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Bu tez çalışmasının materyalini, Bursa Paşalar fosil lokalitesinden 2003 – 2013 yılları arasında ele geçmiş olan hominoid dişleri oluşturmaktadır. Bu çalışmada öncelikle, bu dişlerin anatomik/morfolojik dağılımını belirlemek ve sonrasında da materyalin üst/alt çenedeki konumunun belirlenmesi amaçlanmıştır. Bu belirlemelerden sonra, morfometrik değerlendirmelere göre tür dağılımına bakmak hedeflenmiştir.Yukarıdaki paragrafta belirtilmiş olan değerlendirmeler sonrasında, tüm materyal içerisinde ölçü alınabilenlerden MD – BL ölçüleri alınmış, uygun endisleri hesaplanmıştır. Ölçüleri alınabilmiş olan dişler için SPSS 20 programında `Tanımlayıcı İstatistikler` yapılmıştır. Bu istatistiksel analiz ayrıca Kenyapithecus kizili türüne ait olduğu belirtilmiş dişler, Griphopithecus alpani türüne ait olduğu belirtilmiş dişler ve hangi türe ait olduğu belirtilememiş dişler için de ayrı ayrı yapılmıştır.Bu çalışmanın ana materyalini Bursa Paşalar fosil lokalitesinde 2003 – 2013 yılları arasında bulunmuş hominoid dişleri oluşturmaktadır. Ayrıca literatürde bulunan diğer Miyosen Dönem hominoid türlerinin dişleri ile karşılaştırılmaya çalışılmıştır.Sonuç olarak, yapılan incelemelerden, iki tür arasındaki sayısal farklılığın çok büyük olduğunu ve bu farklılığın önceki çalışmalarla sayısal benzerlik gösterdiğini anlamaktayız. Ayrıca diğer Miyosen Dönem hominoidleri ile Paşalar materyalinin arasındaki kimi farklılık ve benzerlikler görülmüştür. The material of this thesis is the teeth found in Bursa Paşalar fossil locality between 2003 – 2013. Determining the anatomic/morphological distribution of the teeth is the first aim. And then determining the place of the material on upper/lower jaw is the purpose. After these, to the morphometric evaluation, distribution of the species is the aim.After these evaluation, MD – BL dimensions measure and calculate the avaliable indices. For these teeth, `descriptive Statistics` execute on SPSS 20. This statistical analysis execute for the teeth belionging Kenyapithecus kizili, the teeth belonging Griphopithecus alpani and the undetermined teeth.The main material of this thesis is the teeth found in Bursa Paşalar fossil locality between 2003 – 2013. However, this material compares with the other hominoid teeth belonging the other Miocene hominoid species in the literature.As a result of the analyzing, numerical disparity between two species is very large and this disparity shows numerical similarities with former studies. However, there are some similarities and dissimilarities between the thesis material and the other Miocene hominoids. 333

Büyük kuyruksuz maymunlar, insanın yaşayan en yakın akrabalarıdır. Çalışmamızın temel amacı yaşayan büyük kuyruksuz maymun türlerinin evrimsel süreçlerinde rol oynayan dinamikleri araştırmak, türleşme süreçlerine karşılaştırmalı bir bakış açısı getirmektir. Bu doğrultuda şu ana kadar konu üzerine yapılan çalışmalar incelenmiş ve karşılaştırılmıştır.Çalışmamızda şempanze (Pan), goril (Gorilla) ve orangutan (Pongo) popülasyonlarının birbirinden farklılaşmasında etkili olduğu düşünülen paleobiyocoğrafi ve paleoiklimsel süreçler ve bu süreçlerin günümüzdeki popülasyon çeşitliliğine etkisi araştırılmıştır. Bu süreci aydınlatmaya çalışan evrimsel senaryolar incelenmiş, yorumlanmıştır.İki farklı kıtada yayılım gösteren büyük kuyruksuz maymunlar coğrafi bakımdan farklı evrimsel baskılara maruz kalmış olsa da, türleşme süreçlerini etkileyen dinamiklerin benzerlik gösterdiği anlaşılmıştır. Ancak türleşme örüntülerinde davranış farklılığından, fizyolojik farklardan ve rastlantısallıktan kaynaklanan ayrımların olduğu sonucuna varılmıştır. Şempanze ve gorillerde vikaryant türleşme modeli görülürken, iki orangutan türünü ayıran izolasyonun kökeni tartışmalıdır.Üç grubun türleri ve alt türleri arasındaki çeşitlilik düzeyi farkı, onların evrimsel geçmişine ilişkin ipuçları vermektedir. Büyük kuyruksuz maymunların genetik çeşitliliğinin neredeyse bütün türlerde insandan yüksek olduğu görülmüştür. Yine çarpıcı biçimde, bu üç grubun genetik çeşitliliğinin de insan kaynaklı çeşitli baskılar nedeniyle tehlike altında olduğu göze çarpmıştır. Great apes are the closest relatives of humans. The aim of our study is to look at the dynamics which acted in their evolutionary process from a comparative perspective. In this context, we analysed and compared the current studies.We studied the paleobiogeographic and paleoclimatic factors which leaded the population differentiation of chimpanzee (Pan), gorilla (Gorilla) and orangutan (Pongo) species, and the effect of these factors to the current population diversity within these genera. We reviewed and reappraised the evolutionary scenarios targeting to reveal these processes.Although great apes have been exposed to geographically different evolutionary pressures, we realised the dynamics which influenced the speciation process are similar. However, they have differences in their speciation patterns derived from behavioral and physiological differences and randomness. Whereas vicariant speciation is observed in chimpanzee and gorilla, the origin of the isolation resulting in separation of two orangutan species is still controversial. The variation differences between and within the species and subspecies of these three genera give us some clues about their evolutionary history. We saw that the genetic variation of almost all the great ape species is higher than humans. Most strikingly, we realised that the variation within these three genera is threatened by their closest relatives- humans. 163

Bu tez çalışmasında, Sinap, Akkaşdağı ve Kemiklitepe lokalitelerinden ele geçen Hipparion türlerinin diş morfolojileri karşılaştırılmıştır. Belirlenen 25 diş özelliğinin türler ve lokaliteler arasında karşılaştırılması sonucu, Sinap, Akkaşdağı ve Kemiklitepe lokalitelerinden ele geçen Hipparion türlerinin diş morfolojileri arasında belirgin bir fark olmadığı görülmüştür. Bu durum sonucun Hominoid bulunuşluğu açısından anlamlı çıkmadığını göstermektedir. Bu sonucun çıkmasındaki temel etmen eldeki veri eksikliği olabilir. The thesis contains compare and contrast methods which have been practised between types of Hipparion tooth paleoecology found in Sinap, Akkaşdağı and Kemiklitepe. Different types and localities have been analyzed with regard to 25 characteristics of tooth. The comparison between the different types which have been analyzed according to this method proved that there is no great difference between them. This shows that the result is not that meaningful. The main reason for this result might be the limited amount of facts. 97

Hominoidler ilk defa Erken Miyosen Dönem'de görülmeye başlanmıştır. 16 myö ise Afrika'dan Avrasya'ya doğru göç etmişlerdir ve Anadolu bu geçiş noktası olarak kullanılmıştır. Şu ana kadar yapılan çalışmalar sonucu Anadolu'da dört adet fosil hominoid lokalitesi belirlenmiştir ve bu sayı yapılacak olan çalışmalarla artacak ve Anadolu'nun hominoid evrimindeki rolü daha iyi ortaya konulacaktır.Bu tezde öncelikle hominoidlerin ortaya çıkış sürecine kadarki dünyanın ve canlılığın nasıl bir değişim geçirdiği anlatıldı. Daha sonra Hominoid evrimini daha iyi anlamak açısından bilim insanlarının ortaya koymuş oldukları hominoidler arasındaki filetik ilişkiler geçmişten günümüze incelendi. Hominoidleri daha rahat inceleyebilmek açısından taksonomileri verildikten sonra İlk hominoid olan Proconsul'den başlanarak Miyosen Hominoidlerinin genel özellikleri kısaca anlatıldı. Anadolu'nun Hominoid evrimindeki yeri ve önemini ortaya koyabilmek için Anadolu'da yer alan dört adet fosil hominoid lokalitesi ve bu lokalitelerden ele geçirilen fosil hominoidler kronolojik sıraya göre anlatıldı.Sonuç kısmında ise Miyosen Hominoidlerinin dağılımında Anadolu'nun yeri ve önemi ortaya konmaya çalışıldı. Yapılacak olan yüzey ve kazı araştırmaları sonucunda yeni lokaliteler bulundukça Anadolu'nun yeri ve önemi daha da açık hale getirilecektir. Hominoids has started to seen in the Early Miocene. They have passed from Africa to Eurasia through Anatolia 16 ma. There has been four Hominoid fossils localities found in Anatolia and with the further researches it is believed this number will increase. Further researches will prove more information on the Hominoid evolution in Anatolia.In this thesis first describes the evolution of vitality. To understand the Hominoid evulation the taxonomy has been given starting from the first hominoid proconcul and all of other miocene Hominoids. The for Anatolian Homioid have been given with localization, chronology to understand the Anatolian Hominoids.Final part is about Anatolia within Miocene Homioid localization on the earth. Further researches in Anatolia will provide more information on the Miocene Hominoids importance. 100

IV ÖZET Bu çalışmada modern insanın sahip olduğu sese dayalı iletişim yetisinin biyolojik ve sosyal / kültürel yönlerden incelenmesi ve bu bağlamda Homo erectus ve Neanderthal'in sese dayalı olduğu varsayılan iletişim sisteminin benzer ölçütler ışığında karşılaştırılması amaçlanmıştır. Ayrıca, bu çalışmada modern insan, Homo erectus ve Neanderthal kafatasları üzerinde tekrar yapılandırılmış konuşma organları modellerine bilgisayar destekli animasyonlar uyarlanmış ve bu animasyonlar yardımıyla insana özgü sesler temel alındığında Homo erectus' un oldukça kısıtlı, Neanderthal'in ise daha fazla sayıda ses çıkarabildiği saptanmıştır. Sonuç olarak, yaşadıkları dönemde üretmiş oldukları kültürel değerlerle, animasyonlar yardımıyla bulunmuş ses birimlerinin ışığında Homo erectus'un daha ilkel, Neanderthal'in ise öncekine göre biraz daha gelişmiş ancak modern insanla karşılaştırıldığında her iki fosil insanın da proto bir dile sahip olduğu ortaya çıkarılmıştır. Çalışma, aynı zamanda, animasyon tekniğinin antropolojik dilbilim alanında kullanılması bakımından önem taşımaktadır. Bu çalışmanın yazılı metni dışında animasyonların ve animasyonların izlenebilmesini sağlayan Flash adlı programı taşıyan iki adet CD (compact disc) de ekte sunulmaktadır. Aynı zamanda ekler bölümünde animasyon programı ve animasyonların bir bilgisayar yardımıyla nasıl izlenmesi gerektiğine ait görsel nitelikli açıklayıcı bilgi sunulmaktadır. ABSTRACT In this study, it is aimed at examining the vocal based communication system of the modern human beings from the standpoints of biological, social, and cultural peculiarities, in the light of which the assumed vocal communication systems of Homo erectus and Neanderthal are compared and contrasted. Besides, vocal organ models which have been regenerated and designed upon the basicraniums of Homo erectus, Neanderthal, and modern human have been adapted to computer based voiced animations by the help of which we have demonstrated and found that Homo erectus was able to generate a limited number of, but Neanderthal was able to produce more speech sounds which have been based upon the vocal system of modern human beings. In conclusion, in the light of the social and cultural features as well as the visual data which has been gained from the animations, it has been found that when compared to Homo erectus, Neanderthal had a more developed language, but the two fossil humans had a proto language in contrast to modern human beings. This study is also important in terms of the application of a computer based animation technique to the field of anthropological linguistics. Apart from the written chapters for this dissertation, animation files and the animator software called Flash 5.0 are presented in two CDs (Compact Disc). Moreover, visual based explanatory texts oriented towards the installation of the CDs into a computer are submitted in the appendices part. 190

Measurements of the crown base area were made from occlusal photographs. The area of the crown base was measured using by means of a digital planimeter. Linear measurements of the crown were made using vernier callipers. The mean value of the recordings was used. Enlarged occlusal photographs of each tooth served to provide exact planimetric measurements of total occlusal surfaces and individual cusp areas. Overall size measurements and molar crown shapes were recorded. The incidence and expression of the morphological traits were examined as well. The distribution patterns of extra cusps show interesting differences between taxa. The detailed measurements were analysed by a computer program enabling the application of univariate and multivariate analyses. The statistical analyses have suggested the presence of two seperate groups in Paşalar. The distinction between these two groups can not be based on crown shape or morphological traits, but on size. This means, the Paşalar hominoid primates can be divided into two size classes. These groups, however, are morphologically very similar to each other. The Paşalar molar teeth sample is compared with data from the modern orang-utan, chimpanze and gorilla by means of univariate and multivariate analyses. Paşalar' s small specimens appear to be close to the values for the chimpanze (Pan) and the larger ones to the ones for the orang-utan (Pongo). The lower third molar of the large size group in Paşalar comes close to the female gorillas, whereas the same molar of the small size group does not resemble any of the modern hominoids. Secondary cusp areas were not included in the multivariate analyses, because of their low number. 142A general trait of the Factor analyses plots for the modern hominoid specimens is a partial overlap of the male and female size groups. As a result, the different size values constitute roughly a single cluster, with the biggest male specimens at one end and the smallest female specimens at the other. In contrast, the Factor analyses plots of the Paşalar specimens show a different picture: two distinctive clusters are observed. This result is confirmed by the cluster analyses as well. This leads to the following suggestion. If the characteristics of sexual dimorfizm of the modern hominoid molars are valuable for the Paşalar sample, the pairs of clusters shown in the plots of the Paşalar molar sizes would not represent a male and a female group, but two different species. These species are thought to be morphologically identical, however different in size. 143Measurements of the crown base area were made from occlusal photographs. The area of the crown base was measured using by means of a digital planimeter. Linear measurements of the crown were made using vernier callipers. The mean value of the recordings was used. Enlarged occlusal photographs of each tooth served to provide exact planimetric measurements of total occlusal surfaces and individual cusp areas. Overall size measurements and molar crown shapes were recorded. The incidence and expression of the morphological traits were examined as well. The distribution patterns of extra cusps show interesting differences between taxa. The detailed measurements were analysed by a computer program enabling the application of univariate and multivariate analyses. The statistical analyses have suggested the presence of two seperate groups in Paşalar. The distinction between these two groups can not be based on crown shape or morphological traits, but on size. This means, the Paşalar hominoid primates can be divided into two size classes. These groups, however, are morphologically very similar to each other. The Paşalar molar teeth sample is compared with data from the modern orang-utan, chimpanze and gorilla by means of univariate and multivariate analyses. Paşalar' s small specimens appear to be close to the values for the chimpanze (Pan) and the larger ones to the ones for the orang-utan (Pongo). The lower third molar of the large size group in Paşalar comes close to the female gorillas, whereas the same molar of the small size group does not resemble any of the modern hominoids. Secondary cusp areas were not included in the multivariate analyses, because of their low number. 142 A general trait of the Factor analyses plots for the modern hominoid specimens is a partial overlap of the male and female size groups. As a result, the different size values constitute roughly a single cluster, with the biggest male specimens at one end and the smallest female specimens at the other. In contrast, the Factor analyses plots of the Paşalar specimens show a different picture: two distinctive clusters are observed. This result is confirmed by the cluster analyses as well. This leads to the following suggestion. If the characteristics of sexual dimorfizm of the modern hominoid molars are valuable for the Paşalar sample, the pairs of clusters shown in the plots of the Paşalar molar sizes would not represent a male and a female group, but two different species. These species are thought to be morphologically identical, however different in size. 143Measurements of the crown base area were made from occlusal photographs. The area of the crown base was measured using by means of a digital planimeter. Linear measurements of the crown were made using vernier callipers. The mean value of the recordings was used. Enlarged occlusal photographs of each tooth served to provide exact planimetric measurements of total occlusal surfaces and individual cusp areas. Overall size measurements and molar crown shapes were recorded. The incidence and expression of the morphological traits were examined as well. The distribution patterns of extra cusps show interesting differences between taxa. The detailed measurements were analysed by a computer program enabling the application of univariate and multivariate analyses. The statistical analyses have suggested the presence of two seperate groups in Paşalar. The distinction between these two groups can not be based on crown shape or morphological traits, but on size. This means, the Paşalar hominoid primates can be divided into two size classes. These groups, however, are morphologically very similar to each other. The Paşalar molar teeth sample is compared with data from the modern orang-utan, chimpanze and gorilla by means of univariate and multivariate analyses. Paşalar' s small specimens appear to be close to the values for the chimpanze (Pan) and the larger ones to the ones for the orang-utan (Pongo). The lower third molar of the large size group in Paşalar comes close to the female gorillas, whereas the same molar of the small size group does not resemble any of the modern hominoids. Secondary cusp areas were not included in the multivariate analyses, because of their low number. 142A general trait of the Factor analyses plots for the modern hominoid specimens is a partial overlap of the male and female size groups. As a result, the different size values constitute roughly a single cluster, with the biggest male specimens at one end and the smallest female specimens at the other. In contrast, the Factor analyses plots of the Paşalar specimens show a different picture: two distinctive clusters are observed. This result is confirmed by the cluster analyses as well. This leads to the following suggestion. If the characteristics of sexual dimorfizm of the modern hominoid molars are valuable for the Paşalar sample, the pairs of clusters shown in the plots of the Paşalar molar sizes would not represent a male and a female group, but two different species. These species are thought to be morphologically identical, however different in size. 143 738

Paşalar fossil primate specimens were first described by- Andrews and Tobien (1977) and they assigned the small specimens to Ramap_i theçuş wickeri and larger specimens to §üvap_i.thecuş darwi.ni.. Then, Paşalar hominoids were grouped under a single highly dimorphic species as Şi.yapi.theçuş âlpam based on type specimen from Çandır, Turkey (Alpagut, Andrews and Martin, 1990). Recently, Paşalar fossil primates were distinguished from Şi.yap_i. theçuş genus and included in a new genus as Glrafopi theçuş based on their incisive foromen morphology (Martin and Andrews, 1993). The purpose of this study was to reconstruct the locomotor behaviors of Paşalar fossil primates based on the analysis of proximal and middle phalangeal morphology in relation to living and fossil primates. Also, some information were expected for the species problem in Paşalar. Proximal and middle phalangeal morphology of the Paşalar hominoids shows more similarities to general arboreal quadrupedal primates, than to Pongo, Pan and Hylobates. The results of this study suggest that Paşalar fossil hominoids included both arboreal quadrupedal ism and semi suspensory behavior in their locomotor repertoire. In addition, there were at least two primate species in Paşalar based on the differences in their proximal and middle phalangeal morpho 1 ogy. 63 159

SUMMARY Paşalar fossil Primate specimens were first described by Andrews and Tobien (1977) and they assigned the small specimens to Ramapithecus wickeri and larger specimens to Sivapithecus darwini. Then, Paşalar hominoids were grouped under a single highly dimorphic species as Sivapithecus alpani based on type specimen from Çandır, Turkey (Alpagut, Andrews and Martin, 1990). Recently, Paşalar fossil primates were distinguished from Sivapithecus genus and included in a new genus as Girafopithecus based on their incisive foromen morphology (Martin and Andrews, 1993). The aim of this study was to determine wear stages based on the diet from molar teeth of Paşalar fossil primates. In addition, to find out the number of individual, premolar and molar teeth mesial-distal interproximal contact facets were tested. Also, some information were expected for the species problem in Paşalar. 1) On the occlusial surface of the 563 molar teeth of Paşalar fossil hominoids have determined under 16 wear stages. 2) Based on mesial-distal interproximal contact facets 400 premolar and molar teeth contacted from 800 teeth. In this study from premolar and molar teeth, 162 tooth rows, namely 162 individual have obtained 3) The studies on the Paşalar hominoids teeth have indicated that, there are three different forms of teeth size in Paşalar. 4) In addition, there have been at least two primate species in Paşalar based on the studies on wear stages of the molar teeth and tooth rows from mesial-distal interproximal contact facets of premolar and molar teeth. 124 371