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Mammalian biostratigraphy, geochronology, andzoogeographic relationships of the Late MioceneMaragheh fauna, IranRaymond L. Bernor aa College of Medicine, Howard University, Department of Anatomy, Laboratory ofPaleobiology, Washington, D.C., 20059

Available online: 24 Aug 2010

To cite this article: Raymond L. Bernor (1986): Mammalian biostratigraphy, geochronology, and zoogeographicrelationships of the Late Miocene Maragheh fauna, Iran, Journal of Vertebrate Paleontology, 6:1, 76-95

To link to this article: http://dx.doi.org/10.1080/02724634.1986.10011600

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Journal of Vertebrate Paleontology 6(1):76-95, March 1986

MAMMALIAN BIOSTRATIGRAPHY, GEOCHRONOLOGY, AND ZOOGEOGRAPHICRELATIONSHIPS OF THE LATE MIOCENE MARAGHEH FAUNA, IRAN

RAYMOND L. BERNORCollege of Medicine, Department of Anatomy, Laboratory of Paleobiology,

Howard University, Washington D.C. 20059

ABSTRACT- The classical Late Miocene Maragheh fauna has been collectedand studied sporadicallyfor nearly 150years.This study givesa comprehensive account of the entire mammalian fauna recoveredat Maragheh to date and its biostratigraphic, biogeographicand geochronologiccontexts. The sequenceis divided into Lower, Middle and Upper biostratigraphic intervals, ranging from ca. 9.5 my to 7 myin age, based on the first appearance of a potential evolutionary series of hipparionine horses: "Hip­parion" gettyi, Hipparion prostylum, and Hipparion campbe//i. Stratigraphical ranges of individualmammalian species are given, and biostratigraphic intervals are characterized. A comprehensive zoo­geographicanalysisof the entire Maragheh mammalian fauna shows its relationships with late Miocene­early Pliocene "savanna-mosaic" assemblages of Eurasia and Africa. It has been found that the Mar­agheh genera which have the broadest geographic distribution were part of a late early Miocene pan­Eurasian and African dispersal event. Their subsequent diversification may be attributable to biogeo­graphic vicariance with both tectonic and paleoenvironmental factors playing contributing roles.

HISTORICAL BACKGROUND

The fossil vertebrate horizons ofMaragheh, in Azer­baijan, Northwest Iran, are among the most renownedofPalaearctica. Maragheh, the name currently used forthese deposits (in lieu of Maragha), also has long beenused to designate one of the classical "Pontian" sa­vanna-like faunas of Eurasia. Khanikoff, a Russian ex­plorer who visited Iran in 1840, has been credited asbeing the first to collect fossil vertebrates from theMaragheh district. The Khanikoff collection was sentto Dorpat University (now Tortustat University, Es­tonia) and initially studied by Abich (1858) who re­ported the occurrence of a fossil proboscidean tusk,remains offossil " cerv ids" and horses. Further studiesof this collection were made by Brandt (1870) andGrewingk (1881). They presented a more extensivefaunal list , reporting the occurrence of Hipparion,"Rhinoceros tichorhinus", "Mastodon", Helladothe­rium, and "Tragocerus," and suggested a close faunalcorrelation with the Pikermi fauna, near Athens,Greece.

In 1884 an Austrian paleontologist, Pohlig, was in­vited by a merchant from the nearby city of Tabriz tomake a geological tour of Persia. He visited Maraghehand made the first comprehensive paleontological andgeological studies of the deposits. Pohlig (1886: 178)reported: "The fossil bones have been found in thereddish marls at more than six places, at greater or lessdistances from the city (up to 30 miles) and at differenthorizons, which, however, do not differ from each oth­er by age and characteristics ofthe mammalian fauna".Pohlig further noted that there were two stratigraphic

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units, the "Pliocene fossil-bearing beds which were flu­vio-lacustrine in their origin, and overlying unfossilif­erous pebble beds, boulders and blocks ofearliest Plei­stocene age". He reported an extensive mammalianfauna including: "Hipparion gracile" , "Onager" sp .,"Rhinoceros" persiae, "Rhinoceros" blanfordi, "Mas­todon", "Sus" sp., Gazella "brevicornis", Antilope sp.,"Bubalus" sp ., "Cervus" sp .; Helladotherium sp., "Gi­raffa" att ica; Felis "brevicorn is", "Hyaena" cf. exi­mia: "Canis" sp . Pohlig sent most of his collection tothe private museum of Professor Von Fritsch at Halle,Germany.

In 1885 two other Austrians, RodIer and Kittl,visited Maragheh and made an extensive collection offossils under the sponsorship of the Kaiserliche Natur­historisches Hofmuseum Wien. Kittl (1885) reporteda number of vertebrate localities from the Maraghehbasin including Kopran, Kopran-Mescha, Zad Bas­chin, Rasat, and Ketschawa. Their collection was sub­sequently studied and published by Kittl (1887), Rod­Ier (1890), RodIer and Weithofer (1890) and Schlesinger(191 7). These reported fossil occurrences further pro­moted the faunal resemblance of Maragheh with thatof Pikermi.

In 1886, Lydekker wrote a very briefcommunicationon Maragheh fossils collected by Damon of the BritishMuseum (Natural History). Lydekker confirmed theimportance of Maragheh for correlation not only withPikermi and Samos, but also a western extension ofthe Siwalik faunas. In 1893 Forsyth Major, who hadstudied the Samos vertebrate fauna, reported a toothof Orycteropus from these same British Museum (Nat.Hist.) collections. In 1889, Gunther collected a small

© 1986 by the Society of Venebrate Paleontology

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sample for the British Museum (Nat. Hist.) from onelocality, Kirjawa,

In 1897 , the French paleontologist Marcellin Bouleestablished a liaison with J. de Morgan (a foreign del­egate in Persia) and secured permission to conduct apaleontological expedition to Maragheh. In 1904 , agroup of French paleontologists assisted by 12 locallaborers excavated a large sample of Maragheh fossilsfrom a number oflocalities including Kirdjawa on theMurditchai River, Kingir, Kopran, Shollovend, Ker­medjawa, and a hill just west of the town of Maragheh(Mecquenem, 1924-25: 135). R. de Mecquenem, amember of the expedition, undertook the study of thislarge collection, initially publishing two brief com­munications (1905, 1906). In 1908 and 1911 he pub­lished a preliminary inventory of the collection andfinally, having been delayed by World War I , Mec­quenem published a monograph (1924-25) on this veryfine collection. In his monograph, Mecquenem de­scribed 7 families, 26 genera, and 32 species of fossilmammals, and 2 genera of fossil birds. The collectionis still preserved in the Museum National d'HistoireNaturelle, Paris.

More than 50 years elapsed before additional re­ported collections were made at Maragheh. In the au­tumn of 1956, Takai ofTokyo University, Japan, vis­ited Maragheh and collected fossils from a localitynamed Kerjabad (Takai, 1958) . In 1967, Tobien oftheJohannes-Gutenberg University, Mainz, Germany,made some important excavations of the middle por­tion of the Maragheh sequence (Tobien, 1968) . Duringthe 1970's, three scientific groups conducted paleon­tologic, stratigraphic and geochronologic studies of thefossiliferous sequence. A combined Dutch-Germangroup directed by Erdbrink visited Maragheh from Au­gust until November, 1973. They collected a smallsample of Maragheh fossils, a number of volcanic tuffsamples for radiometric dating, and some paleomag­netic samples for local stratigraphic studies. A jointUniversity of Kyoto-Geological Survey of Iran expe­dition directed by T. Kamei visited Maragheh fromSeptember to November, 1973. They made an exten­sive collection of vertebrate fossils from one localityand sampled some nearby volcanic tuffs for fission­track dating. The Lake Rezaiyeh Expedition, a jointUniversity of California at Los Angeles, and NationalHistory Museum ofIran project directed by Campbell,worked at Maragheh during the spring of 1974 andsummers of 1975 and 1976 (Campbell et aI., 1980).The program of the Lake Rezaiyeh Expedition includ­ed studies of mammalian systematics, physical stra­tigraphy, sedimentology, biostratigraphy, biochronol­ogy, paleoecology, and radiometric dating. Another fieldseason was planned for the summer of 1978 to continueall aspects of the program, but was cancelled becauseof pol itical unrest in Iran.

The geological and paleontological investigations ofMaragheh during the past 25 years have appreciablyadvanced our knowledge of this very important lateMiocene fauna. Precise locality information had not

JVP 6( Il, March 1986

been documented prior to these studies. However, in­formation provided both in publications, and offeredby Tobien and Erdbrink, led to the development of thefirst biostratigraphy ofthe Maragheh deposits (Bernor,1978 ; Bernor et al ., 1979b; Campbell et al ., 1980).Nevertheless, the age of the Maragheh fauna has beenan issue of some recent debate. Erdbrink et al. (1976)reported at least four distinct faunal horizons rangingfrom "Vindobonian" to medial Turolian age, and sub­stantiated their claim with a radiometric age bracketof 12.9 to 7.5 my. Kamei et al. (1977) claimed a shortchronologie range of 6.9 to 6.6 my for the Maraghehfossil horizons. Campbell et al. (1980) and Bemor etal. (1980) reported the results of 28 fission track andpotassium-argon determinations ofthe Maragheh For­mation and underlying basal tuff. These included fourof the five horizons dated by Erdbrink et al. (1976),three of the five horizons dated by Kamei et al. (1977),and spanned all but the upper seven meters of fossillocalities found by the Lake Rezaiyeh Expedition. Theradiometric determinations, coupled with biochro­nologic correlations, suggest an age range for this faunaof approximately 9.5 to 7 my .

This paper reports the results of systematic, bio­stratigraphic and zoogeographic studies of the Mara­gheh mammalian fauna undertaken by the author since1974 . Biostratigraphic information has been assem­bled from a combination offield and archival research.Solounias (1981) provided systematic studies of theBovidae and Hyaenidae, and a similar synopsis of theentire vertebrate fauna from Samos and Pikermi, aswell as essential background on classical "Pontian"large mammal faunas. Finally, it should soon becomeapparent to the reader that this paper is not meant tocomplete research at Maragheh. Maragheh remains oneofthe richest Miocene vertebrate localities in the worldfor preservation of diverse and abundant fossil re­mains, long stratigraphic interval, and potential forclose geochronologic control. Completion of researchalong these lines will require years ofstudy by a diversegroup ofgeologists and paleontologists. The Maraghehbasin has the potential for establishing a provincialmammalian systematic and geochronologic standardfor southwest Asia.

Abbreviations and Definitions

AMNH - Department ofVertebrate Paleontology, TheAmerican Museum ofNatural History, New York.

BMNH-Department of Paleontology, British Mu­seum (Natural History), London.

GIU -Geological Institute, Utrecht.JGUM-Institute of Paleontology, Johannes-Guten­

berg Universitat, Mainz.KNHM-Naturhistorisches Museum, Vienna.MNHM-Museum National d'Histoire Naturelle,

Paris.UCR- University of California, Riverside, Depart­

ment of Earth Sciences.hipparionine-an informal taxonomic rank referring

to equid species belonging to any of the following

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~

k» >d HIGHLAND S

FIGURE I. Relationship of the Maragheh Formation to the Mendelassar and Anatolian Transform Faults of the EurasianPlate.

genera: Hipparion, Neohipparion, Nannippus,Cormohipparion, Proboscidipparion, "Cormohip­parion" (Siva/hippus), Stylohipparion.

"Hipparion"-following MacFadden and Woodburne(1982), Bernor and Hussain (1984) and Bernor(1984): Old World hipparionine horses with facialmorphologies that differ from either H ipparion orCormohipparion (includes groups I, 2, and 3 ofBernor and Hussain, 1984 and Bernor, in press).

GEOLOGY

The geological context of the classical Maragheh fos­sil district has only recently been described adequately(Erdbrink et aI., 1976; Campbell et aI., 1980; Bernoret aI., 1980). The Maragheh stratigraphic sequence ac­cumulated on the southern flank of the Mt. Sahandvolcanic massif, which is a large complex structureapproximately 100 km in circumference. Despite itsnearly circular outline, Mt. Sahand is not a single vol­cano, but rather a series of distinct cones arrangedalong an east-west system of fissures and vents, in­cluding several parasitic cones. Erdbrink et al. (1976)observed that eight high points can be seen looking

78

southeast from the city of Tabriz, the highest beingQuch Goli Dagh (3703 m). They further noted that theeastern sector of the massif seems to have been sub­jected to erosion for a longer time than the westernsector. Campbell et al. (1980) and Bernor et al. (1980)described the lithology, stratigraphy and sedimentol­ogy of the Maragheh Basin , which is summarized asan introduction to the biostratigraphy. For more de­tailed presentations of the geology, the reader is re­ferred to these studies.

The late Miocene deposits of the Maragheh Basinconsist ofa thick sequence ofvolcaniclastic continentalstrata with a basal pyroclastic unit on the southern andeastern sides of the Sahand volcanic massif. On thesouthern side , these strata show a virtually unde­formed, very gentle inclination to the west-southwest.To the northeast, where these strata are bound by theAnatolian Transform Fault (Bern or et aI., 1980; Fig.I, this paper), they are thrown into a steep monoclinalfold that dips away from the fault. While Kamei et al .(1977) termed the entire late Miocene section of theMaragheh Formation, Campbell et al. (1980) restrictedthis name to the volcaniclastic series, and distinguished

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FIGURE 3. Stratigraphic array of isotopic age determinations from the Maragheh Formation and Basal Tuff. Refer to Table3 for sample documentations.

(1886) was the first to recognize that fossil bones inthe Maragheh region can be collected from several ho­rizons. Rodier (1887), who collected at least 50 local­ities throughout the Maragheh Basin, did not believethat recording their stratigraphic occurrence was im­portant. Mecquenem (1908) claimed that there weretwo distinct fossil-bearing horizons with no differencesin the faunas. Tobien (1968), and later Erdbrink et al.(1976) have argued for three or four distinct fossil­bearing horizons, with several meters of barren strataseparating them. Use of the "Loose Chips" and BasalTuff marker beds (Campbell et al., 1980) has affordedan accurate correlation of several fossil vertebrate lo­calities in the Maragheh Basin ("Loose Chips" was alsoused by Kamei et al., 1977, but was not related to somany localities as by Campbell et al ., 1980) . However,the "Loose Chips" marker bed is limited in its lateralextent and the Basal Tuffis exposed only in a few places

across the basin. Furthermore, as noted above, bio­stratigraphic information suggests that the northeast­ern-most vertebrate locality of Ilkhchi, in beds thatunconformably rest upon the Basal Tuff, is substan­tially younger than those localities found on the pre­sumed same tuff in the southwestern portion of thebasin.

The mammalian faunal composition compiled here(Table I) is based on a number of sources including:

X: taxa collected by the Lake Rezaiyeh ExpeditionI: taxa collected by the KNHM2: taxa collected by Erdbrink et aI., Laboratory for

Zoological Ecology and Taxonomy, Utrecht Uni­versity

3: taxa collected by Kamei et aI., Department of Ge­ology, University of Kyoto

4: taxa collected by Mecquenem, MNHN

FIGURE 2. Geographic Sites and Fossil Localities of the Maragheh Basin, N .W. Iran. Contours on the upper surface of theBasal Tuff strike N-NW. Data points for elevations are Ilkhchi, Suma Pa'in, Davazim, Eshrat Abad, and Maragheh. Threepoint solutions of attitude from these points indicate a moderate flexure near Maragheh, in the vicinity of the Mendelassartrend, and small faults (not shown) displace the lower Maragheh beds near Kopran. Heavy dashed lines represent the sectionsshown in figure 6 of Bemor et al. (1980). Dots are fossil localities (some represent more than one) . The 7 major localities (1,31 ,37, etc .) are shown by heavy numbers. Circled numbers (1-7) correspond to the stratigraphic columns of figure 6 of Bemoret al. (1980) . Mb: Pre-Tertiary basement. Tev: Eocene volcanic rock. Tmm: Maragheh Formation (Basal Tuff shown in black).Qk: Kerajek Beds (alluvial). Qt: Uppermost terrace gravel.

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(1924-25). The provenance of th is specimen is notprecisely known. Mecquenem (1908:43) reported thathe collected fossils from two levels of I m and 0.6 mthickness each, and only a few meters apart vertically.He reportedly collected these horizons along a coursethat was 5 to 6 km wide, 30 km long, and suppos­edly between 1620 and 1650 m elevation. His prin­cipal collections came from Kirdjawa (- 52 to - 28 minterval of this paper), but also from areas near Shol­lovend, Kingir and Kopran, which probably did notinclude any higher levels, but could have mistakenlyincluded levels as low as the - 115 m level ofthis paper(i.e . upper Kopran). If Mecquenem's observations ofthe stratigraphic provenance of his own collections arecorrect, the best estimate of Mesopithecus pentelici'soccurrence would fall within the - 52 to - 28 m in­terval.

CarnivoraTwelve carnivore taxa are known from Maragheh:

Martes sp. indet. , Promeles palaeattica, Melodon ma­raghanus, Parataxidea polaki, Indarctos maraghanusMetailurus orientalis, Felis att ica, Machairodusaphanistus, Ictitherium viverrinum, Thalassictis won­eu. and Percrocuta eximia. Of these, Rodier's (1887)collection included a Metailurus (= Paramachairodus)orientalis, Felis attica. Thalassict is wongii, Percrocutaeximia from Ildtschi (= our Ilkhchi); and Melodonmaraghanus, Machairodus aphanistus. and Parataxi­dea polaki from Ketschawa. Indarctos maraghanus wascollected by Mecquenem presumably from somewherewithin the middle interval of the Maragheh sequence.The Lake Rezaiyeh Expedition has collected a numberof carnivore species from the Maragheh Basin. Theseinclude a left mandible ofMartes sp. indet. from MMTT7 (- 28 m interval), a nearly complete skeleton of Pro­meles palaeattica from MMTT 13 (-18 m interval);a mandible fragment of Ictitherium viverrinum fromIlkhchi (correlates with - 18 m interval); a partial skulland isolated teeth of Thalassictis wongii from MMTT13 (- 18 m interval) an isolated tooth from MMTT 25(-12 m interval) and skull material from Ilkhchi; askull of Machairodus aphanistus from MMTT 13 (-18m interval), as well as postcrania and isolated teethfrom MMTT 7 and 14 (- 28 m interval) and postcraniafrom MMTT 3 (-40 m interval).

The Ilkhchi specimens are biostratigraphically bestcorrelated with the - 28 to +7 m interval based uponthe co-occurrence ofHipparion cf. campbelli and "Hip­parion" ?m atthewi as well as a 7.4 my date based uponK-Ar and fission track determinations (Campbell etaI. , 1980). These correlations would place the Ilkhchilocality closely equivalent in age to the Lake RezaiyehQuarry at MMTT 13 (- 18 m level) . The Ketschawalocality is believed to be the same as the MMTT lA,IB, and IC localities of this paper (- 52 to - 30 minterval) reported by Campbell et al. (1980) as well asTobien (1968), Erdbrink et al. (1976), and Kamei etal. (1977).

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TABLE 2. Biostratigraphic intervals of Maragheh mam­malian species.

Upper Maragheh -20 to + 7 meters above/below LooseChips:

Vertebrate local it ies (in desc ending stratigraphic order):UCR-MMTT 26 , 39, 24 , 32, 38 , 22, 16, 47,27, 25, 15,49 , 49a, 50, 31 , 33, 2 1, 13,37, 40, 51

Mammalian fauna: Promeles palaeatt ica, Metailurus ori­entalis, Felis attica, Ma chairodus aphanistus, Ictithe­rium viverrinum, Thalassictis wongii, Percrocuta exim ­mia, Orycteropus sp. indet., Choerolophodon pentelici,Deinotherium sp . indet., Hipparion campbelli. "Hippar­ion" ?matthewi, An cylotherium pentelici, Microstonyxerymanthius, Palaeotragus coelophrys, Samotherium sp .indet., M iotragocerus undescr. sp. , Gazella cf. deperdita ,Oioceros atropatenes, Protoryx crassicornis, Urmia­therium polaki

Middle Maragheh - 52 to - 20 meters below Loose Chips:Vertebrate localities: UCR-MMTT 7,14,34, Ic, 6,18,19,

20 , Ib, 2,17, 11,4,35,3, lA, 5Mammalian fauna: Mesopithecus pentelici, Indarctos ma­

raghanus, Mart es sp. indet. , Melodon maraghanus,Parataxidea polaki, Felis attica, Ma chairodus aphanis­tus, Thala ssictis wongii, Percrocuta eximia, Choerolo­phodon pentelici, Hippa rion prostylum, " Hipparion" atf.m oldavicum, "Hipparion" Tm au he wi, Chilotheriumpersiae, Diceros neumayri, Microstonyx erymanthius.Cervi dae gen . and sp . indet., G iraffinae gen. and sp.ind et ., Palaeotragus coelophrys. Samotherium neumay­rio Helladotherium du vernoyi, Miotragocerus sp ., Gazel­la cf. deperdita , Prostrepsiceros houtumschindleri, Pro­tra gelaphus skou zesi , Oioceros rothii, Oi ocer osatropatenes, Protoryx laticeps

Lower Maragheh - 150 meters to - 52 meters below LooseChips:

Vertebrate localities: 8, 42, 45 , 36, 23, 28, 41 , 44 , 43, 9,48

Mammalian fauna: Choerolophodon pentelici, " Hippari­on" gettyi, "Hipparion" Tmatthewi, Chilotherium per­siae, Microstonyx erymanthius. Samotherium neumay­ri , Miotragoceru s amalthea , Gazella deperdita,Prostrepsiceros houtumschindleri, Oioceros atropatenes

TubulidentataThe only known occurrence of a tubulidentate with

stratigraphic provenance is an Orycteropus sp. indet.tibia-fibula from MMTT 13 (-18 m level) .

ProboscideaA species of Proboscidea retrieved by the Lake Re­

za iyeh Expedition is Choerolophodon pentelici fromlocalities I-A (- 52 m level) , MMTT 13 (- 18 m level),and MMTT 37 (= Ilkhchi). Erdbrink et al. (1976) re­ported the occurrence of"Synconolophus" (a synonymof Choerolophodon ; Tassy and Madden, pers. comm.)from the Kerjabad and Ci Anne (- 52 to - 28 m in­terval), and Kamei et al. (1977) reported a completeskull of Choerolophodon pentelici from locality I-B(- 32 m interval). The KNHM collection also containsChoerolophodon pentelici from Kopran Mescha (- 150

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vidual collected from Ilkhchi, and some postcranialelements referable to Chalicotheriidae incertae sedishave been recovered from MMTT 13 (- 18 m level) .A nearly complete skull and lower jaw of Chilotheriumpersiae was recovered from MMTT 7 (- 28 m level) ,but other than this specimen there are only a few post­cranial remains ofRhinocerotidae collected by the LakeRezaiyeh Expedition. The KNHM collection recordsChilotherium persiae from Kopran Mescha (- 150 minterval) and Kara Kend (= l A, 1B, 1C localities, - 52to - 30 m interval). The MNHN collection includesboth Chilotherium persiae and Diceros neumayri, whichwere presumably derived from the - 52 to - 28 minterval.

Five species of hipparionine horses have been rec­ognized from Maragheh: "Hipparion" gettyi, Hippar­ion prostylum, Hipparion campbelli, "Hipparion" affmoldavicum, and " Hipparion" Tmatthewi. "Hippari­on" gettyi is represented by a skull and mandible housedby the KNHM from Kopran Mescha (- 150 to - 115m interval), as well as several cheek teeth from MMTT43 (- 150 m level) , MMTT 41 (- 115 m level) andMMTT 36 (-70 m level). Hipparion prostylum is rep­resented by skulls, dentitions, and postcrania fromMMTT lA (- 52 m level ; JGUM collection), by skullsfrom unknown provenance in the MNHN and AMNHcollections. Hipparion campbelli is represented by skullsand lower jaws from the - 18 m interval, and teethand postcranial remains from the - 20, - 18, - 17, and+7 m intervals. "Hipparion" aff moldavicum is rep­resented by several skulls in the MNHN collection(- 52 to - 28 m interval), a single skull in the BMNHcollection, and some skull fragments in the KNHMcollection derived from Ketschawa (-52 to -28 minterval) and Zad Baschi (unknown provenance)."Hipparion" 'lmatthewi is represented in the Lake Re­zaiyeh collection by teeth from the - 28, - 18, - 10and +7 m intervals, from Kopran II (?-115 m level)in the KNHM collection, and a skull fragment of un­known provenance in the GIU collection.

ArtiodactylaSeventeen species of Artiodactyla are known from

Maragheh including the families Suidae (Microstonyxerymanthiusi, Cervidae (gen. and sp . indet.), Giraffidae(Giraffinae gen . and sp . indet. , Palaeotragus coelo­phrys, Samotherium neumayri, Samotherium sp . in­det., and Helladotherium cf. duvernoyi) and Bovidae(Miotragocerus sp. nov., Miotragocerus sp. indet., Ga­zella cf. rodleri, Gazella cf. deperdita, Prostrepsiceroshoutumschindleri, Protragelaphus skouzesi, Oiocerosrothii, Oioceros atropatenes, Protoryx crassicornis,Protoryx laticeps, and Urmiatherium polaki.

The Suidae and Cervidae are the least evolutionarilydiverse and least abundant families of all the Artio­dactyla. The suid Mi crostonyx erymathius is repre­sented in the Lake Rezaiyeh Expedition collection bya few teeth and postcranial fragments from MMTT 41(- 115 m level) and MMTT 13 (- 18 m level). Thereare more abundant remains of this species in the Mec­quenem collection including skulls, teeth and postcra-

JVP 6(1), March 1986

nial remains, which most likely were collected withinthe - 52 to - 28 m interval. The KNHM collectioncontains Microstonyx erymanthius from Ketschawa(- 52 to - 28 m interval). The Cervidae are representedby one antler fragment with pedicle collected by Tobienfrom MMTT I-A (-52 m level).

The Giraffidae are relatively diverse but the numberof specimens is low in the various Maragheh collec­tions. A Giraffinae gen. and sp. indet. palate was re­trieved by the Lake Rezaiyeh Expedition from MMTT7 (- 28 m level) and is the only specimen ofthis speciesknown. Postcrania and teeth of Palaeotragus coelo­phrys were collected from MMTT 13 (- 18 m level)and are also present in Mecquenem's collection (- 52to - 28 m interval). Samotherium neumayri is foundin the Vienna collection originating from Kopran Mes­cha (- 150 m interval), Kara Kend, and Ketschawa(- 52 to - 30 m interval). Samotherium sp . indet. isrepresented by teeth and postcrania from MMTT 13and 37 (-18 m level and ca . -18 m level respectively).Finally, the large sivatherine Helladotherium cf. du­vernoyi is represented by teeth and postcrania in theMecquenem collection (- 52 m to - 28 m interval).

Remains of Bovidae are relatively abundant and ev­olutionarily diverse in the various Maragheh assem­blages. There are two Miotragocerus species knownfrom Maragheh, a smaller more primitive form, Mio­tragocerus amalthea collected by Rodler (skull, teeth,postcrania) from Kopran (- 150 to - 115 m interval)and a larger, more advanced form Miotragocerus un­descr. sp . collected by the Lake Rezaiyeh Expeditionfrom MMTT 7 (-28 m level) and MMTT 13 (-18 mlevel) as well as Mecquenem (- 52 to - 28 m interval).The gazelles are also represented by two species, Ga­zella rodleri and Gazella cf. deperdita. Gazella rodleriwas collected by the Lake Rezaiyeh Expedition fromMMTT I-A (- 52 m level) and was reported by Pohlig(1886) from Ketschawa (- 52 to - 28 m interval). Ga­zella deperdita has been collected by the Lake RezaiyehExpedition from MMTT 44 (- 115 m interval), MMTT1b (32 m interval), MMTT 13 (- 18 m interval, MMTT25 (- 12 m interval), MMTT 26 and 39 (+ 7 m inter­val). It is also known in the MNHN collection (- 52to - 28 m interval). Prostrepsiceros houtumschindleriis a relatively well known species, having been col­lected by the Lake Rezaiyeh Expedition from MMTTlocalities 43 (- 150 m level), 36 (-70 m level), I-A(- 52 m level) , the - 30 m level, and MMTT 7 (-28m level) . Mecquenem's collection includes a good sam­ple ofskull, tooth, and postcranial remains presumablyfrom the - 52 m to - 28 m interval. Protragelaphusskouzesi is represented in the Lake Rezaiyeh collectionby horn and dental material from MMTT 7 (-28 mlevel) , while being more completely represented by skulland tooth material in the Mecquenem collection. Oio­ceros rothii is represented in the Lake Rezaiyeh Ex­pedition sample by horn material from MMTT 1C and7 (- 30 m and - 28 m levels respectively), as well asMecquenem's collection (- 52 to - 28 m interval).Oioceros atropatenes is the most abundant single speciesin the Lake Rezaiyeh Expedition sample, being rep-

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TABLE 3. Isotopic age determinations from the Maragheh Formation and Basal Tuff.

8.1 ± 0.4 VlIwr6.2 ± 0.6 VlIhb

6.6 730-19 8.7 ± 0.4 IIIwr

6.5 730-17 14.4 ± 0.7 Vwr

7.0 730-10 12.9 ± 0.7 Iwr

7.2 730-20 10.1 ± 0.5 IXwr11.4±0.5 IXbi

+ 100 Korde-deh Pumice(Murdag Chai)

+90 est. Upper Pumice(Ilkchi)

+ 60 est. Middle Pumice(Ilkchi)

+ 15 est. Lower Pumice(Ilkchi)

+ 15 est. Village Pumice(Ilkchi)

+ 7 Layered Marker Pumice(Shollovend)

o Loose Chippings Pumice(Shollovend)

- 50 Giirt Dareseh Pumice(Murdag)

- 110 Ignimbritic Tuff(Murdag)

Basal Tuff(western area)

Basal Tuff(Ilkchi)

Zircon FT K-ArBernor et al.,1980

7.2 ± 0.4 RIO 7.5 ± 0.4 RIOp/

7.3 ± 0.5 R 5 7.7 ± 0.5 R 5p/7.2 ± 0.5 R 6p/

7.4 ± 0.4 R 8 7.9 ± 0.7 R 8p/7.0 ± 0.4 R 85.2 ± 0.3 R 9 5.5 ± 1.0 R 9p/5.3 ± 0.3 R 97.6 ± 0.5 R 2 8.3 ± 0.9 R 2p/

7.7 ± 0.5 RI2 6.4 ± 0.6 R12p/6.9 ± 0.3 M 5hb

7.8 ± 0.4 R11 7.4 ± 0.3 Rllp/

10.6 ± 0.8 R 3 6.4 ± 0.5 R 3p/8.8 ± 0.5 R 3p/8.9 ± 0.5 R 3p/

11.2 ± 0.5 M Ibi

11.2 ± 0.6 R 4 10.1 ±0.5 R abi9.3 ± 0.1 R 4an

12.8 ± 0.5 R 7 11.3 ± 1.0 R 7bi9.3 ± 0.1 R 7an

Zircon FTKamei et al.,

1977

5.2 730-04

K-ARErdbrink et

al.,1976

Measured stratigraphic relationships of dated levels in the Mirdug-Shollovend area and in the Ilkchi area are correlated byprojecting the regional dip of the Maragheh Formation in the central area to Ilkchi . Sample numbers are given to the right ofeach determination. All fission-track (F'T) ages were obtained on zircon crystals; the analytical uncertainty of the F'T agespublished by Kamei et al. (1977:158-159) is stated to be approximately 20% of each measurement. The K-Ar ages wereobtained from materials indicated by the suffixes to the sample numbers, as follows : wr (whole rock), pi (plagioclase), hb(hornblende), bi (biotite) and an (anorthoclase) (from Bernor et al ., 1980) .

sion track and KIAr determinations, Table 3 and Fig­ure 3 show considerable contradictions at each strati­graphic level. However, the implementation ofhipparionine biochronology by Bernor et al. (1980) wasfound to be generally in agreement with the higherquality potassium-argon and fission track determina­tions from the sequence.

The frequent relatively large error ranges within andbetween KIAr and fission track determinations is inpart thought to be the result of inclusions ofaccidentalpebbles and grains of detrital andesite. Whole rockdeterminations reported by Erdbrink et al. (1976) arethought to be particularly suspect because of this sup­posed contamination problem. However, Drake (pers .comm.) reports further problems with the consistentdiscrepancy between anorthoclase KIAr ages on theone hand, and biotite KIAr and zircon fission trackages on the other, especially in the Basal Tuff For­mation. Nevertheless, Campbell et al. (1980) and Ber­nor et al. (1980) believed that a 9.5-7 my interval

JVP 6(1l. March 1986

represents the best estimate for the age range of theMaragheh fauna. This assessment was based on thestage-of-evolution of"Hipparion" gettyi, Hipparion cf.prostylum, and Hipparion campbelli compared to oth­er closely related Eurasian horse species as well as anassessment of the radiometric determinations. Bernoret al. (1980) estimated that Lower Maragheh rangedfrom 9.5 to 8.5 my, Middle Maragheh ranged from8.5 to 8 my, and Upper Maragheh ranged from 8 to 7(or fewer) my in age. A synopsis of the zoogeographicrelationships and geochronologic significance of theMaragheh fauna comprises the remainder of this sec­tion.

The primate Mesopithecus pentelici has been revisedby Delson (1973) from the Turolian of Greece (Piker­mi, Vathylakos), Yugoslavia (Titov Veles), Bulgaria(Kalimanci, Gorna Susica, Kromidovo), the Ukraine(Grebeniki, Taraklia?, Grossulovo?), Germany (Wiss­berg), Italy (Baccinello V3), Hungary (Hatvan; ?Pol­gardi). Of this suite of localities, Pikermi is demon-

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strably the oldest (ca. 9 my after hipparioninebiochronologic correlations by Bernor et aI., 1980).The occurrences of Mesopithecus pentelici in MiddleMaragheh and Vathylakkos are very close in age, butperhaps slightly younger than Pikermi. The age of Ti­tov Veles, the Bulgarian, Ukranian, and German lo­calities is not certain, but may generally be consideredas being medial-late Turolian. The Baccinello fauna issubstantially younger (MN 13; Mein, 1975, 1979), asare the Hungarian faunas ofHatvan and Polgardi (MN12/13, ca. 7 my, Steininger and Papp, 1979). The con­spicuous absence of M. pentelici from the upper bio­stratigraphic interval at Maragheh and the extensiveSamos assemblage, contrasted with its comparativelylate occurrence in the Hungarian localities, suggeststhat its range extended northwestward into eastern andCentral Europe along with the late dispersal ofthe morewooded part of the Turolian savanna-biotope.

The Maragheh carnivore assemblage is extensive andincludes species of the families Ursidae, Mustelidae,Hyaenidae, and Felidae. Generally speaking, all of thesehave a broad Eurasian, and in some cases African andNorth American geographic range at the generic level.The one ursid species, Indarctos maraghanus is uniqueto Maragheh. However, closely related late Mioceneage species are known from China (I. sinensis; Zdan­sky , 1924); Hasnot, the Siwaliks (I. punjabiensis; Ly­dekker, 1886 ; Erdbrink, 1953) ; Greece (I. atticus, Sa­mos and Pikermi; Solounias, 1981); Spain (lndarctosvireti-Can L1obateres, Can Ponsich, Villadecavalls; I.atticus adroveri-Concud; I. atticus-Los Mansuetos;I. sp.-La Fontana; Alberdi , 1974); North Africa (I.atticus-Sahabi; Howell, 1982) .

Martes is a chronologically long ranging and geo­graphically widely distributed genus and is oflittle bio­chronological significance here. Of the three species ofMelinae from Maragheh, Prome/es palaeattica is themost primitive morphologically, but is found in theupper Maragheh interval. This species is also knownfrom Samos and Pikermi, the western U .S.S.R. (Boris­siak, 1962), and together with the Maragheh skeletonsuggests a provincial age range ofca. 9-7 my . Me/odonmaraghanus has been collected from Ketschawa, Mid­dle Maragheh, with an estimated age of 8.5 my. Thisspecies is morphologically very close to Me/odon("Parataxidea") sinensis (Zdansky, 1924 ; pl. X, figs.5-10), and may eventually prove to be the same. Para­taxidea polaki is also from Ketschawa, Middle Mara­gheh, and similarly shows close morphological identityto Chinese species of Parataxidea (Zdansky, 1924).

Hyaenidae are relatively diverse at Maragheh, andbroadly characteristic oflate Miocene Old World largemammal faunas. Ictitherium viverrinum is only re­corded from the Upper Maragheh interval (locality ofIlkhchi) and its morphology is virtually identical tospecimens from Samos and Pikermi (Solounias, 1981).Solounias also reports that this species is similar to aChinese species Ictitherium sp. Thalassictis wongii isa larger ictitheriine hyaenid which has a distributionin late Miocene horizons of Greece (Samos), Turkey

88

(?Garkin, ?Kinik), the western U .S.S.R. (Borissiak,1962) and China (Solounias, 1981). Solounias sepa­rates T. wongii from T. hipparionum on several mor­phological grounds. The latter has a reported geograph­ic occurrence in southwest Europe (Guerin and Mein,1971; Alberdi, 1974) during the late Miocene. WithinSolounias' (1981) synonymy there are contained speciesof the genus Pa/hyaena. Pilbeam et al. (1979) havereported two species ofPalhyaena, from the Nagri andDhok Pathan Formations: P. sivalense (Chronozones7-9) and P. indicum (Chronozone 10). Percrocuta ex­im ia is the largest Maragheh hyaenid, and has a geo­graphic range including Europe, the western U.S.S.R.,southwest Asia, and North Africa.

The felids are a diverse group ofcarnivores at Mara­gheh. Metailurus (= Paramachairodus) orientalis is re­ported from the western U.S.S.R. (Tchobroutchi , Novo­Elisavetovka, Tarkalia; Pilgrim, 1931); Hungary (Pol­gardi; Kormos, 1911), Greece (Veles, Schlosser, 1921 ;Pikermi, Solounias, 1981), and China (Zdansky, 1924).Felis attica has been reported from Pikermi and Samosby Solounias (1981), and from Vathylakkos by Ar­ambourg and Piveteau (1929). Closely related speciesoccur in China (Felis paleosinensis; Zdansky, 1924, pl.32 , fig. 1), the western U.S.S.R. (Felis sp.; Borissiak,1962), the Siwaliks (Felis sp ., Chronozone 10; Pilbeamet aI., 1979; Barry et aI., 1982), Spain (Felis vireti,Alberdi, 1974). Machairodus aphanistus has a lateMiocene pan-European and west Asian distribution(Pilgrim, 1931) , and has closely related species re­ported from China (Machairodus planderi and Ma­chairodus tingii, Zdansky, 1924), and North Africa (Sa­habi; Howell, 1982).

The genus Orycteropus was broadly distributedthroughout Eurasia and seasonal parts ofAfrica duringthis time. Its earliest record extends to the beginningofthe Astaracian (MN 6) ofTurkey (locality of Pasalar;Sickenberg et aI., 1975).

The proboscideans Choerolophodon pentelici andDeinotherium were also broadly distributed through­out much of Eurasia during the middle and late Mio­cene. Choerolophodon is an advanced member of theGomphotheriidae that is first reported from the EastAfrican early middle Miocene locality of Maboko (ca.15 my ; Tassy, 1977), subsequently dispersed and di­versified in western Asia (Sickenberg et aI., 1975), anddemonstrably later in the Siwalik Province (Pilbearnet aI., 1979) . Its occurrence is currently unrecordedfrom Europe and China. Identification of Deinothe­rium (Erdbrink, pers. comm.) is not significant herebecause of its long chronologie (MN 4-MN 13 of Eu­rope) and geographic (Eurasian-African) ranges.

The hipparionine species have proven to be the mostsensitive biochronologic indicators of the Maraghehfauna. The oldest Maragheh hipparionine, "Hippari­on" gettyi, is an advanced Group I species that mostclosely compares with Hipparion melendezi of theSpanish late Vallesian and with a specimen from Pi­kermi (BMNH M42603; Bernor and Hussain, 1984).Furthermore, I have argued (Bernor, in press) that de-

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tails of the facial fossa, dentition and postcrania ofHipparion gettyi are not so morphologically distantfrom Hipparion prostylum as to preclude their closeevolutionary relationship. A 9.5 to 8.5 my age rangeappears to be compatible with "Hipparion" gettyi stage­of-evolution.

Hipparion prostylum has been found in medial Tu­rolian horizons in France (Mt. Luberon), Greece (Va­thylakkos, Pikermi) and Afghanistan (pers. observ.).Hipparion antelopinum from the Middle Siwaliks ofthe Indian Subcontinent appears to be a closely relatedspecies. An 8 my (or slightly older) age estimate forthe Maragheh assemblage of Hipparion prostylum pre­sented by Bernor (in press) corresponds to age esti­mates for Hipparion prostylum and the closely relatedform Hipparion antelopinum from Europe/westernAsia and the Indian Subcontinent. Hipparion camp­belli is the most advanced Group 3 horse currentlyknown. It is closest morphologically in details of theskull, dentition and postcrania to the Samos speciesHipparion dietrichi and an undescribed skull fragmentof a Hipparion sp . from Shansi, China (Bernor andQiu, ms.).

The Group 2 horses "Hipparion" aff. moldavicumand "Hipparion" aff. Tmatthewi are similarly signifi­cant from geochronologic and zoogeographic stand­points. Group 2 horses occur only in western Asia,western U.S.S.R ., and China during the late Miocene." Hipparion" aff moldavicum most closely resemblesthe type "Hipparion" moldavicum series from Tar­kalia, Moldavia (U .S.S.R .), which Mein (1975, 1979)correlates with MN 12 of Europe. The closely relatedspecies "Hipparion" mediterraneum from Pikermi isalso bel ieved to be of MN 12 age (= MN 12a of Bernor,1983, 1984). The potential occurrence of"Hipparion"aff moldavicum from Middle and Upper Maraghehlevels (ca. 8.5-7 my) falls within the known temporalrange of this species in the western U .S.S.R. (i.e . MN12a and 12b, ca . 9-7 my ; Bernor, 1983; in press).Biochronologic and zoogeographic relationships of" Hipparion" Tmatthewi are difficult to assess becauseof the current confusion surrounding the morphologyof the holotype and probable parallel evolution of anumber of separate lineages of dwarf-sized species(Bernor, in press). However, the geographic and chro­nologie ranges of this species may well be limited toeastern Europe, western U.S.S.R. and western Asiaduring MN 12 and possibly MN 13. The occurrenceof "Hipparion" 'tmaithewi from Middle and UpperMaragheh appears concordant with its other presumedoccurrences in Eurasia.

The Maragheh rhinoceroses, Chilotherium persiaeand Diceros neumayri have been recovered only fromthe Lower and Middle Maragheh horizons. Chiloth­erium persiae has only been reported from Iran, al­though the genus was broadly distributed during thelate Miocene in southwest Europe (Spain; Alberdi,1974), the eastern Mediterranean (several species inTurkey, Sickenberg et al., 1975; and Greece, Solounias,1981), the western U .S.S.R. (Borissiak, 1962), the Si-

JVP 6(1l. March 1986

waliks (Pilbeam et al., 1979) and China (Chiu et al.,1979). Diceros neumayri has a reported distributionfrom the late Miocene of Greece (Samos and Pikermi;Solounias, 1981), Turkey (Kayadibi; Heissig, 1982),possibly the western U.S.S.R. (Borissiak, 1962), andthe Basal Pliocene of North Africa (Sahabi; Heissig,1982). The genus has also been reported by Sickenberget al. (1975) from the Vallesian and Turolian age lo­calities of Esme Akcakoy and Kayadibi. The chalico­there Ancylotherium pentelici is an exotic element inlate Miocene faunas and has been reported from Spain(Alberdi, 1974), the Greek localities of Samos and Pi­kermi (Solounias, 1981), Veles (Coombs, 1973), Va­thylakkos (pers. observ.), Kalimanci, Bulgaria, andNovo Ukrainka, U.S.S.R. (Coombs, 1973) .

The only suid known from Maragheh, Microstonyxerymanthius, has a geographic range including south­west Asia and the western U.S.S.R. and is known fromearly Vallesian through Turolian age horizons. It mayshare some close phylogenetic relationships with Di­coryphochoerus reported from Turkey (Sickenberg etal., 1975) and the Siwaliks (Pilbeam et al., 1979).

The ruminants are an especially diversified group atMaragheh. The Giraffinae gen. and sp . indet. is anexotic element in the Maragheh fauna, as it is in lateMiocene faunas of the eastern Mediterranean (Samos,Pikerrni; Solounias, 1981), China (Bohlin, 1926) andNorth Africa (Upper Beglia; Solounias, 1981). It ispossible that giraffines were exotic elements of pan­Eurasian and African middle-late Miocene open coun­try faunas of Eurasia and Africa.

Palaeotragus coelophrys is a large paleotragine speciesthat has been reported from medial Turolian age lo­calities of Garkin and Kinik Turkey (Sickenberg et al. ,1975), Samos (Solounias, 1981), the late Vallesian agelocality of Agha Jari (Thomas et al., 1980) and the lateMiocene of China (Bohlin, 1926) . The genus Palaeo­tragus has a chronologie range from the middle to lateMiocene. Geographically, the genus is recorded earlyfrom East Africa (P. primaevus, Fort Ternan, ca. 14my; Churcher, 1970) , North Africa (P. lavocati, BeniMellal, MN 7; Heintz, 1976) , Turkey (P . sp., Sofca,MN 7; Sickenberg et al. , 1975). Late Miocene speciesof Palaeotragus were broadly distributed in Spain (Pa ­laeotragus sp ., Can Ponsich, MN 9; Alberdi, 1974),France (Palaeotragus sp ., Soblay, MN 10; Guerin andMein, 1971) ; Greece (P. roueni, Samos, Pikerrni; P.coelophrys, Samos; Solounias, 1981); Turkey (P. de­cipiens, Esme Akacoy; P. germaini, Kayadibi; P. quad­ricornis, Kayadibi; P. coelophrys, Garkin, Kinik; Sick­enberg et al., 1975); the western U.S.S.R. (P. roueni;Borissiak, 1962); China (P. m icrodon, P. coelophrys;Bohlin, 1926) . Palaeotragus has never been reportedfrom the Central European or Siwalik Provinces.

The genus Samotherium similarly has a long chro­nologie and broad geographic range. Churcher (1970)reported an early Samotherium, S. africanum, fromFort Ternan while Sickenberg et al. (1975) recoveredanother primitive form Samotherium sp. from themiddle Miocene Turkish locality of Candir. Its pres-

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ence in Turkey is not again found until the late (?me­dial) Turolian locality of Kinik, which contains Sa­motherium "neumayri" (Sickenberg et aI., 1975). FromSpain, Alberdi (1974) reported the possible occurrenceof a Samotherium sp. indet. from Vallesian levels ofHostalets de Pierola. Solounias (1981) recorded twovarieties of Samotherium boisseri from Samos, one ofwhich (var. A) he also identified at Maragheh. Boris­siak (1962) reported Samotherium expeetens from theSarmatian and Samotherium boisseri from the Meo­tian of the western U .S.S.R. The record in China hasan uncertain age range, including Samotherium si­nense, S. neumayri, S. sp. 1 and S. sp. 2 (Bohlin, 1926) .Finally, Harris (1982) noted the occurrence of a prob­able Samotherium species from the early Pliocene lo­cality ofSahabi, Libya. The genus Helladotherium ap­pears to have a more restricted chronologie andgeographic range, being limited to late Miocene hori­zons of the Western U.S .S.R. (Borissiak, 1962) andSubparatethyan Provinces (Solounias, 1981) .

Among the bovids, the genus Miotragoeerus has oneof the broadest geographic and longest chronologieranges known within the Miocene. This genus has beenreported from a number ofmiddle and/or late Miocenelocalities of southwest Europe (Alberdi, 1974; Guerinand Mein, 1971), Central Europe (Thenius, 1959),western U.S.S.R. (Gabunia, 1979); the eastern Medi­terranean (de Bonis et aI., 1979; Solounias, 1981),southwest Asia (Sickenberg et aI., 1975); North China(Bohlin, 1935) ; the Siwaliks (Thomas, manuscript);North Africa (Thomas, 1979; Lehman, 1982); and EastAfrica (Smart, 1976). The great number of species andlack of rigorous revision of the entire group currentlyprohibits any meaningful attempt to discuss its evo­lutionary and zoogeographic history despite the greatquantity of material.

Gazella is another chronologically long ranging andgeographically broadly distributed genus. The earliestreported occurrence of Gazella is from the early Mio­cene locality of Gebel Zelten (Hamilton, 1973) . Itsmiddle Miocene distribution includes Turkey (Candir;Sickenberg et aI., 1975) and East Africa (Fort Ternan;Gentry, 1966). By the late Miocene the genus un­doubtedly was distributed in savanna-like biotopesthroughout Eurasia and Africa. The gazelle G. rodleriis known only from Middle Maragheh. However, G.deperdita has a relatively broad geographic and longchronologie range including the Vallesian ofSpain (AI­berdi, 1974) and Iraq (Thomas et aI., 1980), and sev­eral Turolian localities of France (Guerin and Mein,1971) and Spain (Alberdi, 1974).

The species of Prostrepsieeros and Protragelaphusappear to have limited chronologie and geographicranges. The Agha Jari form "Prostrepsiceros aff. hou­tumsehindleri" (Thomas et aI., 1980) is the smallestand most primitive species of this group. It appearsthat by the early Turolian this group diversified intoat least three species as represented at Pikermi (ca. 9my, Bernor et aI., 1980): Prostrepsieeros houtum­schinderli, "Prostrepsiceros" rotundicornis, and Pro-

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tragelaphus skouzesi. These same three species seemto have ranged through the medial Turolian, duringwhich time they are recorded from Samos and Mara­gheh (Solounias, 1981) . Sickenberg et al. (1975) alsoreport Prostrepsieeros sp. from Garkin (MN 12?), andcf. Protragelaphus sp. from Kayadibi (MN 13). A lateoccurrence of Prostrepsieeros has been reported fromthe early Pliocene Libyan locality ofSahabi by Lehman(1982). It should also be mentioned that Pilbeam etal. (1979) recorded a possible occurrence of cf. Pros­trepsieeros houtumsehindleri from the Siwaliks. Pro­tragelaphus is a monospecific genus found at Pikermi(Solounias, 1981), Maragheh, and during the Meotianof the western U .S.S.R . (Borissiak, 1962) .

The earliest occurrence ofthe genus Oioeeros is fromthe late early Miocene locality of Xieja, China (Chiuet aI., 1979) . Subsequently, Oioeeros appears to havehad a broader geographic range during the middle Mio­cene that included China (0. noverea and O. grangerifrom Tung Gur; Pilgrim, 1934; Chiu et aI., 1979), EastAfrica (0. tanyeeras from Fort Ternan; Gentry, 1971) ,and Turkey (Oioeeros sp. from Candir and Kinik; Sick­enberg et aI., 1975) . The first known late Miocene dis­tribution is limited to Greece, Turkey, and Iran. Ac­cording to Solounias (1981), the late Miocene speciesOioeeros rothii and Oioeeros atropatenes are moreprimitive than the very large species O. wegneri. Oio­eeros rothii is known to occur at Pikermi and in MiddleMaragheh, while Oioeeros atropatenes is known fromPikermi and Maragheh. Oioeeros wegneri has only beenreported from Samos (Solounias, 1981).

Protoryx laticeps and Protoryx crassieornis have bothbeen collected at Maragheh. Protoryx latieeps was re­covered by Mecquenern's expedition, and as arguedearlier here would probably have been collected withinthe Middle Maragheh biostratigraphic interval. Thesmaller and more advanced species, Protoryx crassi­eornis (Gentry, 1971 = his Pachytragus crassieornis) ,has been collected by the Lake Rezaiyeh Expeditiononly in the upper biostratigraphic interval. The ap­parent stratigraphic relationships of these species sug­gest that Gentry's (1971) hypothesis that Protoryxcrassieornis was evolutionarily more advanced thanProtoryx latieeps is supported to some degree by bio­chronologic and geochronologic information given here .If the provenance of the Maragheh specimens and ra­diometric-biochronologic assessments at that localityare proven accurate by future investigations, then theage range of the two species at Maragheh could be asmuch as 1.5 my or more, with Protoryx crassieornisapparently replacing Protoryx latieeps in the 8 to 7 myinterval. The possibility nevertheless remains that thetwo species coexisted provincially for a significant in­terval and that they are not useful for detailed bio­chronologic interpretation. In addition to these local­ities, Protoryx has also been reported from the lateMiocene ofChina (Bohlin, 1935), the middle Mioceneof Turkey (Sickenberg et aI., 1975) and possibly fromSahabi (Lehman, 1982) .

The type locality of Urmiatherium polaki is Ilkhchi,

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TABLE 5. Provincial zoogeographic ranges of Maragheh mammalian taxa of genus or superspecific taxonomic rank.

Province

Taxon 2 3 4 5 6 7 8 Age range estimates

Inda rctos X X X X X 12-5 rna .Martes sp. X X X X X X X X ?- RecentMelodon X X 8.5-? rna .Paratax idea X X 8.5 rna.Thalassictis ? X X X 9-? rna .Metailurus

(= Paramachairodus) X X X X ?9-6 rna.Felis X X X X X ?- RecentOrycteropus X X X X X X X X 15 ma .-RecentDeinot herium X X X X X X X X 18 ma.-?Choerolophodon X X X X 15-?7 rna."Hipparion" Gp. I X X X X X ? 12-7 rna."Hipparion" Gp. 2 X X X 9-7 rna.Hippa rion Gp. 3 X X X 9-7 rna .Chilotherium X X X X X 9-7 rna.Palaeotragus X X X X X X 14-7 rna .Samotherium X X X X X X 14-5 rna.Miotragocerus X X X X X X X X 14-5 rna.Gazella X X X X X X X 18 ma .-RecentProstrepsiceros X X ? 9-5 rna.Protragelaphus X X 9-8 rna.Oioceros X X X 16-7 rna .Protoryx X atf. aff. X 9-7 rna.Urmiatherium X X ca. 7 rna .

Legend-see Legend for Table 4.

lationships. A number of taxa show African-Asian re­lationships including Choerolophodon (15-7 my), Pa­laeotragus (14-7 my), Samotheriu m (14-5 my),Prostrepsiceros (9-5 my), Oioceros (16-7 my) and Pro­toryx (9-7 my) . The three supraspecific groups of hip­parionine horses show distinctly different zoogeo­graphic relationships from one another: Group I isfound in Europe, the Subparathethyan Province, prob­ably North China, but not in the Siwaliks or East Af­rica . Group 2 hipparionines are restricted to the Sub­paratethyan, Western U.S.S.R. , and Chinese Prov­inces . Group 3 hipparionines are found in a relativelynarrow west-east latitudinal belt from southern andwestern Europe to Indopakistan.

Some interesting generalized patterns emerge fromthis analysis. First, cosmopolitan taxa usually have thelongest chronologie ranges, and include genera thatwere inv olved in the initial late Orleanianl Astaraciandeployment of African or trans-African/Subpara­tethyan mammals throughout many sectors of Eurasia.Species of these groups may have evolved by vicari­ance subsequent to the ir early migration into non-sea­sonal subtropical regions. The mechanism isolatingtheir populations could have been the diachronoustransgression of progressively more seasonal environ­ments (Bern or, 1983, 1984) . Second, the Maraghehmammals that show close evolutionary relationshipswith North and East African mammals are first knownto occur in the late Miocene. Thomas et al. (1982)argued for the origin ofmany ofthese species in Eurasia

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and Bernor (1982, 1983, 1984) has cited th is as abasal Pliocene "regression" of the "Pontian" chrono­fauna. Third, the zoogeographic relationships ofMara­gheh taxa with those of the Siwalik Province appearto be phyletically and chronologically more distant fromthe ones with western Europe, Central Europe, thewestern U .S.S.R., China, and North Africa. Fourth,the Maragheh mammal fauna shows striking speciesand genus/superspecific group zoogeographic relation­ships within the Subparatethyan and Western U.S.S.R .Provinces. This zoogeographic analysis largely verifiesBernor's (1978,1983,1984) and Bernor et al.'s (1979a)earlier reconstruction of the general evolution and zoo­geographic relationships of the classsic "Pontian" largemammal faunas .

SUMMARY

The Maragheh mammalian assemblage represents asavanna-mosaic chronofauna (sensu J. A. H. Van Cou­vering, 1980) that shows a long and complicated evo­lutionary history. The geochronologic evidence citedhere suggests that during the interval of time repre­sented by the Maragheh fauna, little change took placein its community structure; there were no major faunalturnovers. The characterization and rough definitionof biostratigraphic intervals using a potential lineageof hipparionines, combined with radiometric dating,has allowed broad provincial and interprovincial zoo­geographic comparisons with late Miocene mammal­bearing deposits throughout Eurasia and Africa.

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Acknowledgements-This paper is an outgrowth of re­search conducted by the Lake Rezaiyeh Expeditionbetween 1974 and 1976. I would like to thank theDirector of the project, Professor Bernard G. Camp­bell, for providing me with the opportunity and con­stant encouragement to study this fauna . I thank Dr.Michael O. Woodbume for providing direct guidanceofthis research while I was a student, and considerablepostdoctoral support continuing this and closely re­lated research. I am also extremely grateful to Mr.Gordon Getty of the L. S. B. Leakey Foundation forproviding virtually all ofthe funds for conducting boththe Lake Rezaiyeh Expedition and my personal re­search on various facets of the mammalian fauna . Iam also grateful to Drs . John Van Couvering, NikosSolounias, Daryl Domning, Eric Delson, Richard Ted­ford, William Akersten, and Mr. George Jefferson forreading earlier drafts of this manuscript. I also thankDrs . Bruce MacFadden and Robert Emry for providingreviews and editorial comments that shaped the manu­script into its current form. Finally I would like tothank the dozens of colleagues from natural historymuseums and universities in London, Paris, Lyon, Vi­enna, Uppsala, Mainz, Frankfurt, Ludwigsberg, Stutt­gart, and New York who provided advice, hospitality,and countless hours of after-hours enjoyment duringthe eleven years of this study.

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Received 5 September 1984; accepted 20 February 1985.

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