This art icle was downloaded by: [ I zzet Hoşgör] On: 12 Novem ber 2013, At : 07: 28 Publisher: Taylor & Francis I nform a Lt d Regist ered in England and Wales Regist ered Num ber: 1072954 Regist ered office: Mort im er House, 37- 41 Mort im er St reet , London W1T 3JH, UK Journal of Vertebrate Paleontology Publicat ion det ails, including inst ruct ions for aut hors and subscript ion informat ion: ht t p:/ / www.t andfonline.com/ loi/ uj vp20 Late Miocene fishes from the Eurasian/ Afro-Arabian contact zone in Eastern Anatolia, Turkey a Alison M. Murray & Izzet Hoşgör b a Depart ment of Biological Sciences , Universit y of Albert a , Edmont on , AB , T6G 2E9 , Canada b TransAt lant ic Explorat ion Med. Int . Pt y. Lt d.–Viking Int . Lt d. TR-06680 , Ankara , Turkey Published online: 12 Nov 2013. To cite this article: Alison M. Murray & Izzet Hoşgör (2013) Lat e Miocene fishes from t he Eurasian/ Afro-Arabian cont act zone in East ern Anat olia, Turkey, Journal of Vert ebrat e Paleont ology, 33:6, 1292-1299 To link to this article: ht t p:/ / dx.doi.org/ 10.1080/ 02724634.2013.780962 PLEASE SCROLL DOWN FOR ARTI CLE Taylor & Francis m akes every effort t o ensure t he accuracy of all t he inform at ion ( t he “ Cont ent ” ) cont ained in t he publicat ions on our plat form . However, Taylor & Francis, our agent s, and our licensors m ake no represent at ions or warrant ies what soever as t o t he accuracy, com plet eness, or suit abilit y for any purpose of t he Cont ent . Any opinions and views expressed in t his publicat ion are t he opinions and views of t he aut hors, and are not t he views of or endorsed by Taylor & Francis. The accuracy of t he Cont ent should not be relied upon and should be independent ly verified wit h prim ary sources of inform at ion. Taylor and Francis shall not be liable for any losses, act ions, claim s, proceedings, dem ands, cost s, expenses, dam ages, and ot her liabilit ies what soever or howsoever caused arising direct ly or indirect ly in connect ion wit h, in relat ion t o or arising out of t he use of t he Cont ent . This art icle m ay be used for research, t eaching, and privat e st udy purposes. Any subst ant ial or syst em at ic reproduct ion, redist ribut ion, reselling, loan, sub- licensing, syst em at ic supply, or dist ribut ion in any form t o anyone is expressly forbidden. Term s & Condit ions of access and use can be found at ht t p: / / www.t andfonline.com / page/ t erm s- and- condit ions Journal of Vertebrate Paleontology 33(6):1292–1299, November 2013 © 2013 by the Society of Vertebrate Paleontology ARTICLE LATE MIOCENE FISHES FROM THE EURASIAN/AFRO-ARABIAN CONTACT ZONE IN EASTERN ANATOLIA, TURKEY ALISON M. MURRAY*,1 and IZZET HOŞGÖR2 Department of Biological Sciences, University of Alberta, Edmonton, AB, T6G 2E9 Canada, ammurray@ualberta.ca; 2 TransAtlantic Exploration Med. Int. Pty. Ltd.–Viking Int. Ltd. TR−06680, Ankara, Turkey, Izzet.Hosgor@viking-intl.com Downloaded by [Izzet Hogör] at 07:28 12 November 2013 1 ABSTRACT—Two fossil fish were recently recovered from late Miocene freshwater deposits of the Muş Basin in Eastern Anatolia, Turkey. This area is in the collision zone of the Afro-Arabian and Eurasian plates, and therefore is a biogeographic contact zone or ecotone, where different biotas have been brought together. Only one fish was previously known from the area, the leuciscine cyprinid Leuciscus (Palaeoleuciscus) oeningensis. Renewed collecting at this locality provides the potential to determine much more information about the composition of the biogeographic contact zone. The two fishes described here are not named; one is placed in the family Clupeidae and the other is left as an indeterminate teleost. The fishes from the Muş Basin locality indicate that the fauna, or at least the fish portion, of the ecotone has an overall Eurasian origin, with little or no Afro-Arabian component. Recent field work in Tertiary successions of the Muş Basin area have resulted in new fossils being collected bed by bed from the Contact zones between formerly isolated land masses form a late Miocene–early Pliocene-aged sediments. The fish reported type of ecotone, an area of transition between different environhere come from the Keçidere section of the Zırnak Formation, ments that are of particular interest as places where faunal compolocated in the northeastern part of the Muş Basin (Fig. 2). sition may change abruptly (Schilthuizen, 2000). Eastern Anatolia is such a biogeographic contact zone, in the collision zone of the Geologic Setting Afro-Arabian and Eurasian plates. This area has been shown to During the Neogene, both strike-slip and extensional regimes have a reasonably high species diversity associated with it, based on studies of amphibians (Fritz and Rahbek, 2012). The present gave rise to a number of fault-bounded basins in eastern Turkey study focuses on two fossil fishes recovered from freshwater de- (Fig. 1) (Şengör et al., 1985; Hüsing et al., 2009). Tectonic evoposits of the Muş Basin in Eastern Anatolia, Turkey (Fig. 1). Al- lution was largely influenced by the convergence of the Arathough poorly preserved, they add to our knowledge of the ichthy- bian and Anatolian plates (Fig. 1). ENE–WSW-oriented folds ofauna of the area in the late Miocene, at a time when Eurasia and and thrusts developed as a result of a NNW–SSE compressional Afro-Arabia contacted each other, ending over 100 million years regime (Şengör et al., 1985; Şaroğlu and Yılmaz, 1986; Hüsing of African isolation (e.g., Otero, 2010). Fish diversity in the fresh et al., 2009). Shallow-marine carbonates, as well as fluvial and waters of the Eastern Anatolian contact zone may be the result lacustrine deposits, were deposited in eastern Turkey during the of immigrants from either plate, or invaders from marine waters. Oligocene to early Pliocene (Sancay et al., 2006; Batı and Sancay, Although only three fishes are known from the area, one previ- 2007; Hüsing et al., 2009; Murray and Hoşgör, 2012). The Muş basin is an elongated structure located north of the ously described and two noted here, with renewed collecting at this locality, there is a potential to determine much more about Bitlis–Pötürge Massif and east of the North and East Anatolian faults (Fig. 1). According to previous studies (Ünal, 1970; Şaroğlu the composition of the biogeographic contact zone. The stratigraphy and depositional paleoenvironments of the and Yılmaz, 1986; Sancay et al., 2006), the basin contains upper Muş Basin were described earlier by Ünal (1970), Sakınç (1982), Eocene to lower Miocene limestones, marls, and turbiditic sandAkay et al. (1989), Sancay et al. (2006), Batı and Sancay (2007), stones, with marine sedimentation continuous from the Oligocene Hüsing et al. (2009) and Özcan et al. (2010). During recent de- to Aquitanian (early Miocene). These deposits overlie an upper tailed stratigraphic work by one of us (I.H.) on the Paleogene- Cretaceous ophiolitic melange. Şaroğlu and Yılmaz (1986) sugNeogene successions in the Eastern Anatolia area, some inver- gested that lower Miocene limestones are widespread in the northtebrate and vertebrate remains were found in the Oligocene and ern part of the Muş area, whereas middle Miocene strata were Miocene sequences (Hoşgör and Okan, 2009; Murray and Hoşgör, not found. These sequences are unconformably covered by proba2012; Szyndlar and Hoşgör, 2012). Prior to this work, other than ble upper Miocene and younger continental clastics and volcanics a brief note of remains of a cyprinid fish (Leuciscus (Palaeoleu- based on palynomoprhs (Ünal, 1970; Şaroğlu and Yılmaz, 1986; ciscus) oeningensis Agassiz, 1832), reported by Rückert-Ülkümen Sancay et al., 2006). The Zırnak Formation, studied herein, is composed of conglom(1987), no Neogene fishes had been described from eastern erate, siltstone, marl, clayey limestones, tuff, and basalt towards Turkey. the top. Generally, yellowish siltstones, sandstones, and beige to light-yellow tuffs of the Zırnak Formation unconformably overlie the older units. There are conglomerates at the bottom of the *Corresponding author. section, followed by a very thick succession of light green marl and INTRODUCTION 1292 Downloaded by [Izzet Hogör] at 07:28 12 November 2013 MURRAY AND HOŞGÖR—MIOCENE FISHES FROM EASTERN ANATOLIA 1293 FIGURE 1. Outline of schematic geological map of Southeastern Anatolia in southeastern Turkey with major tectonic structures. Note the fossil locality indicating the studied areas: Muş Basin (drawn after Hüsing et al., 2009). yellowish to gray siltstone alternations. Towards the top of the section, thicknesses of the dark green tuff, light white lacustrine, macrofossiliforous (congeria beds) clayey limestone, and marl layers increase, with yellowish siltstone interbeds seeming to dominate. MATERIALS AND METHODS The fossil material reported here is housed in the Middle East Technical University Geology Museum, Ankara, Turkey (METUGM), cataloged under the numbers METUGM İZ11MŞ01 and METUGM İZ11-MŞ02. The latter is preserved as a carbon film on the rocks with very little relief, but the former was preserved as a partial mould allowing a silicone peel to be made of that specimen. Photographs were taken with a Nikon DXM 1200C digital camera mounted on a Zeiss Discovery.V8 stereo microscope. SYSTEMATIC PALEONTOLOGY Subdivision TELEOSTEI Müller, 1845 Subcohort CLUPEOMORPHA Greenwood, Rosen, Weitzman, and Myers, 1966 Order CLUPEIFORMES Bleeker, 1859 Suborder CLUPEOIDEI Bleeker, 1859 Family CLUPEIDAE Cuvier, 1817 (Figs. 3, 4) Material—METUGM İZ11-MŞ01, a small fish preserved in lateral view, missing most of the skull and the distal portion of the caudal fin rays. Description—This fish is small and moderately deep-bodied (Fig. 3). The anterodorsal part of the head is missing, as is the posterior portion of the tail fin. The preserved portion is 65 mm long, and we estimate the standard length (SL) would have been about 65 mm, and the total length perhaps 75–80 mm. The preserved part of the head, which includes part of the lower jaw, is 20 mm, just under one-third of estimated SL, and the body depth is 24 mm, slightly more than one-third estimated SL. Few bones of the head and pectoral girdle are preserved, and few details can be determined. The opercle is ornamented with 13 striations that radiate ventrally from the point of articulation with the hyomandibula, similar to those on some species of Alosa. The posterior portions of both the left and right anguloarticulars are preserved; these are fairly narrow, and the right one shows a deep open sensory canal on the ventral edge of the lateral surface. The limits of the right retroarticular are difficult to discern, but it appears that the retroarticular was small and confined to the ventral portion of the posterior tip of the lower jaw. The cleithrum curves gently towards the ventral margin, but the majority of the bone forms the upright portion behind the opercle. There is a long rod-like postcleithrum present. Other details are not preserved. The dorsal fin origin is closer to the head than the tail and the fin has a short base. There are 17 branched rays preceded by two unbranched rays and the fin is supported by 18 or 19 pterygiophores. The anal fin origin is posterior to the level of the dorsal fin insertion, and the fin base is about twice as long as the dorsal fin base. The anal fin is disrupted so counts are tentative, but there are about 22 branched rays preceded by two small unbranched rays. Remains of 19 pterygiophores are preserved. The pectoral fin is low on the body and contains 17 rays, and the pelvic fin is positioned below the midpoint of the dorsal fin; it contains perhaps one unbranched and six or seven branched rays. There are 40 centra visible, including one ural centrum. However, based on remains of neural spines and intermuscular bones, Downloaded by [Izzet Hogör] at 07:28 12 November 2013 1294 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013 there are an additional seven centra hidden under the opercular region, so the total number of vertebrae (including the ural centrum) is 47. The anterior few visible centra are shorter anteroposteriorly than the more posterior ones. At least two sets of intermuscular bones are present throughout the column. Dorsally, those identified based on position as the epineurals are present from the first centrum through to the caudal fin skeleton. Those more ventrally located are present on the eighth centrum through to the caudal fin skeleton; we cannot determine if ventral intermuscular bones were present on the obscured first seven centra. The ventral series of bones are associated with the centra anteriorly in the series, but the position is moved ventrally more posteriorly in the series so that by the 24th centrum the intermuscular bones are associated with the hemal arches or spines. The anterior intermuscular bones associated with the centra or articular heads of the ribs are likely from the epicentral series, as found in Clupea (Patterson and Johnson, 1995); both epicentrals and epipleurals are expected in clupeomorphs (Patterson and Johnson, 1995), but we cannot distinguish the two from one another in the fossil. There are 19 pairs of ribs, which would be associated with centra 5 through 24. Abdominal scutes are present; there is no evidence of dorsal scutes. There are four abdominal scutes anterior to the pectoral fin, and 13 scutes between the pectoral and pelvic fins, for a total of 17 prepelvic scutes. There are an additional eight postpelvic scutes preserved, but there is space in the preserved part of the fossil that potentially could have accommodated one or two more postpelvic scutes anterior to the anal fin origin. The caudal fin is better preserved than the head, but some details are still not clear, and our interpretation is somewhat tentative (Fig. 4). The first preural centrum has a neural spine that is broader but shorter than the preceding neural spines. This centrum is fused with the long, robust first uroneural, and bears the parhypural. The proximal base of the triangular first hypural does not reach any centrum, as in many clupeomorphs (e.g., Grande, 1985; Fujita, 1990). The first ural centrum is much smaller than the first preural centrum, and is fused with the narrow, stick-like second hypural. The robust second uroneural does not reach the first ural centrum. It is possible that there is a second ural centrum (not included in the vertebral counts above), but this cannot be determined with confidence. Six hypurals would be expected in a clupeid, but the bone in this area is difficult to determine and hypurals 4 through 6 cannot be distinguished. The epurals also cannot be clearly distinguished, but there are at least two, possibly three, very narrow ones present. The presence of a parhypurapophysis is indicated by the broken raised ridge on the parhypural. A count of principal fin rays is not possible because the distal portions of the rays are missing; however, if there were 19 principal rays, as expected in a clupeid, then the remains of one ventral and seven dorsal procurrent rays are preserved. Relationships—This specimen is identified as a member of the Clupeoidei based on the presence of abdominal scutes with developed ascending arms, the fusion between the first uroneural and first preural centrum, and lack of odontoids (Grande, 1985). Following characters given by Grande (1985), it is excluded from the Engrauloidea, Pristigasteroidea, Chirocentridae, Dussumeriinae, and Pellonulinae (the latter are two subfamilies of Clupeidae) by lacking characters of those groups, including lack of a strongly inclined suspensorium, presence of pelvic fins, fewer than 57 anal pterygiophores, presence of a complete series of abdominal scutes, and the first preural and first ural centra not fused. The presence of FIGURE 2. A, location map of the study area; B, the Keçidere section at least one rod-like postcleithrum further indicates that it may be from which the fish samples came. related to the Clupeidae (Grande, 1985). Inclusion in the Clupeidae is also indicated by characters of the caudal skeleton listed by Downloaded by [Izzet Hogör] at 07:28 12 November 2013 MURRAY AND HOŞGÖR—MIOCENE FISHES FROM EASTERN ANATOLIA 1295 FIGURE 3. Specimen METUGM İZ11MŞ01 and interpretive drawing, Clupeidae. Hatched lines indicate area where fossil has been damaged and only matrix remains. The right fin is preserved in articulation showing through the left side of the fossil. Abbreviations: aa, anguloarcular; apt, anal pterygiophore; as, abdominal scute; cl, cleithrum; op, opercle; pcl, postcleithrum; pop, preopercle; ptt, postemporal; q, quadrate; r, right; ra, retroarticular; scp, sensory canal of preopercle. Scale bar equals 1 cm. Chang and Maisey (2003) at various levels of their cladogram: first ural centrum much smaller than first preural centrum (in Clupea and Knightia; Clupeidae), fusion of second hypural with first ural centrum (in all taxa examined except outgroups; Clupeomorpha), and first uroneural fused to first preural centrum (in Clupea and Knightia; Clupeidae). Within the Clupeidae, the remaining three subfamilies to which this fish might belong are the Clupeinae, Alosinae, and Dorosomatinae. Characters given by Whitehead (1985) to distinguish these groups are primarily features of the jaws, which are not preserved in the fossil. However, the number of fin rays in the anal fin being about 22 indicates that it is more likely to be a clupeine or alosine rather than a dorosomatine (Whitehead, 1985). Subdivision TELEOSTEI Müller, 1845 (Fig. 5) FIGURE 4. Drawing of the caudal fin of a silicone peel of METUGM İZ11-MŞ01, Clupeidae; anterior to right. Abbreviations: ep, epural; ?ep, possible remains of a third epural; hy1–3, hypurals 1–3; ib, intermuscular bones; npu1, neural spine of first preural centrum; phy, parhypural; pu1–2, preural centra 1–2; u1–2, ural centra 1–2; un1–3; uroneural 1–3. Scale bar equals 1 mm. Material—METUGM İZ11-MŞ02, the anterior portion of a fish in lateral view; missing the tip of the head and the posterior part of the body. Description—The fossil is preserved as a carbonized film on the rock, with no clear details of the osteology; however, we can make a few counts and determine the overall morphology. The fish was long and relatively thin; the preserved portion is about 63 mm from the tip of the head to a point anterior to the caudal fin, but only 15 mm deep throughout its preserved length. The head depth and maximum body depth are almost the same (15 mm for the head and 16 mm for the body). The head is longer than deep, measuring 20 mm from the anterior tip to the presumed posterior border of the opercle. The preserved portion probably ends just after the anal fin insertion; whether or not an anal fin pterygiophore is preserved on the fossil is difficult to determine, but we consider one present. The jaws are terminal, and seem to have the articulation point with the quadrate in front of the orbit. Downloaded by [Izzet Hogör] at 07:28 12 November 2013 1296 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013 FIGURE 5. Specimen METUGM İZ11MŞ02 and interpretive drawing, Teleostei suborder indeterminate. Abbreviations: ? apt, bone impressions possibly of anal pterygiophores; cl, cleithrum; ?hs, presumed hemal spines; op, opercle; pop, preopercle; rr, ribs of right side. Scale bar equals 1 cm. The dorsal fin insertion is situated at a level posterior to the posterior edge of the pelvic girdle, but above the pelvic fins themselves. The dorsal fin is small, having maybe 10 branched rays and one leading unbranched ray. The first dorsal pterygiophore angles anteroventrally to insert between the posterodorsally angled neural spines of centra 11 and 12, but the fin origin is situated above the 14th centrum. The first two pterygiophores each bears a single ray, although there could have been one or more procurrent rays associated with the first pterygiophore that are not preserved. There are probably eight pterygiophores supporting the dorsal fin. There are 15 or more pectoral rays, and the fin is positioned low on the body. There are about 11 pelvic rays preserved, indicating that six or more were present on each side. The left and right pelvic girdles seem to meet in the midline or be quite close to one another. There are a total of 24 centra preserved, which are slightly longer than high but have a strong tall neural arch attached. Ribs start on the third centrum and continue to centrum 14, for a total of 11 pairs. The ribs are long, reaching almost to the ventral body wall. Intermuscular bones are associated with the first through 12th or 13th neural spines. Relationships—The poor preservation of this fish makes it difficult to positively identify it. However, it is a teleost based on the sculptured centra (Patterson and Rosen, 1977; Arratia, 1997) and a one-to-one ratio of median fin rays with supports (Patterson, 1992), and, combined with the abdominally placed pelvic fin, long robust ribs, and overall shape of the body, we consider it to be a fairly primitive teleost. water deposits (Grande, 1985; Reichenbacher, 1988). The majority of these Miocene fishes have been assigned to the extant clupeid genera Alosa, Clupea, and Clupeonella, although Grande (1985) indicated that some might be better assigned to other previously named or new genera with renewed study. At least some of these Tethyan and Paratethyan species have been interpreted as endemic to smaller areas (e.g., Rückert-Ülkümen, 2006). Clupeids, particularly those of the genus Alosa, are not uncommon visitors to fresh waters, particularly during spawning, and may also be found in landlocked fresh waters (Nelson, 2006). However, fossil clupeids from freshwater deposits are much less common than in marine deposits, as riverine deposits rarely contain identifiable clupeid elements (Murray et al., 2005) and lake forms are rare (with some notable exceptions such as those from the Green River lacustrine deposits). The Miocene marine clupeids previously reported from Turkey (e.g., Rückert-Ülkümen, 2006) are from regions much further west (north of the Sea of Marmara) than the Muş locality. Miocene Paratethyan clupeids were reported as coming from freshwater deposits in Germany (Meyer, 1851; Grande, 1985); however, these sediments were thought to have a marine influence (Meyer, 1851) and more recent study shows them to be brackish water (Reichenbacher, 1988). Two clupeids are known from these deposits, Clupeonella humillis and C. cornuta, the latter based on otoliths (Reichenbacher, 1988, 1993, 2012). The Miocene freshwater clupeid noted here may, therefore, be an undescribed species. In the middle to late Miocene, the Mediterranean region was affected by the reduction and eventual disappearance of its connections with the Atlantic Ocean through the Betic (Western Europe) and Rifean (Northern Africa) basins, prior to forming DISCUSSION a new connection in the Pliocene through the modern Strait of Many clupeids have been reported from marine deposits of Gibraltar (e.g., Duggen et al., 2003). The Paratethys also became Miocene age in the Tethys and Paratethys, but none from fresh- separated from the Mediterranean during the late middle Miocene MURRAY AND HOŞGÖR—MIOCENE FISHES FROM EASTERN ANATOLIA 1297 Downloaded by [Izzet Hogör] at 07:28 12 November 2013 FIGURE 6. The relative locations of the Tabriz Basin of Iran, and the fossil locality in the Muş Basin, Eastern Anatolia, Turkey. (Serravallian), and the waters of the Paratethys became isolated in smaller basins resulting in a gradual change to brackish and eventually fresh waters (Rögl, 1998; Orszag-Sperber, 2006). In the isolated Paratethyan basins, an endemic fauna arose, adapted to the brackish to fresh waters (Rögl, 1998; Orszag-Sperber, 2006). The lacustrine fishes reported here would likely have been associated with one of those isolated basins, as would have those fishes reported from the Tabriz Basin of Iran, about 400 km to the east of the Turkish locality (Fig. 6). The Tabriz Miocene locality has produced skeletons of silversides (Atherina atropatiensis), a cyprinodont (Aphanius (or Brachylebias) persicus), and an indeterminate cyprinid, as well as teeth that have been assigned to three different cyprinids (Carnevale et al., 2011; Gaudant, 2011; Reichenbacher et al., 2011). No clupeid remains have been reported from there. The Muş Basin in Eastern Anatolia has produced a cyprinid (RückertÜlkümen, 1987), an indeterminate teleost, and a clupeid. Other than the cyprinids, both faunas comprise fishes that can be considered secondary freshwater fishes, indicating that they may well have arisen from ancestors trapped in the isolated basins of the Paratethys. The cyprinids, on the other hand, in both localities have all been allied with the Leuciscinae, a subfamily that has a wide distribution around the Mediterranean (Perea et al., 2010). This area (the Balkan Peninsula, Anatolia, and Iran) has been proposed as an important area from which leuciscine cyprinids colonized the rest of Europe in the late Paleogene (Perea et al., 2010). If correct, this indicates that the leuciscines were present in the area long before the late Miocene. Although cyprinids are normally considered to be intolerant of saline waters, this should not be taken as absolute, as some have been found in brackish waters and others may be quite tolerant of higher salinities (Coad, 1987, 1996). However, it seems that the leuciscines were part of the freshwater fauna of Eurasia. Cavender (1991) noted that cyprinids are thought to have arisen in Asia, and migrated to Europe from there, mainly based on the great diversity of living forms in Asia. Additionally, some of the earliest cyprinids, of possible Eocene age, are known from Sumatra, Indonesia. Although the majority of the Sumatran species were placed in living genera (Sanders, 1934), preliminary study of new collections from the site indicates that many of the species will be reassigned to new fossil genera (A.M.M., pers. observ.). Perea et al. (2010) suggested that the leuciscines first colonized Europe in the early Oligocene from southwestern Asia using a land mass made up of the Balkans, Anatolia, and Iran. They further proposed that the leuciscines then diversified in this land mass, and only arrived in Central Europe in the mid-Miocene, about 20 Ma, when the marine barrier separating the landmass from the rest of Europe closed, resulting in the separation of the Tethys and Paratethys. It is quite possible that the older freshwater fauna of the Balkan/Anatolian/Iranian block containing leuciscines was augmented by marine invaders (Atherina and the clupeid) from the Paratethys. Based on the fishes now known from the Muş Basin, the ichthyofauna is a mix of elements with an overall Eurasian origin, sharing little with Afro-Arabia (e.g., see Coad, 1996). New discoveries from the Anatolian and Iranian regions are increasing our understanding of this less-studied part of the Paratethys. The area is a critical collision zone between the AfroArabian and Eurasian plates, and is therefore important as an area of faunal exchanges. Reichenbacher et al. (2011) noted that there was likely a marine connection to the area around Tabriz, Iran, and suggested that the connection was to the southern part of the Caspian Sea to the east. However, they noted that fossil evidence was lacking for this connection based on the fish fauna. The new discoveries of fishes from the Eastern Anatolian Muş Basin indicate a potential new direction for investigation, to the west and perhaps south. These two sites are expected to provide more information, with renewed collections providing more fish taxa with which to unravel biogeographic connections in an area that is marked with questions on recent maps (e.g., Popov et al., 2006:fig. 4). ACKNOWLEDGMENTS Our thanks to T. Prikryl and B. Reichenbacher for very helpful reviews, and editorial comments from M. Friedman that improved the manuscript. We thank A. Lindoe for making the silicone peel of the specimen and R. B. Holmes for comments on earlier drafts of the manuscript. A.M.M. is supported by National Science and Engineering Research Council of Canada Discovery Grant 327448. 1298 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013 Downloaded by [Izzet Hogör] at 07:28 12 November 2013 LITERATURE CITED Agassiz, L. 1832. Untersuchungen über die fossilen Süsswasser-Fische der tertiären Formationen. Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde, Heidelberg 1832:129–138. Akay, E., E. Erkan, and E. 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