Palaeobio Palaeoenv (2016) 96:373–381

DOI 10.1007/s12549-016-0234-3

ORIGINAL PAPER

The first hominoid from the Maragheh Formation, Iran
Gen Suwa 1 & Yutaka Kunimatsu 2 & Majid Mirzaie Ataabadi 3 & Zahra Orak 4 &
Tomohiko Sasaki 1 & Mikael Fortelius 5

Received: 29 December 2015 / Revised: 18 April 2016 / Accepted: 27 April 2016 / Published online: 8 June 2016
# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2016

Abstract Miocene hominoid fossils are known from Africa
and Eurasia, in the latter ranging widely from western Europe
to Anatolia and from South Asia to Southeast/East Asia. Iran
is located between the known western and eastern Eurasian
hominoid distributions and is potentially important in understanding Miocene hominoid dispersal patterns. Maragheh is a
late Miocene fossil locality in northwestern Iran, well known
since the nineteenth century for its abundant mammalian fossils. However, until now, the only primate fossils reported
from Maragheh or Iran were the Old World monkey
Mesopithecus pentelicus. Recent field research at Maragheh
has changed this situation by the discovery of the first hominoid fossil from Iran, a maxillary fragment with wellpreserved second and third molars. Here, we provide a detailed description of this new specimen, comparing it with
other similarly large-sized Eurasian late Miocene hominoids,
Ouranopithecus, Ankarapithecus, Sivapithecus, and
Indopithecus. Molar morphology of the Maragheh hominoid
is similar to that of these Eurasian Miocene genera, with only
This article is a contribution to the special issue BThe late Miocene
Maragheh mammal fauna; results of recent multidisciplinary research^
* Gen Suwa
gsuwa@um.u-tokyo.ac.jp

1

The University Museum, The University of Tokyo, Hongo,
Bunkyo-ku, Tokyo, 113-0033, Japan

2

Ryukoku University, Fushimi-ku, Kyoto, 612-8577, Japan

3

Department of Geology, Faculty of Science, University of Zanjan,
Zanjan, 45371/38791, Iran

4

National Museum of Natural History (MMTT), Department of
Environment, Tehran, Iran

5

Department of Earth Sciences and Geography, University of
Helsinki, Helsinki, FIN-00014, Finland

minor differences in morphology and wear pattern. Based on
the presently available materials, we tentatively prefer the interpretation that the Maragheh hominoid may be related more
closely to either Ankarapithecus or Sivapithecus rather than to
Ouranopithecus, but the fragmentary nature of the fossil
makes evaluations difficult. Future discoveries of this
Iranian hominoid are needed to determine its phylogenetic
position with more certainty.
Keywords Late Miocene . Large hominoid . Eurasia .
Maragheh

Introduction
The Maragheh Formation fauna is well known as representing
the western part of the classic late Miocene Pikermian
chronofauna (Eronen et al. 2009). The Pikermian fauna is
considered an open-habitat adapted palaeobiome, which
climaxed at about 7–8 Ma when it ranged from the Balkans
and Greece to Afghanistan and further into central Asia and
China. Until now, the only primate reported from Maragheh
was Mesopithecus pentelicus from the Bmiddle^ Maragheh
levels (Mirzaie Ataabadi et al. 2013). Here, we report the first
known occurrence of a large-sized species of hominoid from
the Maragheh Formation, northwestern Iran, a maxillary fragment with second and third molars, from the lowest Bmiddle^
Maragheh levels.
The hominoid fossil was discovered in 2004 at locality
MMTT II in Dareh Gorg. The location of this locality and
its stratigraphy and chronology are presented in detail in several papers of this issue (Mirzaie Ataabadi et al. in prep.; Sakai
et al. 2016, this issue; Salminen et al. 2016, this issue; Sawada
et al. 2016, this issue). The hominoid fossil comes from a
stratigraphic level ca. 8 m below the Middle Pumice (Sakai

Casanovas-Vilar et al.1 Ma (Kaya et al.’s (1977) BUpper Pumice^. this issue.5 mm. 2013). Materials and methods The new hominoid fossil from Maragheh catalogued MMTT 3453 is housed at the National Natural History Museum (or Muze Melli Tarikh Tabiei (MMTT) in Persian). Sawada et al. 2001. The M2 BL is 16. RPl-128. Both M2 and M3 root systems are of typical 3-rooted form with well-developed mesiobuccal and distobuccal roots. and replicated on research casts by GS and YK.5 mm as worn) and BL 17. XIR-1. we provide the basic morphological information of the well-preserved maxillary molars and make initial comparisons with other 7–10-Ma large-sized hominoid fossil teeth known from eastern Europe to southern Asia. RPl-775. its molars as large as the largest known Eurasian Miocene examples. The Middle Pumice and the hominoid level. The Middle Pumice is considered correlative of the marker bed known as the Gürt Dareseh Pumice (Bernor et al. respectively (Mirzaie Ataabadi et al. Metrics of AS95- . The latter value was used in the metric analysis. suggest a ∼7. 2000.g. Udabnopithecus.5 Ma. Mirzaie Ataabadi et al. in turn. 1980. 1) Ouranopithecus macedoniensis In our comparisons. most often attributed to Dryopithecinae (e. Gabunia et al.9 Ma. Cambell et al. 2016.16 and ∼7. which would lead to a ∼8. Koufos and de Bonis 2006). Because the Maragheh hominoid is large. is known from Georgia and considered ∼8. Using the Cande and Kent (1995) palaeomagnetic calibration scale. we used the MD and BL diameters taken by GS on the following original fossils housed at the Aristotle University of Thessaloniki: RPl-78.8 mm compensating for interproximal wear. According to the chronology based on a series of 40Ar/39Ar ages spanning the lower to upper Maragheh levels (Swisher 1996. The specimen is a left maxillary posterior alveolar fragment containing the upper second and third molars (M2 and M3) (Fig. However. Iran. 1980. Both crowns are well preserved. a much smaller-sized taxon. 2) Ouranopithecus turkae Casts of the CO-205 maxilla were observed. The age of these are considered to range from 9. Metrics are from Güleç et al. Inset is a CT scan section at around mid-height of the M3 roots. 3) Ankarapithecus meteai Casts of MTA 2125 were observed. this issue) or Pumice Bed 2 (Mirzaie Ataabadi et al. Begun 2002). The damaged M2 roots are partially exposed. The M2 MD is 14. RPl-90. the Maragheh hominoid would be ∼7. and the M3 roots are enclosed in the alveolar bone.6– 8. 2016. This taxon is known from a single fragmentary maxilla with P4 and M1.’s (1977) BLower Pumice^ (Swisher 1996). the latter the same unit as Kamei et al. Department of Environment. Ouranopithecus and Ankarapithecus from Turkey. (1977) BUpper Pumice^ levels were interpolated as ∼8. the Gürt Dareseh Pumice (=the Kamei et al.7 Ma (Sen et al. Here. Mirzaie Ataabadi et al. 2013). we restrict our comparisons to similar-sized taxa including Ouranopithecus from Greece. 2016. Palaeobio Palaeoenv (2016) 96:373–381 Fig. (2007).6 to 8. Scale bar is 10 mm Mesiodistal and buccolingual crown dimensions were measured following established methods (see Suwa et al. 1 Occlusal view of the Maragheh hominoid MMTT 3453.4 mm as worn and preserved and estimated as 14. The MMTT 3453 crown dimensions were measured with callipers on the original specimen by MMA and MF. Its age is considered 7. Mesiodistal length (MD) was taken sagittally along the midline of the tooth and buccolingual breadth (BL) perpendicular to this. The following comparative materials that preserve the M2 and M3 crowns were considered in the morphological assessments. 2016). 2011).5 mm.6-Ma age for the hominoid fossil calibrated to GTS 2012 (Salminen et al. Tehran. RPl-80/ 81. 2013). 2001. this issue).5–9 Ma (Gabunia et al. (1977) BLower Pumice^) and the Kamei et al.374 et al. 1). The chronological ages cited below based on magnetostratigraphy are calibrated to the Cande and Kent (1995) geomagnetic polarity time scale. 2013). Pardisan Park. this issue). combined with new palaeomagnetic data bracketing the Middle Pumice/hominoid levels with a normal–reverse– normal magnetic interval transition. and Sivapithecus and Indopithecus from the Indo-Pakistan area. Aside from these. 2009). which occurs ∼30 m higher in the section than the Middle Pumice. underlies Kamei et al.0-Ma age of the hominoid fossil.0 mm (14. The corresponding M3 dimensions are MD 15. recent K-Ar dates (Sawada et al. and NKT-89. 2016.

meteai (MTA 2155) M3s exhibit a larger metacone resulting in a squarer crown shape. A similar but stronger expression is present in S. there appears to be some differences in the details of feature expression. O. and are considered to be approximately 10. and Kelley (1988) for the following specimens: SM individual I. but its M3 is considerably larger than in MMTT 3453. Stereo pair renderings based on the micro-CT imagery are shown in Fig. These were used for qualitative assessments of occlusal morphology and wear pattern. meteai. and larger than in A. (1996) and Begun and Güleç (1998). the buccal cusps are less worn and more salient than the lingual cusps. O. . The MMTT 3453 M3 crown has a small metacone resulting in a rounded buccal to distal occlusal view crown contour. although an alternative possibility is 7. parvada. The age of this specimen is considered most likely 8. One characteristic of MMTT 3453 is the M3 > M2 size relationship. a tendency common to all of the taxa compared. respectively (Kappelman et al. described below. 2005). 3. while other O.5-Ma time interval (Kelley 2002. occlusal view crown shape is broader (buccolingually) than long in MMTT 3453. macedoniensis homologues. broadly comparable in size to those of O. Kelley 2005a). GSP 11708. Mesiolingual groove expression tends to be lacking in the upper molars of Sivapithecus species and. meteai individuals and more expressed in the single specimen of O. we used the metrics of Prasad (1969). especially in BL dimension. macedoniensis and S. considered approximately 9. they tend to differ from MMTT 3453 in buccal crown face morphology in their stronger buccal grooves associated with more mesiodistally rounded buccal cusp faces. von Koenigswald (1983). GSI-D1 (type of Paleopithecus sivalensis) is also considered to come from the Potwar Plateau U-level (Kelley 2005a). macedoniensis and A. GSP 9977. and GSP 20450. Occlusal view crown contour of the MMTT 3453 M2 is a rounded trapezoid. 6) cf. Pillans et al.3 Ma (Barry et al. and GSP 15000.Palaeobio Palaeoenv (2016) 96:373–381 50 and MTA 2125 were taken from Alpagut et al. or S. 2005). turkae. The single known M2 of O. at 50-μm voxel resolution. This condition is not matched in known O. The M2 shows a sizeable (∼1. turkae has a more squarish crown outline.0 Ma (Barry et al.9 Ma. 2002). These specimens come from the U-level of the Nagri Formation. (2005). Some O. macedoniensis M3s show a similar crown shape.5-mm transverse diameter) protocone dentine exposure that extends buccally along the mesial protocone crest. Buccal and lingual cingular (or cingulum-derived) features. We used the GSI-D196 and GSI-D299/300 metrics available in Gregory et al.6 and 9. VPL/HD I(1). macedoniensis. 2005a. respectively. The hypocone shows no dentine exposure but is worn relatively flat with weak mesial and distal wear slopes. 2003). However. Lingually.2–9. GSI-D196 and GSI-D299/300 (initially K29/ 466 + 617) come from the Haritalyangar area and can be considered to generally lie within the 8. The MMTT 3453 M2 and M3 both express weak but distinct parastylar grooves on the mesiobuccal crown margin. turkae M3 shows a complex cusp pattern with a partially subdivided large protocone. Crown dimension metrics of the comparative materials are summarised in Table 1 and compared with MMTT 3453 (Fig. Selected casts were scanned by micro-CT at The University Museum. Description and comparisons Both M2 and M3 crowns of MMTT 3453 are well preserved. macedoniensis and A. The M3 is less worn than the M2 with no dentine exposure in any of the cusps. macedoniensis M3s also tend to exhibit tuberculated distal crown portions. 5) Sivapithecus sivalensis The following representative maxillary dentitions with well-preserved M2 and M3 were included in the metric comparisons. b. with greater wear in the M2 than the M3. similar to that of A. Indopithecus giganteus A large M2. O. are weak or ill developed in MMTT 3453. (1980). turkae and most O. (1980) and Pilbeam (1982). The O. The two 375 main buccal cusps are occlusally worn and combine buccolingually concave wear surfaces with salient cusp tips and well-defined paracone and metacone occlusal margins. The ages of AS95-50 and MTA 2125 are considered 9. and O. macedoniensis M2s tend to exhibit a somewhat bilobed shape with rounded buccal cusp contours and lingually bulging hypocone. and a tuberculated distal margin. was attributed to Indopithecus by Pillans et al. parvada upper molars. when present. 2002. For GSI-D1. The Potwar Plateau specimens.7–9 Ma. meteai (MTA 2125) and Sivapithecus species. well-developed hypocone and metacone. 4) Sivapithecus parvada Descriptions and metrics are available in Pilbeam et al. Both M2 and M3 crowns are large sized. Sivapithecus species tend to lack such enhancements of the buccal crown face. The University of Tokyo. 2).5–9. Occlusal wear is more advanced in the mesial than the distal crown. are described (with metrics) in Pilbeam et al.7–8 Ma (Pillans et al. However. All specimens come from locality Y311 of the Nagri Formation. In both M2 and M3. a short but distinct mesiolingual groove occurs on the MMTT 3453 M3 and possibly on the M2 (albeit obscured by occlusal wear). turkae M2 is also comparably sized. GSP 10500. macedoniensis M2s and M3s tend to exhibit a weak parastyle expression similar to the MMTT 3453 condition. (1938). The enlarged O. Potwar Plateau. As with the M2. more so than in O. parvada. Casts of GSP 9986 (M1) and GSP 10500 (M2) were observed. takes the form of weak and short groove(s).

characterised by salient buccal cusps. suggesting taxonomic distinction.4 13.90 0.1 0.93 0.92 0. The occlusal wear pattern of MMTT 3453.031 CO-205 MTA 2125 15. (1980) Pilbeam et al.0 17.6–8.96 0. GSP 9977.93 0.2 14.96 Pilbeam (1982) Prasad (1969) Gregory et al. S. turkae A.95 0.96 0. sivalensis Specimen UM2 UM3 Crown shape Relative size UM3/UM2 Source MD length BL breadth MD length BL breadth UM2 MD/BL UM3 MD/BL MD BL 14. BL is buccolingual breadth of the crown in mm.6-Ma Maragheh hominoid.94 0.89 0. AS95-500. despite its large size. This wear pattern is shared with A.88 0. and estimations that correspond to our methods were adopted *MD and BL values are estimates corrected for wear.2 14.90 0. A.82 0.8 12.98 0.6 Ma) also considerably predates the Iranian hominoid. macedoniensis (8.0 0.031 0.4 13. turkae (Güleç et al.99 Pillans et al.9 12.93 Kelley (1988) 0.97 1. parvada predate the Maragheh hominoid by ≥2 million years and are geographically located east and west of Maragheh.039 0. which also shows a suite of other derived features probably associated with enhanced postcanine mastication (Güleç et al.06 This study* Mean (n = 7) 15. Right and left sides averaged in MTA 2125.84 0.2 13.4 14. Both M2 and M3 crowns are endowed with well-developed.87 0. (1980) GSP 15000 GSI D1 GSI D196 GSI D299/300 12. the M2 is comparatively square shaped.0 16.96 0.7– 9.5 0. The M3.8 14. meteai. (2005) Pilbeam et al.0 15.35 12. especially the M3.5 15. and other Sivapithecus species.01 Güleç et al. and GSP 15000. turkae is larger sized.15 17.068 0.99 Alpagut et al.900 0.96 0.94 0. Pilbeam et al.9 0. O. (2007) Begun and Güleç (1998) AS95-500 SM individual I GSP 10500 12. bulbous main cusps. meteai S. . The two share a M3 > M2 size relationship.96 1. The Greek O.92 0.95 18.6 11.97 0.5 0. and a buccolingually concave occlusal wear plane.2 13.5 13.973 0. turkae. The latter is common in the M3 of many species and represents a generalised condition.87 0.3 13. parvada cf. (n = 7) 1.85 0. Among the comparative taxa. 2007).5 13.88 0.0 13. but the younger end of its documented chronological range is closer to the Maragheh fossils in age.890 0. The better preserved side was measured Discussion and conclusions The morphology of the MMTT 3453 molars is broadly comparable to those of all the above compared taxa but differs from each in some details or in the combination of features.93 0. (1996) von Koenigswald (1983) Pilbeam et al.9 13. 2007).2 14.88 0.91 0. although large. parvada. probably a primitive retention. Indopithecus S. MMTT 3453 probably lacked the postcanine enhancements apparent in O.5 13.86 1.25 16.5 9. sharp buccal occlusal margins.4 13.05 14. turkae differs from MMTT 3453 considerably.9 14. These features are not seen in the Maragheh hominoid.0 GSP 20450 13.2 12. These were cross-checked. 2012).77 0.80 HD I-1 GSP 9977 GSP 11708 13.88 0.05 1.87 0. turkae (7.90 0. (1938) MD is mesiodistal length of the crown in mm. see the text for further details.6 12.35 17.6 16.10 1. but this condition is much more exaggerated in O.7 11.376 Table 1 Palaeobio Palaeoenv (2016) 96:373–381 Crown dimension metrics of the Maragheh hominoid and comparative specimens Taxon Maragheh hominoid O. To the contrary. Compared to MMTT 3453.8 16.0 12.3 12.4 11.4 14. suggests masticatory differences from O.00 Suwa unpublished** St. the MMTT 3453 molars are relatively broader buccolingually.1 13. The above range of observations is sufficient to infer that.55 0.95 0. Despite its temporal and geographical proximity to the Iranian hominoid.91 0.94 0. A younger (∼7 Ma) isolated maxillary P4 from Bulgaria has recently been suggested to represent a small individual or taxon related to Ouranopithecus (Spassov et al. **MD and BL values are estimates corrected for wear and/or damage. O.101 0. (1980) MMTT 3453 15. turkae.6 16. meteai and S.65 17. (1938) Gregory et al. macedoniensis O.90 0.96 0. and the M3 is distally hypertrophied.8 11.dev.11 1.01 1. O.9 0. has a reduced metacone.8 11.1 Ma) is probably temporally the closest to the ∼7. (1980) provided actual and estimated crown dimensions.

8 ad ea e m ur et ka A. macedoniensis (i. in each species. they tend to be mesiodistally expanded towards the O. In the first three plots (mesiodistal length. macedoniensis M2s as large as in O.9 species. the two might have diverged prior to the known time range of O. macedoniensis upper molars tend to wear with a weaker buccolingual wear gradient. macedoniensis examples share a buccolingually concave occlusal wear pattern.7 ns is O . turkae. m et ea S.1 0.t ur ka e A. Such a scenario may imply a parallel acquisition of large postcanine size at the eastern and western ends of the emerging Pikermian palaeobiome.9 1. m et ea S. O. Because of the apparent morphological specialisations of the O. 2 Box plot comparison of the Maragheh hominoid MMTT 3453 upper M2 and M3 with the other taxa.m . buccolingual breadth.0 1. some distinctions can be pointed out. macedoniensis posterior molars.2 1. Upper right: buccolingual crown breadth.Palaeobio Palaeoenv (2016) 96:373–381 Mesiodistal length UM2 UM3 Buccolingual breadth UM2 UM3 19 20 17 18 15 16 13 14 11 12 9 10 on ie ns is O . Lower left: crown shape. macedoniensis. 2003).6 Ma). si va le ns is 0. crown shape). To the contrary. if MMTT 3453 was phylogenetically related to O. Andrews and Alpagut 2001. pi do In le ec th rv pa S. si va le ns is h he ag ar M ac ed ns is .m ac S. 1996. i pa rv a da In do pi th ec us S. macedoniensis also shares with O. turkae condition. macedoniensis are greater than with O. with limited information available regarding the morphologies discussed above (Alpagut et al. turkae. the M2 and M3 plots are shown on the left and right sides. although MMTT 3453 and some of the O.0 0. The relatively horizontal occlusal wear in O. si va le ns is O O O . meteai is difficult because there are only two maxillary dentitions known. Upper left: mesiodistal crown length. Lower right: relative M3 size. m et ea S. the MD and BL box plots are shown on the left and right sides.m ac ed M ar on ie ag he is h 0. respectively. One is that O. macedoniensis. macedoniensis suggests a somewhat stronger emphasis of postcanine crushing function in that ns us a i 0. i pa rv ad In a do pi th ec us S. the M3 size dominance seen in MMTT 3453 is not matched in O.t ur ka e A. before ∼9.e. ns . turkae a tendency for distal M3 complexity. In the relative M3 size plot. macedoniensis also appears to express postcanine expansion more so than in MMTT 3453. As was the case . M3 mesiodistal or buccolingual diameters divided by the corresponding M2 diameters. Comparison with A. O. Another difference is that. Not only are O. Kappelman et al. respectively 377 Crown shape (MD/BL) UM2 UM3 Relative M3 size (M3/M2) BL MD 1.t ur ka e A. i pa rv a d In a do pi th ec us S. Although the general crown size and shape similarities of MMTT 3453 with O. Begun and Güleç 1998.m ac ed M ar on ie ag he h mm Fig. mesiodistal length divided by buccolingual breadth.t O O .8 h he is on ie ag ar ed M va si O .

macedoniensis XIR-1.378 Fig. 3 Stereo pair image comparisons of the Maragheh hominoid MMTT 3453 with selected comparative specimens. A. This was done by first standardising mesial crown breadths so that fitting becomes easier and then. O. right M2 (mirrored). after the fitting. right M2 (mirrored) and left M3. RPl-193. meteai MTA 2125. right M2 (mirrored) and left M3. right M2 and M3 (both mirrored). parvada GSP 10500. Scale bar is 15 mm. S. Surface polygon data were oriented relative to the Maragheh homologues by visual best fitting of the mesial crown halves. and GSP Palaeobio Palaeoenv (2016) 96:373–381 9986 right M1 (mirrored). RPl-128. scaling each specimen back to their original sizes . left M2.

turkae that seemingly enhanced the O. meteai acquisition of such slightly derived postcanine function (Fig. meteai lineage. In these slight differences. Our analysis suggests that it is possible that the Maragheh hominoid was related to (1) O. meteai AS95-50 upper molars appear to have minimal buccolingual wear gradient (fig. for example. meteai would necessitate less complication in terms of morphological transitions. parvada is also quite possible. If the Iranian hominoid was a phylogenetic descendent of the geographically proximate A. However. macedoniensis A. That the Iranian molars appear more derived than those of Sivapithecus spp. meteai. indicus S. parvada molars are similar in morphology and wear pattern. 9.6– 9. A. the MTA 2125 molars are smaller. such a transition is not impossible. Such a hypothesis would explain the general resemblances with Sivapithecus species. the minor morphological and wear pattern differences discussed above may actually be insignificant at the species or lineage level. parvada 15 Ma adapted fauna of Maragheh and the more closed-habitat fauna of the Siwaliks contradicts a Siwalik origin of the Maragheh hominoid. (2) A. S. wear differential between the buccal and lingual cusps is weaker. meteai molars are similar to the MMTT 3453 condition. The hypothesis that the Maragheh ape was related to A. However. meteai. macedoniensis. parvada that preserves both M2 and M3 exhibits a M2 > M3 size relationship. of the Siwaliks. macedoniensis or related taxon. Two phylogenetic hypotheses that do not require a reversal of masticatory adaptations are shown. the emerging (proto-) Pikermian fauna. Alternatively. meteai O. An alternative possibility would be a close cladistic relationship with the Sivapithecus clade but with divergence extending deep into the middle Miocene (Fig. although the single individual of S.1 of Andrews and Alpagut 2001). meteai (9. macedoniensis. (3) S. The above brief comparisons show that the morphological and phylogenetic affinities of MMTT 3453 are difficult to assess. sivalensis S.Palaeobio Palaeoenv (2016) 96:373–381 379 with O. 4). The MMTT 3453 and known S. 4 Temporal positions of the Maragheh hominoid and other fossils discussed in this study. parvada exhibits the primitive condition shared with other Sivapithecus species. there may have been a secondary shift towards a dietary adaptation with somewhat less emphasis in crushing. giganteus Turkey Iran India/Pakistan . is concordant with such a hypothesis. parvada. the largest species of Sivapithecus. a simple increase of fossils may result in a blurring of differences. then some postcanine features characteristic of O. macedoniensis-like masticatory adaptation makes such an opposite transition somewhat unlikely. However. Given the subtle morphological features under consideration. the strong contrast between the openFig. this would have occurred within.9 Ma). a phylogenetic relationship between MMTT 3453 and S. macedoniensis. This hypothesis would entail a postcanine size increase to have taken place within the Ankarapithecus clade or in the A. or S. without invoking immigration from the late Miocene Siwaliks. such as a direct derivation of the Maragheh hominoid from either O. See the text for further details and other possibilities. Sethi Nagri. parvada. macedoniensis would have been secondarily modified. the broadly contemporary presence of O. The female A. despite an increase in postcanine size. If the Maragheh hominoid was a late surviving descendent of O. Geographically. it appears that the chronologically older S. or (4) Sivapithecus in general. turkae Maragheh hominoid S. In such a case. but not sampled by. Based on the limited available fossils and morphologies. with a greater representation of tragulids. parvada molars also seem to show a consistently greater buccolingual wear gradient than seen in MMTT 3453. 10 Ma 5 Ma Miocene Middle Late Greece O. This species is so far recorded only from a single ∼10-Ma locality. the Maragheh hominoid could have split prior to the A. Kelley (1988) suggested that this locality may be sampling a fauna somewhat different from that of the other Sivapithecus localities and may represent a relatively closed-habitat fauna. or adjacent to. This hypothesis would in turn necessitate independent acquisition of large size and a long-standing ghost lineage in the western Asian palaeobiome that was juxtaposed to. parvada ? I. and the distal M3 crown is better developed resulting in a less trapezoidal outline. the general morphology and wear pattern of the MTA 2125 male A. The fourth comparison is with S. meteai and the Maragheh hominoid. Given the meagre fossil evidence currently available for both A. 4).

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