The additional presence of Skoufotragus schlosseri (=Pachytragus crassicornis) is possible but not substantiated by the current revision. We also recognize the presence of Hystrix in Maragheh based on material in MNHN. In summary, the Maragheh fauna has a chronologic range of nearly 9 Ma to less than 7.4 Ma, but the bulk of


555

Table 25.1 Summary of the Isotopic Age Determinations from the Maragheh and Basal Tuff Formations Ar-Ar

+ 100 + 90 est. + 60 est. +15 est. + 15 est. +7

0

−50

−110

Korde-deh Pumice (Murdag Chai) Upper Pumice (Ilkchi) Middle Pumice (Ilkchi) Lower Pumice (Ilkchi) Village Pumice (Ilkchi) Layered Marker Pumice (Shollovend) Loose Chippings Pumice (Shollovend) Gürt Dareseh Pumice (Murdag) Ignimbritic Tuff (Murdag) Basal Tuff (western area) Basal Tuff (Ilkchi)

Weighted Mean

Isochron Age

Sample

7.420 ± 0.107

7.536 ± 0.111 (11 pt.)

R 10

7.579 ± 0.106

7.550 ± 0.071 (8 pt.)

Zircon FT

K-Ar

Sample

Sample

7.2 ± 0.4

R 10

7.5 ± 0.4

R 10pl

7.3 ± 0.5 R

R5 R8 R8 R9 R9 R 12

7.7 ± 0.5 7.2 ± 0.5 7.9 ± 0.7

R 5pl R 6pl R 8pl

5.5 ± 1.0

R 9pl

6.4 ± 0.6

R 12pl

7.592 ± 0.094

7.642 ± 0.033 (8 pt.)

R 12

7.4 ± 0.4 7.0 ± 0.4 5.2 ± 0.3 5.3 ± 0.3 7.7 ± 0.5

7.748 ± 0.136

7.787 ± 0.139 (8 pt.)

R 11

7.8 ± 0.4

R 11

7.4 ± 0.3

R 11pl

8.667 ± 0.040

8.635 ± 0.029 (9 pt.)

R3

10.6 ± 0.8

R3

10.391 ± 0.024

R4

11.2 ± 0.6

R4

10.432 ± 0.020

R7

12.8 ± 0.5

R7

6.4 ± 0.5 8.8 ± 0.5 8.9 ± 0.5 10.1 ± 0.5 9.3 ± 0.1 11.3 ± 1.0 9.3 ± 0.1

R 3pl R 3pl R 3pl R 4bi R 4an R 7bi R 7an

R8

Source: After Bernor (1986), Swisher (1996); Ar-Ar (Swisher 1996); Zircon FT and K-Ar (Campbell et al. 1980).

fossil material is from the middle and upper parts of the fossiliferous section (“Middle and Upper Maragheh”). The “Lower Maragheh” fauna is mostly composed of taxa with very long time distributions that cannot be used in fi ne time-resolution interregional comparisons (e.g., Kostopoulos and Bernor, 2011). For the bestknown “Middle Maragheh” localities (1A, 1B, and 1C), where several groups have collected fossils, the current oldest age is for locality 1A, 8.16 Ma (see table 25.2 [estimated]), and the youngest localities (“Upper Maragheh”) in the Shol’avand area (MMTT 26) date to 7.68 Ma (see table 25.2 [estimated]). To the east, the youngest localities of Ilkhchi (MMTT 37) would appear to be ca. 7.4 Ma.

The biochronological correlation of Maragheh with Samos and Pikermi, the classical mammal fossil localities of the Eurasian Late Miocene, is still open to discussion. Although this problem has been previously addressed (Bernor et al. 1996) and local biostratigraphy and geochronology of Maragheh and Samos have been greatly improved in the past years (Bernor et al. 1996; Swisher 1996; Kostopoulos, Sen, and Koufos 2003), the age of Pikermi remains badly resolved, being dated only indirectly. According to the latest magnetostratigraphic correlation, the Samos fauna ranges from ca. 7.8 Ma to 6.7 Ma, but the core of the Samos fauna referred to as the Samos Dominant Faunal Assemblage (DMAS) is dated

556


Table 25.2 Locality Ages of UCR– MMTT Fossil Localities of the Maragheh Formation, Northwest Iran, Inferred from the Estimated Sedimentation Rates

UCR– MMTT Locality 26 30 39 24 32 38 22 16 47 27 46 25 50 31 33 13 21 51* 40 7* 52 6 1C 18 19 20 34 1B 2 4 17 35 3 1A 5 8* 42* 36* 23

Level (m) from Loose Chippings 7 4 4 0 −1 −2 −3 −5 −6 −6 −10 −12 −12 −15 −17 −17 −18 −18 −20 −20 −28 −28 −29 −30 −30 −30 −30 −30 −32 −32 −35 −35 −38 −40 −52 −52 −60 −60 −70 −72

District in Maragheh Area Sholavand Sholavand Sholavand Sholavand Sholavand West Sholavand West Sholavand East Sholavand West Sholavand West Sholavand Sholavand West Sholavand West Gort Daresi Sholavand West Sholavand Sholavand West Sholavand West Moghanjeq Sholavand Sulu Dere Sholavand West Gort Daresi Gort Daresi Sholavand West Sholavand West Sholavand West Sholavand West Gort Daresi Gort Daresi Gort Daresi Sholavand West Sargezeh Gort Daresi Gort Daresi Gort Daresi Aliabad Aliabad Aliabad Gort Daresi

Estimated Age (Ma) 7.68274 7.70728 7.70728 7.74* 7.74818 7.75636 7.76454 7.7809 7.78908 7.78908 7.8218 7.83816 7.83816 7.8627 7.87906 7.87906 7.88724 7.88724 7.9036 7.9036 7.96904 7.96904 7.97722 7.9854 7.9854 7.9854 7.9854 7.9854 8.00176 8.00176 8.0263 8.0263 8.05084 8.0672 8.16536 8.16536 8.2308 8.2308 8.3126 8.32896

28

−80 −110 −120 −150

Moghanjeq Kopran II Kopran I

8.3944 8.64* 8.7216 8.967

Note: Ages with * are isotopic age determinations of zero level, which corresponds to “Loose Chippings,” and –110 level, which corresponds to Mordaq (Murdag) Ignimbritic tuff (see also figure 25.5 and table 25.1). Localities with * have an estimated level with respect to “Loose Chippings.”

between 7.2 Ma and 6.9 Ma (Kostopoulos, Sen, and Koufos 2003; Koufos, Kostopoulos, and Vlachou 2009). Th is chronology (figure 25.6) would make “Middle and Upper Maragheh” generally correlative with the lower fossil horizons (PMAS) at Samos (7.8– 7.4 Ma), but the Maragheh localities, as currently understood, appear somewhat older than those producing the Samos Intermediary (IMAS) and Dominant Mammal Assemblages (7.4– 6.9 Ma; Koufos, Kostopoulos, and Vlachou 2009). However, this conclusion is not consistent with the evolutionary stage of several mammalian taxa and their combined presence in both Samos and Maragheh (i.e., Melodon [=Parataxidea] maraghanus, Hyaenictitherium wongii, Adcrocuta eximia, Hippotherium brachypus, “Cremohipparion” cf. C. matthewi, Diceros neumayri, Skoufotragus laticeps, Miotragocerus ex. gr. valenciennesi, Gazella capricornis, Prostrepsiceros fraasi, Tragoportax, Palaeoryx, Protragelaphus, Palaeotragus, Samotherium), which dates to the 7.4– 6.9 Ma interval of Samos but appears to be older in “Middle and Upper Maragheh” intervals (for a detailed discussion, see Kostopoulos and Bernor 2011). The Maragheh mammal association as it is shown in figure 25.4 is in part constructed from the old provenance data of the Vienna (NMW) and the Paris (MNHN) collections developed in Bernor (1986). However, the Tobien collection (BSP) does have stratigraphic provenance and was collected from the same part of the stratigraphic sequence where MMTT localities 1A–1C were collected (H. Tobien, pers. comm.). One of us (RLB) correlates “Middle Maragheh,” 8.1– 7.9 Ma (estimated), with Pikermi based principally on the occurrence of Hippotherium brachypus, which is very similar to the Pikermi form, and Cremohipparion moldavicum, which is the sister taxon to Cremohipparion mediterraneum, also known from Pikermi (Koufos 1987). Another (DSK) holds that the apparent faunal relationships between Pikermi and Maragheh, also evidenced by several artiodactyl taxa, do not confi rm such ages, as the

Plate 3.1 The stratigraphy of the Tieersihabahe section, Junggar Basin, Xinjiang. Modified from Meng et al. (2006). The blue arrows and line point to the base of the Xiejian defined by the lowest occurrence of Democricetodon sp. nov. The red arrows and line indicate the base of the Xiejian defined by the polarity timescale (base of C6Cn.2n). The black arrow shows the occurrence of Metexallerix junggarensis. For a complete list of fauna, see Meng et al. (2006). Plate 7.1 Summary of mesowear, hypsodonty, and stable isotope data for herbivores from localities 49 and 30. HYP = hypsodonty, mswr = mesowear, with “+” indicating a more abrasion- dominated pattern (roughly corresponding to more grazing) and “ ” indicating a more attritiondominated pattern (roughly corresponding to more browsing). Green shading indicates pure C3 diets based on carbon isotopic analyses of tooth enamels, and red shading indicates a clear C4 dietary component. Block arrows depict animals that occur both at Locality 49 and Locality 30, while rectangles depict animals that occur only at one of the localities. Occurrence data (%ocr) are taken from Kurtén (1952) and are calculated as the number of identifiable individuals from a taxon divided by the total number of individuals recovered from the locality. Relative stratigraphic positions are based on results presented in this chapter. Data are from Passey (2007) and our unpublished work.

Plate 22.1 Insectivora and Rodentia from the Tagay and Saray I sections of Olkhon Island (Lake Baikal, Siberia), Kalagay Formation. To facilitate comparisons, all right side teeth of small mammals are figured as mirror images, and their figure numbers are underlined. Desmaninae gen. et sp. indet.—Tagay section, Middle Miocene (MN 7/8): (1) right M1, NHMW2009z0067/0001; (2) left M3, NHMW2009z0067/0002; (3) left M3 (fragmentary), NHMW2009z0067/0003. Erinaceidae gen. et sp. indet.—Tagay section, Middle Miocene (MN 7/8): (4) right m2 (fragmentary), NHMW2009z0068/0001 Sciurinae gen. et sp. indet.—Tagay section, Middle Miocene (MN 7/8): (5) left P4, NHMW2009z0069/0001. Miodyromys sp. (Gliridae)—Tagay section, Middle Miocene (MN 7/8): (6) right D4/P4, NHMW2009z0070/0001; (7) right M1/2, NHMW2009z0070/0002. Keramidomys aff. mohleri Engesser 1972 vel Keramidomys aff. fahlbuschi, Qiu 1996 (Eomyidae)—Tagay section, Middle Miocene (MN 7/8): (8) left M2, NHMW2009z0071/0001; (9) right M3, NHMW2009z0071/0002; (10) left m1/2, NHMW2009z0071/0003; (11) left m1/2, NHMW2009z0071/0004; (12) left m3, NHMW2009z0071/0005; (13) right m1/2, NHMW2009z0071/0006; (14) right m1/2, NHMW2009z0071/0007; (15) right m1/2, NHMW2009z0071/0008. Eomyops oppligeri Engesser, 1990 (Eomyidae)—Tagay section, Middle Miocene (MN 7/8): (16) left m1/2, NHMW2009z0072/0008. Democricetodon sp. (Cricetidae)—Tagay section, Middle Miocene (MN 7/8): (17) left M2, NHMW2009z0073/0001; (18) left m3, NHMW2009z0073/0002; (19) left m1, NHMW2009z0073/0003. Eozapus intermedius Bachmayer and Wilson, 1970 (Zapodidae)—Saray I section, Late Miocene: (20a) right mandible with m1–3 (occlusal), NHMW2009z0074/0001; (20b) right mandible with m1–3 (labial), NHMW2009z0074/0001. Magnifications: 25×; (all specimens in the NHMW collection Inv. Nr. NHMW2009z0067– 074).

Plate 22.2 Gastropoda and ectothermic vertebrates from the Tagay and Saray I sections of Olkhon Island (Lake Baikal, Siberia); Kalagay Formation. Scale bar for figures 1–10 is 1 mm; scale bar for figures 11–17 is 500 μm. (1) Leuciscinae indet., pharyngeal tooth, Tagay; (2) Palaeotinca sp., pharyngeal tooth, Tagay; (3) Palaeocarassius sp., pharyngeal tooth, Tagay; (4) Palaeocarassius sp., pharyngeal tooth, A1 position, Tagay; (5) Palaeocarassius sp., pharyngeal tooth germ, Tagay; (6) Esox sp., palatine, Tagay; (7) Esox sp., tooth, Tagay; (8) Rana sp. (R. temporaria group), right ilium, Tagay; (9) Bufo aff. calamita (Laurenti 1768), left ilium, Saray I; (10) ?Chalcides nov. sp., right dental, Tagay; (11) Vallonia subcyclophorella (Gottschick 1911), apical view, Saray I-140899- 8; (12) Vallonia tokunagai Suzuki, 1944, apical view, Saray I-140899-11; (13) Vertigo (Ungulidenta) ancata Steklov 1967, apertural view, Saray I-140899- 8; (14) Gastrocopta (Kazachalbinula) cf. ukrainica Steklov 1966, apertural view, Saray I-140899- 8; (15) Carychium sp., apertural view, Saray I-140899-10; (16) Gastrocopta (Sinalbulina) intorta Steklov 1967, apertural view, Saray I-140899- 8; (17) Radix sp., lateral view of an incomplete juvenile shell, Saray I-140899-11.

Plate 26.4 Early Miocene Kaplangı 1–2. Megacricetodon primitivus: (1) M1. Debruijnia sp.: (2) m2. Mirabella tuberosa: (3) M2. Democricetodon cf. franconicus: (4) M1. Cricetodon tobieni: (5) M1. Eumyarion sp.: (6) M1. Anomalomys cf. aliveriensis: (7) M1. Keseköy. Megacricetodon sp: (8) M1; (9) m1. Debruijnia arpati: (10) M1. Mirabela crenulata: (11) M1. Enginia gertcheki: (12) M1. Enginia djanpolati: (13) M1. Democricetodon doukasi: (14) M1. Cricetodon kasapigili: (15) M1 Eumyarion montanus: (16) M1. Eumyarion intercentralis: (17) M1. Vallaris zappai: (18) M1; (19) m1. Harami 1. Mirabella anatolica: (20) M1. Deperetomys intermedius: (21) M1. Democricetodon doukasi: (22) M1. Eumyarion microps: (23) M1. Eumyarion carbonicus: (24) M1.

Kılçak 0–3. Enginia beckerplatini: (25) M1. Enginia djanpolati: (26) M2. Meteamys alpani: (27) M2. Deperetomys anatolicus: (28) M1. Spanocricetodon sinuosus: (29) M1. Democricetodon anatolicus: (30) M1. Cricetodon sp.: (31) M1. Cricetodon versteegi: (32) M1. Eumyarion microps: (33) M1. Eumyarion cf. carbonicus: (34) M1. Heterosminthus cf. nanus: (35) m1; (36) m2; (37) m2; (38) M1; (39) M2. Kargı 2. Enginia djanpolati: (40) M1. Meteamys alpani: (41) M1. Deperetomys anatolicus: (42) M1. Spanocricetodon sinuosus: (43) M1. Cricetodon versteegi: (44) M1. Eumyarion sp.: (45) M1. Melissiodon sp.: (46) M3. Muhsinia sp.: (47) M1. Heterosminthus cf. firmus/lanzhouensis: (48) m1; (49) m2; (50) M1.

Plate 26.3 Middle Miocene Bag˘içi. Myocricetodon cf. liui: (1–2) M1; (3–5) m1; (4) M2. Cricetulodon aff. sabadellensis: (6) M1; (7) M2; (8) M3; (9) m1. Megacricetodon aff. collongensis: (10) M1; (11) m1. Pliospalax sp.: (12) M1. Byzantinia ozansoyi: (13) M1. Heterosminthus gansus: (14) M1; (15) M2; (16) M3; (17) m1; (18) m2; (19) m3.

Zambal. Megacricetodon cf. collongensis: (20) M1; (21) m1. Megacricetodon cf. gregarius: (22) m1. Cricetodon cf. pasalarensis: (23) M1. Cricetodon cf. hungaricus: (24) M1. Eumyarion sp.: (25) m2. Heterosminthus cf. mongoliensis: (26–27) M1; (28) M2. Pas˛alar. Megacricetodon andrewsi: (29) M1. Pliospalax marmarensis: (30) M1. Democricetodon brevis: (31) M1. Cricetodon pasalarensis: (32) M1. ?Protalactaga sp.: (33) M2.

Plate 26.2 Late Miocene Süleymanlı 2. Pseudomeriones abbreviatus: (1) M1; (2) M2; (3) M3; (4) m1; (5) m2; (6) m3. Hansdebruijnia neutrum: (7) M1; Apodemus gudrunae: (8) M1. Apodemus gorafensis: (9) M1.

Düzyayla. cf. Rhinocerodon sp.: (21) M1; (22) M2; (23) m1; (24); m2. Pseudomeriones sp.: (25) M1. Apodemus/Parapodemus sp. 1: (26) M1. Apodemus/Parapodemus sp. 2: (27) M1. Castromys littoralis: (28) M1; (31) m1. Hayranlı 1. cf. Rhinocerodon sp.: (50) M1; (51) M2; (52) M3; (53) m1; (54) m2; (55) m3. Pseudomeriones sp.: (56) M3. Parapodemus lugdunensis: (57) M1; (58) M2. Parapodemus n. sp.: (59) M1; (60) M2. Altıntas¸. cf. Calomyscus sp.: (70) M2; (71) m1; (72) m2; (73) m3. Abudhabia n. sp.: (74) M1; (75) M2; (76) M3; (77) m1; (78) m2; (79) m3. Progonomys cathalai: (80) M1; (81) M2; (82) M3; (83) m1; (84) m2; (85) m3. Myocricetodon n. sp.: (86) M1; (87) M2; (88) M3; (89) m1; (90) m2; (91) m3. Cricetulodon hartenbergeri: (92) M1; (94) m1.

Plate 26.2 Late Miocene (continued) Süleymanlı 2. Allocricetus sp.: (10) M1; (11) M2; (12) M3; (13) m1; (14) m2; (15) m3. Calomyscus delicatus: (16) M2; (17) m1; (18) m2. Pliospalax sp. 1: (19) M2. Pliospalax sp. 2: (20) M2. Düzyayla. Castromys littoralis: (29) M2; (30) M3. “Karnimata” provocator: (32) M1; (33) M2. Parapodemus lugunensis: (34) M1; (35) M2. Allocricetus sp.: (36) M1; (37) M2; (38) M3; (39) m1; (40) m2; (41) m3. Pliospalax sp.: (42) M1. Byzantinia hellenicus: (43) m1. Prospalax sp.: (44) M1. Eozapus intermedius/similis: (45) M1; (46) M2; (47) M3; (48) m1; (49) m2.

Hayranlı 1. Apodemus/Parapodemus sp.: (61) M1; (62) M2. Cricetulodon cf. sabadellensis: (63) M1; (64) M2; (65) M3; (66) m1; (67) m2; (68) m3. Byzantinia hellenicus: (69) M1. Altıntas¸. Cricetulodon cf. hartenbergeri: (93) M2; (95) m2; (96) m3. Allocricetus sp.: (97) M1; (98) M2; (99) M3; (100) m1; (101) m2; (102) m3. Pliospalax aff. marmarensis: (103) M1. Byzantinia uenayae: (104) M1. Byzantinia pikermiensis: (105) M1. Protalactaga minor: (106) P4; (107) M1; (108) M2; (109) M3; (110) m1; (111) m2; (112) m3.

Plate 26.1 Pliocene Hamamayag˘ı. Allophaiomys deucalion: (1) m1. Kalymnomys major: (2) m1. Allocricetus sp: (3) m1. Apodemus atavus: (4) M1. Apodemus dominans: (5) M1. Yenice. Pliomys greacus: (6) m1. Orientalomys similis: (7) M1; (8) m1; (9) M2; Apodemus dominans: (10) m1. Ortalıca. Mimomys gracilis: (11) m1. Pliomys sp.: (12) M3. Mesocricetus primitivus: (13) M2. Apodemus dominans: (14) M1. Ig˘deli. Promimomys insuliferus: (15) m1. “Cricetus” lophidens: (16) M1. Mesocricetus primitivus: (17) M2. Pseudomeriones n.sp.: (18) M1; (19) M2; (20) m1; (21) m2. Allocricetus cf. bursae: (22) M1. Micromys bendai/tedfordi: (23) M1. Apodemus gorafensis: (24) M1. Occitanomys debruijni: (25) M1.

Plate 29.1 Dice similarity index values at the genus level (Dice FRI) of the Eurasian localities compared to Tunggur-Moergen (A– G) and their mean ordinated hypsodonty (Hypso) values (H– N) mapped as a grid for MNEQ 3 to MNEQ 10 time intervals. (A) Dice FRI- MNEQ 3; (B) Dice FRIMNEQ 4; (C) Dice FRI- MNEQ 5; (D) Dice FRI- MNEQ 6; (E) Dice FRI- MNEQ 7+8; (F) Dice FRI- MNEQ 9; (G) Dice FRI- MNEQ 10. Yellow/red patterns in A– G indicates high similarity to Tunggur locality, while blue represents low similarity; (H) Hypso- MNEQ 3; (I) Hypso- MNEQ 4; (J) Hypso- MNEQ 5; (K) Hypso- MNEQ 6; (L) Hypso- MNEQ 7+8; (M) Hypso- MNEQ 9; (N) Hypso- MNEQ 10. Yellow/red pattern in H– N indicates increased aridity, while blue represents high humidity. SS = Sihong-Songlinzhuang, China; CC = Córcoles, Spain; BZ = Bézian, France; EMF = Esvres– Marine Faluns, France; KL = Kalkaman Lake, Kazakhstan; SW = Shanwang, China; TD = Tongxin- Dingjiaergou, China; HL = Hezheng- Laogou, China; S = Sansan, France; CB = Catakbagyaka, Turkey; AM = Ayibaligi Mevkii, Turkey; TM: Tunggur- Moergen, China; CM = Chiang Muan, Thailand; CP = Can Ponsic I, Spain; CL = Can Llobateres I, Spain.

Plate 29.2 Dice similarity index values at the genus level (Dice FRI) of the Eurasian localities compared to Baode locality 49 (A– G) and their mean ordinated hypsodonty (Hypso) values (H– N) mapped as a grid for MNEQ 9 to MNEQ 15 time intervals. (A) Dice FRI- MNEQ 9; (B) Dice FRI- MNEQ 10; (C) Dice FRI- MNEQ 11; (D) Dice FRI- MNEQ 12; (E) Dice FRI- MNEQ 13; (F) Dice FRI- MNEQ 14; (G) Dice FRI- MNEQ 15. Yellow/red pattern in A– G indicates high similarity to Baode loc. 49, while blue represents low similarity. (H) Hypso- MNEQ 9; (I) Hypso- MNEQ 10; (J) Hypso- MNEQ 11; (K) Hypso- MNEQ 12; (L) Hypso- MNEQ 13; (M) Hypso- MNEQ 14; (N) Hypso- MNEQ 15. Yellow/red pattern in H– N indicates increased aridity, while blue represents high humidity. S = Sinap, Turkey; SS = Subsol de Sabadell, Spain; G = Grebeniki, Ukraine; P = Poksheshty, Moldova; K = Karacahasan, Turkey; KT = Kemiklitepe 1–2, Turkey; G = Garkin, Turkey; T = Thermopigi, Greece; NE = Novaja Emetovka, Ukraine; L = Lantian, China; B = Baode locality 49, China; Y = Yushe, China.


557

Figure 25.5 Stratigraphic array of isotopic age determinations from the Maragheh Formation and Basal Tuff. After Bernor (1986) and Swisher (1996).

mammal association of Pikermi appears to be more advanced than those from Vathylakkos-2, Prochoma, and Perivolaki (Greece), magnetostratigraphically dated between 7.4 Ma and 7.2 Ma (Koufos et al. 2006). In addition, Hippotherium brachypus and Cremohipparion moldavicum are also present in Akkaşdaği (Turkey; dated 7.1 Ma), while the type locality of Cr. moldavicum is Taraklija of the late Turolian. Clearly, a conclusive correlation between the fauna of Maragheh, Samos, and Pikermi will have to await resolution of the apparent mismatch between the available geochronological and biochronological evidence. Without anticipating what the resolution will be, we note that the discrepancies can be explained by one or a combination of four possibilities: • Mistakes in the radiometric or magnetostratigraphic dating • Mismatches in the correlation of old localities with new stratigraphic evidence • Major diachrony in the occurrence of species between these localities • A mistaken attribution of ecomorphs to chronospecies

PALEOBIOGEOGRAPHY AND PALEOECOL OGY

We follow recent investigations on paleobiogeographic analysis by Bernor, Fortelius, and Rook (2001) and Fortelius et al. (1996) by undertaking genus-level faunal resemblance index (GFRI) studies using both the Simpson (1943) and the Dice (Sokal and Sneath 1963) indices. Dice FRI is highly recommended by Archer and Maples (1987) and Maples and Archer (1988) and is calculated as 2C/(A + B), where C is the number of shared taxa between two faunas and A and B are the total number of taxa present in fauna 1 and fauna 2, respectively. Simpson’s FRI also has a long tradition of use (Bernor 1978; Flynn 1986; Bernor and Pavlakis 1987) and is calculated as C/smaller of (A or B). Figure 25.7 illustrates the plot of GFRI in pairwise comparisons between “Middle and Upper Maragheh” and 10 other localities under consideration. Among the early Vallesian (MN 9) localities compared to Maragheh—the Western and Central European localities—all plot with an index value of less than 20% for both Dice and Simpson’s GFRI. However, Sinap loc.12 (hominid zone), Turkey, has nine genera in common with Maragheh and a Dice GFRI

Table 25.3 Mammalian Species, Biostratigraphic Intervals and Their Estimated Ages, and UCR-MMTT Fossil Localities of the Maragheh Formation, Northwest Iran Biostratigraphic Interval Distance from Zero Level (i.e., Loose Chippings) Estimated Age MN-equivalent zone UCR-MMTT fossil localities

Order Primates Linnaeus, 1758 Family Cercopithecidae Gray, 1821 Mesopithecus pentelici Wagner, 1839 Order Carnivora Bowdich, 1821 Family Ursidae Gray, 1825 Indarctos maraghanus Mecquenum, 1924 Family Mustelidae Swainson, 1835 Promeles palaeattica Weithofer, 1888 Melodon maraghanus Kittl, 1887 Parataxidea polaki Kittl, 1887 Martes sp. Family Hyaenidae Gray, 1869 Ictitherium viverrinum Roth and Wagner, 1854 Hyaenictitherium? wongii (Zdansky, 1924) Adcrocuta eximia (Kaup, 1828) Family Felidae Gray, 1821 Metailurus orientalis Zdansky, 1924 Felis attica Wagner, 1857 Amphimachairodus aphanistus (Kaup, 1832) Order Tubulidentata Huxley, 1872 Family Orycteropodidae Bonaparte, 1850 Orycteropus sp. Order Proboscidea Illiger, 1811 Family Gomphotheriidae Cabrera, 1929 Choerolophodon pentelici Gaudry, 1862 Family Deinotheriidae Bonaparte, 1845 Deinotherium gigantissimum Stefanescu, 1892 Order Perissodactyla Owen, 1848 Family Equidae Gray, 1821 “Hipparion” gettyi Bernor, 1985 Hippotherium brachypus Hensel, 1862 Hipparion campbelli Bernor, 1985 Cremohipparion aff . C. moldavicum Gromova, 1952 Cremohipparion aff . C. matthewi Kormos, 1911 Family Chalicotheriidae Gill, 1872 Ancylotherium pentelici (Gaudry, 1862) Family Rhinocerotidae owen, 1845 Diceros neumayri Mecquenem, 1905

“Lower Maragheh” (−150m to −52m) 8.96– 8.16 Ma. 10/11 8, 42, 45, 36, 23, 28, 41, 44, 43, 9, 48

“Middle Maragheh” (−52 m to −20 m) 8.16–7.9 Ma. 11 7, 14, 34, 1c, 6, 18, 19, 20, 1b, 2, 17, 11, 4, 35, 3, 1A, 5

“Upper Maragheh” (−25 m to +7 m) 7.9–7.4 Ma. 11/12 26, 39, 24, 32, 38, 22, 16, 47, 27, 25, 15, 49, 49a, 50, 31, 33, 21, 13, 37, 40, 51

X

X X X X X

X X

X X X

X X

X X X

X

X

X

X X

X X X X

X X

X X

X

Biostratigraphic Interval Distance from Zero Level (i.e., Loose Chippings) Estimated Age MN-equivalent zone UCR-MMTT fossil localities

Chilotherium persiae Pohlig, 1887 Iranotherium morgani (Mecquenem, 1908) Rhinocerotidae gen. and sp. nov. Order Artiodactyla Owen, 1848 Family Suidae Gray, 1821 Microstonyx major (Gervais, 1848) Family Cervidae Gray, 1821 Cervidae gen. and sp. indet. Family Giraffidae Gray, 1821 Bohlinia attica (Gaudry and Lartet, 1856) Palaeotragus coelophrys Rodler and Weithofer, 1890 Samotherium neumayri Rodler and Weithofer, 1890 Helladotherium duvernoyi (Gaudry and Lartet, 1856) Family Bovidae Gray, 1821 Miotragocerus cf. M. maius (Meladze, 1967) Miotragocerus sp. Gazella capricornis (Wagner, 1848) Gazella cf. G. ancyrensis Tekkaya, 1980 Demecquenemia rodleri (Pilgrim and Hopwood, 1928) Prostrepsiceros cf. P. rotundicornis Weithofer, 1888 Prostrepsiceros houtumschindleri (Rodler and Weithofer, 1890) Prostrepsiceros fraasi (Andree, 1926) Prostrepsiceros cf. P. vinayaki (Pilgrim, 1939) Protragelaphus skouzesi Dames, 1883 Skoufotragus laticeps (Andree, 1926) Tragoportax cf. T. amalthea (Roth and Wagner, 1854) Urmiatherium polaki Rodler, 1889 Oioceros atropatenes Rodler and Weithofer, 1890 Oioceros rothii Wagner, 1857 Samokeros minotaurus Solounias, 1981 Palaeoryx sp. Nisidorcas sp. cf. Palaeoreas sp. Order Rodentia Bowdich, 1821 Family Hystricidae Burnett, 1830 Hystrix sp. Source: Modified after Bernor (1986).

“Lower Maragheh” (−150m to −52m) 8.96– 8.16 Ma. 10/11 8, 42, 45, 36, 23, 28, 41, 44, 43, 9, 48

“Middle Maragheh” (−52 m to −20 m) 8.16–7.9 Ma. 11 7, 14, 34, 1c, 6, 18, 19, 20, 1b, 2, 17, 11, 4, 35, 3, 1A, 5

X

X X X

X

X

“Upper Maragheh” (−25 m to +7 m) 7.9–7.4 Ma. 11/12 26, 39, 24, 32, 38, 22, 16, 47, 27, 25, 15, 49, 49a, 50, 31, 33, 21, 13, 37, 40, 51

X

X

X X

X

X X X X X X X X X X X X X X X X X X X X X X X

X

X X X

X X

X X X X X X

560


Figure 25.6 Chronostratigraphic position of Maragheh (MAR), Samos (SAM), and Pikermi (PIK) fossiliferous sites and faunal assemblages according to data presented in this chapter and Koufos, Kostopoulos, and Vlachou (2009).

of 30% and Simpson’s GFRI of 47%, which is the highest among these early Vallesian localities. The shared genera between Sinap loc. 12 and Maragheh are Chilotherium, Choerolophodon, Criotherium, Gazella, Hipparion, Orycteropus, Palaeoreas, Palaeotragus, and Tragoportax. We also used ungulate tooth crown height (Bernor, Fortelius, and Rook 2001) to show the contrast between the localities under consideration. The three-part subdivision of crown height includes the following: brachydont, whereby crown length of the upper second molar

(M2) is greater than its crown height; mesodont, whereby M2 crown length is roughly the same as crown height; and hypsodont, whereby M2 crown height is more than two times that of crown length. All data have been downloaded from the NOW (Neogene of Old World) database on September 30, 2009 (Fortelius 2009; http://www.helsinki.fi /science/now/). Our analyses here consider only large mammals because small mammal records vary greatly across the faunas as a result of taphonomic and sampling bias.


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Figure 25.7 Genus-level Faunal Resemblance Indices (GFRI); pairwise comparison of Maragheh with localities under consideration. MN 9: Can Llobateres, Spain; Eppelsheim, Germany; Rudabanya, Hungary; Sinap loc. 12, Turkey. MN 12: Samos and Pikermi, Greece; Baode loc. 49, China. MN 13 equivalent: Baode loc. 30, China; Sahabi, Libya, and Middle Awash, Ethiopia.

Figure 25.8 Crown height diagrams for Maragheh and Vallesian/Turolian localities under consideration. MN 9: Can Llobateres, Spain; Eppelsheim, Germany; Rudabanya, Hungary; Sinap loc. 12, Turkey. MN 12: Samos and Pikermi, Greece; Maragheh, Iran; Baode loc. 49, China. MN 13 equivalent: Baode loc. 30, China; Sahabi, Libya and Middle Awash, Ethiopia.

Analysis of crown height (figure 25.8) also demonstrates that Western and Central European (MN 9) localities of Rudabanya, Eppelsheim, and Can Llobateres have low incidence of mesodont and hypsodont forms (5–10% of mesodont and 5–10% hypsodont taxa) com-

pared to more than 85% of brachydonts. Th is suggests that Central and Western Europe during the Vallesian was forested with warm climates and low seasonality. The hypsodonty in these faunas is only due to occurrence of the invasive species of hipparionine horses (Hippotherium)

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that show adaptation to closed environments with a significant amount of browsing diets (Bernor, Kordos, and Rook 2003). In remarkable contrast is Sinap loc. 12, showing stronger mesodonty and hypsodonty signals. Sinap loc. 12 exhibits 35 % brachydonts, 28% mesodonts, and 35% hypsodont forms. As demonstrated before (Fortelius et al. 2003), later Vallesian Western Eurasian faunas are very similar in their community structure to regional Turolian faunas. They might be called “proto-Pikermian,” showing the origin of the “Pikermian Chronofauna,” which is best characterized by the Maragheh–Pikermi– Samos triad (Eronen et al. 2009). Maragheh’s closest resemblance is to the Greek locality of Samos. Maragheh has 28 genera (22 species) in common with Samos, and its GFRI is 68% for Dice and 74% for Simpson. The next closest relationship of Maragheh is clearly to Pikermi, Greece. Maragheh shares 28 genera (20 species) with this locality, and its GFRI is 61% for Dice and 63% for Simpson. These classical localities of the “Pikermian Chronofauna” show stable community structures. Maragheh and Samos have crown-height percentages among its ungulate taxa as follows: about 45– 35% of brachydont, 30% of mesodonts, and 25–35% of hypsodonts (see figure 25.8). On the other hand, Pikermi has a lower percentage of hypsodont forms, which reflects a more wooded setting than Samos and perhaps much of Maragheh (Solounias et al. 1999). Recently, Kostopoulos (2009) argued that the paleoecological profi le in Pikermi includes the tree-dwelling semi-terrestrial primate Mesopithecus, diversified felids and mustelids, and browsedependent proboscideans, rhinos, giraffes, and bovids. In contrast, Samos has no primate and browse-dependent taxa and has diversified hipparionine horses and gazelles with grazing rhinos, giraffes, and bovids. Maragheh paleoecological profi le is remarkable in this context by having a mixture of Samos and Pikermi characters (see also Koufos, Kostopoulos, and Merceron 2009:fig.7). In addition to primates, there are diversified felids, mustelids and hipparionine horses present in Maragheh. Browsing and grazing proboscideans, rhinos, giraffes, and bovids also occur in Maragheh. Therefore, although Maragheh is somewhat more similar to Samos than it is to Pikermi (see figure 25.7), environmentally it indicates more wooded settings in its dominant grass/bushy vegetation. For this reason, as mentioned by Strömberg et al. (2007), an east to west climatic and vegetational gradient across the Greco-Iranian province is not clearly evident. In the Middle Turolian, great intercontinental dispersion of large carnivores and ungulates of the “Pikermian chronofauna” evidently occurred (Kostopoulos 2009), and some of the core Pikermian genera extended their

geographic range so that a sizable number of genera are shared among western Eurasian members of the “Pikermian chronofauna” (Eronen et al. 2009), Chinese MN 13–equivalant faunas (Mirzaie Ataabadi et al., chapter 29, this volume), and the terminal Miocene northern and eastern African faunas (Bernor and Rook 2008, Bernor, Rook, and Haile-Selassie 2009). Baode loc. 30 (MN 13 equivalent in China) has 13 genera in common with Maragheh and a Dice GFRI of 40% and Simpson’s GFRI of 56%. Baode loc. 49 (MN 12 equivalent in China) has 11 genera in common with Maragheh and its Dice GFRI is 32% and Simpson GFRI 41%. North African (Libyan) locality of Sahabi shows a similar level of similarity. Sahabi has 9 genera in common with Maragheh, and its Dice GFRI is 27% and Simpson’s GFRI 37%. The Baode loc. 49 and 30 are considered to be eastern extensions of the “Pikermian Chronofauna” (Mirzaie Ataabadi et al., chapter 29, this volume), having exhibited Pikermian-type community structures. The MN 12–equivalent locality of Baode 49 is similar to Pikermi, with about 12% hypsodont taxa, 30% mesodonts, and 59 % brachydonts. On the other hand, the MN 13–equivalent locality of Baode 30 is very similar to Maragheh and Samos, with about 33% hypsodont taxa (see figure 25.8). Th is locality is also more similar to Maragheh than locality 49 in terms of GFRI (see figure 25.7). Taxa shared between Baode localities and Maragheh are as follows: Adcrocuta, Amphimachairodus, Chilotherium, Cremohipparion, Felis, Gazella, Hipparion, Ictitherium, Indarctos, Metailurus, Palaeoryx, Palaeotragus, Parataxidea, Samotherium, Hyaenictitherium, and Urmiatherium. The large elasmothere rhinocerotid, Iranotherium morgani, also occurs in the early Late Miocene of the Linxia Basin, northwestern China (Deng 2005). Th is species has apparently fi rst appeared in northwestern China during the Vallesian and immigrated to Maragheh later in the Turolian. The terminal Miocene Libyan locality of Sahabi is also similar in community structure to Maragheh. A high percentage of hypsodont taxa (32%) occurs, with 23% mesodont and 45% brachydont forms in this locality, which is as similar to Maragheh, as are the Chinese Baode localities (see figure 25.7). Taxa shared between Sahabi and Maragheh are as follows: Adcrocuta, Amphimachairodus, Ceratotherium (Diceros), Cremohipparion and possibly Hipparion, Gazella, Indarctos, Prostrepsiceros, Samotherium, and Tragoportax. The Late Miocene Middle Awash fauna (Ethiopia) is less than 20% similar to Maragheh in terms of both Dice and Simpson GFRIs. However, it has a somewhat similar community structure, but with fewer hypsodont and more brachydont taxa. These African faunas clearly had


“Pikermian” elements that were vicariant and evolved independently subsequent to an early–middle Turolian extension (Bernor and Rook 2008; Bernor, Rook, and Haile-Selassie 2009).

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and helpful comments. Z. Davoudifard extensively helped in preparation of several illustrations for this paper.

REFERENCES CONCLUSION

Apart from widely distributed “Pikermian Chronofauna” taxa (e.g., Adcrocuta, Amphimachairodus, Deinotherium, Felis, Hipparion, Hyaenictitherium, Hystrix, Ictitherium, Indarctos, Metailurus, Palaeotragus, and Samotherium), the giraffid Bohlinia, bovids such as Demecquenemia rodleri, the large Palaeoryx, and Miotragocerus cf. M. maius, and hipparionine horses such as Cremohipparion moldavicum indicate affi nities of the Maragheh fauna to the Northern Black Sea region. Some taxa, such as Prostrepsiceros cf. P. vinayaki, show relationships with the western Asian (Arabia, Afghan istan, and the Indian Subcontinent) region, whereas Hippotherium brachypus, Hipparion campbelli (sister taxon to Samos Hipparion dietrichi), Cremohipparion matthewi, Gazella cf. G. ancyrensis, Prostrepsiceros fraasi/houtumschindleri, Samokeros, and Skoufotragus suggest affi liations with those from Anatolia and Samos. Urmiatherium and Iranotherium are taxa in common with China. Regardless of the absence of real-time resolution and the mismatch between some geochronological and biochronological data, it is evident that the Maragheh area was affected by biogeograph ically distinct Late Miocene areas, representing the crossroads of several Old World provinces. AC KNOW LEDG MENTS

We appreciate the efforts of Dr. D. Najafi Hajipour and Dr. S. Montazami and their predecessors in Iran’s Department of Environment for reinitiating and supporting new excavations in the Maragheh fossiliferous area. We also thank the Governor, the Mayor, and the local DOE office managers in Maragheh and Tabriz for their hospitality and assistance during INSPE fieldworks. MMA, MF, and fieldwork in Maragheh were partially supported by the Academy of Finland, RHOI project, and the Sasakawa Foundation. RLB wishes to acknowledge the National Science Foundation, including EAR- 0125009 (grant to R. L. Bernor and M. O. Woodburne), BCS- 0321893 (grant to F. C. Howell and T. D. White), and the Sedimentary Geology and Paleobiology Program (GEO: EAR: SEP) for supporting his research on this project. We thank Zhaoqun Zhang and the volume editors for their reviews

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