New Evidence from the Middle Miocene Locality Damiao, Inner Mongolia, China

Palaeoenvironments of the Pliopithecoid primates - New evidence from the middle Miocene locality Damiao, Inner Mongolia, China

LEENA SUKSELAINEN

ACADEMIC DISSERTATION To be presented, with the permission of the Faculty of Science of the University of Helsinki, for public examination in Auditorium XII, University main building, on December 10th 2016, at 12 noon.

DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 / HELSINKI 2016 © Leena Sukselainen (synopsis and Paper III) © Elsevier (Paper I) © Vertebrata PalAsiatica (Paper II) Cover photo: Leena Sukselainen

Author´s address: Leena Sukselainen Department of Geosciences and Geography P.O.Box 64, 00014 University of Helsinki, Finland OHHQDVXNVHODLQHQ#KHOVLQNL¿

Supervised by: Professor Mikael Fortelius Department of Geosciences and Geography University of Helsinki, Finland

Professor Terry Harrison Center for the Study of Human Origins New York University, USA

Dr. Anu Kaakinen Academy research fellow Department of Geosciences and Geography University of Helsinki, Finland

Reviewed by: Professor Yutaka Kunimatsu Faculty of Business Administration Ryukoku University, Japan

Docent Suvi Viranta-Kovanen Institute of Biomedicine University of Helsinki, Finland

Opponent: Dr. Lawrence J. Flynn Assistant director Peabody Museum, Harvard University, USA

ISSN 1798-7911 ISBN 978-951-51-1365-8 (paperback) ISBN 978-951-51-1366-5 (PDF) KWWSHWKHVLVKHOVLQNL¿

Helsinki University Print Helsinki 2016 Sukselainen L., 2016. Palaeoenvironments of the Pliopithecoid primates - New evidence from the middle Miocene locality Damiao, Inner Mongolia, China. Helsinki University Press. Helsinki. 33 SDJHVDQG¿JXUHV

Abstract

Pliopithecoidea is an extinct and diverse super- isotope analyses. The aim is to understand the family of primitive catarrhine primates with no presence of humid-favouring pliopithecoid pri- NQRZQ GHVFHQGDQWV 7KH\ ¿UVW DSSHDU LQ WKH mates in central Asia after the middle Miocene fossil record in the late early Miocene China, climatic optimum (MMCO; ca. 17–15 Ma). The ca. 18–17 million years ago (Ma). They were reason this is interesting is because it seems to widely distributed throughout Eurasia between contradict the general trend of strengthening cli- FDDQG0DDQGZHUHDPRQJWKH¿UVWSUL- matic zonality and increasing aridity from the mates to be discovered and described. Despite early Miocene onwards. their wide distribution, pliopithecoids are rarely The results show that pliopithecoids inhabit- found together with the contemporaneous and ed more humid environments than other contem- equally widely distributed hominoid primates. porary primate groups, suggesting an inability The latest known occurrence of pliopithecoids to adapt to changing environmental conditions. is also from China where they co-existed with The conservative nature of pliopithecoid adapta- hominoids in the late Miocene ca 6.9–6.2 Ma. tions seems to have restricted the co-occurrence Continuing climatic deterioration and dispersal of pliopithecoids and hominoids, and has been of cercopithecoid primates from Africa may have therefore rarely documented. The study also sug- contributed to their demise. gests that direct ecometric analysis gives a bet- The main objective of this study is to pro- ter separation of the ecological preferences of duce new information on pliopithecoids, their these primates than do analyses of taxonomi- environments, as well as on environmental con- cally based community structure. ditions in Eurasia during the Miocene (23–5.3 The results in Damiao support previous infer- Ma), with a special focus on the middle Miocene ences concerning the presence of locally humid Inner Mongolian locality of Damiao. These goals environments within the increasingly arid sur- DUHDSSURDFKHG¿UVWE\LQVSHFWLQJWKHGLIIHUHQFHV roundings that characterized Central Asia. En- between pliopithecoid localities and other con- vironments within the DM01 locality may have temporaneous localities with particular focus on been more humid and possibly more forested the rare localities of co-occurring pliopithecoid and wooded, as suggested by hypsodonty, esti- and hominoid primates. To do this we used both mated mean annual precipitation (MAP), local traditional ecological diversity analysis as well as sedimentology and large mammal fossils. We direct ecometric method based on hypsodonty in compared our results with the adjacent fossil-rich mammalian herbivores. A closer examination of middle Miocene Tunggur localities. However, the Inner Mongolian pliopithecoid locality, Da- the small mammal fauna and isotope data are miao, will follow, using small mammal tapho- consistent with a mosaic of forest and grassland nomy, faunal similarity, ecometrics, and stable environment for all Damiao localities. Based on DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 our results, Tunggur may have been too season- FXPXODWHGE\SUHGDWRUVDQGGHSRVLWHGLQDÀXYLDO DORUQRWVXI¿FLHQWO\KXPLGIRUSOLRSLWKHFRLGV VHWWLQJ6RPHUHZRUNLQJE\ÀXYLDOSURFHVVWRRN This idea is supported by the higher mean hyp- place in DM01 and DM02. DM16 represents sodonty and lower predicted MAP estimates, as WKHGLVWDOSDUWRIWKHÀRRGSODLQ'0SRUWUD\V ZHOODVVOLJKWO\KLJKHUį13C values. We suggest DFKDQQHO¿OODQG'0LVDUHVXOWRIDQHSL- that DM01, the driest known Asian pliopithecoid VRGLFÀRRGGLVFKDUJHWRWKHÀRRGSODLQ:HDOVR locality, may have been a more humid refugium show that systematic excavation for small mam- within a generally drier regional context. mals is possible, and allows for the reduction of The bone material in Damiao was mainly ac- some of the damage collecting always causes.

4 Tiivistelmä

Pliopithecoidea on sukuputtoon kuollut mon- korkeuteen. Sisä-Mongolian pliopithecoid- imuotoinen yläheimo primitiivisiä kapeanenäap- löytöpaikan, Damiaon, paleoympäristoöjä inoihin (Catarrhini) kuuluvia kädellisiä, joilla tutkitaan tarkemmin piennisäkkäiden tafo- ei ole tunnettuja jälkeläisiä. Ensimmäiset plio- nomian, nisäkäslajiston samankaltaisuuden, pithecoidit fossiiliaineistossa on löydetty Kiinas- ekometrian sekä hapen ja hiilen pysyviä ta ja ne ajoittuvat varhaisen mioseeniepookin isotooppien avulla. Työn tavoitteena on lopulle noin 18–17 miljoonan vuoden taakse ymmärtää syitä kädellisten esiintymiselle (Ma). Tämän kädellisryhmän edustajat olivat Damiaossa: kosteita alueita suosivien plio- laajasti levinneet läpi Euraasian noin 17–7 Ma pithecoid-kädellisten esiintyminen Keski- ja olivat ensimmäisten löydettyjen sekä kuvat- Aasiassa mioseeniepookin ilmasto-optimin tujen kädellisten joukossa. Huolimatta laajasta (noin 17–15 Ma) jälkeen näyttäisi olevan levinneisyydestään, pliopithecoideja on harvoin ristiriidassa mioseenin yleisen ilmaston kehi- löydetty yhdessä samanaikaisten ja levinnei- tyksen kanssa, jota leimasi vähittäinen keski- syydeltään vastaavanlaisten hominoidien kans- leveysasteiden kuivuminen ja ilmaston muut- sa. Myös viimeisimmät tunnetut pliopithecoidit tuminen vyöhykkeisemmäksi mioseenin tulevat Kiinasta, myöhäismioseenin (noin 6,9– alkupuolesta lähtien. 6,2 Ma) Lufengista, joka oli myös yksi har- Tulokset osoittavat, että pliopithecoidit voista pliopithecoidien ja hominoidien yhteise- asuttivat kosteampia ympäristöjä kuin muut siintymistä. Ilmasto-olojen jatkuva muutos sekä saman ajan kädellisryhmät, mikä viittaa ni- vanhan maailman häntäapinoiden (Cercopithe- iden kyvyttömyyteen sopeutua muuttuviin coidea) leviäminen Afrikasta saattoivat osaltaan ympäristöolosuhteisiin, mikä puolestaan vaikuttaa pliopithecoidien sukupuuttoon. näyttääkin olleen rajoittava tekijä niiden Tämän tutkimuksen päätarkoituksena ja hominoidien yhteisesiintymisissä,. Tut- on tuottaa uutta tietoa pliopithecoideista, kimuksessa selvisi myös, että suora ekomet- niiden elinympäristöistä sekä Euraasian rinen analyysi erottelee paremmin ekologiset ympäristöolosuhteista mioseeniepookissa mieltymykset kuin taksonomiaan perustuva (23–5,3 Ma). Tutkimuksen painopiste on yhdyskuntarakenneanalyysi. keskimioseenisessa Damiaon löytöpaikassa Damiaosta saadut tulokset tukevat aikais- Kiinan Sisä-Mongoliassa. Työssä tarkastel- empia päätelmiä, joiden mukaan Keski-Aa- laan ensin pliopithecoid-löytöpaikkojen ja sian jatkuvasti kuivuvassa ympäristössä oli muiden samanikäisten löytöpaikkojen eroja paikallisesti kosteita alueita. Sisä-Mongolian keskittyen erityisesti pliopithecoidien ja pliopithecoid-löytöpaikka, DM01, on saat- hominoidien harvinaisiin yhteisesiintymi- tanut olla kosteampi, rehevämpi ja metsäis- siin. Löytöpaikkojen eroavaisuuksia tutki- empi hypsodontia-analyysien ja niiden pe- taan käyttämällä perinteistä ekologista di- rustella arvioidun keskimääräisen vuotuisen versiteettianalyysiä sekä suoraa ekometristä sademäärän (MAP), paikallisen sedimentolo- menetelmää, hypsodontiaa, joka perustuu gian sekä suurnisäkäslajiston perusteella kuin kasvinsyöjänisäkkäiden poskihampaiden ympäröivät alueet. Saatuja tuloksa verrattiin

5 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 viereiseen fossiilirikkaaseen keskimioseeni- ovat pääasiassa petojen kasaamia ja kerros- seen Tunggurin löytöpaikkaan. Piennisäkäsl- WXQHHW ÀXYLDDOLVHVVD \PSlULVW|VVl 1XRULP- ajisto ja isotooppiaineisto viittaavat mosaiik- milla löytöpaikoilla (DM01 ja DM02) on kimaisen metsä-ruohomaa ympäristön olleen jälkiä virtaavan veden uudelleenmuokkauk- vallitseva kaikilla Damiaon löytöpaikoilla. sesta. Vanhin löytöpaikka DM16 edustaa Tutkimustulosten perusteella Tunggur saat- tulvatasangon distaalia osaa; keskimioseenin toi olla ilmastoltaan liian kausiluontoinen pliopihtecoid-löytöpaikka DM01 edustaa tai ei ollut riittävän kostea pliopithecoideille. uoman täytettä; ja DM02 on episodimaisesti Tätä ajatusta tukee Tunggurin korkeampi purkautuneen tulvan muodostama. Tafono- keskimääräinen hypsodontia ja vastaavasti miset tutkimukset Damiaossa osoittivat, että alhaisempi MAP sekä hieman korkeam- piennisäkkäiden systemaattiset kaivaukset PDWį13C arvot. Vaikka työ osoittaa DM01:n ovat mahdollisia. Kaivausten yhteydessä olleen Aasian kuivin pliopithecoid-löytöpai- fossiileille aiheutuu aina vahinkoa, mutta kka, se oli kuitenkin ilmeisimmin ympäröiviä systemaattinen kaivaus aihettaa sitä toden- alueita kosteampi refugio. näköisesti vähemmän kuin perinteinen seu- Damiaon piennisäkäsfossiiliakumulaatiot lontamenetelmä.

6 Acknowledgements

First and foremost I want to thank my supervi- fellow PhD student Radoslaw Michallik who do- sors; Mikael Fortelius, Terry Harrison, and Anu nated his precious time and expertise and pro- Kaakinen. The original idea for this study came duced a set of perfect SEM pictures of tapho- from Terry, and together with Mikael they further nomic features. cultivated the project and where kind enough to I am grateful to my pre-examiners, profes- think of me when they were looking for a suit- sor Yutaka Kunimatsu and docent Suvi Viranta- able person to take the task. Anu, who was also Kovanen for their useful comments and sugges- responsible for supervising my master’s thesis tions that helped to improve the thesis. has been extremely patient in guiding me through This thesis was funded by Emil Aaltonen WKHMRXUQH\LQWURGXFLQJWRQHZ¿HOGVRIVWXG\ Foundation, Nordenskiöld Samfundet i Finland. pushing me forward, and providing invaluable and Dissertation completion grant. Also sever- help with many practical matters regarding fund- al travel grants from Doctoral School in Geol- ing applications and other thesis related practi- ogy and Chansellor’s travel grant helped reach- calities. Also I am grateful to all my supervisors ing the goal. for providing me with the opportunity to visit 3DVWDQGSUHVHQWRI¿FHPDWHVDVZHOODVDOO unusual and exiting places, meet with interesting the people in the Department of Geosciences and people and learn new things on the way. Geography are thanked for providing friendly I want to thank all my co-authors; professor and pleasant working atmosphere. Especially I Zhang Zhaoqun, Hannele Peltonen, Dr. Benja- want to thank Dr. Seija Kultti and Dr. Mia Ko- min H. Passey, and Dr. Jussi T. Eronen, for let- tilainen; without you I would have been lost so ting me exploit their knowledge and expertise many times. and for giving me support and valuable advice The only other member of a very exclusive on this journey. peer-support group, Elina Hernesniemi, thank I wish also to acknowledge my fellow pa- you for all the whining and wining and some oc- leopeople and colleagues whose time and knowl- cational dining sessions. Those were lifesavers. edge was consumed during different stages of the My dear friends from the department; Paula process; especially Kari Lintulaakso providing Niinikoski, Niina Kuosmanen, Elina Sahlstedt, PH ZLWK YDOXDEOH GDWD VRXUFH IRU WKH ¿UVW DU- Elina Lehtonen, Marttiina Rantala, and all the ticle, Ferhat Kaya for taphonomic support, Dr. members of the GeoDoc community, thank you Majid Mirzaie Ataabadi for advice with similar- for many fun moments and peer-support. Spe- ity analyses, and Dr. Tang Hui for helping with cial thanks to Paula, and Niina for all the laughs, some recent environmental datasources. Not to support, gossip, and friendship. forget Aleksis Karme, Mikko Haaramo and Su- Last but not least, my dear family and friends, sanna Sova for their help with data collecting thank you for your love, support in all forms, and software related matters. I must express my and occational ass-kicking, all very much needed gratitude to Dr. Julien Louys who kindly shared and appreciated. KLV$IULFDQGDWDZLWKPHIRUWKH¿UVWSDSHUDQG

7 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

Contents

Abstract ...... 3 Tiivistelmä ...... 5 Acknowledgements ...... 7 List of original publications ...... 9 Author’s contribution to the publications ...... 9 /LVWRI¿JXUHV ...... 10 1 Introduction ...... 11 1.1 Palaeogeography of the Miocene ...... 11 1.2 Miocene climatic and vegetation history in Eurasia ...... 11 1.3 Pliopithecoids ...... 13 1.4 Damiao, Inner Mongolia ...... 15 2 Aims of the study ...... 16 3 Material ...... 17 4 Summary of the original publications ...... 18 4.1 Paper I ...... 18 4.2 Paper II ...... 19 4.3 Paper III ...... 19 5 Discussion ...... 20 6 Conclusions ...... 23 References ...... 24

Appendices: Publications I-III

8 List of original publications This thesis is based on the following publications:

I Sukselainen, L., Fortelius, M., Harrison, T., 2015. Co-occurrence of pliopithecoid and hominoid primates in the fossil record: an ecometric analysis. Journal of Human Evo- lution. 84 25-41. DOI:10.1016/j.jhevol.2015.04.009

II Sukselainen, L., Peltonen, H., Kaakinen, A., Zhang, Z. Small mammal taphonomy of three Miocene localities from Damiao, Inner Mongolia. Vertebrata PalAsiatica. (in press).

III Sukselainen, L., Kaakinen, A., Eronen, J.T., Passey, B.H., Harrison, T., Zhang, Z., For- telius, M., Palaeoenvironment of the middle Miocene locality Damiao, Inner Mongolia (submitted).

The publications are referred to in the text by their roman numerals.

Author’s contribution to the publications

The research project was planned by L. Sukselainen, M. Fortelius, T. Harrison, and A. Kaakinen. I The study was designed by T. Harrison, and M. Fortelius. Material was collected, and analysed by L.Sukselainen and interpreted by all authors. Writing the article was done jointly by all authors with the main responsibility on L. Sukselainen. II The study was designed by H. Peltonen, M. Fortelius, A. Kaakinen and Z.-Q. Zhang. $QDO\VHVZHUHGRQHE\+3HOWRQHQ$OODXWKRUVMRLQHGWKHUHODWHG¿HOGZRUN5HVXOWV were interpreted by L. Sukselainen. Writing the article was done jointly with A. Kaaki- nen and other authors commented the article. Main responsibility was on L. Sukse- lainen. III The study was planned by L. Sukselainen, M. Fortelius, and A. Kaakinen. Stable isotope sampling, analysing and interpreting was done by B.H. Passey. Other analyses were done jointly by L. Sukselainen and J.T. Eronen. L. Sukselainen prepared the manuscript with the contributions from the other authors.

9 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

/LVWRI¿JXUHV

Fig 1 Compilation of the important events during the Miocene, page 12 Fig 2 Map of Chinense pliopithecoid localities, page 15 Fig 3 General views of the Damiao area, page 16 Fig 4 Three studied Damiao localities, page 17 Fig 5 Mean annual precipitation (MAP) in the Miocene Eurasia, page 21

10 1 Introduction clude the opening of the Greenland-Scotland ridge, that allowed cold, deep Arctic waters in- WRWKH$WODQWLF :ULJKW WKH¿QDOFORVXUH 1.1 Palaeogeography of the Miocene RIWKH0HGLWHUUDQHDQ,QGRSDFL¿FFRQQHFWLRQLQ Recognition of the modern continents would eas- the early late Miocene (Rögl, 1999); the closing ily have been possible already in the early Mio- of the Panama Isthmus connecting the Americas FHQHDOWKRXJKWKHUHZHUHVRPHVLJQL¿FDQWGLI- (Duque-Caro, 1990; Coates and Obando, 1996); ferences: the Americas were separated from each DQGWKHHQGLQJRIWKHJOREDOHTXDWRULDOÀRZDV other; Greenland was connected to Canada; the well as initiating the birth of the modern Gulf Tethyan Ocean from India and Pakistan stretched Stream (Potter and Szatmari, 2009). At the latest to the Atlantic with what is southern continental Miocene, the Mediterranean connection to the Europe today as an archipelago. In addition, a $WODQWLFDW*LEUDOWDUZDVEULHÀ\GLVFRQQHFWHG shallow sea covered much of southern Russia; (Hsü, 1983; Krijsman et al., 1999; Potter and very different river systems existed, as well as Szatmari, 2009). other features such as islands, lower mountain belts with wider valleys, and lesser mountain gla- 1.2 Miocene climatic and ciations (Potter and Szatmari, 2009). A warmer vegetation history in Eurasia and globally uniform climate prevailed, with pos- The Neogene (23–2 Ma) of Eurasia can be char- sibly only a small ice sheet covering Antarctica acterized by three climatically distinct periods. (Potter and Szatmari, 2009). 7KH¿UVWZDVWKHZDUPDQGKXPLGHDUO\0LRFHQH One of the most noticeable tectonic events (23–17 Ma), with weak seasonality and low tem- that has affected the Eurasian topography was perature gradients (Utescher et al., 2000; Mos- the collision of the Indian Plate with Eurasia. The brugger et al., 2005; Bruch et al., 2007, 2011). timing of the collision is debated; however, most The second was the middle Miocene climatic palaeogeographic reconstructions suggest it oc- optimum that occurred ca. 17–15 Ma, and was a curred at 55 ±10 Ma (Wang et al., 2014 and ref- globally warm and humid period with weakened erences therein). It is generally recognised that a seasonality and regional differentiation (Flower spatially and chronologically varying (Pan, 1999; and Kennett, 1994; Fortelius et al., 2002; Böhme, Zhang et al., 2008; Wang et al., 2014 and refer- 2003; Bruch et al., 2004; Kaandorp et al, 2005; ences therein) multi-stage process (e.g. Pan et Liu et al., 2009). The third period involved the al., 1990; Li, 1995; Zhong and Ding, 1996; Yin late Miocene cooling that lead to a trend towards et al., 1999) characterized the uplift of the Ti- more arid and open habitats (Zachos et al., 2001, betan Plateau. Intense stages of uplift took place 2008; Fortelius et al., 2014; Song et al., 2014). in both the early (ca. 23 Ma) and late Miocene The Miocene was the most crucial period (ca. 13–8 Ma) (Zhang et al., 2008) (Figure 1). for the initiation of the East Asian monsoon The world we know today has resulted from system (EAMS) (e.g. Guo et al., 2002; Liu et ongoing tectonic changes originating from the al., 1998; Sun and Wang, 2005). The climate middle Miocene (Potter and Szatmari, 2009). in eastern Asia was dominated by the planetary Important tectonic events affecting global oce- wind system, with zonal pattern of atmospher- anic circulation and heat transportation in the ic circulation that was steered by the latitudi- Northern Hemisphere during the Miocene in- nal pressure belts, throughout most of the Mio-

11 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

Compilation of important events that took place during the Miocene; solid precipitiation curve for plio-

Figure 1. pithecoid localities, dashed line depicts the general trend of mean annual precipitation (MAP) in Eurasia during the studied interval (MNEQ04-MNEQ12).The estimates MAP shown are based on the method of Liu et al. (2012).The deep-sea oxygen isotopes, and the derived tempereature data from Zachos et al. (2001).

12 cene, with dry summers and a distinctive dry Eder et al., 2008; Ivanov et al., 2011). Major belt in the mid-latitudes. This lasted until the vegetation changes of the area occur in the late latest Miocene, when the uplift of the Tibetan Miocene, with the subtropical region shrinking strengthened the regional monsoon system (An VLJQL¿FDQWO\DVWHPSHUDWHGHFLGXRXVDQGPL[HG et al., 2001; Passey et al., 2009; Liu et al., 2009; mesophytic forests spread, and open environ- Tang et al., 2011, 2013). The climate in China ments became more common (Kovar-Eder et was relatively humid, showing no clear pattern al., 2008; Ivanova et al., 2011). From the mid- of latitudinal gradient during 17–13.5 Ma (Liu GOH0LRFHQHRQZDUGVWKHZHVWHUQ(XUDVLDQÀRUD et al., 2009). Between 13.5 Ma and 11.1 Ma, UHÀHFWHGDFOHDUODWLWXGLQDOJUDGLHQWZLWKWHP- the mid-latitudes experienced more arid condi- peratures increasing and precipitation decreas- tions, while in the north and south, humid condi- ing from north to south (Fauquette et al., 2007). tions prevailed (Flower and Kennett, 1994; Liu Stable isotopes of herbivorous mammals’ et al., 2009). There is no consensus on whether enamel as well as soil carbonates have revealed North China was humid during this time, but that at the end of the Miocene there was glob- within the generally arid environment, regional expansion of C4 grasses, the warm season ally humid climate systems may have prevailed grasses, (Cerling et al., 1997; Strömberg, 2011), (Kaakinen et al., 2015; Wang and Zhang, 2011). which indicates the spread of more open envi- Pollen data from early Miocene China sug- ronments. In northern China, C4 grasses became gests that some areas of western inland China a dominant part of the vegetation ca. 8–7 Ma may still have been affected by the Tethys cli- (e.g. Passey et al., 2009; Zhang et al., 2009). It PDWHZLWKSDODHRÀRUDKDYLQJVPDOODPRXQWVRI was not a uniform change, however, but had a tropical and subtropical elements (Zhao et al., ODWLWXGLQDOSUR¿OHZLWKPRUHRSHQHQYLURQPHQWV 1982; Wang et al., 1990; Wang, 2006). These el- LQWKHQRUWKDVWKHQRUWKZDUGVLQFUHDVLQJį13C ements disappeared during the middle Miocene, suggest (Passey et al., 2009). DQGWKHLQODQGSDO\QRÀRULVWLFUHJLRQH[WHQGHG into Inner Mongolia (Wang, 1994; Wang, 2006). Several palaeovegetation studies from China 1.3 Pliopithecoids have shown that there was, in correspondence Pliopithecoidea is an extinct and diverse super- with the trend of general global cooling and arid- family of primitive catarrhine primates with no L¿FDWLRQDQLQFUHDVHGRSHQLQJRIYHJHWDWLRQLQ known descendants, involving approximately 20 North China (Wang, 1994; Sun and Wang, 2005; LGHQWL¿HGVSHFLHV VHH%HJXQ:XHWDO Jiang and Ding, 2009; Liu et al., 2011; Jacques et 2003; Alba et al., 2010; Alba and Moyà-Solà, al., 2013). In South China, however, no notable 2012). They ranged widely throughout Eurasia change in vegetation can be detected (Jacques between ca. 17 and 7 Ma (Andrews et al., 1996; et al., 2013). The contrast between north and Harrison and Gu, 1999; Begun, 2002; Alba et al., south in the Pliocene is even more pronounced DQGZHUHDPRQJWKH¿UVWIRVVLOSULPDWHVWR (Jacques et al., 2013). be discovered and described (e.g. Lartet, 1837; In western Eurasia early and middle Miocene Biedermann, 1863; Hofmann, 1893; Ginsburg vegetation cover was rich and diverse with sub- and Mein, 1980; Ginsburg, 1986; Andrews et tropical forests in the western central Europe and al., 1996; Harrison and Gu, 1999; Begun, 2002; more temperate forests in the eastern part (Kovar- Alba et al., 2010; Harrison, 2013). Their estimat-

13 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 ed size-range was ca. 6–20 kg (Fleagle, 2013; pithecoids diverged, most likely in the late Oli- Harrison, 2013), with dietary adaptations vary- gocene in Africa (Harrison, 2013). Possible sup- ing between generalized frugivory and highly port for the African origin comes from discover- restricted folivory (Ungar and Kay, 1995; Un- ies from Uganda in the form of Lomorupithecus gar, 1996; Begun, 2002). The locomotive adap- harrisoni (Rossie and MacLatchy, 2006). If this tations of pliopithecoids were also diverse: the indeed belongs to the Pliopithecoidea, it would smaller species were more monkey-like above- be the oldest and only non-Eurasian member of branch quadrupeds, while the larger ones might the superfamily. Begun (2002) has suggested that have been more suspensory in their locomotor VRPHVSHFLPHQVIURP.HQ\DLGHQWL¿HGDVLim- adaptation, similar to South American monkeys nopithecus could in fact be pliopithecoids. How- (e.g. Zapfe, 1960; Simons and Fleagle, 1973; ever, the geographic and phylogenetic origins of Szalay and Delson, 1979; Fleagle, 1983; Lang- pliopithecoids is still waiting to be solved. don, 1986; Rose, 1994; Begun, 2002). 7KH¿UVWFRQ¿UPHGRFFXUUHQFHRISOLRSLWKH- Pliopithecoidea is most often divided in- coids is from the late early Miocene (ca. 18–17 to three taxonomic groups (Harrison and Gu, Ma) of Sihong, Jiangsu Province, China, where 1999; Moyà-Solà et al., 2001; Begun, 2002; Har- the two oldest and the most primitive genera, rison, 2005; Alba et al., 2010). For the purpos- Dionysopithecus and Platodontopithecus are re- es of this study, one family, Pliopithecidae with corded (Li, 1978; Harrison and Gu, 1999; Begun, three subfamilies, Pliopithecinae, Crouzeliinae, 2002). These early primitive Asian forms prob- and Dionysopithecinae is used, following Moyà- ably developed into more specialised pliopith- Solà et al. (2001), Alba et al. (2010), and Alba ecines, later dispersing into Europe. Previously, et al. (2012). the more derived and late-surviving pliopithe- Based on some similarities pliopithecoids coids, the crouzeliines, were believed to have were once considered hylobatids (e.g. Gervais, originated from these European forms (Harrison 1849; Hoffmann, 1893; Hürzeler, 1954; Zapfe, and Gu, 1999; Moyà-Solà et al., 2001; Begun, 1958; Simons 1972; Simons and Fleagle, 1973; 2002; Alba et al., 2010) but, new evidence of Wu and Pan, 1985; Fleagle, 1988; Meldrum and primitive crouzeliines from Fanchang, east China Pan, 1988), however, they retain such primitive implies that they may have originated in Asia pri- features that predate the cercopithecoid-homi- or to entering Europe (Harrison, 2013). The latest noid divergence and are therefore currently con- known occurrence of pliopithecoids is also traced sidered primitive catarrhines, possibly sister tax- to China, where they co-existed with hominoids on to the living crown catarrhines (Andrews et at ca. 6.2–6.9 Ma (Dong and Qi, 2013) from the al., 1996; Begun, 2002; Harrison, 2013). late Miocene locality Shihuiba, Lufeng, Yunnan Catarrhines originated during the Paleogene, Province. Other Chinese pliopithecoid localities late Oligocene ca. 29–32 Ma ago and did not are from MNEQ06; Hezheng-Laogou, Tongxin, spread into Eurasia until the early Miocene in and Junggar-Tieersihabahe (Figure 2). Pliopithe- Afro-Arabia (Andrews et al., 1996; Harrison and coid diversity declined during the middle and late Gu, 1999; Harrison, 2005). Unfortunately, the Miocene, and only crouzeliines survived beyond African evolution of early catarrhines is poorly the mid-Vallesian in Europe. In Asia they sur- known, with a gap in the fossil record ca. 20–24 vived in the subtropical forests and woodlands Ma (Harrison, 2013). During this interval plio- until ca. 7 Ma (Andrews et al., 1996). Continu-

14 ing climatic deterioration and dispersal of cerco- ferences in the palaeodemography of pliopithe- pithecoids from Africa may have contributed to coids and hominoids in different sediments, sug- WKHLU¿QDOH[WLQFWLRQ +DUULVRQ gesting that the different ecological requirements Despite their wide distribution, pliopithe- affected their distribution. coids are known to have co-existed with the penecontemporaneous and equally broadly distributed hominoids at only 7/101 localities that 1.4 Damiao, Inner Mongolia were studied (see Paper I for description). Inad- The widespread fossil-rich Neogene terrestrial HTXDWHVDPSOLQJZDVEULHÀ\GLVFXVVHGDVDUHDVRQ depostis of Central Inner Mongolia have attracted for this by Andrews et al. (1996), but was ruled the attention of palaeontologist since the early out, as various well-sampled and rich localities years of the last century (e.g. Andersson, 1923; have yielded only one of the clades, while some Teilhard de Chardin, 1926a,b; Andrews, 1932). poor localities have yielded both. Andrews et al. Inner Mongolia contains some of the richest ver- (1996) and Harrison et al. (2002) suggested dif- tebrate-producing fossil localities in the world. ferences in habitat preferences that would have The fossil-rich sediments of this tectonically sta- allowed only minimal overlap in clade distribu- ble region are commonly spread in small ba- tion. This has proven to be the case at Rudabánya sins with a limited distribution, and stratigraphic by Armour-Chelu et al. (2005), who found dif- thickness lacking direct superpositional relation-

Figure 2. Map showing locations of Damiao and Tunggur, and other Chinese Miocene pliopithecoid localities; 1. Sihong, 2. Fanchang, 3. Hezheng-Laogou, 4. Tongxin, 5. Junggaar-Tieersihabahe, and 6. Lufeng

15 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 ships (Kaakinen et al., 2015; Wang et al., 2009; open adapted carnivores (e.g. Felis manul, Lynx Qiu et al., 2013). Inner Mongolian Miocene fos- lynx, Vulpes corsac, Vulpes vulpes, Mustela al- sil mammal localities are mainly concentrated in taica, Mustela eversmanii, Vormela peregusna) a triangular area bounded by three major cities; (Wozencraft, 2008). Xilinhot, Erenhot, and Zhangjiakou (Qiu et al., One of the most continuous sequences of 2013). Damiao, however, is located outside of strata in Inner Mongolia with early, middle, this triangle, in the central region of Inner Mon- and late Miocene fossil faunas present occurs golia, along the Sharamurun river valley near the in Damiao (Zhang et al., 2011; Kaakinen et al., Damiao village of Shiziwang Banner, ca. 65 km 2015). The Damiao mammal locality in Siziwan- north of the town of Wulanhua. The undulating gqi, Inner Mongolia was discovered in 2006 by terrain has elevations ranging between 1250 and professor Zhang Zhaoqun from the Institute of 1350 m a.s.l. marking the landscape (Kaakinen Vertebrate Paleontology and Paleoanthropology et al, 2015) (Figure 3). ,933 'XULQJWKHIROORZLQJWKUHH¿HOGVHDVRQV The modern setting is arid, with a mean an- three main fossil horizons with rich mammalian nual precipitation of 237 mm, and a mean an- fauna were discovered (Figures 4). nual temperature of 4.7°C (Harris et al., 2014). The Damiao sequence is characterized by a Biogeographically, modern Damiao belongs to ÀXYLDOHQYLURQPHQWZKHUHWKHEXONRIWKH¿QH the Inner Mongolian arid steppe and desert grass- grained deposits in the sequence represent well- land region (Xie et al., 2004). The area is occu- GUDLQHGÀRRGSODLQVDQGVRPHHSKHPHUDOODFXV- pied by small mammal fauna consisting mainly trine environments. The western part portrays of open and arid adapted forms of dipodid and PRUHGLVWDOIDFLHVRIWKHÀXYLDOV\VWHPUHSUH- cricetid rodents (Smith, 2008; Lunde and Smith, VHQWLQJDÀRRGEDVLQHQYLURQPHQWZKHUHDVLQ 2008). The modern large mammal fauna includes the eastern section the sedimentological evidence both closed, forest/woodland-adapted forms of SRLQWVWRDFRQWLQXDOLQÀXHQFHRIDÀXYLDOV\VWHP cervid artiodactyls (Alces americanus, Cervus of low sinuosity (Kaakinen et al., 2015). nippon), as well as more open adapted forms of Capreolus pygargus, Cervus elaphus, bovid artiodactyls (Gazella subgutturosa, Procapra gut- 2 Aims of the study turosa) (MacKinnon, 2008), equid perissodactyl (Equus hermionus) (Smith, 2008), and mostly The main objective of this study is to produce

Figure 3. General view of the area at Damiao fossil localities. A. View from the western locality DM16 toward the north, B. basal sands, at the eastern section, C. view towards east, facing the localities DM01 and DM02.

16 Figure 4. A. DM16 the early Miocene locality with systematic excavation, B. DM01 the middle Mio- cene pliopithecoid locality, and C. The adjacent early late Miocene locality of DM02. new information about pliopithecoids, their ecol- and Harrison (2008). ogies as well as about environmental conditions in Eurasia during the Miocene, with a special focus on the middle Miocene Inner Mongolian lo- 3 Material cality of Damiao. This locality provides valuable information on the Miocene of Inner Mongolia, Palaeontological data for Paper I and III was having the rare occurrence of early, middle, and downloaded from the NOW database (Fortel- late Miocene fossil faunas in sequence (Zhang et LXVKWWSZZZKHOVLQNL¿VFLHQFH al., 2011; Kaakinen et al., 2015). Damiao also has now). When trophic or locomotor data were not the latest occurrence of pliopithecoid primates in available to use for Paper I in NOW, comple- the late middle Miocene of Central Asia. mentary data were sourced from the Palaeobi- 7R DFKLHYH WKLV JRDO ZH ¿UVW WDNH D ORRN ology database (PBDB; http://www.fossilworks. at the pliopithecoid localities in general: What org), and from literature (Baudelot and Collier, were their localities like, and how did they differ 1978; Qi, 1979; Piller et al., 2000; Ye et al., 2000; from other contemporaneous localities? Were the Ginsburg, 2001; Moyà-Solà et al., 2001, 2009; rare co-occurrences of pliopithecoids and homi- Harrison et al., 2002; Bernor et al., 2004; Xiang noids both temporal and spatial? Subsequently, et al., 2004; Alba et al., 2006, 2010; Sun et al., we study the “unlikely” locality in north Chi- 2007). The data from NOW included all mam- na; we consider this location to be unlikely, as mal localities between 18 and 7 Ma range. Fol- the presence of pliopithecoid primates in Cen- lowing Mirzaie Ataabadi (2010), ad hoc MN tral Asia after the middle Miocene climatic opti- equivalents (MNEQ) are used instead of tradi- mum is not in accordance with the general trend tional MN units, to make Eurasian mammal bio- of strengthening climatic zones and increasing stratigraphy provisionally uniform. The MNEQ aridity since the early Miocene (Kaakinen et al., is computed from minimum and maximum age 2015). Why were there pliopithecoids in Damiao, estimates for the locality. The assignment to a but not in the adjacent, seemingly similar fossil- VSHFL¿F01XQLWLVEDVHGRQDORFDOLW\¶VWHPSR- rich locality of Tunggur? Did they live in warm, ral range midpoint: it is assigned when this falls humid forest habitats, or did they occupy rela- within the corresponding traditional MN unit’s tively dry grassland or woodlands under strong range, and has a duration equal to or less than seasonality, a question brought up during the de- the said MN unit (Mirzaie Ataabadi, 2013). For scription of the Damiao pliopithecoid by Zhang Paper I small mammal orders (Chiroptera, Eu-

17 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46 lipotyphla (Insectivora), Lagomorpha and Ro- lysed. From the late Miocene DM02, analysis dentia) were excluded from the study, following material comprised 158 fossil specimens: mainly Fortelius et al. (1996), Eronen and Rook (2004), teeth, a few jaw fragments and no postcranial and Nargolwalla (2009), because they are col- material. For taphonomical analysis, the sample lected by different procedures and are subject sizes were relatively small, and for DM02, they to different taphonomic processes compared to were strongly biased against postcranial mate- large mammals (Behrensmeyer et al., 1979; An- rial. However, samples from DM16 and DM01 drews, 1990; Behrensmeyer, 1991; Fernández- consisted of a good selection of different skel- Jalvo, 1995). All primates were also excluded etal elements (Paper II) with clear taphonomic from the analysis, to avoid circularity in deter- features indicative of similar taphonomic history mining the palaeoecological context (Fortelius for each locality. and Hokkanen, 2001; Eronen and Rook, 2004). Material for the stable isotope analysis in The ABSOLUT6 database (Damuth, 2010) and 3DSHU ,,, ZHUH FROOHFWHG GXULQJ WKH ¿HOG VHD- literature (Louys, 2008 and references therein; sons in 2007 and 2008. For the analysis enamel Louys pers.comm. 2014) are sources for the re- from 79 teeth of large herbivorous fossil and cent comparative data in Paper I in the form of modern mammals were studied. Of these teeth the 101 Eurasian and African national parks and 52 were perissodactyls (rhinos and equids), 25 reserves from which the data is derived. DUWLRGDFW\OV ERYLGVFHUYLGVRQHJLUDI¿GDQG For the purposes of this study, the fossil local- DIHZXQLGHQWL¿HGUXPLQDQWV DQGWZRSURERV- LWLHVZHUHJURXSHGLQWR¿YHVXEVHWVSOLRSLWKHFRLG cideans (gomphotheriids). The fossil material localities (P), hominoid localities (H), localities comes from twelve early to late Miocene local- with co-occurring pliopithecoids and hominoids LWLHV7KHPRGHUQVDPSOHLQYROYHV¿YHERYLG (CO), cercopithecoid localities (C), and fossil lo- specimens. calities without primates (O). If a locality con- tained two or more of the above-mentioned primate groups it was included in multiple subsets, 4 Summary of the except for the co-occurrence of pliopithecoids original publications and hominoids, which was always recognized as a separate group. 4.1 Paper I Fossil specimens for the taphonomic analy- Co-occurrence of pliopithecoid and hominoid sis of Paper II were collected from Damiao dur- primates in the fossil record: an ecometric anal- LQJWKHWKUHH¿HOGVHDVRQV±)RVVLOV ysis from the early Miocene DM16 consist of 360 specimens from the systematic excavation site, Paper I considers the rarity of co-occurrence of of which 255 were available for taphonomical pliopithecoids and hominoids in the Miocene analyses. Half of the analysed specimens are with Eurasia, and offers possible explanatory reasons. full grid reference (i.e. co-ordinate and depth in- The possibility that differences in their habitat formation). The majority of DM16 material are preferences allowed only a minimal overlap in postcrania, with few teeth and jaws. Total of 185 their distribution has been suggested by Andrews fossil specimens from the middle Miocene local- et al. (1996) and Harrison et al. (2002), but this ity DM01, both teeth and postcrania, were ana- suggestion had not been further explored. We

18 studied the ecological context of the co-occur- in this way it may be possible to reduce some of rence of pliopithecoid and hominoid primates the damage collecting always produces. using taxonomically based primate-free palaeoecological diversity, as well as direct ecometric analysis based on hypsodonty in mammalian 4.3 Paper III herbivores. The results show that pliopithecoids The palaeoenvironment of the middle Miocene had more persistently humid occurrence contexts pliopithecoid locality in Damiao, Inner Mon- compared to the other groups studied, suggesting golia a more conservative climate niche. This paper also shows that direct ecometric analysis gives Paper III sheds light on the presence of humid- a better separation of the ecological preferences favouring pliopithecoids in the increasingly arid of primate clades than analyses of taxonomically Central Asia, in the middle Miocene in Damiao, based palaeoecological diversity analysis, at least Inner Mongolia, as well as their absence from when the analyses themselves exclude primates. the seemingly similar adjacent locality of Tung- gur. At the same time, we inspect the environmental history of the whole Damiao sequence 4.2 Paper II using faunal similarity, ecometrics, and stable Small mammal taphonomy from three Miocene isotope analyses. The data from the Damiao plio- localities from Damiao, Nei Mongol, China pithecoid locality (DM01) supports previous no- tions of the presence of locally humid environ- Paper II presents the sedimentary context and ta- ments within the increasingly arid surroundings phonomic features of small mammal burials and in a generally drying Central Asia, as proposed LGHQWL¿HVWKHDJHQWVWKDWDUHUHVSRQVLEOHIRUWKH by Kaakinen et al. (2015). Hypsodonty, and the fossil accumulations in Damiao, Inner Mongolia. estimated mean annual precipitation suggest a We examine three different aged Miocene locali- more humid and possibly more closed environ- WLHVZLWKGLIIHUHQWÀXYLDOVXEHQYLURQPHQWVUDQJ- ment for DM01. However small mammal fauna ing from early Miocene to early late Miocene (ca. and isotopes suggest a mosaic environment for 21–11.6 Ma). Following Andrews (1990), we the whole Miocene for Damiao. The absence of scored taphonomic features on an ordinal scale, pliopithecoids from Tunggur may result from GHSHQGLQJRQWKHGHJUHHRIWKHPRGL¿FDWLRQ,Q the locality being seasonal or simply not humid addition, we calculated the ratios of major distal enough for pliopithecoids, as evidenced by lower and proximal elements, as well as the ratios of predicted mean annual precipitation estimates, as cranial to postcranial elements. The results reveal ZHOODVVOLJKWO\KLJKHUį13C values. We suggest that predation was the primary means of accu- that DM01, the driest-known Asian pliopithecoid mulation of small mammals at all three Damiao locality, may have been a humid refugium within localities, although the type of predator differed. a drier regional context, just as some of the late The data suggest overprinting of other means late Miocene hominoid localities of western Asia RIDFFXPXODWLRQVXFKDVÀXYLDOWUDQVSRUWDWLRQ seem to have been (Kaya et al., 2016, Mirzaie as well as possible trampling for the youngest Ataabadi et al., 2016). locality. This paper also shows that systematic excavation of small mammals can be done, and

19 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

5 Discussion we can assume that co-occurring localities had the most diverse environmental conditions, possibly comprising a mosaic of habitats between The results of the ecometric analysis in Papers I riparian or gallery forest and surrounding forest. and III support the results of Eronen and Rook Evidence of a body of water in the vicinity of (2004), as hominoids and pliopithecoids in Eur- all co-occurrence localities is provided by the asia were restricted to more humid areas. Addi- higher proportion of aquatically adapted mam- tionally, the increasing aridity towards the end mal fauna there. of Miocene pushed their range southwards (Fig- Differences in ecological preferences of pli- ure 5). It seems however that neither group of opithecoids and hominoids are detected in the primates was able to cope with drying environ- Hungarian locality Rudabánya. In this location, ments: Pliopithecoids disappeared from the Eu- the relative abundance and age distribution of the ropean fossil record by the end of MNEQ10, two groups varied in different sediments, indicat- hominoids somewhat later in MNEQ12 or even ing different core areas for the groups (Armour- MNEQ13, and in Asia both had disappeared by Chelu et al., 2005). In Rudabánya, the possible the end of MNEQ13. differences in the home range sizes of pliopithe- In Paper I we show that the ecological diver- coids and hominoids, may indicate the co-exis- VLW\DQDO\VLVWKDWZDV¿UVWDSSOLHGWRWKHIRVVLO tence of pliopithecoids and hominoids only at record by Andrews et al. (1979), makes a dis- the peripheries of their ranges and are therefore tinction between broad categories such as open rarely documented (Begun et al. 2010). Seasonal versus closed habitats, when recent habitat types overlap may be one explanation of the rare co- are observed. However, this method does not occurrence of pliopithecoids and hominoids in separate different types of fossil localities from the fossil record. High levels of humidity are each other, nor group them with any type of re- also known to determine the co-occurrence of cent habitat type. This may result from incom- modern crested mangabeys and yellow baboons plete sampling, incorrectly assigned trophic and/ in Kenya, whose home ranges and diets overlap or locomotor adaptation, or uneven sample sizes. during high rainfall and rarely during the dry sea- Also comparisons to recent material may cause son (Wahungu, 1998). It is argued that among problems, as modern counterparts to these Mio- modern primates, co-occurrence is most likely in cene localities may not exist, due to the change areas where necessary, spatially restricted basic in faunal communities over time (Andrews and resources, such as sources of water and sleep- Humphrey, 1999). As a method, this may be too ing sites, exist (Wahungu, 1998). Examples of coarse-grained for the purposes of this study, in such areas include riparian and gallery forests. which the differences may be more subtle. The Closer inspection of an Asian pliopithecoid direct ecometric approach of hypsodonty shows locality Damiao, Inner Mongolia (Papers II and that in only the most humid areas pliopithecoids III) supports the results of Paper I. Indication of and hominoids co-occurred. Mean hypsodonty closed and humid environment for the middle values for pliopithecoid localities overlap little Miocene locality DM01 comes from the pres- with those of hominoid localites, but are gen- ence of the humid favouring pliopithecoid pri- erally lower. Based on the combined results of mate (Zhang and Harrison, 2008) as well as an- ecological diversity and hypsodonty in Paper I, chitheriine horse and the cervid Euprox alticus

20 Figure 5. MAP maps for different MNEQ of Eurasia, with pliopithecoid localities marked by stars.

(Kaakinen et al., 2015; Wang and Zhang, 2011). 2015). The occurrence of dipodids and ochoto- Sedimentological evidence further supports this, nids together with Sciuridae (Atlantoxerus) could with abundant goethite and manganese occur- even indicate a semidesert environment as sug- rence, as does the predicted MAP indicating a gested for Tunggur by Qiu (1996). However, more humid climate for DM01. However, the large mammal fauna from Damiao has relative- small mammal fauna in the entire sequence is ly stable environmental pattern through time dominated by rodents, and the presence of only with cervoids as the dominant group suggest- relatively few insectivores might indicate a more ing mainly closed rather than open habitats for dry and open environment (Van den Hoek Os- the sequence (Kaakinen et al., 2015). tende, 2001; Van Dam, 2006; Klietmann et al., The isotope data from 12 Damiao fossil lo-

21 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

FDOLWLHVVKRZYDULDWLRQLQį13C values between conditions, with C3 plants displaying more en- -4.5 ‰ and -13.4 ‰ indicating that large herbiv- ULFKHGį13C values (Farquhar et al., 1989; Cerling orous mammals in Damiao were predominantly et al., 1997a; Wang et al., 2008a). feeding on C3 plants or water-stressed C3 plants, The sedimentological data from the Damiao while no pure C4 diets were consumed, much VHTXHQFHVXJJHVWZHOOGUDLQHGÀRRGSODLQVDQG like in Tunggur (Zhang et al., 2009). All the groundwater evaporation, as evidenced by abun- į13C values higher than -7 ‰ are from the late GDQWUHGEURZQ¿QHJUDLQHGGHSRVLWVZLWKFRQ- late Miocene locality UH Museum, suggesting centrations of nodular carbonate. Abundant goe- WKDW&YHJHWDWLRQZDVQRWDVLJQL¿FDQWSDUWRI thite and manganese staining in the stacked car- KHUELYRUHGLHWVLQ0LRFHQH'DPLDRį13C com- bonate nodule conglomerate at DM01 point to position variation between taxa (orders/families) periodic wetting and drying of the sediments (cf. PLJKWPHDQPL[HGKDELWDWVZLWKVPDOOHUį13C Kraus and Hasiotis, 2006) suggesting relatively values suggesting forests, while higher values humid conditions locally. The abraded soil nod- VXJJHVWJUDVVODQGV7KHORZHVWį13C values out ule clasts, however, arise from reworking of un- of all the studied Damiao localities, were found derlying overbank deposits, indicating that cli- in the primate-bearing DM01, indicating the rel- mate was seasonally dry, (cf. Van Itterbeeck et atively most humid conditions for this primate- al., 2007; Kaakinen et al., 2015). bearing locality. Locally mixed habitats are like- 7KHÀXYLDOHQYLURQPHQWFKDUDFWHULVLQJWKH O\DVLQGLFDWHGE\WKHZLGHUDQJHRIį13C values. Damiao sequence is evident also in the small Comparison of the MAP estimates of Damiao mammal fossil remains that show selective loss localities also suggests that the area was more and/or destruction of elements as indicative of humid during the pliopithecoid occupancy than ÀXYLDOWUDQVSRUWDWLRQ HJ$QGUHZV%HK- HLWKHUEHIRUHRUDIWHU7KHZLGHUDQJHRIį18O val- rensmeyer, 1975; Behrensmeyer et al., 2000; ues for DM01, and other middle Miocene locali- .RUWK +RZHYHUWKHÀXYLDOWUDQVSRUWD- ties at Damiao may indicate seasonal aridity, with tion has not been extensive for any of the locali- high values suggesting more dry, while low val- ties, as the taphonomy and preservational state of ues more humid environmental conditions. The specimens suggest. Predators, most likely owls, KLJKHVWYDOXHVRIį18O are seen in the Artiodac- for early Miocene DM16, and diurnal birds of tyla, possibly because they were the least water- prey for DM01 and DM02, were the main ac- dependent and were able to subsist on plant water cumulators of small mammals in Damiao. How- in times of seasonal aridity, as are several species ever, mammals are not fully excluded from con- today (e.g. Western, 1975; Braun, 1999a, b, c). tributing to DM01 fossil assemblage. ,WVHHPVWKDW&ELRPDVVZDVQRWDVLJQL¿- It seems that the environment in DM01 was cant component of these central Inner Mongolian already extreme for pliopithecoid survival, as, ecosystems until the Pliocene. This is consistent compared to any other pliopithecoid locality of a with other studies from central Inner Mongolia similar age in Asia, it had the lowest MAP values. (Zhang et al., 2009), which date the onset of C4 As the majority of Tunggur localities had lower expansion to the late Miocene. However, in some than predicted MAP values than Damiao, with cases, the samples approach the upper limits of only two similar localities, it seems that Tung- pure C3 diets, indicating either the presence of a gur as a whole was too dry for pliopithecoids. As small fraction of C4 vegetation or water-stressed suggested in Paper I, pliopithecoids were highly

22 conservative in their habitat selection, preferring in the increasingly arid surroundings that char- humid environments, and being unable to adapt acterized Central Asia, as proposed by Kaaki- to changing conditions. However, evidence from nen et al. (2015). Evidence supporting a more Damiao suggests that some pliopithecoids could humid and possibly more forested and wood- cope with some level of seasonal dryness. ed environment for DM01 includes estimated MAP, combined with local sedimentology and the taxonomic composition of large mammals. 6 Conclusions However, the presence of small mammal fauna indicates a more dry and open environment, as In comparison to other contemporary primate the dominance of rodents would suggest (Van JURXSVWKHGLVWULEXWLRQRISOLRSLWKHFRLGVUHÀHFWV den Hoek Ostende, 2001; Van Dam, 2006; Kliet- an ecological preference for more persistently mann et al., 2015), as well as the similarity with humid habitats. They did not adjust their habitat Tunggur fauna (Qiu, 1996). The stable isotope preferences in response to the global trend for data are consistent with a mosaic of forest and increasing aridity through the Miocene, but ef- grassland environment for all Damiao localities. fectively remained in their ancestral ecological Small mammals in Damiao were mainly ac- and climatic niche. FXPXODWHGE\SUHGDWRUVDQGGHSRVLWHGLQDÀX- The conservative nature of pliopithecoid ad- vial setting. Accumulating predators were likely aptations to a progressively diminishing range to have been owls for DM16, and diurnal birds of humid habitats appears to have restricted the of prey for DM01 and DM02. However, mam- co-occurrence of hominoids and pliopithecoids. mals are not fully excluded from contributing to The ecological context of localities experiencing DM01 fossil assemblage. Some reworking by co-occurrence of pliopithecoids and hominoids, ÀXYLDOSURFHVVWRRNSODFHLQWKHPLGGOHDQGODWH as data tentatively indicate, were even more hu- Miocene localities, DM01 and DM02. mid than of localities with pliopithecoids alone. The primate-bearing DM01 was the most hu- The detected differences between co-occurrence mid Damiao locality, although seasonally dry. localities and localities with only hominoids or This suggests that although as a clade, pliopithe- SOLRSLWKHFRLGV DUH QRW VWDWLVWLFDOO\ VLJQL¿FDQW coids were humid-favouring, the possibility of however, and it is impossible to determine de- some taxa being adapted to more arid or seasonal ¿QLWLYHO\ZKHWKHUWKHUHPLJKWKDYHEHHQRWKHU environment cannot be excluded. special conditions that were favourable to co- Based on the results in Paper III, Tunggur occurrence. PD\KDYHEHHQWRRVHDVRQDORUQRWVXI¿FLHQW- Small home-ranges of pliopithecoids (Begun ly humid for pliopithecoids. This inference is et al., 2010) and seasonal overlap as in some supported by the higher mean hypsodonty and recent primates (Wahungu, 1998), may further lower predicted MAP estimates, as well as the explain the rare recordings of co-occurrence of VOLJKWO\KLJKHUį13C values. DM01, the driest- pliopithecoids and hominoids. known Asian pliopithecoid locality, may have Our analyses of the palaeoecology of the late been a more humid refugium within a generally- middle Miocene pliopithecoid locality DM01 drier regional setting, similar to some of the late support previous inferences concerning the late Miocene hominoid localities of western Asia presence of locally humid environments with- (Kaya et al., 2016, Mirzaie Ataabadi et al., 2016).

23 DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A46

Main reason for the success of the direct sula based on new material from the Vallès- ecometric approach of mean hypsodonty over Penedès Basin. American Journal of Physi- traditional, taxonomically based palaeoecological Anthropology, 147, 135-140. cal analysis is that it is less sensitive to the ex- Alba, D.M., Moyà-Solà, S., Malgosa, A., Casa- clusion of the primates from the analysis. This novas-Vilar, I., Robles, J.M., Almécija, S., type of procedure shifts the hypsodonty value Galindo, J., Rotgers, C., Bertó Mengual, towards the higher (drier) end, but has a seri- J.V., 2010. A new species of Pliopithecus ous impact on the relative weight of the arbo- Gervais, 1849 (Primates: Pliopithecidae) real and frugivorous categories of the ecologi- from the Middle Miocene (MN8) of Abo- cal diversity analysis. Thus, ecometric traits are cador de Can Mata (els Hostalets de Pierola, inherently less prone to problems of circularity Catalonia, Spain). Am. J. Phys. Anthropol. and sampling bias than taxon-based analyses of 141, 52-75. community structure. An, Z.S., Kutzbach, J.E., Prell, W.L., Porter, Systematic sampling for small mammals in S.C., 2001. Evolution of Asian monsoons DM16 resulted in a wider range of skeletal ma- and phased uplift of the Himalaya-Tibetan terial, however it did not produce taxonomically plateau since Late Miocene times. Nature, richer sample than the more traditional excava- 411, 62–66. tion methods used at DM01 and DM02. Break- Andersson, J.G., 1923. Essays on the Cenozoic age was less pronounced in the systematic sam- of Northern China. Mem. Geool. Surv. Chi- ple, but the difference was not statistically signif- na Ser. A 3, 1-152. icant and comparative, tradtitionally excavated Andrews, R.C., 1932. The new conquest of Cen- material from the same locality is needed for tra Asia, a narrative of the explorations of GH¿QLWLYHFRQFOXVLRQV the Central Asiatic Expeditions in Mongolia and China, natural history of Central Asia. Am. Mus. Nat. History 1, 1.678. Andrews P., 1990. Owls, Caves and Fossils: Pre- References dation, Preservation, and Accumulation of Small Mammal Bones in Caves, with an Abe, M., Kitoh, A., Yasunari, T., 2003. An evolu- Analysis of the Pleistocene Cave Faunas tion of the Asian summer monsoon associ- from Westbury-sub-Mendip, Somerset, UK. ated with mountain uplift—Simulation with British Museum of Natural History, London. the MRI atmosphere–ocean coupled GCM. Andrews, P., Lord, J.M., Nesbitt-Evans, E.M., J Meteor Soc Jpn, 81, 909–933. 1979. Patterns of ecological diversity in fos- Alba, D.M., Moyà-Solà, S., 2012. A new plio- sil and modern mammalian faunas. Biol. J. pithecoid genus (Primates: Pliopithecoidea) Linn. Soc. 11, 177-205. from Castell de Barberà (Vallès-Penedès Andrews, P., Harrison, T., Delson, E., Bernor, Basin, Catalonia, Spain). Am. J. Phys. An- R. L., Martin, L., 1996. Distribution and thropol. 147, 88-112. biochronology of European and Southwest Alba, D.M., Moyà-Solà, S., Robles, J., M., Galin- Asian Miocene catarrhines. In: Bernor, R. do, J., 2012. Brief communication: The old- L., Fahlbusch, V., Mittmann, H-W. (Eds.), est pliopithecoid record in the Iberian Penin- The evolution of Western Eurasian Neo-

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