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Zeitschrift/Journal: Beiträge zur Paläontologie

Jahr/Year: 2009

Band/Volume: 31

Autor(en)/Author(s): Konidaris George E., Koufos George D.

Artikel/Article: The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection - 8. Proboscidea 139-155 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Beitr. Paläont., 31:139-155, Wien 2009

The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 8. Proboscidea

by

George E. Konidaris1* & George D. Koufos^

Konidaris, G.E. S c Koufos, G.D., 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 8. Proboscidea. — Beitr. Palaont., 31:139-155, Wien.

Abstract geographische und stratigraphische Verteilung dieser zwei Gattungen diskutiert. Die biochronologischen Daten, die Recent excavations in the Mytilinii basin of Samos Island, anhand der beiden Proboscidier-Arten festgestellt wurden, Greece, unearthed a rich mammal fossil collection. Among entsprechen der übrigen Fauna im Mytilinii Becken und the new Samos material, there are some proboscidean deuten das mittlere Turolium als Alter, MN 12 (7.1 - 7.0 remains which are described in the present article. The Ma) für die studierten Proboscidier an. Samos proboscidean assemblage is comprised of a few dental remains, nevertheless two species were identified: Schlüsselworte: Obermiozän, Samos, Griechenland, Choerolophodon pentelici and Zygolophodon turicensis. The Mammalia, Proboscidea, Systematik. material is compared with other known Miocene probos­ cideans from Greece and Eurasia, and the geographic and stratigraphic distribution of these two genera is discussed. 1. Introduction The biochronological data offered by the two proboscide­ ans corresponds with the other faunal data from Mytilinii The mammal localities of Samos Island have been well- basin and suggests a Middle Turolian age, MN 12 (7.1 - 7.0 known since the second half of the 19th century(F orsyth Ma) for the studied proboscideans. M ajor, 1888,1894). Later on, several palaeontologists or fossil dealers excavated on the island and numerous fossils Keywords: Late Miocene, Samos, Greece, Mammalia, have been unearthed. The Samos mammalian fossils are Proboscidea, Systematics. housed in several European and American museums and institutions today. More data about the excavations and the mammal collections from Samos is given in Koufos (this Zusammenfassung volume), and for the fossiliferous sites in Kostopoulos et al. (this volume). Neue Ausgrabungen im Mytilinii Becken der Insel Samos, Among the collected material from Samos the probos­ Griechenland, brachten eine reiche Fossilienansammlung cideans are relatively rare. Two proboscidean taxa are von Säugetieren hervor. In dieser neuen Ansammlung mentioned in the first Samos collection: “Mastodon (= gibt es einige Proboscidier, die im vorliegenden Artikel Choerolophodon) pentelici and “Mastodon” (= Zygolophodon) beschrieben werden. Die Proboscidier-Funde von Samos turicensis (Forsyth M ajor, 1894). In the Samos faunal enthalten wenige dentale Reste, nichtsdestoweniger wur­ lists given by Solounias (1981:tab. IV, V), the species den zwei Arten identifiziert: Choerolophodon pentelici und }Mammut borsoni (= Zygolophodon turicensis or Mastodon Zygolophodon turicensis. Das Material wird mit anderen tapir aides), Stegotetrabelodon grandincisivus (= Tetralophodon Proboscidiern des Miozäns verglichen, welche aus Grie­ longirostris), Choerolophodon pentelici and Deinotherium chenland und aus Eurasien bekannt sind, und es wird die cf. giganteum are mentioned, but the individual locality records include only Tetralophodon longirostris, Choerolopho­ don pentelici and Deinotherium cf. giganteum (Solounias, 1981 :tab. VII). Some years later, Bernor et al. (1996) men­ :) MSc. George E. K o n id a r is and Prof. George D. K o u f o s , tioned the taxa Mammut borsoni, Tetralophodon longirostris, Aristotle University of Thessaloniki. Department of Geology, Choerolophodon pentelici and Deinotherium giganteum from Laboratory of Geology and Palaeontology, GR-54124 Thes­ the Main Bone Beds of Mytilinii Formation. In the lists saloniki, Greece, e-m ail: [email protected]; koufos@ of NOW (2007) Deinotherium giganteum is noted from geo.auth.gr Samos and Samos A l, while Choerolophodon pentelici is ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

140 Beitr. Palàont., 31, Wien, 2009

listed from Samos A l. Bearing all the above mentioned root. Moreover, choerodonty and ptychodonty are well- facts in mind, there is no certain and clear opinion about expressed. The maxillary fragment M TLB-126 preserves the Samos proboscideans. The material has never been the right M l and both M 2 which are just erupted (PI. 1, studied systematically, except for a more detailed study of fig. 2a). The M l is broken buccally, lingually and distally. C. pentelici, housed at the N H M W and at the HNHM The morphology of the left M 2 is not well-preserved, so a (ScHLESINGER, 1917, 1922). description is not possible. The palate is relatively narrow The newly collected material is relatively scanty and and shallow. The anterior part of the choanae is situated includes very few dental remains, which are described at the distal part of the M2. in the present article. In this study an isolated tooth The persistent right M l (PI. 1, fig. 2b) has a rectangular (PMMS - 53) from the collection of Prof. J. Melentis shape with three transverse lophs. The protocone and the (Thessaloniki, Greece) was also included. The material is paracone, as well as the conelets between these two main described and compared with other known remains from cusps of the first loph, are not well visible because of wear, Greece and Eurasia. The nomenclature of the bunodont which has merged them. In the transverse valley between teeth is according to Tassy (1983) and of the zygodont the first and second loph, there is a rather strong anterior teeth according to Tobien (1975). A ll the studied material pretrite central conule of the second loph. In the second is housed at NHMA. loph, the two main cusps are partly broken buccally and lingually, respectively. The posttrite halfloph is formed by Abbreviations: two conelets, the pretrite by one larger conelet. In the distal HNHM = Hungarian Natural History Museum, Budapest wall of the hypocone there is a small and low cusplet, that N H M A = Natural History Museum of the Aegean, Mytilinii, closes the lingual opening of the transverse valley between Samos Island, Greece the second and the third loph. In the middle of the second MNHN = Muséum National d’Histoire Naturelle, Paris transverse valley, there are four central conules (the central N H M W = Naturhistorisches Museum Wien is larger), dividing the transverse valley into two parts. The M RG = Maragheh, Iran conules are situated in a semicircle and connect the second M TLA = Mytilinii - 1A, Samos Island, Greece and third loph. In the third loph, the two main cusps are MTLB = Mytilinii - IB, Samos Island, Greece broken buccally and lingually. In the distal part of the PIK = Pikermi, Greece tooth some additional cusplets are developed, forming a PM M S = Palaeontological Museum of Mytilinii, Samos Island, cingulum, which is partly broken. Greece The right M 2 (PI. 1, fig. 2c) has three lophs. The mesial SAM = Samos, old collections cingulum is almost semi-circular and consists of a series SLQj= Salonique, Axios Valley, Greece (Arambourg-Collection, of small cusplets, which decrease toward the buccal side. housed in MNHN) The cusplet of the mesial cingulum situated most buccally is large, but still smaller than the lingual ones. In the first loph the protocone is larger than the paracone. The 2. Systematics posttrite halfloph is formed by the paracone and three conelets, which are all about the same size and fused Family Gomphotheriidae Hay, 1922 to each other, as well as with the paracone; the pretrite Subfamily Choerolophodontinae G aziry, 1976 halfloph consists of a large protocone and a single and more isolated conelet. In the transverse valley between Genus Choerolophodon S chlesinger, 1917 the first and second loph, there is a small posterior pretrite central conule of the first loph. In the second loph the Choerolophodon pentelici (Gaudry & Lartet, 1856) hypocone is larger and situated well behind the metacone. (PI. 1, figs. 1, 2) The three posttrite conelets are of the same size and more or less fused to each other, as well as to the metacone. Locality: M ytilinii-lB (MTLB), Mytilinii Basin, Samos, There are two pretrite conelets, the inner being larger Greece. than the other. In the second transverse valley there are Age: Middle Turolian, MN 12 (Late Miocene); 7.1 - 7.0 three central conules of medium size and a larger fourth Ma. one. In the third loph, the posttrite main cusp and the Material: DP2 sin, MTLB-11; maxilla with M l - M2 two conelets are of the same size and fused to each other, dex and M 2 sin, M TLB-126. while the pretrite halfloph is formed by a stronger main Description: The DP2 has a triangular occlusal outline cusp and two smaller conelets. In the distal cingulum the with a distolingual enlargement (PI. 1, fig. 1). The para- cusplets are connected to each other and situated almost cone is slightly larger than the protocone and both are on a straight line. Their size decreases from the lingual joined, forming a large cusp in the mesial part of the to the buccal side. tooth. The metacone is much smaller than the mesial Discussion: The oldest choerolophodons known from main cusp, whereas the hypocone is very weak. There the Early - Middle Miocene of Africa are referred to as is a strong mesial-mesiolingual cingulum, as well as a Afrochoerodon (Pickford, 2001). This genus includes the distal cingulum, which is more developed on the lingual following Early - Middle Miocene African and Eurasian side. The tooth bears one mesial and one larger distal choerolophodont species (Pickford, 2001; Sanders, 2003): ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. &K oufos, G.D., Proboscidea. 141

Figure 1: Second upper decidu­ ous premolars of Choerolophodon pentelici from various localities. a. DP2 sin, MTLB-11; b. DP2 dex (reverted), MNHN-PIK 3665, Pikermi, Greece; c. DP2 dex (reverted), NHMW-1914 no. 13, Samos, Greece d. DP2 sin, MNHN-SLQ4124, Axios Valley, Greece; e. DP2 sin, AK2 - 300, Akka^dagi, Turkey (taken from Tassy, 2005); f. DP2 sin, NHMW-MRG-A 4868, Ma- ragheh, Iran.

mA. kisumuensis (M cInnes, 1942), Eady - Middle Miocene. upwards. Concerning the morphology of the teeth, the It is known from the localities of Maboko and Cheparawa permanent cheek teeth o f A. kisumuensis lack choerodonty in Kenya and from Wadi Moghara in Egypt. and ptychodonty and they are slightly covered with cement % A. palaeindicus (Lydekker, 1884), Early Miocene, known in the valleys. In contrast, choerodonty and ptychodonty from the Bugti beds and possibly from Siwaliks (Pakistan). are well expressed in the permanent cheek teeth of the According to Ta ssy (1990), it is possible thatA. palaeindicus Eurasian choerolophodons, which are usually heavily and A. kisumuensis are conspecific. Recent data from Bugti covered with cement. suggests that the first occurrence of choerolophodons in In Europe, the oldest known choerolophodon is Choerolo­ Eurasia is dated between 17.0 - 16.0 Ma, corresponding phodon chioticus (Tobien , 1980) found in the locality to the end of Early Miocene (Steininger, 1999). Thymiana of Chios Island, Greece (Tobien, 1980). The mA. ngorora (M aglio, 1974), Middle - Late Miocene (13.0 locality is dated to Late Orleanian, MN 5; the palaeo- - 9.5 Ma), known from Fort Ternan and Ngorora in Kenya. magnetic record suggests an age of ~15.5 M a for the fos- A. zaltaniensis (G az ir y , 1987) from North Africa could siliferous levels of Thymiana (B onis et al., 1997a, b, 1998; be a synonym oiA. ngorora (P ickfo rd , 2001). K oufos et al., 1995; K ondopoulou et al., 1993). Based The type species of the genus Afrochoerodon kisumuensis on more similarities of C. chioticus with. Afrochoerodon than and the Late Miocene Eurasian choerolophodons differ with the more derived Eurasian choerolophodons, P ick­ in various characters. P ickfo rd (2001) mentions that ford (2001) transferred it to Afrochoerodon. The distinction A. kisumuensis is characterized by a short and high skull between the Middle and Late Miocene choerolophodons with a steeply inclined facial region, whereas the skull of is clear; the first choerolophodons have more primitive the Eurasian choerolophodons is elongated and flattened. characteristics while the later forms are highly specialized. Moreover, the orbits in A. kisumuensis are situated above T obien (1980) notes that A. chioticus belongs to a more the M3s and at about % of the height of the skull, while primitive evolutionary stage with regard to C. pentelici in the Eurasian choerolophodons the orbits are behind owing to the narrow temporal region, the elongated skull the rear of the M3s near the top of the skull. The tusks base and the incipient choerodonty, ptychodonty and 01 A. kisumuensis emerge downwards and then curve cementodonty of the Chios specimen. forwards, whereas in the Eurasian choerolophodons the During Late Miocene the genus Choerolophodon was a tusks emerge at a sub-horizontal angle and then curve common representative of the proboscideans in the faunas ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

142 Beitr. Palàont., 31, Wien, 2009

Figure 2: Scatter diagram (length /breadth) comparing the DP2s of Choerolophodon from various localities.

30 - +

? X £ + X « + * Î X ■a 25 " « \* X )-0) 09 □ Q.

20 - of the Eastern Mediterranean □ □ region, distributed across Eurasia from Southeastern Europe to China, while it is absent in Western and Cen­ 15 tral Europe and Northern 20 25 30 35 40 Asia. The type species C. Length (mm) pentelici is known from sev­ eral Greek localities, such as ♦ C. pentelici, Samos, MTLB-11 ■ C. pen telici, Sam os (NHMW) Pikermi (type locality), Axios A C. p en telici, Pikermi (MNHN) X C. pentelici, Akkasdagi (TASSY, 2005) Valley, Samos, Kerassia, Ma- ramena and Nikiti (Koufos, X C. pentelici, Axios valley (MNHN) % C. pen telici, M aragheh (MNHN, NHMW) 2006) . It is also very common + C. corrugatus, Siwaliks (TASSY, 1983) — C . anatolicus, Sinap (SANDERS, 2003) in the Late Miocene faunas of Bulgaria, Turkey, Iran □ C. anatolicus, Kayadibi (GAZIRY, 1976) and Afghanistan (NOW, 2007) . Contemporaneous with C. pentelici is C. corruga­ the mandible. The symphyseal angulation exists in C. tus (Pilgrim, 1913), which is present in the Nagri and anatolicus, though in a lesser degree than in C. corrugatus Dhok Pathan Formations of middle Siwaliks, Pakistan. (Sanders, 2003). Moreover, the deciduous premolars of The choerolophodont material from the Sinap Formation C. anatolicus are smaller than those of the other two Late (Turkey) was assigned byO zansoy (1965) to C. anatolicus. Miocene choerolophodons. Sanders (2003) notes that the Sanders (2003) proposed that the subspecies “C.pente­ molars of C. anatolicus from Sinap differ from those of C. lici lydiensis” also belongs to C. anatolicus and thus the pentelici in having simpler crowns with choerodonty and material from Esme Akçakôy, Kayadibi, Kemiklitepe-D ptychodonty weaker expressed, therefore being closer to and Gôkdere was included in this species. Furthermore, A. ngorora, A. chioticus and A. kisumuensis. he suggested that C. anatolicus existed during the whole There are no mandibular remains of Choerolophodon in Vallesian and then was replaced byC. pentelici. the new Samos material which could allow a certain The distinction between the three Late Miocene species determination of the species. The old material described is more complicated. Tassy (1983) notes the difficulty in by Schlesinger (1917:Taf. XXV, Abb. 1) as Mastodon distinguishing isolated teeth of C. pentelici from those of (Choerolophodon) pentelici, does not show angulation and C. corrugatus, if there is no indication of the geographic the mandibles are similar to the type material from Piker­ location. The main distinguishing characteristics among mi. Concerning the size of the deciduous premolars, the the two species are the angulation between ascending and DP2 from Samos is in fact larger than that of C. anatolicus horizontal ramus and the angulation of the symphyseal (Fig. 2). This is also supported by the DP2 dimensions in rostrum on the mandibular corpus, which is strong in C. the old collection from Samos. Additionally, choerodonty corrugatus. On the other hand, C. pentelici is characterized and ptychodonty are visibly present (Fig. 1). by the absence of the angulation between ascending and The DP2 of the holotype from Pikermi (Fig. lb) is worn horizontal ramus, and the symphysis is situated at the and distally compressed, owing to the close contact with extension of the horizontal ramus. the DP3. However, some basic characteristics can be The distinguishing characteristics between C. anatolicus observed: the tooth forms two bunodont lophs; the two and C. pentelici are also related to the morphology of mesial cusps are connected; the metacone is well-separated ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. Sc Koufos, G.D., Proboscidea. 143

Figure 3: First and second upper molars of Choerolophodon from various localities.

a. M l dex, C. pentelici, MTLB- 126, Samos, Greece; b. M l dex, C. pentelici, NHMW-1913 no 12, Samos, Greece; c. M l dex, C. pentelici, MNHN-SLQT122, Axios Valley, Greece; d. M l dex, C. pentelici, MNHN-MRG-nn, Maragheh, Iran; e. M l dex, C. corrugatus, GSP 15001, Siwaliks, Pakistan (taken from T a s s y , 1983); f. M2 dex, C. pentelici, MTLB-126, Samos, Greece; g. M2 sin (reverted), C. pentelici, NHMW-MRG-nn, Maragheh, Iran; h. M2 dex, C. corrugatus, GSP 15001, Siwaliks, Paki­ stan (taken from T a ssy , 1983).

from the hypocone and it is larger-sized; the tooth has a veloped on the lingual side of the tooth. The distolingual mesial cingulum and a distolingual enlargement. These enlargement is less in NHMW-SAM-1914 no. 13, while features are similar to those of Samos DP2, but that of in NHMW-SAM-A 4355 it is more expressed. Pikermi is slightly larger in size (Figs. 1, 2). In Arambourg’s collection from Axios Valley (Macedo­ In the old collection of Samos housed at NHMW, there nia, Greece), housed at MNHN, there are two DP2s are two skulls (N HM W -SAM -1914 no. 13 and NH M W - of C. pentelici (Fig. Id). Their direct comparison to the SAM-A 4355) which were originally described by S ch- studied DP2 from Samos suggests similar morphology l e sin g e r (1917), and each of them retains both DP2s. and size (Figs. 1, 2). However, the distolingual enlarge­ The morphology of these deciduous premolars fits in well ment of the DP2s from Axios Valley is less intense. A with that of MTLB-11 (Figs, la, c). The first loph bears DP2 of C. pentelici is described by T a ss y (2005) from the typical massive cusp, while in the second loph the the Turkish locality Akka^dagi (Fig. le). In this tooth, metacone and the hypocone are well separated. In NH- the choerodonty and ptychodonty are less expressed M \V-SAM-A 4355, these two distal cusps are larger and than in the Samos material, probably due to the stage the sulcus between them is slightly deeper. However, the of wear. Concerning the size, the Akka^dagi DP2 is ratio of the size between the metacone and the hypocone wider than that of Samos (Fig. 2), nonetheless the distal remains the same. The mesial cingulum is well-expressed, enlargement of the tooth is of the same degree. Both while there also is a distal cingulum, which is more de­ teeth generally share the same morphology (Figs, la, e). ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

144 Beitr. Palaont., 31, Wien, 2009

M l Figure 4: Scatter diagrams (length/breadth) comparing the Mis and M2s of the sub-family Choerolophodontinae from vari­ ous localities.

♦ C. pentelici, Samos, MTLB-126 O C. pentelici, Samos (NHMW) □ C. pentelici, M aragheh (MNHN) + C. pentelici, Axios valley (MNHN) • C. corrugatus, Siwaliks (TASSY, 1983) ■ C. anatolicus, Kayadibi (GAZIRY, 1976)

M2

♦ C. pentelici, Samos, MTLB-126 ■ C. pentelici, M aragheh (NHMW) A C. corrugatus, Siw aliks (TASSY, 1983) 0 A . kisumuensis, Cheparawa (PICKFORD, 2001) — C. anatolicus, K ayadibi (GAZIRY, 1976)

Second deciduous premolars ofC. pentelici are also known boscidean of trilophodont grade. The choerodonty of Ml is from Maragheh (Iran). Some choerolophodont remains of similar to that of the Late Miocene forms ofChoerolopho- this locality are housed at N H M W and were originally don from Axios Valley, Maragheh and Siwaliks (Fig. 3). In described by S chlesinger (1917). The best preserved the old collection of Samos, M l is known from a skull of specimen is a maxilla preserving all the deciduous teeth C. pentelici (N H M W -SAM -1913 no. 12) which bears both in both sides (N H M W -M RG-A 4868). The DP2s are DP4-M1 series. The morphology of M l (Fig. 3b) is similar in an advanced stage of wear, though the morphological to that of the studied M l (Figs. 3a, b). The comparison of characteristics can be observed. As all DP2s, the teeth bear the first loph is not possible because it is too worn in the two lophs, the second one with a distolingual enlargement M l of MTLB-126, nonetheless, they seem to be similar. which is of the same degree as in M TLB-11. A ll their Both teeth have a strong anterior pretrite central conule morphological features fit in well with those of the Samos of the second loph. Moreover, there is one conelet in the specimen (Figs, la, f). The only difference seems to be the internal part of the hypocone. They differ, however, in the less intense mesial and distal cingulum, but this could be internal part of the metacone, which bears two conelets in due to attrition. Concerning the size, there is a resemblance M TLB-126 instead of one in the N H M W -SAM -1913 no. between Samos and Maragheh specimens (Fig. 2). The 12. In the transverse valley between the second and third studied maxilla (MTLB-126) obviously belongs to a pro­ loph of the N H M W -SAM -1913 no. 12, there are some ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

K onidaris, G.E. &K oufos, G.D., Proboscidea. 145

weak conules, which do not block the transverse valley. Samos the valleys are wider, especially on the buccal side In both specimens the sulcus of the third loph is wide and of the tooth. The distal cingulum seems to be the same deep. The M l of NHM W -SAM -1913 no. 12 is strongly size, as is the overall tooth size, too (Fig. 4). covered with cement. Concerning the M2, Schlesinger (1917) described a The M l from Axios Valley (MNHN-SLQH122) differs left M2 of C. pentelici coming from Maragheh (N H M W from the M TLB-126 in the placement of the conelets be­ MRG-nn). In this tooth (Fig. 3g) the second and third tween the second and the third loph (Fig. 3c). The conelets posttrite halflophs, as well as, the distal cingulum are of the third loph of MNHN-SLQH122 are situated in broken. The morphology of the rest of the tooth fits in a straighter line than those of MTLB-126 (Figs. 3a, c). quite well with the studied M2 (Fig. 3f). Moreover, there is a posterior pretrite central conule of the In the M2 of C. corrugatus, from Siwaliks (Fig. 3h) pty- second loph of MNHN-SLQH122 that is absent in the chodonty is more expressed, while in the Samos one it is tooth from Samos. Apart from that, the morphology of absent. A ll three lophs of the M 2 from Siwaliks are broader the second loph is the same in both samples, showing one than those of M TLB-126 (Tab. 1 & Fig. 4). There is a strong anterior pretrite central conule of the second loph, variation concerning the length, as the length of C. cor- two conelets in the inner part of the metacone and one rugatus varies from 110 mm up to 130 mm (Tassy, 1983). larger in the inner part of the hypocone. The distal cingu­ Apart from this, both teeth morphologically resemble lum in the two specimens seems to be equally sized. each other. The studied material from Samos has been directly com­ The above mentioned comparison of the teeth indicates pared with the sample of C. pentelici from Maragheh, that the morphology and size between the various speci­ housed at MNHN. The M l from Maragheh (Fig. 3d) mens of C. pentelici is similar; the small size differences shares the same morphological features with the Samos are probably due to sexual dimorphism, variability and one. They differ in the presence of the small and low cus- wear stage. plets situated on the lingual side of the first and second transverse valley that are entirely missing in the Samos Family Mammutidae Hay, 1922 specimen. The Maragheh M l is larger than M TLB-126 Subfamily Mammutinae Hay, 1922 (Fig. 4). Although the general morphology of the Samos M l is Genus Zygolophodon Vacek, 1877 similar to that of C. corrugatus from Siwaliks, Pakistan (Fig. 3e), there are some distinctive characteristics. In Zygolophodon turicensis (ScHlNZ, 1824) comparison to Samos, the anterior pretrite central conule (PI. 1, fig. 3) of the second loph is also strong, but the conelet in the inner part of the hypocone seems to be rather weak. In Locality: The studied specimen belongs to the collection the transverse valley between the second and the third of Prof. J. Melentis, housed at NHMA. It was collected loph there is one central strong conelet but there are not from M T L A site in Adrianos ravine (see Koufos, this additional conules connecting the second and third loph as volume). in the Samos specimen. Furthermore, the two transverse Age: M iddle Turolian, MN 12 (Late Miocene); 7.1 - 7.0 valleys are narrower, meaning that the three lophs of the M a tooth are closer to each other, whereas in the M l from Material: dp4 dex, PMMS-53.

Figure 5: Fourth lower deciduous molars of zygodons from various localities.

a. dp4 sin (reverted), Zygolophodon turicensis, MNHN-Si 11, Simor- re, France, (taken from Tassy, 1977); b. dp4 dex, Zygolophodon turicensis, PMMS-53, Samos, Greece; c. dp4 dex, M am?nut cf. borsoni, MNHN-PIK-3613, Pikermi, Greece. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

146 Beitr. Palaont., 31, Wien, 2009

.20

Figure 6: Geographic distribution of the sub-family Choerolophodontinae in Africa, Southwestern Asia and Europe.

Legend: 1. Bugti, India; 2. Maboko, Kenya; 3. FortTernan, Kenya; 4. Cheparawa, Kenya; 5. Ngorora, Kenya; 6. Nakali, Kenya; 7. Gebel Zelten, Libya; 8. Wadi Moghara, Egypt; 9. Thymiana, Greece; 10. Axios valley (Pentalophos, Xirochori, Ravin de la Pluie, Ravin des Zouaves, Vathylakkos, Prochoma, Dytiko), Greece; 11. Nikiti, Greece; 12. Pikermi, Greece; 13. Samos, Greece; 14. Titov Veles, FYR of Macedonia; 15. Various sites in Bulgaria; 16. Varnitsa, Moldova; 17. Kalfa, Moldova; 18. Tiraspol, Moldova; 19. Chobruchi, Moldova; 20. Novaja Emetovka, Ukraine; 21. Eldari I, Georgia; 22. Yulafli, Turkey; 23. Batalcesme, Turkey; 24. Kuciikcesmece, Turkey; 25. Giilpinar, Turkey; 26. Gokdere, Turkey; 27. Eminova, Turkey; 28. Inonu 2, Turkey; 29. Sinap, Turkey; 30. Corakyerler, Turkey; 31. Sofca, Turkey; 32. Akin, Turkey; 33. Garkin, Turkey; 34. Kemiklitepe, Turkey; 35. Esme Ak9akoy, Turkey; 36. Kayadibi, Turkey; 37. Akka^dagi, Turkey; 38. Dadasun, Turkey; 39. Duzyala, Turkey; 40. Maragheh, Iran; 41. Siwaliks, Pakistan. The data was taken from NOW (2007).

Description: The right dp4 is trilophodont, rootless and of the lingual halflophids is vertical. In the first lophid has a rectangular shape (Fig. 5). The halflophids of the there is an anterior and a weaker posterior crescentoid first lophid are situated opposite each other, whereas the of the first pretrite. The metaconid bears at least three halflophids of the second and third lophid are slightly conelets and the protoconid two conelets. These conelets oblique to the longitudinal axis of the tooth, pinted are laterally merged and situated in a straight line. There mesiolingually - distolabially. The number of conelets is is a weak zygodont crest in the posttrite halflophid. The higher in the posttrites, so that the median sulcus is shifted second lophid is worn, and as a result the conelets cannot labially. The lophids are separated by transverse valleys be clearly distinguished. The anterior crescentoid of the which are open lingually and buccally. The buccal wall of second pretrite is weak due to wear. In the third lophid, the buccal halflophids is inclined, while the lingual wall the tip of the posttrite main cusp and the conelets are ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. &K oufos, G.D., Proboscidea. 147

Figure 7: Geographic distribution of Zygolophodon species in Africa, Southwestern Asia and Europe.

Legend: 1. Sansan, France; 2. La Romieu, France; 3. Esvres-Marine Faluns; 4. Pontlevoy, France; 5. Simorre, France; 6. Poudenas, France; 7. Malartic, France; 8. Bourg-Saint-Bernard, France; 9. Montbrun-Bocage, France; 10. Castelnau, France; 11. Saint-Gaudens, France; 12. El Arquillo 1, Spain; 13. Concud, Spain; 14. Benavente, Spain; 15. Las Casiones, Spain; 16. Cerro delaGarita, Spain; 17. Gau-Weinheim, Germany; 18. Esselborn, Germany; 19. Wartenberg, Germany; 20. Breitenbrunn, Germany; 21. Dinotheriensande, Germany; 22. Bermersheim, Germany; 23. Elgg, Switzerland; 24. Devinska Nova Ves, Slovakia; 25. Novalky, Slovakia; 26. Gravitelli, Italy; 27. Lapushna, Moldova; 28. Chimishlija, Moldova; 29. Taraklia, Moldova; 30. Borshchi, Ukraine; 31. Novo-Elizavetovka, Ukraine; 32. Axios valley, Greece; 33. Halmyropotamos, Greece; 34. Samos, Greece; 35. Akqahisar 1, Turkey; 36. Dera Bugti 6, Pakistan; 37. Wadi Moghara, Egypt. The data was taken from NOW (2007).

broken. There is a mesial and a stronger distal cingulum. t is , 1967; K oufos, 1980). The mammutid material from The mesial cingulum continues to the buccal side of the Pikermi (MN 12) housed at MNHN includes a maxilla first lophid. The distal cingulum is badly preserved and it with DP2 - DP4 dex and sin, associated with the mandible is difficult to trace its morphology. with dp3 sin and both dp4, and is referred to as M ammut Discussion: The family Mammutidae is characterized cf. borsoni (G audry, 1862-67; T a ssy , 1985). T a ss y (1977) by the zygodont pattern (yoke-like transverse crests) of described a left dp4 of Zygolophodon turicensis from the the intermediate molars and M3. The first representative Middle Miocene of France (Simorre). Morphologically, of the family is the primitive genus Eozygodon, which is the dp4 PMMS-53 from Samos and those from Simorre the smallest zygodon, known from the Early Miocene of and Pikermi present the same zygodont characteristics Kenya (Ta ssy & P ickfo rd , 1983). In Eurasia the family (Fig. 5). The dp4 of Z. turicensis from Samos differs from is represented by two genera only, Zygolophodon and its that of Simorre in having a semicircular mesial cingulum descendant Mammut. The “Zygolophodon turicensis group” whereas in the Simorre specimen, the cusplets of the mesial includes four Eurasian species: the type species Z. turicensis cingulum are arranged in a straight line. Moreover, the (Schinz, 1824) from Europe, Z. atavus from Kazakhstan, cingulum appears all around the tooth from Simorre, being Z. metachinjiensis from India and Z. gobiensis from Mon­ particularly strong on the buccal side. A lingual cingulum golia and China (Ta ssy , 1996). Recently,Zygolophodon has is not observed in the teeth from Samos. The distal cingu­ been discovered in Wadi Moghara, Egypt, with the new lum of the dp4 from Samos seems to be slightly stronger. species Z. aegyptensis, dated to Early Miocene (Sanders The dp4 from the three mentioned localities are different &. M iller, 2002). The zygodons generally seem to be a in size. The teeth from Pikermi show larger dimensions, conservative group with limited evolutionary changes, while the teeth from Samos and Simorre have similar especially in comparison to the bunodons, who appear dimensions (Tab. 2). more diversified. Summarizing the above mentioned data and since the tran­ In Greece, Zygolophodon turicensis is less common than sition from Zygolophodon to Mammut is not clear (especially Choerolophodon pentelici. The species is known by a skull in the milk teeth features), the studied dp4 is assigned to with DP2 - DP4 dex and sin from Ravin des Zouaves 5 Z. turicensis. The discovery of new material with perma­ (Axios Valley, MN 11) and a maxilla with DP2 - DP4 nent dentition will allow more detailed comparisons and dex and sin from Halmyropotamos (MN 12) (M elen- w ill probably lead to more certain taxonomic conclusions. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

148 Beitr. Paläont., 31, Wien, 2009

Figure 8: Chronology and biostratigraphy of the sub-family Choerolophodontinae and the family Mammutidae. The data was taken from NOW (2007), Sanders (2003), Sanders et al. (2002), Pickford (2001). ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. & K oufos, G.D., Proboscidea. 149

3. Geographic and Stratigraphic Distribution choerolophodons are divided into three different species. C. anatolicus is typical for Vallesian, while C. pentelici is its The oldest known Eurasian choerolophodons were found descendant and occurs in Turolian. The genus Choerolo­ in Bugti (Pakistan) and they are assigned to Choerolophodon phodon disappeared at the end of Miocene. Based on the palaeindicus, a species that has been recently transferred similarities oftheM TLB Choerolophodon with C. pentelici to Afrochoerodon (Tassy, 1990; Pickford, 2001). The from Pikermi, Axios Valley and other localities, a Turolian Bugti Beds are dated to Early Miocene (-20.0 Ma) by age can be proposed for the MTLB fauna. Z. turicensis Tassy (1990). Earlier Barry et al. (1985) considered the has a great stratigraphic range (Fig. 8) and cannot be Bugti fauna as belonging to the end ofM N 3 (-18.3 Ma). significant for biochronology. However, the other faunal Recent data from Bugti date the presence of choerolopho­ data from M T L A and MTLB suggests a Middle Turolian dons to the late Early Miocene at about 17.0 - 16.0 Ma age, and the palaeomagnetic record indicates an age of (W elcomme et al., 2001). On the other hand, the oldest 7.1 - 7.0 Ma (Koufos et al., this volume). known choerolophodon of Africa is A. kisumuensis dated to 18.0 - 17.0 Ma (Sanders & M iller, 2002). Thus, the original hypothesis of Tassy (1983) for a south Asian 4. Acknowledgements origin of choerolophodons changed and an African origin was proposed for them. It is suggested that they arrived in The excavations on Samos have been supported by the Pre­ Eurasia across the “Gomphotherium-C&ndfond.gz at -18 Ma fecture of Samos and the “Konstantinos and Maria Zimalis” together with the gomphotheres and deinotheres(T assy, Foundation.The Municipality of Mytilinii provided generous 1990; Rogl, 1999). The first arrival of proboscideans in help, too. The Natural History Museum of the Aegean of­ Europe is known as “Proboscidean-datum” and it is well- fered us the premises for the preparation and storage of the traced in Lesvos Island (Greece), whereProdeinotherium fossils. The first author G.E.K. thanks the SYNTHESYS was recognized in levels older than 18.5 Ma (Koufos et al., program for supporting his visit to NHMW (AT-TAF-3825). 2003). After their arrival the choerolophodons dispersed in G.E.K. thanks very much Dr. U. Göhlich for granting him Europe and Asia (Fig. 6). In the Eastern Mediterranean re­ access to the collection of NHMW, as well as for the help gion the oldest choerolophodon isA. chioticus known from and hospitality during his visit at the museum. The second the locality of Thymiana (Chios Island, Greece), dated to author, G.D.K., also thanks the SYNTHESYS program for MN 5 (-15.5 Ma). The genus Afrochoerodon covers a long supporting his visits to NHMW (AT-TAF-702) and MNHN time span, as it is known from the Early Miocene (MN (FR-TAF-3102). G.D.K. thanks Dr. M. Harzhauser and Dr. 4) to the Vallesian (MN 10) of Africa (Fig. 8). The genus G. Daxner-Höck very much for granting him access to the Choerolophodon appeared in Vallesian and persisted during collection at their disposal, as well as for their great hospitality the whole Late Miocene. A t the end of the Miocene the at NHMW. We thank Prof. P. Tassy and Dr. S. Sen very much choerolophodons disappeared, having their last occurrence for granting us access to the collection of MNHN and Dr. C. in the locality of Maramena (Greece), dated to the end of Argot (MNHN) for her help in searching the fossils, as well the Turolian, MN 13/14 (Schmidt-K ittler et al., 1995) as for their great hospitality during our visit to MNHN. We (Fig. 8). In spite of their great geographic expansion, the would also like to thank Dr. U. Göhlich for reviewing this choerolophodons stopped in the Balkans and Eastern article. We also thank all colleagues and students who helped Europe and did not enter Central and Western Europe, with the excavations and the preparation of the fossils. probably because of the different palaeoecological condi­ tions (Koufos, 2003). During the “Proboscidean-datum”, the first zygodons ar­ 5. References rived in Eurasia. Their ancestors were found in Kenya and are referred to as Eozygodon (Tassy &, Pickford, 1983). Barry, J.C., J ohnson, N.M., Raza, S.M. & Jacobs, The sole known European representative ofZygolophodon L.L., 1985. Neogene mammalian faunal change in is Z. turicensisy with numerous occurrences (Fig. 7) and a southern Asia: correlations with climate, tectonic, and great stratigraphic expansion, covering the Middle and the eustatic events. — Geology, 13:637-640. Late Miocene of Europe (Fig. 8). The first occurrences of Bernor, R.L., Solounias, N., Swisher, C.C. & van this species are traced in the locality La Romieu, France, C ouvering, J.A., 1996. The correlation of three and in the locality Akfahisar, Turkey, both dated to MN4 classical “Pikermian” mammal faunas - Maragheh, (NOW, 2007). Its last appearances are known from the Samos and Pikermi - with the European MN unit localities El Arquillo 1, Spain, and Gravitelli, Italy, both system. — [in:] Bernor R.L., Fahlbusch, V. &c dated to MN 13 (NOW, 2007). Its descendant, M ammut M ittmann, H.W. (eds). The evolution of Western borsoni, is represented in the Pliocene faunas of Europe Eurasian Neogene mammal faunas, p. 137-154, New and existed until Early Pleistocene. In the Late Miocene York (Columbia University Press). of Europe it is rare and is known from Moldova, Ukraine, Bonis, L. de., K oufos, G.D. ¿S en, S., 1997a. Agiraffid Hungary, Bulgaria and Pikermi, Greece (NOW, 2007). skull and mandible from the middle Miocene of the In spite of the rarity of the proboscideans in the new island of Chios (Aegean Sea, Greece). — Palaeon­ collection of Samos, they offer some biochronological tology, 40:121-133, London. data. According to Sanders (2003), the Late Miocene B onis, L. de, Koufos, G.D. & Sen, S., 1997b. The ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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sanitheres (Mammalia, Suoidea) from the middle Prodeinotherium bavaricum (Proboscidea, Mammalia) Miocene of Chios Island, Aegean Sea, Greece. — from Lesvos Island, Greece; the appearance of dei- Revue Paleobiologique, 16:259-270, Genève. notheres in the Eastern Mediterranean. — Geobios, Bonis, L. de, Koufos, G.D. ôcSen, S., 1998. Ruminants 36:305-315, Lyon. (Bovidae and Tragulidae) from the middle Miocene K oufos, G.D., Kostopoulos, D.S. &c Vlachou, T.D., (MN 5) of the island of Chios, Aegean Sea (Greece). this volume. The Late Miocene Mammal Faunas of — Neues Jahrbuch für Geologie und Paläontologie, Samos Island, Greece: New Collection. 16. Chro­ 210:399-420, Stuttgart. nology. — Beiträge zur Paläontologie, 31:397-408, Forsyth M ajor, C.J., 1888. Sur un gisement d’ossements Wien. fossiles dans l’île de Samos, contemporains l’âge de Lydekker, R., 1884. Additional Siwalik Perissodactyla Pikermi. — Comptes Rendus Académie Sciences and Proboscidea. — Memoirs of the Geological Paris, C X V II:608-610, Paris. Survey of India, Palaeontologia Indica, (X), III, Forsyth M ajor, C.J., 1894. Le gisement ossifère de Part 1:1-34. M ytilini et catalogue d’ossements fossiles. - Georges M acI nnes, D.G., 1942. Miocene and post-Miocene Bridel &Cie Editeurs, pp. 1-51, Lausanne. Proboscidea from East Africa. — Transactions of the G audry, A., 1862-67. Animaux fossiles et géologie de Zoological Society of London, 25:33-106, London. l’Attique. - Editions Savy, Paris. M aglio, V.J., 1974. A new proboscidean from the late G audry, A . & Lartet, E., 1856. Résultats des recher­ Miocene of Kenya. — Palaeontology, 17:699-705. ches paléontologiques entreprises dans l’Attique sous M elentis, J.K., 1967. Studien über fossile Vertebraten les auspices de l’Académie. — Comptes Rendus de Griechenlands. 19. Die Pikermifauna von Halmyro- l’Académie des Sciences, 43:271-274, Paris. potamos (Euböa, Griechenland). I Teil: Odontologie G aziry, A.W., 1976. Jungtertiäre Mastodonten aus Ana­ und Kraniologie. — Annales Géologiques des Pays tolien (Türkei). — Geologisches Jahrbuch, 22:3-143, Helléniques, 18:283-411, Athens. Hannover. NOW, 2007. Database with the Neogene localities and G aziry, A.W., 1987. New mammals from the Jabel Zaltan their faunal lists, www.helsinki.fi/science/now/ site, Libya. — Senckenbergiana Lethaea, 68:69-89, database.htm Frankfurt a. Main. O zansoy, F., 1965. Etude des gisements continentaux K ondopoulou, D., Bonis, L. de, K oufos, G.D. & et des mammifères du Cénozoïque de Turquie. Sen, S., 1993. Palaeomagnetic and biostratigraphic — Mémoires de la Société Géologique de France, data from the middle Miocene vertebrate locality of 102:1-92. Thymiana (Chios Island, Greece). — Proceedings 2nd Pickford, M., 2001. Afrochoerodon nov. gen. kisumu- Congress Geophysical Society of Greece, 2:626-635, ensis (M acInnes) (Proboscidea, Mammalia) from Athens. Cheparawa, Middle Miocene, Kenya. — Annales K ostopoulos, D.S., Koufos, G.D., Sylvestrou, I.A., de Paléontologie, 87:99-117. Syrides, G.E. ôcTsombachidou, E., this volume. Pilgrim, G.E., 1913. The correlation of the Siwaliks with The Late Miocene Mammal Faunas o f Samos Island, mammal horizons of Europe. — Records, Geological Greece: New Collection. 2. Lithostratigraphy and Survey of India, 43:264-326. Fossiliferous Sites. — Beiträge zur Paläontologie, Rögl, F., 1999. Circum-Mediterranean Miocene Paleo- 31:13-26, Wien geography. — [in] Rössner, G. ScH eissig, K. (eds). Koufos, G.D., 1980. Palaeontological and stratigraphical The Miocene Land Mammals of Europe. - 39-48, study of the Neogene continental deposits of the basin Munich (Pfeil Verlag). of Axios river. — Scientific Annals Faculty Physics Sanders, W. J., 2003. Proboscidea. — [in:] Fortelius, &c Mathematics, Univ. Thessaloniki, 19:1-322, M., K appelman, J., Sen, S., B ernor, R. (eds.). Thessaloniki. Geology and Paleontology of the Miocene Sinap Koufos, G.D., 2003. Late mammal events and biostra­ Formation, Turkey. - 202-219, New York (Columbia tigraphy in the Eastern Mediterranean. — Deinsea, University Press). 10:343-371, Rotterdam. Sanders, W.J. ôcM iller, E., 2002. New proboscideans K oufos, G.D., 2006. Contribution to the Turolian from the early Miocene of Wadi Moghara, Egypt. — chronology and palaeoecology of Greece. — Hellenic Journal of Vertebrate Paleontology, 22:388-404. Journal of Geosciences, 41:183-214, Athens. Schinz, H.R., 1824. Naturgeschichte und Abbildungen K oufos, G.D., this volume. The Late Miocene Mammal der Säugethiere. — Brodtmanns Lithographische Faunas of Samos Island, Greece: New Collection. 1. Kunstanstalt, Zürich, vi + 417, Atlas, 177 pits. History of the Samos Fossil Mammals. — Beiträge Schlesinger, G., 1917. Die Mastodonten des K.K. Na­ zur Paläontologie, 31:1—12, Wien. turhistorischen Hofmuseums. — Denkschriften des K oufos, G.D., Bonis, L. de & Sen, S., 1995. Lophocyon K.K. Naturhistorischen Hofmuseums, Geologisch- paraskevaidisi a new viverrid (Carnivora, Mammalia) Paläontologische Reihe, 1:1-230, Wien. from the middle Miocene of Chios Island (Greece). S c h l e s i n g e r , G., 1 9 2 2 . Die Mastodonten der Budape- — Geobios, 28:511-523, Lyon. ster Sammlungen. — Geologica Hungarica, ed. sep. Koufos, G.D., Z ouros, N. &M ourouzidou, O., 2003. 2(1):1—2 8 4 , Budapest. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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Schmidt-K ittler, N., B ruijn, H. de & D oukas, C., European Neogene mammal chronology. - 237-252, 1995. The Vertebrate locality Maramena (Macedonia, New York (Plenum Press). Greece) at the Turolian/Ruscinian boundary (Neo­ Tassy, R, 1996. W ho is among the Proboscidea? — [in:] gene). 1. General Introduction. — Münchner Geowis- Shoshani, J. & Tassy, P. (eds.). The Proboscidea: senschaftliche Abhandlungen, 28:9-18, Munich. Evolution and Palaeoecology of Elephants and S o l o u n i a s, N., 1981. The Turolian fauna from the their Relatives. - 39-48, Oxford (Oxford University island of Samos, Greece. — Münchner Geowissen- Press). schaftliche Abhandlungen, Contributions Vertebrate Tassy, R, 2005. Proboscideans (Mammalia) from the late Evolution, 6:1-232, Basel. Miocene Akkajdagi, Turkey. — [in:] S en, S. (ed.). Steininger, F.F., 1999. Chronostratigraphy, Geochro­ Geology, mammals and environments at Akkajdagi, nology and Biochronology of the Miocene “European late Miocene of Central Anatolia. - 707-714, Paris Land Mammal Mega-Zones” (ELMMZ) and the (Geodiversitas). Miocene “Mammal-Zones (MN-Zones)” — [in:] Tassy, P. ôcP ickford, M., 1983. Un nouveau mastodonte Rössner, G. & H eissig, K. (eds). The Miocene zygolophodonte (Proboscidea, Mammalia) dans le Land Mammals of Europe. - 9-24, Munich (Pfeil Miocène inférieur dAfrique orientale: systématique Verlag). et paléoenvironment. — Geobios, 16:53-57, Lyon. Tassy, R, 1977. Découverte de Zygolophodon turicensis Tobien, H., 1975. The structure of the mastodont molar (Schinz) (Proboscidea, Mammalia) au Lieu-Dit (Proboscidea, Mammalia). Part 2: The zygodont and Malartic à Simorre, Gers (Vindobonien moyen); zygobunodont patterns. — Mainzer Geowissen- implications paléoécologiques et biostratigraphiques. schaftliche Mitteilungen, 4:195-233, Mainz. — Géobios, 10:655-669, Lyon. Tobien, H., 1980. A note on the skull and mandible of Tassy, P., 1983. Les ElephantoideaMiocènes du Plateau a new choerolophodont mastodont (Proboscidea, du Potwar, Groupe de Siwalik, Pakistan. Ile Partie: Mammalia) from the middle Miocene of Chios (Ae­ Choerolophodontes et Gompothères. — Annales de gean Sea, Greece). — [in:]J acobs, L.L. (ed.). Aspects Paléontologie (Vert.-Invert.), 69:235-297, Paris. of vertebrate history: Essays in honor of Edwin Harris Ta s s y, R, 1985. La place des mastodontes miocènes de Colbert. — 299-307, Flagstaff, Arizona. l’ancien monde dans la phylogénie des Proboscideans W elcomme, J.L., Benammi, M., C rochet, J.Y., M ari­ (Mammalia). — Unpublished Thèse Doctorat des vaux, L., M etais, G., A ntoine, P.O. & Baloch, Sciences, Université Pierre et Marie Curie, Paris. L, 2001. Himalayan forelands: palaeontological Tassy, R, 1990. The “Proboscidean Datum Event:” how- evidence for Oligocène detrital deposits in the Bugti many proboscideans and how many events? — [in:] Hills (Baluchistan, Pakistan). — Geological Maga­ Lindsay, E.H., Fahlbusch, V , M ein, P. (eds.). zine, 138:397-405, London. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

152 Beitr. Palaont., 31, Wien, 2009

^ E U CN, O eu >> _ 12 CO Bi < 0 - PQ PQ Sd| < s 2 O Oi w S2 2 zQ ® < G CO

feo

< o H C-

2 2 2 V, 2 a 5 2 > 2 2 1 V, 2 a 5 2

R

~R

8

2

PQ

JQ ^ Ho 2 g- J22

pq PQ 21 PQ PQ PQ PQ

Table 1: Dental measurements of the sub-family Choerolophodontinae from various localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. &K oufos, G.D., Proboscidea. 153

Zygolophodon turicensis M a m m u t cf. b orson i Table 2: Measurements of the Mytilinii-1 Simorre Pikermi dp4 of Zygolophodon turicen- (? MTLA) sis from Samos and Simorre PMMS-53 MNHN-Si 11 MNHN-PIK-3613 and M a m m u t cf. b orson i from G oehlich original measurements Pikermi. The measurements of (pers. comm.) the Simorre specimen were given dex sin dex sin to us by Dr. U. Göhlich. [dp4 ______L 67.9 70 75.6 [68.1] B 1st lophid 36.9 35 42.7 42.5 B 2nd lophid 38.8 41 51.6 48.7 B 3rd lophid 37.9 41 47.5 — ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

154 Beitr. Palaont., 31, Wien, 2009

PLATE 1

Choerolophodonpentelici, M ytilinii- IB (MTLB), Samos, Greece, Middle Turolian (MN 12)

Fig. 1. DP2 sin, MTLB - 11; a. occlusal, b. buccal, and c. lingual view.

Fig. 2. a. Maxilla with M l - M 2 dex and M 2 sin, MTLB - 126; occlusal view

b. M l dex of the maxilla MTLB — 126; occlusal view

c. M 2 dex of the maxilla MTLB - 126; occlusal view.

Zygolophodon turicensis, possibly Mytilinii - 1 (? MTLA), Samos, Greece, Middle Turolian (MN 12)

Fig. 3. dp4 dex, PM M S - 53; a. occlusal, b. lingual, and c. buccal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Konidaris, G.E. &.K oufos, G.D., Proboscidea. 155

PLATE 1