Nishioka+ (2019) Myanmar Bovids

E
Palaeontographica, Abt. A: Palaeozoology – Stratigraphy Article

Vol. 314, Issues 1–3: 11–68
Stuttgart, June 2019
Bovidae (Mammalia, Artiodactyla) from the Neogene

Irrawaddy beds, Myanmar
Yuichiro Nishioka, Masanaru Takai, Takehisa Tsubamoto, Naoko
Egi, Takeshi Nishimura, Reiko Kono, Shinkai Ogino, Thaung-Htike,
Zin-Maung-Maung-Thein and Chavalit Vidthayanon
With 9 plates, 7 text-figures, 13 tables and 2 appendices
Abstract
The evolution of the Neogene bovids (Mammalia, Artiodactyla, Ruminantia) in central Myanmar was examined, with de-
scriptions of new specimens of caprine and bovine bovids from the upper Miocene to Pliocene part of the Irrawaddy beds.
In the early late Miocene of central Myanmar, bovids had a relatively low diversity and were represented by Helicoportax sp.
and Selenoportax vexillarius. Thereafter, they rapidly diversified until the latest Miocene (c. 6 Ma), including a caprine species
(Caprinae gen. et sp. indet.) with spiral horn cores and seven bovines – Sivaportax dolabella, Tragoportax sp., Selenoportax vexil-
larius, Selenoportax falconeri, Pachyportax latidens, Pachyportax giganteus, and Proleptobos birmanicus. The latest Miocene/early
Pliocene bovids are characterised by the coexistence of the Irrawaddy-endemic species (Sivaportax dolabella and Proleptobos
birmanicus) and the other genera/species shared with the Nagri/Dhok Pathan faunas of the Siwaliks of Indo-Pakistan (Tra-
goportax, Selenoportax vexillarius, Selenoportax falconeri, Pachyportax latidens, and Pachyportax giganteus). Among them, only
the endemic species had survived throughout the Pliocene. The occurrence of Duboisia (Pleistocene bovids in Java, Indonesia)
from the upper part (upper Pliocene) of the Irrawaddy beds also supports the dominance of endemic species in Southeast Asia.
Through the late Neogene, no typical faunal turnovers were demonstrated in the Irrawaddy bovids with respect to those in the
Siwalik bovids. This relatively low turnover rate was possibly affected by the progress in the faunal isolation of central Myanmar
from the fauna of Indo-Pakistan during the late Pliocene owing to the formation of biogeographical barriers between central
Myanmar and South Asia.
Keywords: Artiodactyla, Mammalia, Miocene, Pliocene, Southeast Asia
Contents
Abstract �� 11 3.2 Measuring methods and nomenclature �� 17
1. Introduction �� 12 3.3 Abbreviations �� 17
2. Geological setting �� 13 4. Systematic palaeontology �� 17
2.1 General geology �� 13 5. Discussion �� 37
2.2 Regional geology and mammalian fauna of the fossil 6. Conclusion �� 38
localities �� 15 Acknowledgements �� 38
3. Material and methods �� 16 References �� 39
3.1 Material �� 16 Explanation of the plates �� 43
Authors’ addresses:
Y. Nishioka (corresponding author), Waseda Institute for Advanced Study, Waseda University, Nishi-Waseda 1– 6 –1,
Shinjuku-ku 169-8050, Japan; e-mail: nishioka@aoni.waseda-u.ac.jp
M. Takai, N. Egi and T. Nishimura, Primate Research Institute, Kyoto University, Kanrin 41– 2, Inuyama 484-8506, Japan
T. Tsubamoto, Graduate School of Science and Engineering, Ehime University, Bunkyo-cho 2 – 5, Matsuyama 790-8577, Japan
R. Kono, Faculty of Letters, Keio University, Hiyoshi 4 –1–1, Kohoku-ku, Yokohama 223-8521, Japan
S. Ogino, Tamba Dinosaurs Fossil Studio, Sannan-cho, Tamba 669-3198, Japan
Thaung-Htike, Department of Geology, Meiktila University, Meiktila Township, Myanmar
Zin-Maung-Maung-Thein, Department of Geology, Magway University, Magway Township, Myanmar
C. Vidthayanon, Independent Consultant on Biodiversity, Bangkok 10900, Thailand
© 2019 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/pala/2019/0088 0375-0442/2019/0088 $ 26.10
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12 Yuichiro Nishioka et al.
1. Introduction Miocene Chinji Formation, which ranges from 14.2

Oxen and antelopes, belonging to the family Bovidae to 11.2 Ma (Gentry et al. 2014). These early bovines
(Mammalia, Artiodactyla, Ruminantia), are currently have been traditionally included in a non-monophy-
widely distributed terrestrial mammals. Their phy- letic ‘boselaphine’ group, with living Boselaphini in
letic origin usually traces back to the early Miocene India (e.g. Pilgrim 1939, Table 1). In recent years,
(c. 18 Ma), when the most primitive bovid genus Eo- such Neogene ‘boselaphines’ have been replaced by an
tragus appeared in northern South Asia and Europe extinct monophyletic clade, Tragoportacini, including
(Solounias et al. 1995, Gentry et al. 1999, Bibi et Tragoportax and Miotragocerus (Bibi et al. 2009). The
al. 2009). Palaeontological and molecular biological full mitochondrial genome data by Hassanin et al.
studies have revealed that two clades (or Bovinae and (2012) and fossil calibrations of ruminants by Bibi
the others) derived from the early bovids dispersed (2013) indicated that the divergence time between
into Eurasia and Africa, and then diverged into more Boselaphini and Bovini stems from the middle Mio-
than ten major lineages, such as tribes Bovini and cene. However, fossils of early species in each tribe
Caprini, since the middle Miocene (Hassanin & have been poorly recorded, and there is a wide gap be-
Douzery 1999, Bibi et al. 2009, Bibi 2013). tween molecular and palaeontological studies on the
The Siwalik Hills is a well-known fossil locality in evolution of Neogene bovines in Asia.
South Asia, yielding a large amount of Neogene bovid The Neogene Irrawaddy beds of Myanmar are
fossils (Pilgrim 1937, 1939). Early bovines are ‘ante- closely located to the Siwalik Group (= Siwaliks) in
lope-sized’ small species (e.g. Strepsiportax and Heli- the geographical distance and geological age. John
coportax) that have been recovered from the middle Crawfurd, who was a British colonial administrator of
Table 1. Previous and our classifications of bovid fossils in the Irrawaddy fauna. The previous faunal lists were based on materi-
als from the following localities: Yenangyaung (no horizon data) by early studies (Buckland 1829, Noetling 1895), several
areas (e.g. Ava, Chaingzauk) by Pilgrim (1939), several areas (e.g. Magway, Seikpyu) by Colbert (1943), Magway by Chit
Sein (2006); Chaingzauk by Zin-Maung-Maung-Thein et al. (2011), Chaungtha (C), the basal beds in Yenangyaung (Y),
and Magway (M) by Chavasseau et al. (2013).
Early studies Pilgrim (1939) Colbert (1943) Chit Sein (2006)
Antilopinae ‘Boselaphinae’ ‘Caprine?’ Caprinae
‘Antelope’ sp. Tragoportax cf. islami ‘Hippotragine?’ Ovibovini
‘Boselaphinae’ Sivaportax dolabella ‘Boselaphine?’ Plesiaddax simplex
Boselaphus sp. Pachyportax latidens var. ‘Gazellinae’ Bovinae
Bovinae dhokpathanensis ?Gazella sp. Boselaphini
Bos sp. Bovinae Bovinae Tragoportax cf. amalthea
Bubalus sp. Proleptobos birmanicus Hemibos triquetricornis Tragoportax cf. rugosifrons
?Proamphibos cf. lachrymans ?Bibos cf. sondaicus Selenoportax vexillarius
Selenoportax sp.
Bovidae gen. et sp. indet.
Zin-Maung-Maung-Thein et al. (2011) Chavasseau et al. (2013) This study

Bovinae Incertae sedis Caprinae
Boselaphini Low-crowned form (C) Gen. et sp. indet.
Cf. Tragoportax sp. Several indet. species (Y) Bovinae
Cf. Selenoportax vexillarius Antilopinae Tragoportacini
Cf. Selenoportax lydekkeri Antilopini Sivaportax dolabella
Gen. et sp. indet. (M) Helicoportax sp.
Bovidae gen. et spp. indet. (M) Tragoportax sp.
Boselaphini
Duboisia sp.
Bovini (+ stem Bovini)
Selenoportax vexillarius
Selenoportax falconeri
Pachyportax latidens
Pachyportax giganteus
Proleptobos birmanicus
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Bovid fossils from Myanmar 13
India in the 19th century, collected mammalian fossils Since 2001, the Myanmar-Japan Palaeontologi-
from the Irrawaddy beds of central Myanmar (Clift cal Expedition Team has obtained many bovid skulls,
1828, Buckland 1829). His collection, currently horn cores, and teeth from different horizons of the
housed at the National History Museum (London), Irrawaddy beds in central Myanmar. Our recent stud-
includes some bovid skulls and teeth. One of the bo- ies revealed fossil evidence of geographical barriers,
vid fossils was originally reported as ‘a mandible of such as the Indo-Burma ranges, that influenced the
deer’ by Clift (1828). Later, this fossil was assigned isolation of rodents and primates in the Pliocene of
to Proleptobos birmanicus by Pilgrim (1913), with a Myanmar (Nishioka et al. 2015, Takai et al. 2016).
hornless skull. Bovid fossils from the Irrawaddy beds However, we do not know whether large-sized mam-
were commonly known before 1900 (e.g. Buckland mals such as bovids in the Irrawaddy fauna temporally
1829, Noetling 1895), but the knowledge of extinct changed throughout the late Neogene. The bovid fos-
Irrawaddy bovids had not yet been established (Table sils in the present study provide new information re-
1). garding the development of geographical barriers, or
Colbert (1938) examined the Irrawaddy mam- the increase of endemic taxa, from the early late Mio-
malian fossils collected by Barnum Brown (the Amer- cene to late Pliocene in central Myanmar.
ican Museum of Natural History, New York) in 1923.
Colbert (1938) divided the Irrawaddy fauna into 2. Geological setting
two biostratigraphical groups: the Lower and Upper
2.1 General geology
Irrawaddy fauna. Regarding bovids, he described two
species (Pachyportax latidens and Proleptobos birman- The Irrawaddy beds are widely distributed along
icus) from the Lower Irrawaddy and proposed an in- the Irrawaddy (Ayeyarwady) and Chindwin Rivers in
determinate bovid species from the Upper Irrawaddy, central Myanmar (Text-fig. 1). These sediments con-
basically following identification by Pilgrim (1913, sist mainly of fluvial deposits, and their thickness has
1937). Later, Pilgrim (1939) systematically described been estimated to be 2000 – 3000 m (Bender 1983).
the bovid fossils from the Irrawaddy beds (strictly the The base of the sediments unconformably overlies the
Lower Irrawaddy), including the material from the Oligo–Miocene Pegu Group, which consists of shal-
Chaingzauk area, central Myanmar, and Colbert low marine/freshwater deposits (Stamp 1922). The
(1943) reported on the Upper Irrawaddy fauna (Table uppermost part of the Irrawaddy beds is overlain by
1). the middle to upper Pleistocene river terrace deposits
After the contributions by Pilgrim and Colbert, or alluvium.
bovid fossils of central Myanmar were not examined The Irrawaddy beds have been subdivided into
for a long time. Some Myanmar researchers provision- lower and upper parts based on palaeontological
ally reported Irrawaddy bovid fossils in their master’s and lithological criteria (Stamp 1922, Colbert
or doctoral dissertations (e.g. Moe Nyunt 1987, 1938, Bender 1983, see also Text-fig. 2). Referring
Chit Sein 2006, Zin-Maung-Maung-Thein to the paleomagnetic stratigraphy of the Siwaliks
2010), where they classified bovids into several taxa (Barry et al. 2013), the Lower Irrawaddy beds cor-
(Table 1) without detailed comparisons with bovid respond primarily to the Nagri Formation (11.5 – 9.0
taxa discovered from Siwalik and other neighbouring Ma) and Dhok Pathan Formation (9.8 –c. 3.5 Ma) in
areas. Recently, several indeterminate bovid species, the Middle Siwalik (Colbert 1938; Chavasseau
including a brachydont form, were reported from the et al. 2006, 2013; Takai et al. 2006; Zin-Maung-
base of the Irrawaddy beds or the middle to lower up- Maung-Thein et al. 2011). In central Myanmar,
per Miocene of central Myanmar (Chavasseau et several middle Miocene mammalian fossils, which are
al. 2006, 2013). Zin-Maung-Maung-Thein et al. correlated with the Chinji fauna (14.0 –11.4 Ma) of the
(2011) described an outline of the latest Miocene/ Lower Siwalik, have been found. Previous studies have
early Pliocene mammalian fauna in the Chaingzauk traditionally considered that the sediments yielding
area, recognizing three bovid species based on tooth middle Miocene fossils in central Myanmar are the
size: cf. Tragoportax sp., cf. Selenoportax vexillarius, ‘Freshwater Pegu Beds’, i.e., the uppermost part of the
and cf. S. lydekkeri (Table 1). However, he examined Pegu Group (e.g. Bender 1983). However, Chavas-
neither cranial nor horn morphology, which are pres- seau et al. (2006) suggested that the basal Irrawaddy
ently considered to be the most important characters beds could extend back to the middle Miocene in the
for bovid classification. Chaungtha area, central Myanmar.
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Text-fig. 1. Geological map of central Myanmar (after Myanmar Geosciences Society 2014), showing fossil localities
examined in this study. Abbreviations for fossil localities are listed in the sentence.
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Text-fig. 2. Stratigraphical correlation of the Siwalik Group and the Irrawaddy beds and characteristic taxa of mammalian fos-
sils from central Myanmar (source: Zin-Maung-Maung-Thein et al. 2010, 2011, Chavasseau et al. 2013, Nishioka et al.
2015, Takai et al. 2015, 2016, also data from our recent excavation in 2016 – 2017).
Therefore, the Lower Irrawaddy fauna potentially (1) the Yenangyaung area included Salé, Nyaung-Do,
has a relatively long range from the upper middle Mio- and Nyaung-Hla; (2) the Chaingzauk area included
cene to lower Pliocene, and the Upper Irrawaddy fauna Chaingzauk, Myokhinthar, and Palaung; and (3) the
is correlated to the Tatrot or Pinjor fauna (< 3.5 Ma) Gwebin area included Gwebin, Than-bin-gyaung,
in the Upper Siwalik, ranging from the upper Pliocene and Peikswe.
to lower Pleistocene (Colbert 1938, Bender 1983, The Yenangyaung area is located along the east Ir-
Zin-Maung-Maung-Thein et al. 2008, Chavas- rawaddy River (Text-fig. 1), and the sediments distrib-
seau et al. 2013, Nishioka et al. 2015). uted in the area are designated for the type locality of
the Irrawaddy beds, showing both its basal and upper
2.2 Regional geology and mammalian fauna of the parts (Stamp 1922, Colbert 1938). The Irrawaddy
fossil localities beds in this area yield wood fossils in both the basal
The fossil specimens newly collected by the Myan- and upper parts. The basal Irrawaddy beds uncon-
mar-Japan Palaeontological Expedition Team were formably overly the Kyaukkok Formation of the Pegu
from the three areas of central Myanmar: Yenang- Group (e.g. Bender 1983, Ridd & Racey 2016)
yaung, Chaingzauk, and Gwebin. Each area contains and yields vertebrate fossils dominated by crocodiles
several collection localities (or GPS codes) which and turtles. It consists of red residual soil horizons
were named after neighbouring villages (Text-fig. 1): (‘red beds’), reddish brown-coloured conglomerates,
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and coarse-grained sandstones with cross beddings other fossil mammals in this area also indicate a late
and abundant sand concretions. The basal Irrawaddy Pliocene or early Pleistocene age (Moe-Nyunt
fauna is represented by Hipparion (Equidae, Hippari- 1987, Thaung-Htike et al. 2006, Tsubamoto et
oni) (Colbert 1938), which first occurred 10.7 Ma al. 2006, 2012, Zin-Maung-Maung-Thein et al.
in Pakistan (Barry et al. 2002). Chavasseau et al. 2008, Egi et al. 2011).
(2013) updated the basal Irrawaddy fauna, based on
material from the Yenangyaung area. According to 3. Material and Methods
Chavasseau et al. (2013), the basal Irrawaddy fauna
3.1 Material
consists of rhinocerotids, ‘Hipparion’ sp., Propotamo-
choerus hysudricus, Tetraconodon minor, Microbuno- A total of 899 bovid cranial and dental specimens
don milaensis, Merycopotamus sp., tragulids, and sev- (Appendices 1 and 2) collected from the Irrawaddy
eral bovid species. They considered that the age of the beds in central Myanmar were systematically exam-
lower Yenangyaung fauna approximately ranges from ined. The fossil specimens catalogued as NMMP-
10.7 Ma to 8 Ma. The lower boundary of the upper KU-IR (hereafter abbreviated as ‘IR’) were mainly
part is characterised by a hard-conglomerate bed (e.g. collected in three areas (Yenangyaung, Chaingzauk,
Chhibber 1934, our field observation). and Gwebin) by the Myanmar-Japan Palaeontological
The Chaingzauk area is located near the north- Expedition Team. A horn core, IR 3539, and some
south trending axis of the Salin syncline, the axis of dental specimens were found from Palaung 10 km to
which runs at 94° 30′ E in Pauk Township (Text- the southwest of Chaingzauk, and they were tenta-
fig. 1), where the lower Irrawaddy beds are distributed. tively included in a contemporaneous fauna with the
The Irrawaddy beds in this area are composed primar- fossil assemblage from Chaingzauk area.
ily of weakly consolidated sandstones and siltstones The specimens catalogued as NPT, BAM, MGW,
with clay intercalation and pebbly sandstones. The and PKK are housed at the Naypyitaw Museum (Nay-
mammalian fauna is characterised by abundant fos- pyitaw), Bagan Archaeological Museum (Bagan), Ge-
sils of Hexaprotodon (Hippopotamidae), Sivachoerus ology Museum of the Magway University (Magway),
(Suidae), and Stegodon (Stegodontidae), and is signifi- and Fossil Museum of Botanical Garden (Pyin Oo
cantly younger in geological age than the ‘Hipparion’ Lwyn, Mandalay), respectively. These specimens were
fauna from the Yenangyaung area. The mammalian previously collected from the Irrawaddy beds by Bur-
fossil assemblage from the Chaingzauk area is cor- mese surveys/fossil traders, but lack detailed geologi-
related partially with that from the Dhok Pathan For- cal and locality information. NPT and MGW speci-
mation of the Siwaliks, and is strictly dated as the latest mens do not have labels, but were possibly collected
Miocene to early Pliocene (c. 6 – 4 Ma), based on fauna from Magway area that exposes the basal Irrawaddy
comparisons (Takai et al. 2006, 2015, Zin-Maung- beds corresponding stratigraphically to the lower
Maung-Thein et al. 2008, 2010, 2011, Nishioka horizon in the Yenangyaung area (Chavasseau et
et al. 2011, Ogino et al. 2011, Thaung-Htike et al. al. 2013). BAM specimens include two bovid skulls
2008, Tsubamoto et al. 2006, 2012). from Bagan. One specimen (BAM 558/99) was pos-
The Gwebin area is located along the eastern flank sibly recovered from the upper part of the Irrawaddy
of the Salin syncline, near Seikpyu Township (Text- beds at an archaeological site, associated with Elephas
fig. 1), where the upper Irrawaddy beds are distrib- remains. The labels attached to the PKK collection
uted. The Irrawaddy beds in this area are composed show Pakokku, as a locality name; this is situated be-
of alternating massive siltstones and thin-bedded and tween Bagan and Chaingzauk.
fine-grained sandstones. The sandstones sometimes The specimens catalogued as ONG are Irrawaddy
include planar and trough cross beddings, sandstone fossils donated from a collector, Mr. Ong-Arj, to the
concretions, and caliches. The top of the Irrawaddy Northeastern Research Institute of Petrified Wood
beds is discontinuously overlain by river terrace de- and Mineral Resources (or Khorat Fossil Museum),
posits containing rounded pebbles. Many small/medi- Nakhon Ratchasima Rajabhat University, North-
um-sized mammalian fossils, such as teeth and jaws of eastern Thailand. The PRY and CCZ specimens are
rodents and cercopithecid monkeys, have been found housed in the Sukhothai Museum of Natural History,
primarily from the lowest horizon of the Irrawaddy Sukhothai (Thailand), which were collected by Mr.
beds in this area, which possibly ranges within 4 – 3 Porpan Vachajitpan (Bangkok) and one of the authors,
Ma (Nishioka et al. 2015, Takai et al. 2016). The CV, respectively. According to fossil traders, most Ir-
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rawaddy fossils sold in Thailand may have come from 3.3 Abbreviations

the Magway or Yenangyaung area. Although these Institution: AMNH, American Museum of Natural
collections have no geological/stratigraphical infor- History, New York, USA; BAM, Bagan Archaeologi-
mation, they usually include well-preserved skulls and cal Museum, Bagan, Myanmar; BMNH, British Mu-
horn cores. Therefore, we used such materials to de- seum Natural History (currently, The Natural History
scribe general morphology of each species, and then Museum, London), UK; CCZ, Chavalit Vidthaynon
referred our IR specimens to determine their geo- collection, Sukhothai Museum of Natural History,
graphical and stratigraphic distributions. Sukhothai, Thailand; GSI, Indian Museum, Geo-
logical Survey of India, Kolkata, India; MGW, De-
3.2 Measuring methods and nomenclature
partment of Geology, Magway University, Magway,
Skull and tooth morphometry were measured in mm Myanmar; NMM, National Museum of Myanmar,
using a digital caliper (model no. NTD12P-15C by Yangon, Myanmar; NMMP-KU-IR (= IR), National
Mitutoyo Corp., Japan) and angles of horn cores were Museum of Myanmar, Palaeontology-Kyoto Universi-
measured in degree using a protractor. Measurements ty-Irrawaddy, stored at the Department of Archaeol-
are listed below and shown in Text-fig. 3. Five mea- ogy and National Museum, Yangon, Myanmar; NPT,
surements (2, 3, 4, 6, and 11) were obtained by dou- Naypyitaw Museum, Naypyitaw, Myanmar; ONG,
bling the distance between the interfrontal or sagittal Ong-Arj collection, Northeastern Research Institute
suture and each structure if they lacked one side. In of Petrified Wood and Mineral Resources, Nakhon
teeth, maximum crown length and width were mea- Ratchasima Rajabhat University, Nakhon Ratchasima,
sured (Text-fig. 3). Molars widths were measured at Thailand; PKK, Fossil Museum of Botanical Garden,
the anterior lobe. The crown height was measured be- Pyin Oo Lwyn, Myanmar; PRY, Porpan Vachajitpan
low the protocone or protoconid. Dental terminology collection, Sukhothai Museum of Natural History,
of ruminants follows Bärmann & Rössner (2011) Sukhothai, Thailand.
(see also Text-fig. 4). Classification of bovids in sub- Localities: CHZ, Chaingzauk (21° 30′ 06″ N
families and tribes basically follows Gentry (1992) – 21° 44′ 33″ N, 94° 30′ 57″ E– 94° 31′ 48″ E); Gbn,
with a suggestion of extinct clade, Tragoportacini, Gwebin (20° 58′ 13″ N– 21° 00′ 14″ N, 94° 41′ 10″
proposed by Bibi et al. (2009). E– 94° 42′ 20″ E); MKT, Myokhinthar (21° 30′ 00″
Neurocranium measurements: 1, antero-poste- N– 21° 30′ 54″ N, 94° 29′ 55″ E– 94° 31′ 13″ E); NYD,
rior length of interfrontal suture; 2, distance between Nyaung-Do (20° 23′ 53″ N– 20° 24′ 01″ N, 94° 56′
right and left dorsal orbital rims; 3, distance between 40″ E– 94° 56′ 45″ E); NYL, Nyaung-Hla (20° 25′
right and left supraorbital foramina; 4, minimum dis- 47″ N, 94° 54′ 57″ E); PAL, Palaung (21° 27′ 42″ N,
tance between right and left horn pedicels at base; 5, 94° 23′ 30″ E); Psw, Peikswe (21° 00′ 29″ N– 21° 00′
antero-posterior length of brain case (from bregma 35″ N, 94° 40′ 17″ E– 94° 40′ 21″ E); SAL, Salé (20°
to superior nuchal line); 6, maximum width of brain 33′ 03″ N, 94° 49′ 48″ E); TBG, Than-bin-gyaung
case; 7, dorso-ventral depth of brain case (from the (20° 58′ 51″ N– 21° 00′ 01″ N, 94° 39′ 43″ E– 94° 40′
highest point of temporal crest to mastoid process); 54″ E).
8, height of occipital (from superior nuchal line to Anatomical direction: an., anterior; do., dorsal;
foramen magnum); 9, transverse thickness of basioc- la., lateral; li., lingual.
cipital at anterior tuberosities; 10, transverse thickness
of basioccipital at posterior tuberosities; 11, transverse
distance between occipital condyles. 4. Systematic palaeontology
Horn-core measurements: 12, inclination angle of
the horn core from the frontal plane; 13, divergence Family Bovidae Gray, 1821
angle between right and left horn pedicels; AP, ante- Subfamily Caprinae Gray, 1821
ro-posterior diameter of horn core at base (or usually Caprinae gen. et sp. indet.
distance between anterior and postero-lateral keels); (Plate 1, Figs 1– 3)
ML, medio-lateral diameter of the orthogonal axis to Locality and age: Chaingzauk area (MKT); latest Miocene
the diameter AP. or early Pliocene.
Tooth measurements: L, crown length; W, crown Examined material: A right horn core (IR 3138).
width; H, crown height at protocone/protoconid. Measurements: AP, 44.1 mm; ML, 34.1 mm (at the bottom).
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Text-fig. 3. Measurements in skull and teeth of bovids. A, dorsal view of skull. B, postero-dorsal view of skull. C, right lateral
view of skull. D, ventral view of basioccipital. E, occlusal view of right P4. F, occlusal view of right M3. G, occlusal view of left
P4. H, occlusal view of left M3. Explanations of numbers and abbreviations are given in the sentence. Schematic drawings of the
skull (A and C) were modified from those of Proamphibos kashmiricus by Pilgrim (1939).
Description: IR 3138 is the middle of a right horn the symmetric surfaces. The keels are on the anterior
core strongly twisted anticlockwise, seeing from the and posterior surfaces at the ventral section, rotating
ventral to dorsal directions. The horn core is mas- 180 degrees on the axis at the dorsal section. The sec-
sive and straight with an entire preserved length of tion dimension decreases by half from the ventral to
110 mm. There are two sharp keels spirally twisted on dorsal surfaces. A deep longitudinal groove extends
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Text-fig. 4. Terminology of bovid teeth (after Bärmann & Rössner 2011). A, left P4. B, left M3. C, left P4. D, left M3.
from the antero-lateral to the postero-medial sides lounias 1981; Watabe 1990, Geraads & Güleç
along the anterior keel. The medial surface at the base
1999, Zhang 2003, Bibi & Güleç 2008). Prostrep-
is flat, and the lateral surface is convex; therefore, the
ventral cross section shows a semi-circular outline. siceros vinayaki (Pilgrim 1939) is a spiral-horned cap-
The lateral surface at the dorsal section, which is con- rine from the upper Miocene (8.3 –7.9 Ma by Gentry
tinuous from the medial surface at the ventral section, et al. 2014) of Siwaliks, but its horn core is smaller and
is slightly expanded. weaker in torsion than IR 3138. The horn core of Pros-
Remarks: Miocene spiral-horned caprines, such as trepsiceros differs from IR 3138 in having a shaft open-
Prostrepsiceros and Protragelaphus, were widely distrib- ly spiralled and in lacking a deep longitudinal groove.
uted in the Eurasian Continent (Gentry 1971, So- Protragelaphus, with straight horn cores, is similar to
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IR 3138, although this genus from Europe lacks sharp Table 3. Measurements of horn-core cross sections of Siva-
keels on the anterior and posterior surfaces (Gentry portax dolabella.
1971, Solounias 1981, Geraads & Güleç 1999, AP ML
Roussiakis 2009). Sivaportax dolabella
Regarding two sharp keels and strong tortion, IR 1576 41.3 32.0
IR 3138 is similar to the middle part of the horn core IR 1684 33.6 33.8
of Palaeoreas lindermayeri (Ovibovini) that is com- IR 1729 (3) 32.2 28.2
monly found from the upper Miocene of Greece and IR 1824 39.8 32.6
IR 1832 33.7 27.7
Turkey (Gentry 1971, Solounias 1981, Geraads
IR 2855 35.8 32.1
& Güleç 1999). The upper Miocene of China has IR 3533 38.0 33.4
yielded spiral-horned caprines belonging to Ovibo- IR 3740 37.1 30.5
vini, such as Shaanxispira (Zhang 2003, Shi et al. IR 4026 32.8 27.8
2014). However, Shaanxispira has neither an ante- GSI B791* 39.9 29.6
rior nor posterior keel, which is one of the definitive
*Holotype
characteristics of IR 3138. The present finding is the
first fossil record of Miocene spiral-horned caprines in
Southeast Asia, but the genus and species identifica- Locality and age: Chaingzauk (MKT) and Gwebin (Gbn,
TBG) areas; latest Miocene to late Pliocene.
tion of IR 3138 should be left owing to a fragmentary Examined material: A skull cap (IR 1272); 19 horn cores (IR
material. 1413, IR 1576, IR 1684, IR 1729 (3), IR 1755 (2), IR 1766 (3),
IR 1824, IR 1832, IR 1897, IR 1898, IR 1899, IR 2091, IR 2136,
Subfamily Bovinae Gray, 1821 IR 2766, IR 2855, IR 3161, IR 3313, IR 3533, IR 3740).
Measurements: Shown in Tables 2 and 3.
Genus Sivaportax Pilgrim, 1939
Diagnosis: As for the genus.
Type species: Sivaportax dolabella Pilgrim, 1939
Description: The frontal is preserved in some horn-
Emended diagnosis: Small size among the Bovinae; core specimens (e.g. IR 3533). The dorsal orbital rim
brain case relatively short; fronto-parietal surface ru- is not laterally projected. The anterior surface that ex-
gose, horn cores situated directly over the orbits, being tends to the antero-medial keel of the horn core rises
tilted 45 degrees from the frontal surface, the diver- slightly over the orbit. IR 3533 lacks the postcornual
gence between the right and left horn cores of about fossa, but IR 1684 has a shallow one.
40 degrees, slightly twisted, generally curved inwards, In IR 1272, the parietals, temporals, and occipital
with some deep ditches on the posterior surface, cross are preserved. The brain case is relatively short. The
section subtriangular with a sharp antero-medial keel, parietal crests are prominent, converging posteriorly
a postero-lateral keel, and a strong postero-medial to the lambda. The dorsal surface surrounded by the
angle. right and left parietal crests is depressed and rugose
(Plate 1, Fig. 6), as in primitive bovines, such as Tra-
Sivaportax dolabella Pilgrim, 1939
goportax. Each temporal has two distinct foramina
(Plate 1, Figs 4 –19) above the mastoid process. The supraoccipital surface
Synonym: 1939 – Sivaportax dolabella Pilgrim, p. 163, pl. 4 rises and shows a trapezoidal shape elongated ante-
(fig. 7), pl. 7 (fig. 16). ro-posteriorly. The superior and median nuchal crests
Holotype: A horn core, GSI B791, from Shakwa, Kamedung, are prominent. The superior nuchal crest on the pari-
Myanmar. eto-occipital boundary extends straight. The occipital
Table 2. Measurements of skulls of Sivaportax dolabella and Duboisia sp.

Measurements 2 4 6 8 11 12 13 14 15
Sivaportax dolabella
IR 1272 — — 70.3 34.0 — — — — —
Duboisia sp.
BAM 558/99 124 53.8 90.4 41.6 68.1 40 40 33.3 50.1
eschweizerbart_xxx

surface shows a trapezoidal outline. The parieto-supraoccipital surface in dorsal view and a longer fron-
praoccipital surface crosses with the occipital surface to-parietal region than Sivaportax dolabella.
at almost a right angle. The distance between the right We have not found remarkable differences in ma-
and left mastoid processes is twice as large as the max- terials between Strepsiportax (strictly the holotype of
imum width of the supraoccipital surface. The sagittal Strepsiportax gluten) from the Siwaliks and Sivaportax
suture of IR 1272 is more than 34.5 mm long. from the Irrawaddy beds, although the phylogenetic
The horn cores are situated directly over the or- relationship between these species should be discussed
bits. The main axis makes an angle of 45 degrees with based on sufficient materials. The occurrence range of
the frontal surface and is not tilted backward, as in Sivaportax dolabella is basically restricted to the up-
Tragoportax or Miotragocerus. The right and left horn permost Miocene to Pliocene, which is considerably
cores diverge at c. 40 degrees on the frontals. The ho- younger than that of Strepsiportax (14.0 –11.2 Ma by
lotype (GSI B791) is less curved but the other spec- Gentry et al. 2014). Pilgrim (1939) suggested that
imens are strongly curved inwards, with a slight tor- Sivaportax is related to the extant Boselaphus in India
sion. There are many vertical stripes on the horn-core or Duboisia, which are Pleistocene boselaphins from
surface, and some of them on the posterior side make Java. However, our material does not support his sug-
deep ditches. The cross section is subtriangular, with gestion because the horn-core keels of Sivaportax are
sharp antero-medial and postero-lateral keels. The positioned on the anterior and posterior sides, as in
antero-medial keel is prominent from the base to the tragoportacins rather than in boselaphins.
top, but the postero-lateral keel gradually becomes
blunt near the top. The postero-medial side is strongly Genus Helicoportax Pilgrim, 1937
expanded, as the third triangular corner. The lateral Type species: Helicoportax praecox Pilgrim, 1937.
surface is flat. The parietal crest on the posterior pedi- Diagnosis (only horn core): Horn cores small, mod-
cel connects to the third angle of the horn core. erately long, slender, almost straight, slightly twisted,
Remarks: Sivaportax dolabella was originally pro- medio-laterally compressed, situated directly over the
posed by Pilgrim (1939), based only on a single left orbits; sharp antero-medial and postero-lateral keels;
horn core from Shakwa at the east from Chaingzauk. medial surface between the keels almost flat; cross
The holotype, GSI B791, shows a subtriangular cross section at base subtriangular with lateral third angle
section composed of sharp anterior and posterior keels (after Pilgrim 1939).
and an acute angle on the postero-medial side. Addi- Helicoportax sp.
tional horn-core specimens of Sivaportax dolabella
described here are the same as the holotype in these (Plate 1, Figs 20 – 22)
characteristics. Compared with the tragoportacin Locality and age: Yenangyaung area (NYD, NYL); early late
genera in South Asia, Sivaportax dolabella is similar Miocene.
to Strepsiportax gluten Pilgrim, 1937 (the holotype Examined material: A right horn core (IR 4508); three left
horn cores (IR 4067, 4494, 4502).
skull, AMNH 19746) from the Siwaliks. Sivaportax Measurements: Shown in Table 4.
dolabella is characterised by strong temporal crests
and small horn cores with the subtriangular cross sec- Description: The referred specimens are the basal
tion composed of sharp anterior and posterior keels. part of horn cores with the pedicel. The horn core is
Horn-core structures, including curvature and tor- slender and slightly twisted anticlockwise (left side).
sion, of Strepsiportax gluten are also similar to those of The cross section at the base is triangular, with sharp
Sivaportax dolabella. Tragoportax and Miotragocerus antero-medial and postero-lateral keels. The lateral
from the Siwaliks share cranial characteristics (e.g. ru-
gose surface on the fronto-parietal region) with Siva- Table 4. Measurements of horn-core cross sections of Heli-
portax dolabella. Moreover, Solounias (1981) tenta- coportax sp.
tively considered that Strepsiportax gluten belongs to AP ML
the genus Miotragocerus. However, horn cores of both Helicoportax sp.
Tragoportax and Miotragocerus are larger, more me- IR 4067 36.6 25.3
dio-laterally compressed, and more tilted backward IR 4494 30.3 20.6
than in Strepsiportax gluten and Sivaportax dolabella. IR 4502 29.5 18.5
IR 4508 32.0 23.2
Furthermore, Tragoportax species have a narrower su-
eschweizerbart_xxx

surface is expanded as a third angle, and the medi- The cross section at the base forms a subtriangular
al surface is completely flat. The anterior end of the outline composed of strong antero-medial and poste-
temporal crest connects on the medial surface of the ro-lateral keels. The medial surface is expanded more
pedicel, which is situated at the midpoint between the strongly than the lateral surface. The anterior end of
keels. The pedicel is slightly constricted inwards. The the temporal crest is situated near the posterior keel,
supraorbital foramen is not open on the pedicel, or it and approaches the cervical line between the horn
is situated below the pedicel. core and the pedicel. There are many longitudinal
Remarks: The present horn cores belong to the genus ditches on the basal surface of the horn core, but these
Helicoportax, which have a subtriangular cross section ditches disappear on the upper half of the surface. On
composed of a sharp antero-medial keel, a postero-lat- the anterior keel at the base, there is no rugose struc-
eral keel, and a strong lateral angle. The horn core of ture.
Helicoportax is generally smaller and more slender Remarks: The horn core of IR 3539 is medio-lateral-
than that of other bovines, such as Tragoportax, Mi- ly compressed, similar to a blade. This morphology is
otragocerus, and Selenoportax. Helicoportax, includ- characteristic in the horn cores of Tragoportax or Mi-
ing H. praecox and H. tragelaphoides, were described otragocerus, which differs from the horn cores of oth-
from the Chinji and Nagri formations of the Siwaliks er Miocene bovines, such as Sivaportax, Helicoportax,
(Pilgrim 1937), and additional specimens were also and Pachyportax. IR 3539 is morphologically similar
recovered from the horizon ranging from 12.3 to 11.8 to the horn core of Tragoportax than that of Miotrago-
cerus, in having a subtriangular cross section with a
Ma (Gentry et al. 2014). These species are almost as
medial angle and no antero-basal rugosities along the
large as Strepsiportax gluten and Strepsiportax chinjien-
anterior keel.
sis that are known from the Chinji–Nagri boundary.
Fossil specimens of Tragoportax are commonly
Pilgrim (1937) indicated that Helicoportax differs found in the late Miocene Dhok Pathan fauna of the
from Strepsiportax by a flatter surface of the horn core Siwaliks, and have been classified into three species:
being medial instead of lateral. Gentry (1970) sug- T. salmontanus, T. aiyengari, and T. islami (Pilgrim
gested that this difference could be appeared owing to 1939). These species have minor differences in horn-
a specific variation, but all specimens examined in this core size and angle of their holotypes (AMNH 19467,
study have a lateral angle (no medial angle), similarly GSI B828, and GSI B565). However, such character-
to Helicoportax rather than Strepsiportax. istics are possibly intraspecific variations. Recently,
Genus Tragoportax Pilgrim, 1937 Spassov & Geraads (2004) suggested that T. ai-
yengari and T. islami are synonymised with T. rugosi-
Type species: Tragoportax salmontanus Pilgrim, 1937. frons and T. salmontanus, respectively.
Diagnosis (only horn core): Horn cores relatively Pilgrim (1939) described Tragoportax cf. islami
long, slightly twisted, medio-laterally compressed (e.g. GSI B557) from the Chaingzauk area, based on
(but less compressed than in Miotragocerus); sharp fragmentary horn core and dental specimens, but did
antero-medial and postero-lateral keels; lateral sur- not compare those specimens with the horn cores or
face flattened; cross section at the base triangular to teeth of the other Irrawaddy bovines. Our material, IR
subtriangular with a medial third angle; no or weak 3539, has a more flattened cross section than the horn
anterior rugosities growing downwards from the ante- core of GSI B557.
rior keel at the base (after Pilgrim 1939, Spassov &
Genus Duboisia Stremme, 1911
Geraads 2004).
Type species: Duboisia santeng (Dubois 1891).
Tragoportax sp. Diagnosis (except teeth): Occiput wider relative to
(Plate 1, Figs 23 – 25) its height than that in Boselaphus, with sharp nuchal
Locality and age: Chaingzauk area (PAL); latest Miocene or ridges; horn cores strongly curved, extending laterally
early Pliocene. beyond the orbit and then curving inwards at the tips
Examined material: A left horn core (IR 3593). (after Hooijer 1958).
Measurements: AP, 47.5 mm; ML, 29.0 mm.
Description: IR 3539 is a basal horn core. The horn
core is compressed medio-laterally and almost straight.
eschweizerbart_xxx

Duboisia sp. terior keel may be explained by the intraspecific vari-

(Plate 2, Figs 1– 3) ation in D. santeng, based on the referring specimens
of D. santeng in the Museum Geologi Bandung, Java
Locality and horizon: Bagan; possibly from the Upper Ir-
rawaddy beds or Quaternary terrace deposits.
(the specimen, MGB.Dbs., in the exhibition room)
Examined material: A neurocranium (BAM 558/99). and the Naturalis Biodiversity Centre in Leiden, the
Measurements: Shown in Table 2. Netherlands (T. T. Lim, pers. comm.).
Fossils of Duboisia have been also reported from
Description: BAM 558/99 preserves frontals with
the Quaternary deposits in Tambun, peninsular Ma-
right and left horn-core bases, parietals, temporals,
laysia, although these are fragmentary dental and
and an occipital, but these surfaces are still covered
postcranial specimens (Hooijer 1962). A new fossil
by hard matrix. The dorsal surface of the skull bends
record from the Irrawaddy beds verifies that Duboisia
down in front of the orbits, and is flat on the fron-
existed on continental Southeast Asia, possibly in the
to-parietal region. The frontal surface above the orbits
Upper Irrawaddy, or the Plio-Pleistocene.
rises as a crest, extending to the anterior surface of the
horn core. Genus Selenoportax Pilgrim, 1937
The horn cores are situated slightly behind the Type species: Selenoportax vexillarius Pilgrim, 1937.
orbits, curving up at 40 degrees with the frontal sur-
Emended diagnosis (except teeth): Large size; su-
face. The right and left horn cores have a wide interval
praorbital foramen narrow and elongate; brain case
on the frontals (53.8 mm). The horn-core shaft later-
short and wide; temporal crests very strong; postcor-
ally diverges and slightly curves to the anterior. The
nual region ventrally depressed; occipital relatively
boundary between the pedicel and the horn core is
low and wide; horn cores situated directly above the
not clear but it is near the frontal surface. The cross
orbits, greatly diverged, slightly twisted anticlockwise
section of the horn core has an elliptical medio-lat-
(right horn core), medio-laterally compressed, curved
erally expanded outline. There are sharp medial and
inwards through the upper half, with prominent ante-
lateral keels. The anterior surface is weakly expanded
rior and postero-lateral keels; cross section at base sub-
relative to the posterior surface.
triangular with a lateral angle (after Pilgrim 1939;
Remarks: The genus Duboisia was originally placed Bibi 2009).
with ‘Anoa’ (currently included in Bubalus) by Du-
bois (1891), and later described as an endemic Java- Selenoportax vexillarius Pilgrim, 1937
nese genus (Stremme 1911). The horn cores of Du- (Plate 2, Figs 4 –17)
boisia are different from those of other small bovines 1937 Selenoportax vexillarius Pilgrim, p. 740, figs 2 – 5,
(e.g. Sivaportax and Tragoportax) and living bose- 52 – 54, 56, 80.
laphins (Boselaphus and Tetracerus) with an elliptical 1939 Selenoportax lydekkeri – Pilgrim, p. 172, pl. 7 (fig. 10).
cross section that is antero-posteriorly compressed, Holotype: A skull, AMNH 19748, from the Siwaliks, Paki-
with weak medial and lateral keels. Moreover, the stan.
horn cores of Duboisia are situated behind the orbits, Locality and age of the examined material: Yenangyaung
extending laterally, and curve strongly to the anterior, (NYD) and Chaingzauk (MKT) areas: early late Miocene
as in BAM 558/99. and latest Miocene/early Pliocene.
Homo erectus localities (early to middle Pleisto- Examined material: A frontal with right and left horn cores
cene) of Java have yielded Duboisia santeng and some (IR 3693); three right horn cores (IR 3898, 3901, 4515); three
left horn cores (IR 3903, 3906, 3907); a frontal with horn
other species of Duboisia: i.e., D. problematicus Koe- pedicels (ONG 439).
nigswald (1933) and ?D. sartonoi Geraads (1979) Measurements: Shown in Tables 5 and 6.
(or D. saatensis Koenigswald 1934). The taxo-
nomic status and morphological variation among the Emended diagnosis: A relatively small species of
species of Duboisia are still uncertain, but the size and Selenoportax; horn cores widely diverging at base (c.
horn-core characteristics of BAM 558/99 are similar 100 degrees) and strongly curving throughout their
to those of D. santeng reported by Stremme (1911), course, with sharp antero-medial and postero-lateral
Hooijer (1958) and Aimi & Aziz (1985). D. santeng keels; lateral surface expanded, and medial surface flat
sometimes has a sharp keel on the anterior surface of (after Pilgrim, 1939).
the horn core, but the present specimen from Bagan Description: IR 3693, the best-preserved specimen
completely lacks this structure. The presence of the an- among the hypodigm, is a partial frontal including
eschweizerbart_xxx

Table 5. Measurements of skulls of Selenoportax vexillarius, S. falconeri, Pachyportax latidens, Pachyportax giganteus, and Prolep-
tobos birmanicus.
Measurements 1 2 3 4 5 6 7 8 9 10 11 12 13
Selenoportax vexillarius
IR 3693 — 147 87.0 37.8 — — — — — — — 20 100
AMNH 19748* — 102 60.8 37.9 75.6 88.6 41.9 — — — — 15 110
CCZ 450 145 175 — 27.1 75.4 89.8 40.0 67.9 — — 77.1 25 80
PKK 28 — 159 65.2 26.7 — — — — — — — 25 70
PRY 204 >92.0 174 81.0 29.9 77.0 96.8 54.5 63.2 28.8 51.8 — 20 80
IR 2306 — 171 66.4 28.3 72.9 96.7 51.1 60.3 28.1 49.7 79.4 20 70
Pachyportax latidens
ONG 444 >68.1 146 — 39.9 95.4 93.8 >25.0 53.4 40.9 45.6 66.1 35 80
ONG 588 >57.5 146 — 48.3 94.9 95.9 31.0 54.3 — — 75.5 — —
ONG 604 >76.7 144 64.0 55.3 84.0 90.8 48.3 58.8 23.9 46.1 78.6 25 70
GSI B488 >75.7 131 65.7 54.6 92.7 93.2 53.5 61.4 28.1 44.5 86.2 30 80
MGW 63 >97.8 203 77.5 33.0 106 102 64.1 70.5 31.9 56.3 90.5 30 50
BAM 555/99 >86.0 176 90.4 36.6 76.9 88.6 55.8 66.6 25.9 51.9 80.6 20 30
CCZ 447 >81.0 176 90.1 45.0 — — — — — — — 20 30
IR 1456 >94.0 133 60.8 42.3 73.7 77.6 53.6 48.0 — — 73.7 20 30
IR 1705 (1) — 181 72.1 43.6 — — — — — — — 20 25
BMNH M10909* 101 134 76.8 — 81.5 97.3 64.5 54.2 23.6 — — — —
*Holotype.
lower half parts of right and left horn cores. The fron- Remarks: Selenoportax is characterised by large horn
tal surface has two large supraorbital foramina, elon- cores curving inwards, similarly to a crescent shape,
gated antero-posteriorly, on the anterior surface be- with sharp antero-medial and postero-lateral keels.
low the pedicels. In ONG 439, the dorsal surface of This genus is placed within a stem group of the tribe
the frontal is smooth, without rugose structures. The Bovini, and is distinguished from the other bovines, or
brain case is small relative to the size of horn cores. strictly Tragoportacini, Boselaphini, and Tragelaphi-
The horn cores are situated directly above the or- ni, in having a single large supraorbital foramen sit-
bits. The horn-core shaft at the base inclines backward uated below the pedicel, no rugose structures on the
and diverged laterally (the divergence angle between fronto-parietal region, and a horn core with the an-
the right and left horn cores is at a right angle). The terior keel inserting medially (Bibi 2009). Relative to
right horn core curves slightly inwards and is twisted Tragoportax sp. described in this study, Selenoportax
anticlockwise. The cross section at the base is subtri- has larger horn cores with a lateral surface projected
angular with sharp antero-medial and postero-lateral more strongly.
keels. The medial surface is flat. The lateral surface Bovini is composed of three or four extant genera
is strongly expanded as a third angle in IR 3693 and (Bos/Bison, Bubalus, and Syncerus) and some extinct
IR 3907, but its extent varies in each specimen. Thus, genera belonging to Bubalina and Bovina, and these
the cross section of some horn cores (e.g. IR 3901, IR taxa have diverged from a stem group including Sele-
3906) is nearly elliptical, compressed medio-laterally, noportax and Pachyportax (Bibi 2007, 2009). Proam-
rather than subtriangular. IR 3906 is a middle part of phibos (P. lachrymans and P. kashmiricus), found from
a left horn core, and has a distinct ditch on the medial the Siwaliks (probably from the Pliocene), is consid-
surface along the postero-lateral keel. ered as an early bubalin and a link to living Bubalus
eschweizerbart_xxx

Table 6. Measurements of horn-core cross sections of Seleno- more backward from the orbits, similarly to that of
portax vexillarius, S. falconeri, Pachyportax latidens, Pachypor- Proamphibos.
tax giganteus and Proleptobos birmanicus. In Asia, the stem group of Bovini includes four
AP ML species: Selenoportax vexillarius, Selenoportax fal-
Selenoportax vexillarius coneri, Selenoportax giganteus (Pachyportax giganteus
IR 3693 55.4 41.5 in this study), and Pachyportax latidens (Gentry et
IR 3907 62.1 45.0 al. 2014). Compared with these species, the present
IR 4515 51.2 34.9
specimens from the Irrawaddy beds were assigned to
AMNH 19748* 55.3 34.8
Selenoportax vexillarius, owing to relatively smaller
Selenoportax falconeri horn cores with a lateral expansion (Table 6). This
CCZ 450 63.2 51.8 species is also characterised by the antero-medial keel
IR 2306 62.3 47.4 at the base directing more medially, owing to a wide
IR 3905 66.9 56.6 divergence angle between the horn cores. The cra-
PKK 28 72.5 51.1
PRY 204 61.3 53.5
nium (IR 3693) is slightly larger than the holotype
of S. vexillarius (AMNH 19748) (Table 5), but this
Pachyportax latidens is treated as an intraspecific variation on the basis of
NM-YNG 004 54.5 37.0 sharing non-metric cranial characteristics between
ONG 604 57.3 39.3 them.
ONG 444 49.6 44.4
GSI 488 53.0 42.3 Selenoportax falconeri (Lydekker 1886)
(Plate 3, Figs 1–15, Plate 6, Figs 1– 2)
IR 3691 93.1 50.6 1884 Boselaphus sp. Lydekker, p. 114, figs 1– 4, 7, 8, 11,
IR 3900 76.2 57.4 pl. 13.
IR 3908 80.5 59.1 1886 Strepsiceros(?) falconeri Lydekker, p. 8, figs 2, 2a,
MGW 063 78.0 47.3 pl. 2.
NPT 2015/01/2356 87.1 63.8 1910 Boselaphus lydekkeri Pilgrim, p. 70.
1937 Selenoportax lydekkeri — Pilgrim, p. 745, figs 55,
Proleptobos birmanicus 57– 59.
BAM 555/99 53.0 53.2 1939 Selenoportax lydekkeri — Pilgrim, p. 173, pl. 7
CCZ 447 52.6 50.0 (fig. 11).
IR 1079 62.5 54.2 1939 Perimia falconeri — Pilgrim, p. 166, fig. 15.
IR 1080 66.0 55.0
IR 1705 (1) 53.4 52.2 Holotype: A skull, BMNH 37262, from the Siwaliks, Perim
IR 3770 55.4 51.8 Island, Gulf of Cambay, India.
Locality and age of the examined material: Chaingzauk
*Holotype area (MKT); latest Miocene or early Pliocene.
Examined material: A skull with upper cheek teeth and right
and left horn cores (PRY 204); a neurocranium with right and
(Pilgrim 1939). The horn core of Proamphibos has a left horn cores (CCZ 450); a neurocranium (IR 2306); a right
diamond-shaped cross section with a prominent me- horn core (IR 3905); a left horn core (IR 3665).
dio-lateral expansion. Proamphibos is more derived as Measurements: Shown in Tables 5 –7.
later bovins than Selenoportax, with longer pedicels Emended diagnosis: A relatively large species of Sele-
inserting behind the orbits and supraorbital foramina noportax. Horn cores moderately diverging (c. 70 – 80
situated more anteriorly. degrees) and curving inwards; cross section expanded
Ugandax is also classified into early bubalins pos- medio-laterally, with sharp antero-medial and poste-
sibly linking to Syncerus in Africa (Gentry & Gen- ro-lateral keels; upper molars hypsodont with a strong
try 1978, Vrba 1987), although phylogenetic analy- median basal pillar (after Pilgrim 1939).
ses indicated that Ugandax is placed at the basal clade Description: PRY 204 is the only skull preserving
composed of African bovins: Simatherium, Pelorovis, almost complete horn cores and cheek teeth. The na-
and Syncerus (Geraads 1992, Bibi 2009). The horn so-frontal surface and premaxillae are broken away,
core of U. coryndonae Gentry, 2016 from the Plio- and a part of the right side (around the pedicel) has
cene Hadar Formation of Ethiopia is similar to that of been incorrectly artificially reconstructed. The left
Pachyportax from the Siwaliks, but the former shifts side is composed of the original bones preserving an
eschweizerbart_xxx

Table 7. Measurements of upper cheek teeth of Selenoportax falconeri and Proleptobos birmanicus. Abbreviations: L, tooth
crown length; W, tooth crown width.
P2 P3 P4 M1 M2 M3
L W L W L W L W L W L W
PRY 204 18.06 12.26 16.43 12.74 15.48 15.33 19.87 18.36 22.27 — 27.45 —
CCZ 447 — — 16.86 13.14 15.19 18.38 21.31 22.18 25.14 21.36 29.91 21.61
IR 1456 16.68 12.28 15.95 13.45 13.16 16.87 18.87 20.68 23.92 20.62 23.18 18.40
BMNH M10909* 24.11 17.80 18.66 18.70 14.65 22.13 23.56 26.45 26.99 28.05 28.34 28.13
*Holotype
orbit, a zygomatic arch, a jugal, maxilla, and a pedi- The horn cores are about 400 to 500 mm long,
cel. The orbit is circular rather than trapezoidal. The curving slightly inwards, and are marginally twisted
maxillary tuberosity moderately projects above the in an anticlockwise direction (right horn core). The
boundary between P4 and M1. The pedicel is about horn cores insert the frontal surface at an angle of
50 mm in depth, situated immediately above the or- about 20 – 25 degrees, and then they bend up from the
bit. The supraorbital foramen is antero-posteriorly halfway to the tip. The divergence angle between the
slender and situated on the anterior surface below the horn cores is acute (70 – 80 degrees). There are longi-
pedicel. A small pit opens on the posterior of the main tudinal streaks on the horn-core surface. Through the
supraorbital foramen in IR 2306. The frontal face is horn-core body, the antero-medial and postero-lateral
bent downward on the basicranial axis at an angle of c. keels are prominent. The cross section is rounded sub-
35 degrees. The postcornual fossa is present and very triangular, or diamond shaped with a well-expanded
shallow. lateral surface. The medial surface near the base is also
The neurocranial part is observed in PRY 204, expanded, similarly to the lateral surface. The upper
CCZ 450, and IR 2306. The parietal has strong tem- half of the horn core is compressed medio-laterally.
poral crests, forming a rectangular-shaped ridge on In PRY 204, right P2 –M3 and left P3 –M3 are
the dorsal surface of the parieto-supraoccipital region. preserved, but the occlusal surfaces of these teeth are
This ridge is elongated laterally. The dorsal surface still covered by hard sandstones. All cheek teeth are
of the postcornual region is clearly depressed, except characterised by well-developed styles and ribs on the
around the bregma. The temporal fossa is dorso-ven- labial side. The labial cone on P3 is located near the
trally deep, and antero-posteriorly short relative to anterior style, forming a deep fold. P4 has a square oc-
the width of the brain case. The brain case does not clusal surface and a vestigial posterolingual fold. The
expand laterally, and the squamosal shelves on the styles and ribs on the labial side of P4 are basically
mastoid processes are not strongly concave. The su- prominent, except the rib from the labial cone. The
praoccipital is laterally elongated, showing a trapezoi- upper molars are moderately hypsodont. Their labial
dal shape in the dorsal view. The posterior surface of surfaces between each style are prominent but weak
the occipital is broad and deep, and crosses with the relative to those of the premolars. The upper molars
parieto-supraoccipital surface at a right angle. The have a mesostyle as large as the parastyle and the meta-
nuchal lines on the occipital surface are prominent. style, forming deep folds on the labial side. The ento-
The mastoid processes are well projected laterally. style is robust, reaching the occlusal surface. M3 has a
The basioccipital bone has a weak or no median keel fossetta and no metaconule fold.
and well-developed mascular tuberosities on both the Remarks: The presence of strong temporal crests, a
anterior and posterior sides. The anterior tuberosities depressed postcornual surface, and subtriangular cross
develop like a nodule, extending antero-posteriorly. section of the horn cores, with a lateral convex, indi-
Most of the auditory parts are missing in all materi- cates that the present specimens are closely related to
als. The interval between the right and left mastoid early species of Asian bovins (Selenoportax, Pachypor-
processes is as wide as that between the right and left tax, Proamphibos, Proleptobos, and Leptobos) rather
orbits. than extant lineages (Bos and Bubalus). The antero-me-
eschweizerbart_xxx

dial keel inserting more medially at the base is one of S. lydekkeri and P. falconeri are a maxilla (GSI B231)
the diagnostic characters of Selenoportax, which dis- and a skull (BMNH 37262) of young individuals, re-
tinguish the present specimens from the horn cores of spectively. The diagnosis of both species based only on
Proamphibos, Proleptobos, and Leptobos. Furthermore, the young individuals does not completely elucidate
these three genera differ from Selenoportax and Pachy- the allocation of our specimens to S. falconeri. How-
portax in having a dorso-ventrally shallower temporal ever, the adult horn core of S. ‘lydekkeri’ described by
fossa and stronger temporal crests and a dorsal ridge Pilgrim (1937, 1939) shares apomorphic characteris-
in the parieto-supraoccipital region. The upper tooth tics, as listed in the diagnosis, with the present speci-
row of PRY 204 is much smaller and has less well-de- mens.
veloped labial ribs/foldings than that of Proamphibos
lachrymans (cast of the holotype, BMNH M26576) Genus Pachyportax Pilgrim, 1937
and Leptobos falconeri (holotype, BMNH 40887). Type species: Pachyportax latidens (Lydekker 1876).
The holotype of Pachyportax latidens is an iso- Emended diagnosis (except teeth): Large size; su-
lated M3 (GSI B219) from the Siwaliks, and has a praorbital foramen small, extending from the anterior
larger crown with a flatter labial surface than M3 of keel on the pedicel; brain case long relative to Seleno-
PRY 204. The cranial and horn-core characteristics
portax; temporal crests very strong; postcornual region
of P. latidens were described in detail based on a neu-
ventrally depressed; occipital moderately high; horn
rocranium (GSI B488) of P. l. var. dhokpathanensis by
cores situated directly above the orbits, less diverged
Pilgrim (1937). The brain case of P. latidens is almost
square in dorsal view, and it is longer than that of Sele- (50 – 80 degrees), slightly twisted anticlockwise (right
noportax. Thus, the dorsal ridge on the parieto-supra- horn core), medio-laterally compressed, with promi-
occipital surface is also elongated antero-posteriorly nent anterior and postero-lateral keels; anterior keel
in P. latidens. The basioccipital of IR 2306 is different with basal rugosities, facing more anteriorly; cross
from that of P. latidens, GSI B488, with anterior tu- section at the base subtriangular with a medial angle;
berosities developed not as keels, but as condyles. The lateral surface flattened (after Pilgrim 1939 and Bibi
basioccipital of P. latidens is possibly characterised by 2009).
the presence of the additional tuberosities in front of Pachyportax latidens (Lydekker 1876)
the anterior tuberosities (see also the description of (Plate 4, Figs 1–7)
P. latidens below); however, IR 2306 does not have
such structures. The most remarkable difference be- 1876 Cervus latidens Lydekker, p. 65, figs 7, 10, pl. 8.
tween Pachyportax and Selenoportax appears on the 1878 Capra sp. Lydekker, p. 171.
horn core. The antero-medial keel of P. latidens directs 1884 Oreas(?) latidens – Lydekker, p. 111.
1885 Capra sp. Lydekker, p. 18.
more anteriorly, and its extension on the pedicel con- 1937 Pachyportax latidens – Pilgrim, p. 766, fig. 18.
nects with the supraorbital foramen. In P. latidens, the 1937 Pachyportax latidens var. dhokpathanensis Pilgrim,
medial surface of the horn core at the base is expanded, p. 768.
as a third angle, and the lateral surface is completely 1939 Pachyportax latidens – Pilgrim, p. 197.
flat. These characteristics of P. latidens have not been 1939 Pachyportax latidens var. dhokpathanensis – Pilgrim,
found in the present specimens. p. 198, figs 19 – 20, pl. 7 (fig. 5).
The present specimens are assigned to Selenopor- Holotype: An isolated M3, GSI B219, from the Siwaliks, Has-
tax falconeri, which is an advanced form of the genus. not, Pakistan.
S. falconeri shares many cranial characteristics with Locality and age of the examined material: Locality infor-
S. vexillarius, but the right and left horn cores of the mation is not attached to the examined specimens.
former are close to each other and are less compressed Examined material: Three neurocrania (ONG 444, ONG
medially. S. falconeri had been originally described as 588, ONG 604); a right horn core (NM-YNG 004).
Measurements: Shown in Tables 5 and 6.
Boselaphus lydekkeri by Pilgrim (1910), and was later
combined with Selenoportax lydekkeri by Pilgrim Diagnosis: As for the genus.
(1937). Gentry et al. (2014) and the other recent Description: The neurocrania, ONG 444, ONG
studies have suggested that S. lydekkeri is synonymized 588, and ONG 604, are as large as that of P. latidens
with Strepsiceros(?) falconeri (Lydekker 1886), or var. dhokpathanensis (GSI B488). NM-YNG 004, a
Perimia falconeri Pilgrim 1939, and have used the right horn core, is almost as large as the basal horn
name of Selenoportax falconeri. The holotypes of core of GSI B488.
eschweizerbart_xxx

ONG 604 and the other crania preserve a neuro- dens var. dhokpathanensis from the Siwaliks, which
cranial part with horn pedicels. All specimens are not was proposed by Pilgrim (1939). The comparisons
young individuals, because their sutures on the dorsal between Pachyportax and the other bovines have been
surface have been basically agglutinated. The frontal mentioned in the section of Selenoportax. Pilgrim
is comparatively broad and bends slightly down in (1939) documented that the horn core of Parabos
front of the bregma. The dorsal surface between the (Mio-Pliocene bovines from Europe) is similar to that
orbits is slightly depressed. The orbits are well devel- of Pachyportax in cross section, but the latter has dis-
oped and project laterally. The supraorbital foramina tinct keels and is more compressed medio-laterally.
are small and elliptical and are situated on the exten- Alephis is also a Pliocene bovine from Europe, but its
sion lines of the antero-medial keels of the horn cores. horn core is expanded laterally rather than medially,
The postcornual fossa is present but very shallow. The similarly to that in Selenoportax (Gromolard 1980).
coronal sutures rise dorsally around the bregma. In the Some horn-core specimens from the Middle
parieto-supraoccipital region, the dorsal surface is sur- Awash (Pliocene) of Ethiopia, which had been previ-
rounded by strong temporal crests extending from the ously described as Ugandax sp. by Haile-Selassie et
postcornual to the superior nuchal lines on the occip- al. (2009), are similar to the horn cores of Pachyportax
ital, and form a square-shaped ridge. The lambda and latidens (Bibi 2009). Except for these specimens, horn
bregma on this ridge bulge. cores of Ugandax basically have a more rounded cross
The brain case is relatively long and expanded lat- section, with less-developed anterior and postero-lat-
erally. The ratio of the length to the width of the brain eral keels than Pachyportax. The cranium of Ugandax
case is 0.86 in ONG 604. The parieto-temporal re- is also more advanced, being close to the extant Afri-
gion is not compressed dorso-ventrally. The mastoid can buffalo, Syncerus (Gentry 2016).
processes project laterally and form broad squamosal
shelves. The shape of the supraoccipital is a laterally Pachyportax giganteus Akhtar, 1995
elongated diamond or isosceles triangular in dorsal (Plate 4, Figs 8 – 21)
view, and is connected with the parietal by closed
1995 Pachyportax giganteus Akhtar, p. 337.
lambdoidal sutures. The posterior surface of the oc-
cipital falls vertically down from the supraoccipital Holotype: A neurocranium with left horn core, PMNH
surface, and then curves toward the posterior. The oc- No. 87/323, from the Siwaliks, Dhok Pathan, Chakwal, Paki-
stan.
cipital surface is roughened by a median keel and some Locality and age of the examined material: Chaingzauk
small ridges. The outline of the foramen magnum is area (MKT); latest Miocene or early Pliocene.
oval and dorso-ventrally compressed. Examined material: A neurocranium with right horn core
The occipital condyles are robust and the dis- (MGW 063); a neurocranium (NMY no number); two
tance between them is virtually as long as the width frontal fragments (IR 1840, IR 2303); a right horn core (IR
of the brain case. The basioccipital bone is slender and 3900); four left horn cores (IR 3690, IR 3691, IR 3908, NPT
deeply projected. The anterior mascular tuberosities 2015/01/2356).
moderately protrude as slender ridges, and develop Measurements: Shown in Tables 5 and 6.
small nodules on the anterior end. The posterior mas- Diagnosis: Relative to P. latidens, P. giganteus is larger
cular tuberosities are very large with a triangle outline. and has horn cores medio-laterally compressed (after
There is a vestigial slender keel at the centre between Akhtar 1995).
the anterior and posterior mascular tuberosities. The Description: The cranial morphology is basically
auditory bullae are large. same with P. latidens described above, but both brain
The right and left horn cores diverge at 70 – 80 case and horn cores are considerably larger than P. lati-
degrees (ONG 444, ONG 588, and ONG 604) at dens. The supraorbital foramina of MGW 063 is more
the base of the horn cores or pedicels. NM-YNG 004 distinct than that of ONG 604, and separates a small
curves faintly inwards. The cross section is subtriangu- circular pit from the dorsal end of the main pit. On
lar, composed of two sharp keels on the anterior and the basioccipital, the centre line on the basal bone is
postero-lateral sides. The medial surface is expanded, absent in MGW 063. A non-catalogued specimen
as with the pedicel of ONG 604. The lateral surface displayed in the palaeontological exhibition room at
weakly curves but does not project. the National Museum in Yangon is also referred to the
Remarks: Both skull and horn-core characteristics of same form as MGW 063. Although the specimens (IR
the present specimens are compared to those of P. lati- 1840 and IR 2303) that we collected directly from the
eschweizerbart_xxx

Irrawaddy beds in Chaingzauk are fragmentary fron- sion of horn cores is sufficient apomorphy to define
tals with a pedicel, but these correspond to MGW 063 P. giganteus.
in the shape and size. Genus Proleptobos Pilgrim, 1913
A nearly complete horn core is represented by
the specimen, NPT 2015/01/2356. The basal horn Type species: Proleptobos birmanicus Pilgrim, 1913.
core and frontal characters of this specimen are sim- Emended diagnosis: Medium size; nasals narrow
ilar to those of MGW 063. The horn core of NPT in the posterior, indenting the frontals at an acute
2015/01/2356, is approximately 600-mm long. It is in- angle; ethmoidal vacuity deep dorso-ventrally; fron-
serted to the frontal plane at an angle of 35 degrees in tals strongly arched behind the orbits; supraorbital
lateral view. The right and left horn cores diverge at an foramina extremely large and deep in male, but rela-
angle of 50 degrees (MGW 063) or 70 degrees (NPT tively weak in female; postcornual region ventrally de-
2015/01/2356) through the lower half, and then curve pressed; postcornual fossa very deep and long; brain
faintly inwards, making a crescent shape. The clock- case long relative to Selenoportax; temporal crests very
wise torsion in the left horn core is relatively stronger strong; parieto-supraoccipital region prominently ris-
than that of Selenoportax since the keels rotate 180 de- es as a small square ridge; temporal fossa started grow-
grees from the base to the tip. The cross section at the ing; occipital deep, curving posteriorly; basioccipital
base is a medio-laterally compressed triangle, which triangular with well-developed posterior tuberosities;
is composed of the anterior keel, postero-lateral keel, the anterior end of choana as far as the posterior lobe
and medial angle. The keels are sharp throughout of M3 in occlusal view; hornless in female; horn cores
the horn core, and the medial angle becomes weaker situated immediately above the orbits, straight, no tor-
through the upper half. The lateral surface is com- sion, diverging at an angle of 25 – 30 degrees, inclined
pletely flat. The horn-core surface is covered by many at an angle of 20 degrees from the frontal surface, no
longitudinal grooves, and its base is roughened along prominent keel; cross section totally circular, with
the anterior keel. IR 3690, IR 3691, IR 3900, and IR weak medio-lateral compression; cheek tooth rows
3908 are basal fragments of large horn cores, and re- less curved; teeth large relative to the size of skull; P3
ferred to the same form as NPT 2015/01/2356. NPT much longer than P4; upper molars quadrate, with
2015/01/2356 is 170 % larger in the antero-posterior strong folds and ribs; basal pillars of molars promi-
diameter than that of NM-YNG 004. nent (after Pilgrim 1939).
Remarks: The genus Pachyportax includes two spe- Proleptobos birmanicus Pilgrim, 1913
cies, P. latidens and P. giganteus. Pilgrim (1939) de- (Plate 5, Figs 1–17, Plate 6, Figs 3 –7)
scribed the smallest species, P. nagrii, and the medi- 1913 Proleptobos birmanicus Pilgrim, p. 304.
um species, P. l. var. dhokpathanensis, from the Nagri 1939 Proleptobos birmanicus — Pilgrim, p. 309, fig. 35,
and Dhok Pathan formations, respectively. P. nagrii is pl. 6 (fig. 3).
based on a partial brain case without horn cores, and it Holotype: A skull, BMNH M10909, from the Irrawaddy
is considered to possibly be a female skull of P. latidens beds, Ava, Myanmar.
by Gentry (1974) and Bibi (2009). Akhtar (1995) Locality and age of the examined material: Chaingzauk
classified the largest fossils of Pachyportax from the (CHZ, MKT) and Gwebin (Gbn, TBG) areas; latest Mio-
Dhok Pathan Formation as an independent species, cene to late Pliocene.
P. giganteus, although this species has been informal- Examined material: Four skulls with horn cores and upper
teeth (CCZ 447, CCZ 448, CCZ 449, PKK 17); a neurocra-
ly combined with the genus Selenoportax or possibly nium with right and left horn cores (PRY 205); a neurocrani-
synonimised with P. latidens (Bibi 2009, Gentry et um, associated with upper teeth (IR 1456); a neurocranium
al. 2014). Our examined material from the Irrawaddy (BAM 555/99); three frontals with right horn core/pedicel
beds undoubtedly includes two forms belonging to (IR 1705 (1), IR 2850, IR 3770); three frontals with left horn
the genus Pachyportax: the smaller form is assigned core/pedicel (IR 0716, IR 2844, IR 3312); a parieto-supraoc-
to P. latidens and the larger form is compared to P. gi- cipital (IR 3430); a right horn core (IR 2845); two left horn
cores (IR 1079, IR 1080); 22 horn core fragments (IR 0204,
ganteus. P. giganteus is not significantly different from
IR 0212, IR 0539 (2), IR 1081, IR 1249, IR 2156, IR 2293, IR
P. latidens, except in size, but we tentatively agree with 2305, IR 2645, IR 2646, IR 2647, IR 2765, IR 2796, IR 2846,
Akhtar (1995) for the following reasons: (1) there is IR 2847, IR 2848, IR 2849, IR 2851, IR 2853, IR 3044, IR
no intermediate size in skulls and horn cores between 3162, IR 3185).
P. latidens and P. giganteus; (2) medio-lateral compres- Measurements: Shown in Tables 5 –7.
eschweizerbart_xxx

Diagnosis: As for the genus. osities are relatively small, forming pisolitic nodules.
Description: CCZ 447, CCZ 448, CCZ 449, and The posterior mascular tuberosities are triangular and
PKK 17 are well-preserved skulls with nasals, maxil- bulge strongly. The median surface on the basioccipi-
lae with cheek teeth, lachrymals, frontals, and horn tal is smooth without keels.
cores. The neurocranial section behind the orbits is The horn cores are situated somewhat behind
preserved in BAM 555/99 and PRY 205. A complete the orbits, with long pedicels, and are inserted to the
horn core is observed by CCZ 449, PRY 205, and IR frontal plane at an angle of 20 degrees in lateral view.
1080. Therefore, the present specimens show almost The right and left horn cores diverge at an angle of
the whole part of the skull, without incisive and zygo- 25 – 30 degrees at those bases and extend in parallel,
matic bones. faintly curving inwards. IR 1080 has a total length of
The nasals converge on the naso-frontal suture, more than 400 mm. The cross section of the horn core
indenting at an angle of about 30 degrees. A large and is completely circular without keels. It is medio-later-
shallow lachrymal fossa is present above the level of ally compressed faintly. The pedicel has a ridge (not
M1, and its posterior margin descends vertically. The keel) on the anterior surface, extending to the supra-
infraorbital foramina are moderately large, situated orbital foramen, but this ridge disappears on the horn
above the anterior root of P2. The ethmoidal vacu- core. The horn-core surface is rough with many longi-
ities are large between the nasal and lachrymal. The tudinal ditches. The postero-medial surface has deep
pterygopalatine fossae have an interval of 26 mm be- grooves from the base to the tip in some specimens
hind M3. The frontals between the orbits are weakly (CCZ 449, IR 1079, and IR 2156).
depressed, and then gently rise between the horn ped- IR 1456 is associated with a right tooth row, in-
icels. The frontal surface between the horn cores is cluding P2 to M3; and CCZ 447 and CCZ 449 also
flat and broad. The supraorbital foramina are large, preserve cheek tooth rows. These teeth are medi-
deeply concaved, and teardrop shaped. Some small um-sized and moderately hypsodont in bovines. The
pits are also present in the posterior of the main pits.
tooth row is weakly curved inwards from P2 to M3.
These foramina are situated on the dorsal surface of
All cheek teeth have prominent ribs and foldings on
the pedicels, along the horn-core direction. The min-
the labial side. P2 is subtriangular and elongated ante-
imum distance between the pedicels is half as wide as
the interval of the supraorbital foramina. The orbits ro-posteriorly. The anterior style does not project la-
project laterally below the pedicels. bially. A labial fold at the anterolabial cone is shallow.
The postcornual fossa is prominently marked. The anterolingual crista curves inwards in front of the
The postcornual surface behind the bregma is smooth lingual cone. The occlusal outline of P3 is not subtri-
and depressed. The parieto-supraoccipital surface has angular but trapezoidal, and has a wide and flat lingual
a square-shaped ridge that is surrounded by strong surface. The fold from the anterolabial cone is slender
temporal crests and superior nuchal line. This ridge and is deeply intruded. The posterolabial crista weakly
is taller but smaller in a plane than that of Selenopor- curves inwards. P4 is rectangular in occlusal view and
tax or Pachyportax. The temporal crests overhang the is antero-posteriorly compressed. The antero-poste-
temporal surface. The brain case is antero-posteriorly rior length of P4 is shorter than that of P3. The labial
longer relative to that of Selenoportax vexillarius. The surface has a weak median rib between the anterola-
temporal fossa becomes strong by high pedicels and bial and posterolabial cones. The fossa on P4 shows
strong temporal crests. The squamosal shelves on the a simple U-letter, with a vestigial central fold. The
mastoid processes are wide, projecting laterally. upper molars are characterised by a quadrate occlusal
The supraoccipital has a trapezoidal outline in outline, a strong entostyle, and well-developed ribs on
dorsal view. The posterior surface of the occipital bone the labial side. The labial surface between the ribs are
vertically descends from the supraoccipital surface, rounded from the base to the top, but do not develop
and immediately bends to the posterior. A median as a rib. The mesostyle projects labially as large as the
keel on the occipital surface is prominent but becomes parastyle and metastyle. The metaconule fold on the
weaker on the ventral side. The foramen magnum is posterior fossa is very weak. There is a small fossetta at
large and elliptic. The condyle processes are also large the centre of the occlusal surface. The posterior end of
and elongated laterally. The basioccipital bone is tri- M3 reaches below the anterior rim of the orbit.
angular, with well-developed anterior and posterior In cranial measurements (Table 5), IR 1456 is
mascular tuberosities. The anterior mascular tuber- smaller than the other skulls, such as BAM 555/99,
eschweizerbart_xxx

although it is not a young individual because it has a Proleptobos birmanicus is morphologically similar
completely erupted M3. Furthermore, the premolars to Leptobos and Proamphibos species from the Upper
of IR 1456 are more compressed antero-posteriorly Siwaliks (Pilgrim 1939). The holotype (BMNH
than those of CCZ 447. 39567) of Leptobos falconeri has rounded horn cores
Remarks: The genus Proleptobos is represented by a and strong temporal crests, but the other horn-core
single species, Proleptobos birmanicus. This species has specimens of L. falconeri diverge more laterally. More-
been known from the Irrawaddy beds in Ava (current- over, the parieto-supraoccipital ridge of L. falconeri is
ly Sagain) (Pilgrim 1939). It was described based on elongated more laterally than that of Proleptobos bir-
a female hornless skull (BMNH M10909), but its cra- manicus. Leptobos brevicornis and L. amplifrontalis are
nial and dental characteristics are identical to those of early Pleistocene species from Gansu Province, China
the present specimens of male individuals. Relative to ( Ji 1975). These are undoubtedly similar to L. fal-
Selenoportax and Pachyportax, Proleptobos birmanicus coneri. All species of Leptobos known from Asia are
has the following eight derived features: (1) larger su- more derived than Proleptobos birmanicus, in having
praorbital foramina situated on the pedicels; (2) lon- a temporal fossa antero-posteriorly longer and dor-
ger pedicels; (3) no keel on the horn cores; (4) less/no so-ventrally shallower. Leptobos species have been also
curvature and torsion of the horn cores; (5) stronger reported from the Plio-Pleistocene of Europe, but they
temporal crests, and the postcornual surface depressed have strongly curved horn cores (Duvernois 1990).
more deeply; (6) more prominent but compact pari- The holotypes of Proamphibos lachrymans (GSI
eto-supraoccipital ridge; (7) deeper and larger post- B556) and Proamphibos kashmiricus (GSI B561) share
cornual fossa; and (8) more developed temporal fossa. many neurocranial characteristics with Proleptobos
However, our dental comparisons do not show clear birmanicus, but they have sharp horn keels on the an-
differences between Proleptobos birmanicus and Sele- terior and posterior sides. Regarding cheek tooth pat-
noportax falconeri in either size or enamel pattern (Ta- tern, the labial surface of Proleptobos birmanicus is rel-
ble 7 and Plate 6). atively flat, whereas that of Proamphibos and Leptobos
Table 8. Measurements of cheek teeth of the small-sized bovids from Chaingzauk and Gwebin areas. Abbreviations: L, tooth
P3 P4 M1/M2/M3
L W L W L W
Chaingzauk
min — — — — 13.19 15.24
max — — — — 18.99 18.40
average — — — — 17.25 16.53
(n=7) (n=7)
Gwebin
min 12.77 12.23 10.12 15.09 15.39 15.73
max 15.42 18.30 12.33 21.67 24.29 22.66
average 14.10 15.27 11.23 18.38 19.31 17.90
(n=2) (n=2) (n=2) (n=2) (n=20) (n=20)
P3 P4 M1 M2 M3
L W L W L W L W L W
Chaingzauk
min — — 12.58 8.43 14.52 8.73 16.75 8.68 22.56 9.52
max — — 16.59 7.28 17.65 11.88 19.96 12.77 24.54 11.95
average 12.73 7.75 13.57 7.93 15.91 9.97 18.69 10.82 23.94 10.45
(n=1) (n=1) (n=5) (n=5) (n=10) (n=9) (n=14) (n=14) (n=6) (n=8)
Gwebin
min — — 13.53 6.82 — — 17.47 11.01 23.33 10.24
max — — 14.41 7.62 — — 18.13 11.49 27.04 12.57
average — — 14.08 7.11 13.69 9.78 17.80 11.25 24.19 11.31
(n=3) (n=3) (n=1) (n=1) (n=2) (n=2) (n=9) (n=9)
eschweizerbart_xxx

develop strong ribs and foldings on each lobe. There- weak; entostyle slender; no fossetta; metaconule
fore, Proleptobos birmanicus had acquired derived neu- fold weak or absent. Lower premolars antero-poste-
rocranial characters, retaining plesiomorphy in teeth riorly compressed; anterior stylid and anterior conid
with Selenoportax and Pachyportax. weakly present on P4; mesolingual conid extending
posteriorly (only posterolingual cristid present); an-
Bovidae gen. et sp. indet. 1 (small-sized) terior valley open; back valley shallow. Lower molars
(Plate 7, Figs 1–7) labio-lingually narrow; ribs on the lingual side weakly
profiled; labial surface sharply triangular; mesostylid
Locality and age of the examined material: Chaingzauk and entostylid slender; metastylid always absent on
area (CHZ, MKT): latest Miocene/early Pliocene. M1 and M2; ectostylids short and slender. The aver-
Examined material: Listed in Appendix 2. age of the hypsodonty index (H/L) of M3 at the early
Measurements: Shown in Table 8.
wear stage is 2.6 (n = 6). The third lobe on M3 con-
Description: P4 rounded rectangular in occlu- tains two morphotypes: morphotype S1 is semi-circu-
sal view; anterior and posterior styles well defined lar, with an isolated back fossa, and morphotype S2 is
throughout the crown; labial ribs and folds weak; cen- crescent-shaped, without lingual cristids (Text-fig. 5).
tral fossa U-shaped; no folding structures. Upper mo-
Remarks: The fossil assemblage from Chaingzauk
lars quadrate; styles on the labial side pronounced and
area includes three small-sized species (or ‘non-bovin’
slender; labial ribs from the paracone and metacone
taxa): Caprinae gen. et sp. indet., Sivaportax dolabella,
and Tragoportax sp. Miocene caprines, such as Palae-
oreas and Protragelaphus from Greece (Solounias
1981, Gentry 1971), differ from Bovidae gen. et sp.
indet. 1, in having upper molars with no or tiny ec-
tostyles and lower premolars with a deep branch be-
tween anterior stylid and anterior conid. Dental speci-
mens of Tragoportax cf. islami described from Chaing-
zauk by Pilgrim (1939) are similar to our small-sized
materials. However, those specimens clearly show P4
without the anterolingual cristid, which differs from
P4 of Trgoportax species (Spassov & Geraads
2004). No teeth associated with a skull of Sivaportax
dolabella have been found from the Irrawaddy beds,
but this species is the only candidate for Bovidae gen.
et sp. indet. 1. The present species is moderately com-
mon in the Chaingzauk fauna (Table 9).
Bovidae gen. et sp. indet. 2 (small-sized)

(Plate 7, Figs 8 –12)
Locality and age of the examined material: Gwebin area
(Gbn, TBG, Psw): late Pliocene.
Text-fig. 5. Occlusal morphotypes of M3 in the small-sized Examined material: Listed in Appendix 2.
bovids. Measurements: Shown in Table 8.
Table 9. The numbers of the bovid dental specimens (maxillae with upper teeth, mandibles with lower teeth, and isolated teeth)
from the Irrawaddy beds in each locality.
Yenangyaung Chaingzauk Gwebin Locality area unknown
Small — 77 83 —
Medium 7 322 67 —
Large — 43 25 1
Total 7 442 175 1
eschweizerbart_xxx

Description: The present small-sized species from noportax vexillarius have been known from both the
Gwebin area is similar to Bovidae gen. et sp. indet. 1 Irrawaddy beds and Siwaliks.
from Chaingzauk area in both size and enamel pat-
terns. In addition, P3 rounded triangular in occlusal Bovidae gen. et sp. indet. 4 (medium-sized)
view; labial cone and posterior style prominent; ribs (Plate 7, Figs 22 – 23, Plate 8, Figs 1– 4)
and folds on the labial side weak.
Locality and age of the examined material: Chaingzauk
Remarks: The fossil assemblage from Gwebin area area (CHZ, MKT, PAL): latest Miocene/early Pliocene.
includes only Sivaportax dolabella as a small-sized Examined material: Listed in Appendix 2.
species. Teeth measurements and enamel patterns of Measurements: Shown in Table 11.
Bovidae gen. et sp. indet. 2 is same basically with those
Description: P3 semicircular in occlusal view, ante-
of Bovidae gen. et sp. indet. 1. The present species is
ro-posteriorly elongated; styles and labial cone prom-
dominant in the fossil assemblage from the Gwebin
inent; central fossa slender, gently bending at the
area (Table 9).
level of the labial cone in an ‘L’ shape. P4 quadrate in
Bovidae gen. et sp. indet. 3 (medium-sized) occlusal view; anterior and posterior styles strongly
projected; labial rib flat; basal cingula on the lingual
(Plate 7, Figs 13 – 21) side distinct; central fossa slender, with a weak central
Locality and age of the examined material: Yenangyaung fold. Upper molars quadrate, moderately hypsodont
area (NYD, SAL): early late Miocene. (H/L = 1.4 –1.8 at the early wear stage); styles on the
Examined material: Listed in Appendix 2. labial side strong and slender; labial ribs prominent;
Measurements: Shown in Table 10.
Description: Upper molars moderately hypsodont;
entostyle tall; styles on the labial side well-developed;
labial ribs from the paracone and metacone moder-
ately expanded. Lower premolars with a wide anterior
valley opening lingually; branching between anterior
stylid and anterior conid indistinct; back valley shal-
low; mesolingual conid on P4 extending posteriorly
(no anterolingual cristid); posterolabial conid weakly
protruding. A lower molar with a prominent meso-
stylid and metastylid; metaconid sharply pointed.
Remarks: The fossil assemblage from Yenangyaung
area includes Helicoportax sp. and Selenoportax vexil-
larius, but all examined materials are medium-sized
(Table 9), and larger than the teeth of Helicoportax
species described by Pilgrim (1939). The upper mo-
lars from the Yenangyaung area are similar to those of
Selenoportax vexillarius from the Siwaliks (Pilgrim
1937, 1939), although no skulls including teeth of Sele-
Table 10. Measurements of cheek teeth of the medium-sized

bovids from Yenangyaung area. Abbreviations: L, tooth
L W
IR 4053 M1/M2 21.28 19.83
IR 4078 M1/M2 19.96 —
IR 4091 M1/M2 22.80 22.29
IR 4092 P3 17.03 9.77
IR 4092 P4 16.65 10.42 Text-fig. 6. Occlusal morphotypes of upper molars in the
IR 4546 M1/M2 18.94 10.31 medium-sized bovids.
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Table 11. Measurements of cheek teeth of the medium-sized bovids from Chaingzauk and Gwebin areas. Abbreviations: L,
tooth crown length; W, tooth crown width.
P2 P3 P4 M1/M2/M3
L W L W L W L W
Chaingzauk
min — — — — — — 19.70 17.83
max — — — — — — 28.85 26.35
average 19.23 14.66 — — 24.92 21.52 26.16 22.93
(n=1) (n=1) (n=1) (n=1) (n=62) (n=56)
Gwebin
min — — 16.36 13.61 13.35 16.66 18.81 16.13
max — — 19.28 16.10 15.71 19.42 28.27 30.16
average — — 17.67 14.87 14.83 18.04 25.08 22.51
(n=5) (n=5) (n=3) (n=2) (n=22) (n=21)
P2 P3 P4 M1 M2 M3
L W L W L W L W L W L W
Chaingzauk
min 10.57 7.50 17.17 9.59 16.99 8.46 15.59 10.31 19.75 10.78 24.71 10.23
max 11.39 7.85 19.96 10.36 21.31 12.39 24.90 16.46 29.96 19.21 40.29 19.22
average 11.11 7.62 18.47 9.91 19.11 11.11 20.75 13.89 24.57 15.16 34.66 15.30
(n=3) (n=3) (n=6) (n=6) (n=18) (n=18) (n=40) (n=38) (n=63) (n=63) (n=78) (n=76)
Gwebin
min 8.32 5.63 15.76 9.03 18.35 9.89 16.25 11.63 22.34 12.64 30.30 12.74
max 13.25 10.20 21.29 13.79 20.11 15.82 20.69 13.19 27.85 19.50 40.39 18.85
average 10.79 7.92 18.53 11.42 19.56 12.13 18.47 12.41 24.95 15.52 35.23 14.79
(n=2) (n=2) (n=3) (n=4) (n=4) (n=7) (n=2) (n=2) (n=9) (n=9) (n=13) (n=14)
ONG 577* 9.44 5.56 15.33 8.41 15.80 9.47 15.70 11.17 20.38 12.89 26.76 13.24
*Morphotype ML1
mesostyle projecting laterally, forming a deep intru- posterior fossae (Text-fig. 6). The posterior fossa is
sion forward; entostyle very tall; fossetta usually ab- usually intruded by a weak metaconule fold. On the
sent. DP4 rectangular in occlusal view; enamel pattern other hand, morphotype MU2 has more complicated
similar to that of the molars described below; meso- anterior and posterior fossae indented from both
stylid and metastylid absent; ectostylids well-devel- the anterior and posterior sides, which show butter-
oped; entostylid sometimes present. Lower premolars fly-shaped foldings (Text-fig. 6). Moreover, the labial
labio-lingually wide and triangular in occlusal view; foldings composed of styles and ribs of the morpho-
anterior and posterior valleys open lingually; branch- type MU2 are stronger than those of the morphotype
ing between anterior stylid and anterior conid weak; MU1.
posterior valley usually closed; back valley shallow/ The lower cheek teeth are subdivided into three
indistinct; mesolingual conid situated postero-lin- morphotypes (Text-fig. 7). Morphotype ML1 is repre-
gually, with a posterolingual cristid; anterolingual sented only by two mandibles with P4 to M3 (ONG
cristid absent. Lower molars rectangular, moderately 577 and MGW 005), and is slightly smaller than the
hypsodont (average H/L of M3 = 2.6 at the early wear other morphotypes (Table 11). The morphotype
stage, N = 45); ectostylid tall; posterior ectostylid on ML1 is characterized by P4 with the mesolingual co-
M3 short/absent; mesostylid and entostylid well-de- nid extending anteriorly, similarly to the anterolingual
veloped as a slender pillar; anterior cingulid weakly cristid. The posterior valley is deep, extending to the
present; third lobe of M3 subtriangular in occlusal base, and the anterior/posterior stylid is well isolated
view, without a fossa. from anterior conid/posterolingual conid, relative to
The upper molars are subdivided into two mor- the morphotypes ML2 and ML3 (Text-fig. 7). On the
photypes by folding patterns. Morphotype MU1 is lower molars, the morphotypes ML1 and ML2 have
characterised by simple and U-shaped anterior and short and slender ectostylids and relatively weak lin-
eschweizerbart_xxx

Text-fig. 7. Occlusal morphotypes of lower

cheek teeth in the medium-sized bovids.
gual ribs. The morphotype ML3 is a minor variation Bovidae gen. et sp. indet. 5 (medium-sized)
by the presence of a stout metastylid extending from
(Plate 8, Figs 5 –13)
the base to the occlusal surface (Text-fig. 7).
Locality and age of the examined material: Gwebin area
Remarks: The fossil assemblage from Chaingzauk (Gbn, TBG, Psw): late Pliocene.
area includes five medium- or large-sized species: Sele- Examined material: Listed in Appendix 2.
noportax vexillarius, Selenoportax falconeri, Pachypor- Measurements: Shown in Table 11.
tax latidens, Pachyportax giganteus, and Proleptobos
Description: Same basically as Bovidae gen. et sp.
birmanicus. The above cheek tooth characteristics
indet. 4. The present species is common in the fossil
indicate that Bovidae gen. et sp. indet. 4 belongs to
assemblage from the Gwebin area (Table 9).
Bovini or its stem group, and is more derived than late
Miocene tragoportacins, such as Tragoportax, in hav- Bovidae gen. et sp. indet. 6 (large-sized)
ing strong entostyles/ectostylids and outer foldings
(Bibi 2007). The derived enamel pattern represented (Plate 9, Figs 1–7)
by the morphotype MU2 is similar with that of extant Locality and age of the examined material: Chaingzauk
Bubalus, but such complex enamel foldings appear on area (CHZ, MKT): latest Miocene/early Pliocene.
teeth of the other fossil bovins, such as Hemibos and Examined material: Listed in Appendix 2.
Leptobos, from the Siwaliks. Most dental materials of Measurements: Shown in Table 12.
bovids from the Chaingzauk area are categorized into Description: P2 trapezoidal in occlusal view, elongat-
medium-sized species (Table 9), but the finding of ed antero-posteriorly; antero-lingual wall slightly con-
different morphotypes suggests that Bovidae gen. et caved; central fossa showing a wavelike fold. P3 round-
sp. indet. 4 is composed of some or all medium-sized ed triangular in the occlusal view, short antero-poste-
species from the Chaingzauk area, which were de- riorly; strong styles and ribs on the labial side; labial
scribed by skulls and horn cores in this study. foldings deep; central fossa showing a V-letter, with a
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Table 12. Measurements of cheek teeth of the large-sized bovids from Chaingzauk and Gwebin areas. Abbreviations: L, tooth
P2 P3 P4 M1/M2/M3
L W L W L W L W
Chaingzauk
min — — — — — — 28.83 21.84
max — — — — — — 35.86 30.11
average 23.03 19.64 21.38 24.63 — — 32.18 26.67
(n=1) (n=1) (n=1) (n=1) (n=8) (n=6)
Gwebin
min 23.46 19.02 — — — — 27.99 24.51
max 25.83 21.56 — — — — 34.12 31.20
average 24.65 20.29 19.53 21.00 21.05 25.44 31.59 27.49
(n=1) (n=2) (n=1) (n=1) (n=1) (n=1) (n=10) (n=9)
IR 0016* — — — — — — 41.46 35.07
P4 M1 M2 M3
L W L W L W L W
Chaingzauk
min 26.53 14.23 26.99 14.71 30.66 15.88 41.36 19.32
max 27.99 17.83 32.02 20.85 31.38 22.18 44.07 20.20
average 27.26 16.03 28.56 17.89 30.92 19.00 42.79 19.72
(n=2) (n=2) (n=4) (n=4) (n=4) (n=4) (n=3) (n=3)
Gwebin
IR 1857 — — — — — — 50.25 25.65
IR 0003 (2)* — — — — 36.00 >27.22 — —
*Largest form.
weak central fold on the posterolingual crista; ante- weak ribs, although tooth difference among Seleno-
ro-lingual wall slightly concaved. Upper molars hyp- portax and Pachyportax species has not been clearly
sodont (H/L of unworn M3 = c. 1.8); fossae simple, defined in this study. Bovidae gen. et sp. indet. 6 is rel-
without crenulations; labial styles and ribs moderately atively rare in the fossil assemblage from the Chaing-
projected; entostyle robust; a small fossetta usually zauk area (Table 9).
present at a moderate wear stage. Lower premolars not
compressed antero-posteriorly; branching between Bovidae gen. et sp. indet. 7 (large-sized)
anterior stylid and anterior conid indistinct; anteri- (Plate 9, Figs 8 –10)
or and posterior valleys on P4 always open lingually;
mesolingual conid simple, without clear cristids to the Locality and age of the examined material: Gwebin area
anterior and posterior directions. Lower molars hyp- (Gbn, TBG): late Pliocene.
sodont; mesostylid weaker and slender; lingual sur- Examined material: Listed in Appendix 2.
face flat, with faintly projected ribs; on M3 entostylid Measurements: Shown in Table 12.
strong. Moreover, the third lobe on M3 connects to Description: A few molar fragments (e.g. IR 0003 (2)
the postero-labial wall at the posthypocristid, forming and IR 0016) from Gwebin area belong to the largest
a deep fold behind the entostylid. species of all Irrawaddy bovids (Tables 12), but most
Remarks: The largest species of all bovids from enamel parts are broken. Apart from those specimens,
Chaingzauk area is Pachyportax giganteus. The tooth the large-sized species from the Gwebin area are similar
pattern of Bovidae gen. et sp. indet. 6 is similar to that to those from Chaingzauk area in both size and shape.
of Bovidae gen. et sp. indet. 4 or 5. In addition to size A remarkable difference is found in the extent of the
resemblance, some large-sized upper molars from the labial ribs of the upper molars. The large-sized species
Chaingzauk area are similar to those of the holotype from Gwebin area have almost flat labial surfaces es-
(GSI B219: right M3) of Pachyportax latidens from the pecially on the posterior lobe. They are also character-
Siwaliks, in having relatively flat labial surfaces with ised by an antero-posteriorly elongated postparacrista,
eschweizerbart_xxx

a large fossetta, and a weak (or no) metaconule fold. The early late Miocene fauna is reconstructed by
Remarks: In the fossil assemblage from Gwebin area, the fossil assemblage from the Yenangyaung area, in-
Proleptobos birmanicus is the largest species, based on cluding Helicoportax sp. and Selenoportax vexillarius.
skull and horn-core sizes, but its teeth are significantly The bovids from the Yenangyaung area are larger and
smaller than those of Bovidae gen. et sp. indet. 7 (Ta- more hypsodont than those from Chaungtha, based
ble 12). Therefore, there were at least three species of on comparing dental materials from the two areas,
bovids in the Gwebin fauna. Bovidae gen. et sp. indet. although small-sized teeth compared with Helicopor-
7 is relatively rare in the fossil assemblage from the tax have not been found from the Yenangyaung area.
Gwebin area (Table 9). There was possibly a somewhat faunal change from
brachydont species to hypsodont species around the
5. Discussion middle–late Miocene boundary. The appearance of
Helicoportax is restricted to a short range in a part of
The Neogene Irrawaddy fauna has been tradition- the Chinji Formation (c. 12.3 –11.8 Ma) (Gentry et
ally divided into two biostratigraphic units, Lower al. 2014). However, the fossil assemblage from Yenang-
and Upper Irrawaddy (e.g. Colbert 1938). How- yaung is characterised by the presence of Hipparion
ever, recent studies have suggested that the Lower Ir- whose first occurrence is dated to 10.7 Ma in Pakistan
rawaddy fauna is strictly subdivided by different fossil (Barry et al. 2002). This time is 1 to 2 Ma younger
assemblages, based on biostratigraphic comparisons than the occurrence range of Helicoportax species in
with Mio-Pliocene faunas in Eurasia (Zin-Maung- the Siwalik fauna.
Maung-Thein et al. 2011, Chavasseau et al. 2013, The bovid skull and teeth fossils from the Chaing-
see also Text-Fig. 2). As we mentioned in sections zauk area show an explosive increase and diversifica-
2 and 4, a total of ten species of bovids were recov- tion of the stem lineage of Bovini, including Seleno-
ered from the Irrawaddy beds of basal Yenangyaung portax and Pachyportax, in the latest Miocene or early
(c. 10.7– 8 Ma), Chaingzauk (c. 6 – 4 Ma), Gwebin (c. Pliocene. In the Siwalik fauna, the earliest species,
4 – 3 Ma), and the others (horizon unknown) (Table Selenoportax vexillarius, occurred from 10.2 to 9.8 Ma,
13). Moreover, Chavasseau et al. (2013) reported after which this species was replaced by S. falconeri
the late middle Miocene fauna from Chaungtha. The around 9.3 to 8.9 Ma and S. giganteus (P. giganteus in
bovid fossils from this locality contain at least a small this study) around 8.5 to 7.3 Ma (Gentry et al. 2014).
and low-crowned form of bovids (Chavasseau et The mammalian fauna from the Chaingzauk area is
al. 2006), but such brachydont species did not ap- composed dominantly of Hexaprotodon and Sivachoe-
pear in the fossil assemblages after the age of the basal rus which appeared in Asia after 7 Ma (Zin-Maung-
Yenangyaung fauna. Maung-Thein et al. 2011). However, in Myanmar
the first appearance datum of Selenoportax vexillar-
ius in the Irrawaddy beds is close in age to that in the
Table 13. Locality and stratigraphic distribution of ten bovid Siwalik fauna, based on the fossil assemblage from
species described in this study. Yenangyaung area, and the last occurrence of Seleno-
portax and Pachyportax is much younger than in the
Locality unknown
Yenangyaung area
Siwaliks, namely the latest Miocene or early Pliocene.

Chaingzauk area
Moreover, the fossil assemblage from the Chaingzauk

Gwebin area
Bagan area
area suggest that the three species (S. vexillarius, S. fal-

coneri, and P. giganteus) coexisted in central Myanmar
during this age. Similarly, the latest Miocene/early
Caprinae gen. et sp. indet. – + – – – Pliocene Irrawaddy fauna includes Tragoportax that
Sivaportax dolabella – + + – – occurred from 8.4 to 6.4 Ma in the Dhok Pathan For-
Helicoportax sp. + – – – –
Tragoportax sp. – + – – –
mation of the Siwaliks (Gentry et al. 2014). Previous
Duboisia sp. – – – + – studies have suggested that the Neogene mammalian
Selenoportax vexillarius + + – – – fauna in central Myanmar is biostratigraphically re-
Selenoportax falconeri – + – – – lated to the Siwalik fauna (e.g. Colbert 1938), but
Pachyportax latidens – – – – + our present data does not support a strict correspon-
Pachyportax giganteus – + – – – dence of bovid biostratigraphy between continental
Proleptobos birmanicus – + + – – Southeast Asia and South Asia.
eschweizerbart_xxx

Although many bovid fossils have been collected cent studies (Chit Sein et al. 2009, Zin-Maung-
from the upper Pliocene in the Gwebin area, only two Maung-Thein et al. 2011), and we demonstrated
endemic species, Sivaportax dolabella and Proleptobos morphological difference between these genera or
birmanicus, have been confirmed by skulls and horn species, based on skull and horn-core materials. In the
cores. These species occur throughout the period of present paper, we firstly described Caprinae gen. et sp.
the Chaingzauk and Gwebin faunas, namely from indet., Helicoportax sp., and Duboisia sp. from the Ir-
the latest Miocene through late Pliocene. We cannot rawaddy beds.
determine the exact locality and age of the specimen The species compositions of the Irrawaddy bo-
of Duboisia sp. (BAM 558/99), which is endemic to vids changed through three chronological stages: early
Southeast Asia. The appearance of the endemic taxa late Miocene (Yenangyaung fauna), latest Miocene/
in Southeast Asia possibly corresponded to the frag- early Pliocene (Chaingzauk fauna), and late Pliocene
mentation from the Siwalik fauna after the latest Mio- (Gwebin fauna) (Table 2). Early bovines, including
cene or Pliocene. Currently there are biogeographical Helicoportax and Selenoportax vexillarius, dispersed
barriers between the Siwaliks and Myanmar, such across the Indo-Burma region by the early late Mio-
as high mountain ranges and rivers, along the west cene. The latest Miocene/early Pliocene Chaingzauk
boundary of Myanmar (Corbet & Hill 1992). The fauna is composed of a mixture of Irrawaddy-domestic
palaeontological and geological data supported the species (Sivaportax dolabella and Proleptobos birman-
formation of these barriers, or the uplifting of the In- icus) and relict species (Tragoportax sp., Selenoportax
do-Burma Ranges, affecting the Pliocene mammals in vexillarius, Selenoportax falconeri, Pachyportax lati-
central Myanmar (Licht et al. 2015; Nishioka et al. dens, Pachyportax giganteus) that possibly dispersed
2015; Takai et al. 2016). As we mentioned in the de- from the Siwaliks during the late Miocene, suggesting
scriptions, the Irrawaddy endemic genera (Sivaportax, the beginning of faunal separation between central
Proleptobos, and Duboisia) are closely related to the Myanmar and South Asia. After the late Pliocene, only
Siwalik genera (Strepsiportax, Proamphibos/Leptobos, Irrawaddy-domestic species were survived in central
and Boselaphus), respectively. Geographical isolation Myanmar, possibly because of the development of bio-
between continental Southeast Asia and South Asia in geographical barriers at the Indo–Burma boundary,
the late Neogene probably resulted in the divergence which caused faunal isolation between central Myan-
of common ancestors of each lineage, and influenced mar and Indo-Pakistan. Although the fossil-bearing
the survival time of Selenoportax and Pachyportax be- horizon of Duboisia sp. is not clear, the record in cen-
tween those areas. tral Myanmar may suggest a faunal interchange be-
tween continental Southeast Asia and the Indonesian
6. Conclusions islands during the late Pliocene or Pleistocene.
The skull and horn-core fossils of bovids from the up-
per Neogene Irrawaddy beds consist of one caprine Acknowledgements
(Caprinae gen. et sp. indet.) and nine bovine species We thank to personnel from the Ministry of Religious Affairs
(Sivaportax dolabella, Helicoportax sp., Tragoportax and Cultural, Myanmar for providing the permission for field
research and assistance. We thank specially to P. Jintasakul
sp., Duboisia sp., Selenoportax vexillarius, Selenopor-
(Nakhon Ratchasima Rajabhat University, Nakhon Ratchasi-
tax falconeri, Pachyportax latidens, Pachyportax gigan- ma, Thailand), S. Juan-Ngam (Sukhothai Museum of Natural
teus, and Proleptobos birmanicus). The isolated teeth History, Sukhothai, Thailand), and P. Vachajitpan (Bang-
and tooth rows in the examined bovid fossils are in- kok, Thailand) for their kindness of providing the ONG and
determinate taxonomically, but they supported the PRY collections for our study. Comparative material referred
existence of small-sized species (tragoportacins/bose- in this study was observed in American Museum of Natural
laphins) and medium- to large-sized species (stem bo- History (New York, USA), Smithsonian Institute National
vins) in the Irrawaddy bovids. Museum of Natural History (Washington, D.C., USA), The
Comparing with previous reports on Irrawaddy Natural History Museum (London, UK), Indian Museum,
Geological Survey of India (Kolkata, India), Panjab Univer-
bovid fossils (Pilgrim 1939, Colbert 1943), we sity (Chandigarh, India), and Museum Geologi Bandung
concur with the presence of Sivaportax dolabella, Pro- ( Java, Indonesia). We appreciate the great help of all members
leptobos birmanicus, and Pachyportax latidens. The of the Myanmar-Japan Palaeontological Expedition Team,
existence of Tragoportax and Selenoportax species in who helped us to arrange the field research and collect fos-
the Irrawaddy faunas have been also indicated by re- sil specimens. The present study was financially supported in
eschweizerbart_xxx

part by Japan Society for the Promotion of Science KAKEN- Chavasseau, O., Kyaw, A.A., Chaimanee, Y., Coster,
HI, Grant-in-Aid for Scientific Research to MT [20405015, P., Emonet, E.-G, Soe, A.N., Rugumbrung, M., Tun,
26304019], NE [15K05330], and YN [16K17828]. S.T. & Jaeger, J.-J. (2013): Advances in the biochronol-
ogy and biostratigraphy of the continental Neogene of
References Myanmar. – In: Wang, X., Flynn, L.J. & Fortelius,
M. (eds): Fossil Mammals of Asia: Neogene Biostratig-
Aimi, M. & Aziz, F. (1985): Vertebrate fossils from the San-
raphy and Chronology. – Columbia University Press,
giran Dome, Mojokerto, Trinil and Sambungmacan Ar-
461– 474, New York.
eas. – In: Watanabe, N. & Kadar, D. (eds): Quater-
Chavasseau, O., Chaimanee, Y., Tun, S.T., Soe, A.N.,
nary Geology of the Hominid Fossil Bearing Formations
Barry, J.C., Marandat, B., Sudre, J., Marivaux, L.,
in Java. – Geological Research and Development Center,
Ducrocq, S. & Jaeger, J.-J. (2006): Chaungtha, a new
Special Publication 4: 155 –197.
Middle Miocene mammal locality from the Irrawaddy
Akhtar, M. (1995): Pachyportax giganteus, new species
(Mammalia, Artiodactyla, Bovidae) from the Dhok Formation, Myanmar. – Journal of Asian Earth Sciences
Pathan, district Chakwal, Punjab, Pakistan. – Pakistan 28: 354 – 362.
Journal of Zoology 27 (4): 337– 340. Chhibber, H.L. (1934): The Geology of Burma. – Macmil-
Bärmann, E.V. & Rössner, G.E. (2011): Dental nomencla- lan, 538 pp., London.
ture in Ruminantia: Towards a standard terminological Chit Sein (2006): Miocene–Pleistocene vertebrate fauna of
framework. – Mammalian Biology 76: 762 –768. central Myanmar with special reference to Kyauksaung-
Barry, J.C., Morgan, L.E., Flynn, L.J., Pilbeam, D., Beh- san, Tebingan, and Kyitsonbwe areas. – Unpublished
rensmeyer, A.K., Raza, S. M., Khan, I.A., Badgley, PhD thesis, 250 pp., University of Yangon, Yangon.
C., Hicks, J. & Kelley, J. (2002): Faunal and environ- Chit Sein, van der Made, J. & Rössner, G.E. (2009):
mental change in the late Miocene Siwaliks of northern New material of Propotamochoerus (Suidae, Mammalia)
Pakistan. – Palaeobiology 18: 1–71. from the Irrawaddy Formation, Myanmar. – Neues Jahr-
Barry, J.C., Behrensmeyer, A.K., Badgley, C.E., Flynn, buch für Geologie und Paläontologie – Abhandlungen
L.J., Peltonen, H., Cheema, I.U., Pilbeam, D., Lind- 251 (1): 17– 31.
say, E.H., Raza, M.S., Rajpar, A.R. & Morgan M.E. Clift, W. (1828): On the fossil remains of two new species of
(2013): The Neogene Siwaliks of the Potwar Plateau, Pa- Mastodon and of other vertebrate animals found on the
kistan. – In: Wang, X., Flynn, L.J. & Fortelius, M. left bank of the Irawadi. – Geological Society of Lon-
(eds): Fossil Mammals of Asia: Neogene Biostratigraphy don, 2nd serie 2: 369 – 376, plates 36 – 44.
and Chronology. – Columbia University Press, 461– 474, Colbert, E.H. (1938): Fossil mammals from Burma in the
New York. American Museum of Natural History. – Bulletin of the
Bender, F. (1983): Geology of Burma. – Gebrüder Borntrae- American Museum of Natural History 74: 255 – 436.
ger, 293 pp., Berlin. Colbert, E.H. (1943): Pleistocene vertebrates collected in
Bibi, F. (2007): Origin, paleoecology, and paleobiogeography Burma by the American Southeast Asiatic Expedition. –
of early Bovini. – Palaeogeography, Palaeoclimatology, Transactions of the American Philosophical Society, new
Palaeoecology 248: 60 –72. serie 32: 395 – 429, plates 19 – 32.
Bibi, F. (2009): Evolution, systematics, and paleoecology of Corbet, G.B. & Hill, J.E. (1992): The Mammals of the
Bovinae (Mammalia: Artiodactyla) from the late Mio- Indomalayan Region: A Systematic Review. – Oxford
cene to the recent. – Unpublished PhD thesis, 469 pp., Univ. Press, 488 pp., Oxford.
Yale University, New Haven. Dubois, E. (1891): Voorloopig bericht omtrent het onder-
Bibi, F. (2013): A multi-calibrated mitochondrial phylogeny zoek naar de Pleistocene en Tertiaire vertebraten-fauna
of extant Bovidae (Artiodactyla, Ruminantia) and the van Sumatra en Java, gedurende het jaar 1890. – Natu-
importance of the fossil record to systematics. – BMC urkundig tijdschrift voor Nederlandsch Indië 519:
Evolutionary Biology 13: 166 (15 pp.). 3 –100.
Bibi, F. & Güleç, E.S. (2008): Bovidae (Mammalia: Artio- Duvernois, M. (1990): Les Leptobos (Mammalia, Artio-
dactyla) from the late Miocene of Sivas, Turkey. – Jour- dactyla) du Villafranchien d’Europe occidentale: systé-
nal of Vertebrate Paleontology 28 (2): 501– 519. matique, évolution, biostratigraphie, paléoéocologie. –
Bibi, F., Bukhsianidze, M., Gentry, A.W., Geraads, D., Documents des laboratoires de géologie, Lyon, no. 113:
Kostopoulos, D.S. & Vrba, E.S. (2009): The fossil 1– 213.
record and evolution of Bovidae: State of the field. – Pal- Egi, N., Thaung-Htike, Zin-Maung-Maung-Thein,
aeontologia Electronica 12.3.10A: 11 pp. Maung-Maung, Nishioka, Y., Tsubamoto, T., Og-
Buckland, W. (1829): Geological account of a series of ino, S. & Takai, M. (2011): A mongoose remain (Mam-
animal and vegetable remains and of rocks collected by malia: Carnivora) from the Upper Irrawaddy sediments,
T. Crawfurd Esq. on a voyage up the Irrawadi to Ava in Myanmar and its significance in evolutionary history of
1826/29. – Geological Society of London, 2nd serie, 2: Asian herpestids. – Journal of Asian Earth Sciences 42:
377– 392. 1204 –1209.
eschweizerbart_xxx

Gaudry, A. (1861) Résultats des fouilles entreprises en Grèce Hassanin, A., Delsuc, F., Ropiquet, A., Hammer, C.,
sous les auspices de l’Académie. – Comptes rendus heb- Jansen van Vuuren, B., Matthee, C., Ruiz-Gar-
domadaires des séances de l’Académie des sciences 52: cia, M., Catzeflis, F., Areskoug, V., Nguyen, T.T.
238 – 241, 297– 299. & Couloux, A. (2012): Pattern and timing of diversi-
Gentry, A.W. (1970): The Bovidae (Mammalia) of the Fort fication of Cetartiodactyla (Mammalia, Laurasiatheria),
Ternan fossil fauna. – In: Leakey, L.S.B. & Savage, as revealed by a comprehensive analysis of mitochondrial
R. J. G. (eds): Fossil Vertebrates of Africa 2. – Academic genomes. – Comptes Rendus Biologies 335 (1): 32 – 50.
Press, 243 – 323, plates 1–7, London. Hooijer, D.A. (1958): Fossil Bovidae from the Malay Ar-
Gentry, A.W. (1971): The earliest goats and other antelopes chipelago and the Punjab. Zool. Verh. Mus. Leiden 38:
from the Samos Hipparion fauna. – Bulletin of the British 1–112.
Museum Natural History (Geology) 20 (6): 231– 296. Hooijer, D.A. (1962): Report upon a collection of Pleisto-
Gentry, A.W. (1974): A new genus and species of Pliocene cene mammals from tin-bearing deposits in a limestone
boselaphine (Bovidae, Mammalia) from South Africa. – cave near Ipoh, Kinta Valley, Perak. – Federation Mu-
Annals of the South African Museum 65: 145 –188. seum Journal 7: 1– 5.
Gentry, A.W. (2016): A new bovine (Bovidae, Artiodactyla) Ji, H.-X. (1975): The lower Pleistocene mammalian fossils of
from the Hadar Formation, Ethiopia. – Transactions of Lantian district, Shensi. – Vertebrata PalAsiatica 13 (3):
the Royal Society of South Africa 61 (2): 41– 50. 169 –177 (in Chinese, English summary).
Gentry, A.W. & Gentry, A. (1978): Fossil Bovidae (Mam- Koenigswald, G.H.R. von (1933): Beitrag zur Kenntniss
malia) of Oldvai Gorge, Tanzania, part I. – Bulletin of der fossilen Wirbeltiere Javas, I. Teil. – Wetenschappeli-
the British Museum (Natural History), Geology 29: jke Mededeelingen 23: 1–127.
289 – 446. Koenigswald, G.H.R. von (1934): Zur Stratigraphie
Gentry, A.W., Rössner, G.E. & Heizmann, E.P.J. (1999): des javanischen Pleistocän. – De Ingenieur in Neder-
Suborder Ruminantia. – In: Rössner, G.E. & Heissig, landsch-Inië 1 (4): 185 – 201.
K. (eds): Land Mammals of Europe. – Verlag Dr. Frie- Licht, A., Reisberg, L., France-Lanord, C., Soe, A.N.
drich Pfeil, 225 – 258, München. & Jaeger, J.-J. (2015): Cenozoic evolution of the cen-
Gentry, A.W., Solounias, N. & Barry, J.C. (2014): Sta- tral Myanmar drainage system: insights from sediment
provenance in the Minbu Sub-Basin. – Basin Research
bility in higher level taxonomy of Miocene bovid faunas
(2015): 1–15.
of the Siwaliks. – Annales Zoologici Fennici 51: 49 – 56.
Lydekker, R. (1876): Notes on the fossil mammalian fauna
Geraads, D. (1979): Un nouveau Bovidé (Artiodactyla,
of India and Burma. – Records of the Geological Survey
Mammalia) du Plio-Pléistocène de Java: Duboisia (?) sar-
of India 9: 86 –106.
tonoi n. sp. – Bulletin du Muséum National d’Histoire
Lydekker, R. (1878): Notices of Siwalik mammals. – Re-
Naturelle, Sections B 1 (4): 371– 376. cords of the Geological Survey of India 11: 64 –104.
Geraads, D. (1992): Phylogenetic analysis of the tribe Bo- Lydekker, R. (1884): Indian Tertiary and post-Tertiary ver-
vini (Mammalia: Artiodactyla). – Zoological Journal of tebrata. Rodents and new ruminants from the Siwaliks,
the Linnean Society 104: 193 – 207. and synopsis of Mammalia. – Memoirs of the Geological
Geraads, D. & Güleç, E. (1999): On some spiral-horned Survey of India 10 (3): 105 –134.
antelopes (Mammalia: Artiodactyla: Bovidae) from the Lydekker, R. (1885): Catalogue of the remains of Siwalik
Late Miocene of Turkey, with remarks on their distribu- Vertebrata contained in the Geological Department of
tion. – Paläontologische Zeitschrift 73: 403 – 409. the Indian Museum, Calcutta. Part I, Mammalia. – Su-
Gray, J.E. (1821): On the natural arrangement of vertebrose perintendent of Government Printing, 116 pp., Calcutta.
animals. – London Medical Repository 15: 296 – 310. Lydekker, R. (1886): Siwalik Mammalia, supplement I. –
Gromolard, C. (1980): Une nouvelle interprétation des Palaeontologica Indica 10 (4): 1– 21, plates 1– 6.
grands Bovidae (Artiodactyla, Mammalia) du Pliocène Moe-Nyunt (1987): Geology and vertebrate fossils of Gwe-
d’Europe occidentale classés jusqu’à présent dans le genre bin area, Seikpyu Township. – Unpublished MSc thesis,
Parabos: Parabos cordieri (De Christol) nov. émend., Pa- 253 pp., University of Yangon (= University of Ran-
rabos? boodon (Gervais) et Alephis lyrix nov. gen. nov. sp. goon), Yangon.
– Geobios 13 (5): 767–775. Myanmar Geosciences Society (2014): Geological Map
Haile-Selassie, Y., Vrba, E.S. & Bibi, F. (2009): Chapter of Myanmar. – Myanmar Geosciences Society, Yangon.
9 – Bovidae. – In: Haile-Selassie, Y. & Wolde Ga- Nishioka, Y., Takai, M., Nishimura, T., Thaung-
briel, G. (eds): Ardipithecus kadabba – Late Miocene Htike, Zin-Maung-Maung-Thein, Egi, N., Tsub-
Evidence from the Middle Awash, Ethiopia. – University amoto, T. & Maung-Maung. (2015): Late Pliocene
of California Press, 277– 330,California. rodents from the Irrawaddy sediments of central Myan-
Hassanin, A. & Douzery, E.J.P. (1999): The tribal radia- mar and their palaeogeographical significance. – Journal
tion of the family Bovidae (Artiodactyla) and evolution of Systematic Palaeontology 13: 287– 314.
of the mitochondrial cytochrome b gene. – Molecular Nishioka, Y., Zin-Maung-Maung-Thein, Egi, N., Tsu-
Phylogenetics and Evolution 13 (2): 227– 243. bamoto, T., Nishimura, T., Ito, T., Thaung-Htike
eschweizerbart_xxx

& Takai, M. (2011): New Hystrix (Mammalia, Roden- of the evolutionary history of cercopithecid monkeys in
tia) from the late Miocene/early Pliocene of Myanmar. Southeast Asia. – Historical Biology 28 (1-2): 171–187.
– Journal of Vertebrate Paleontology 31 (4): 919 – 924. Takai, M., Thaung-Htike, Zin-Maung-Maung-
Noetling, F. (1895): The development and sub-division of Thein, Soe, A.N., Maung-Maung, Tsubamoto,
the Tertiary system in Burma. – Records of the geologi- T., Egi, N., Nishimura, T.D. & Nishioka, Y. (2015):
cal survey of India 28: 59 – 86. First discovery of colobine fossils from the Late Mio-
Ogino, S., Egi, N., Zin-Maung-Maung-Thein, cene/Early Pliocene of central Myanmar. – Journal of
Thaung-Htike & Takai, M. (2011): New species of Human Evolution 84: 1–15.
Agriotherium (Mammalia, Carnivora) from the late Mi- Thaung-Htike (2008): Paleontological analysis of Suidae
ocene to early Pliocene of central Myanmar. – Journal of and Hippopotamidae (Mammalia, Artiodactyla) from
Asian Earth Sciences 42: 408 – 414. the Neogene of Central Myanmar. – Unpublished PhD
Pilgrim, G.E. (1910): Notices of new mammalian genera thesis, Kyoto University, 171 pp., Kyoto.
and species from the Tertiaries of India. – Records of the Thaung-Htike (2012): Review on the taxonomic status of
geological survey of India 40: 63 –71. Hexaprotodon iravaticus (Mammalia, Artiodactyla, Hip-
Pilgrim, G.E. (1913): The correlation of the Siwaliks with popotamidae) from the Neogene of Myanmar. – Shwebo
the mammal horizons of Europe. Records of the geologi- University Research Journal 3: 94 –110.
cal survey of India 43: 264 – 326. Thaung-Htike, Zin-Maung-Maung-Thein & Hnin-
Pilgrim, G.E. (1937): Siwalik antelopes and oxen in the Hnin-Htay (2008): New materials of Tetraconodon
American Museum of Natural History. – Bulletin of the malensis (Mammalia, Artiodactyla, Suidae) from the
American Museum of Natural History 72 (7): 729 – 874. Middle Miocene of Central Myanmar. – Universities
Pilgrim, G.E. (1939): The fossil Bovidae of India. – Palaeon- Research Journal 1: 65 –76.
tologica Indica 26: 1– 356. Thaung-Htike, Tsubamoto, T., Takai, M., Egi, N.,
Ridd, M.F. & Racey, A. (2016): Onshore petroleum geology Zin-Maung-Maung-Thein, Chit-Sein & Maung-
of Myanmar: Central Burma Depression. – Geological Maung (2006): Discovery of Propotamochoerus (Artio-
Society of London Memoirs 45: 21– 50. dactyla, Suidae) from the Neogene of Myanmar. – Asian
Roussiakis, S.J. (2009): Prostrepsiceros and Protragelaphus Paleoprimatology 4: 173 –185.
Tsubamoto, T., Thaung-Htike, Zin-Maung-Maung-
(Artiodactyla, Mammalia) from the Late Miocene local-
Thein, Egi, N., Nishioka, Y., Maung-Maung &
ity of Chomateri (Attica, Greece). – Annales de Paléon-
Takai, M. (2012): New data on the Neogene anthra-
tologie 95: 181–195.
cotheres (Mammalia; Artiodactyla) from central Myan-
Shi, Q.-Q., He, W. & Chen S.-Q. (2014): A new species of
mar. – Journal of Vertebrate Paleontology 32: 956 – 964.
Shaanxispira (Bovidae, Artiodactyla) from the upper
Tsubamoto, T., Zin-Maung-Maung-Thein, Thaung-
Miocene of China. – Zootaxa 3794 (4): 501– 513. Htike, Egi, N., Chit-Sein, Maung-Maung &
Solounias, N. (1981): The Turolian fauna from the island Takai, M. (2006): Discovery of chalicothere and Dor-
of Samos, Greece. – Contributions to Vertebrate Evolu- cabune from the upper part (lower Pleistocene) of the
tion 6: 1– 232. Irrawaddy Formation, Myanmar. – Asian Paleoprimatol-
Solounias, N., Barry, J.C., Bernor, R.L., Lindsay, E.H. ogy 4: 137–142.
& Raza, S.M. (1995): The oldest bovid from the Siwa- Vrba, E.S. (1987): A revision of the Bovini (Bovidae) and
liks, Pakistan. – Journal of Vertebrate Paleontology 15: a preliminary revised checklist of Bovidae from Maka-
806 – 814. pansgat. – Palaeontologia Africana 26 (4): 33 – 46.
Stamp, L.D. (1922): An outline of the Tertiary geology of Wagner, A. (1848) Urweltliche Säugethier-Überreste aus
Burma. – Geological Magazine 59: 481– 501. Griechenland. – Abhandlungen der Bayerischen Akada-
Stremme, H. 1911. Die Säugetiere mit Ausnahme der Pro- demie der Wissenschaften 5: 333 – 378.
boscidier. – In: Selenka, L. & Blanckenhorn, M. Watabe, M. (1990): Fossil bovids (Artiodactyla, Mammalia)
(eds): Die Pithecanthropus-Schichten auf Java. – Wil- from Maragheh (Turolian, late Miocene), Northwest
helm Engelmann, 82 –150, plates 16-20, Leipzig. Iran. – Annual Report of the Historical Museum of
Spassov, N. & Geraads, D. (2004): Tragoportax Pilgrim, Hokkaido 18: 19 – 56.
1937 and Miotragocerus Stromer, 1928 (Mammalia, Bovi- Zhang, Z.-Q. (2003): A new species of Shaanxispira (Bovi-
dae) from the Turolian of Hadjidimovo, Bulgaria, and a dae, Artiodactyla, Mammalia) from the Bahe Formation,
revision of the late Miocene Mediterranean Boselaphini. Lantian, China. – Vertebrate PalAsia 41: 230 – 239.
– Geodiversitas 26 (2): 339 – 370. Zin-Maung-Maung-Thein (2010): Plaeoenvironmental
Takai, M., Saegusa, H., Thaung-Htike & Zin-Maung- analysis of the Chaingzauk mammalian fauna (late Neo-
Maung-Thein (2006): Neogene mammalian fauna in gene, Myanmar) using stable isotopes of tooth enamel.
Myanmar. – Asian Paleoprimatology 4: 143 –172. – Unpublished PhD thesis, 78 pp., Kyoto Univ., Kyoto.
Takai, M., Nishioka, Y., Thaung-Htike, Maung- Zin-Maung-Maung-Thein, Takai, M., Tsubamoto, T.,
Maung, Kyaw-Khaing, Zin-Maung-Maung- Thaung-Htike, Egi, N. & Maung, M. (2008): A new
Thein, Tsubamoto, T. & Egi, N. (2016): Late Pliocene species of Dicerorhinus (Rhinocerotidae) from the Plio-
Semnopithecus fossil from central Myanmar: rethinking Pleistocene of Myanmar. Palaeontology 51: 1419 –1433.
eschweizerbart_xxx

Zin-Maung-Maung-Thein, Thaung-Htike, Tsubamoto, from the Irrawaddy sediments. – Journal of Asian Earth
T., Takai, M., Egi, N. & Maung-Maung (2006): Sciences 37: 105 – 206.
Early Pleistocene Javan rhinoceros from the Irrawaddy Zin-Maung-Maung-Thein, Takai, M., Uno, H., Wynn,
Formation, Myanmar. – Asian Paleoprimatology 4: J.G., Egi, N., Tsubamoto, T., Thaung-Htike, Soe,
197– 204. A.N., Maung, M., Nishimura, T. & Yoneda, M.
Zin-Maung-Maung-Thein, Takai, M., Tsubamoto, T., Egi, (2011): Stable isotope analysis of the tooth enamel of
N., Thaung-Htike, Nishimura, T., Maung, M. & Chaingzauk mammalian fauna (late Neogene, Myan-
Win, Z. (2010): A review of fossil rhinoceroses from the mar) and its implication to paleoenvironment and pale-
Neogene of Myanmar with description of new specimens ogeography. – Palaeogeology Palaeoclimatology Palaeo-
ecology 300: 11– 22.
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Explanation of the plates
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Plate 1
Fig. 1. Caprinae gen. et sp. indet., IR 3138, right horn core from Chaingzauk area (MKT); anterior view.
Fig. 2. Ditto; posterior view.
Fig. 3. Ditto; cross section at the bottom.
Fig. 4. Sivaportax dolabella, IR 1272, skull cap from Chaingzauk area (MKT); left lateral view.
Fig. 5. Ditto; dorsal view.
Fig. 6. Ditto; schematic drawing in dorsal view
Fig. 7. Sivaportax dolabella, IR 3533, right horn core from Chaingzauk area (MKT); antero-dorsal view.
Fig. 8. Ditto; anterior view.
Fig. 9. Ditto; lateral view.
Fig. 10. Sivaportax dolabella, IR 1824, left horn core from Chaingzauk area (MKT); antero-dorsal view.
Fig. 12. Ditto; medial view.
Fig. 14. Ditto; cross section of the top.
Fig. 15. Sivaportax dolabella, IR 1576, left horn core from Gwebin area (TBG); anterior view.
Fig. 20. Helicoportax sp., IR 4067, left horn core from Yenangyaung area (NYL); antero-dorsal view.
Fig. 23. Tragoportax sp., IR 3593, left horn core from Chaingzauk area (PAL); anterior view.
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Plate 1. Nishioka
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Plate 2
Fig. 1. Duboisia sp., BAM 558/99, neurocranium with right and left horn-core bases from Bagan; left lateral view.
Fig. 4. Selenoportax vexillarius, IR 4515, right horn core from Yenangyaung area (NYD); anterior view.
Fig. 6. Selenoportax vexillarius, IR 3693, frontal with right and left horn cores from Chaingzauk (MKT); right lateral view.
Fig. 8. Ditto; antero-dorsal view.
Fig. 9. Ditto; cross section of the right horn-core base.
Fig. 10. Selenoportax vexillarius, IR 3907, left horn core from Chaingzauk area (MKT); lateral view.
Fig. 13. Ditto; cross section of the base.
Fig. 14. Selenoportax vexillarius, IR 3906, left horn core from Chaingzauk area (MKT); lateral view.
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Plate 2. Nishioka
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Plate 3
Fig. 1. Selenoportax falconeri, PRY 204, skull with right and left horn cores and teeth from central Myanmar; left lateral view.
Fig. 4. Ditto; cross section of the left horn-core base.
Fig. 5. Selenoportax falconeri, CCZ 450, neurocranium with right and left horn cores from central Myanmar; left lateral view.
Fig. 7. Ditto; cross section of the left horn-core base.
Fig. 8. Selenoportax falconeri, IR 3905, right horn core from Chaingzauk area (MKT); anterior view.
Fig. 12. Selenoportax falconeri, IR 2306, neurocranium from Chaingzauk area (MKT); right lateral view.
Fig. 15. Ditto; ventral view.
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Plate 3. Nishioka
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Plate 4
Fig. 1. Pachyportax latidens, ONG 604, neurocranium from central Myanmar; right lateral view.
Fig. 4. Pachyportax latidens, resin cast of NM-YNG 004, right horn core possibly from Magway; lateral view.
Fig. 8. Pachyportax giganteus, MGW 063, neurocranium with right horn core possibly from Magway; right lateral view.
Fig. 13. Ditto; cross section of the right horn-core base.
Fig. 14. Pachyportax giganteus, resin cast of NPT 2015/01/2356, left horn core from central Myanmar; lateral view.
Fig. 18. Pachyportax giganteus, IR 3691, left horn core from Chaingzauk area (MKT); lateral view.
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Plate 4. Nishioka
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Plate 5
Fig. 1. Proleptobos birmanicus, PRY 205, neurocranium with right and left horn cores from central Myanmar; antero-dorsal
view.
Fig. 2. Ditto; postero-dorsal view.
Fig. 3. Proleptobos birmanicus, CCZ 449, skull with right and left horn cores and teeth from central Myanmar; left lateral view.
Fig. 5. Proleptobos birmanicus, CCZ 447, skull with teeth from central Myanmar; left lateral view.
Fig. 7. Ditto; cross section of the left horn pedicel.
Fig. 8. Proleptobos birmanicus, BAM 555/99, neurocranium from central Myanmar; left lateral view.
Fig. 12. Proleptobos birmanicus, IR 1456, right maxilla from Gwebin area (TBG), associated with Fig. 13; lingual view.
Fig. 13. Proleptobos birmanicus, IR 1456, neurocranium from Gwebin area (TBG), associated with Fig. 12; left lateral view.
Fig. 15. Proleptobos birmanicus, IR 1080, left horn core from Chaingzauk area (MKT); medial view.
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Plate 5. Nishioka
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Plate 6
Fig. 1. Selenoportax falconeri, PRY 204, skull with right P2 –M3 and left P3 –M3 from central Myanmar; occlusal view.
Fig. 2. Ditto; left lateral view (labial view of the left upper tooth row).
Fig. 3. Proleptobos birmanicus, CCZ 447, skull with right P3 –M3 and left P4 –M3 from central Myanmar; occlusal view.
Fig. 4. Ditto; labial view of the left upper tooth row.
Fig. 5. Proleptobos birmanicus, CCZ 449, skull with right and left M2 –M3 from central Myanmar; occlusal view.
Fig. 6. Proleptobos birmanicus, IR 1456, right maxilla with P2 –M3 from Gwebin area (TBG); occlusal view.
Fig. 7. Ditto; labial view.
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Plate 6. Nishioka
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Plate 7
Fig. 1. Bovidae gen. et sp. indet. 1 (small-sized), IR 2684, right P4 from Chaingzauk area (MKT); occlusal view.
Fig. 2. Bovidae gen. et sp. indet. 1 (small-sized), IR 2788, left M1/M2 from Chaingzauk area (MKT); occlusal view.
Fig. 3. Bovidae gen. et sp. indet. 1 (small-sized), IR 3721, right M3 from Chaingzauk area (MKT); occlusal view.
Fig. 4. Bovidae gen. et sp. indet. 1 (small-sized, morphotype S2), IR 1729 (13), right mandible with P4 –M3 from Chaingzauk
area (MKT); occlusal view.
Fig. 5. Bovidae gen. et sp. indet. 1 (small-sized, morphotype S2), IR 1096, right mandible with M1–M3 from Chaingzauk area
(MKT); occlusal view;
Fig. 7. Ditto; lingual view.
Fig. 8. Bovidae gen. et sp. indet. 2 (small-sized), IR 0063 (5), right P2 from Gwebin area (Gbn); occlusal view.
Fig. 9. Bovidae gen. et sp. indet. 2 (small-sized), IR 0063 (2), right maxilla with P4 –M1 from Gwebin area (Gbn); occlusal
view.
Fig. 10. Bovidae gen. et sp. indet. 2 (small-sized), IR 2567, right maxilla with M2 –M3 from Gwebin area (Gbn); occlusal view.
Fig. 11. Bovidae gen. et sp. indet. 2 (small-sized), IR 3993, right mandible with P4 –M2 from Gwebin area (Psw); occlusal view.
Fig. 12. Bovidae gen. et sp. indet. 2 (small-sized, morphotype S2), IR 2590, left M3 from Gwebin area (Psw); occlusal view.
Fig. 13. Bovidae gen. et sp. indet. 3 (medium-sized, morphotype MU1), IR 4053, right M1/M2 from Yenangyaung area (SAL);
occlusal view.
Fig. 16. Bovidae gen. et sp. indet. 3 (medium-sized), IR 4092, right mandible with P3 –P4 from Yenangyaung area (NYD);
occlusal view.
Fig. 19. Bovidae gen. et sp. indet. 3 (medium-sized), IR 4546, left mandible with M1/M2 from Yenangyaung area (NYD);
occlusal view.
Fig. 22. Bovidae gen. et sp. indet. 4 (medium-sized, morphotype ML1), resin cast of ONG 577, right mandible with P4 –M3 from
central Myanmar; occlusal view.
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Plate 7. Nishioka
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Plate 8
Fig. 1. Bovidae gen. et sp. indet. 4 (medium-sized, morphotype MU1), IR 2649, right maxilla with P2, P3, dP3, dP4, M1–M3
and left maxilla with P2, P3, dP3, dP4, M1–M2 from Chaingzauk area (MKT); occlusal view.
Fig. 2. Bovidae gen. et sp. indet. 4 (medium-sized, morphotype MU1), IR 1423, left maxilla with M1–M3 from Chaingzauk
area (CHZ); occlusal view.
Fig. 3. Bovidae gen. et sp. indet. 4 (medium-sized, morphotype ML2), IR 2184, right mandible with P2 –M3 from Chaingzauk
area (MKT); occlusal view.
Fig. 5. Bovidae gen. et sp. indet. 5 (medium-sized), IR 2564, left maxilla with P3 –P4 from Gwebin area (TBG); occlusal view
Fig. 6. Bovidae gen. et sp. indet. 5 (medium-sized, morphotype MU1), IR 4027, left M1/M2 from Gwebin area (Psw); occlusal
view.
Fig. 7. Bovidae gen. et sp. indet. 5 (medium-sized), IR 1843, left M3 from Gwebin area (TBG); occlusal view.
Fig. 10. Bovidae gen. et sp. indet. 5 (medium-sized), IR 3975, left P4 from Gwebin area (TBG); occlusal view.
Fig. 11. Bovidae gen. et sp. indet. 5 (medium-sized), IR 2045, right M3 from Gwebin area (TBG); occlusal view.
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Plate 8. Nishioka
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Plate 9
Fig. 1. Bovidae gen. et sp. indet. 6 (large-sized), IR 0749, right P2 from Chaingzauk area (CHZ); occlusal view.
Fig. 2. Bovidae gen. et sp. indet. 6 (large-sized), IR 1680, right P3 from Chaingzauk area (MKT); occlusal view.
Fig. 3. Bovidae gen. et sp. indet. 6 (large-sized), resin cast of IR 0131, left M1/M2 from Chaingzauk area (CHZ); occlusal view.
Fig. 4. Bovidae gen. et sp. indet. 6 (large-sized), IR 3891, left mandible with M1–M2 from Chaingzauk area (MKT); occlusal
view.
Fig. 5. Bovidae gen. et sp. indet. 6 (large-sized), IR 1086, right mandible with P3 –M3 from Chaingzauk area (MKT); occlusal
view.
Fig. 8. Bovidae gen. et sp. indet. 7 (large-sized), IR 1846, left M3 from Gwebin area (TBG); occlusal view.
Fig. 10. Bovidae gen. et sp. indet. 7 (large-sized), IR 0016, left upper molar from Gwebin area (Gbn); occlusal view.
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Plate 9. Nishioka
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Appendix 1. List of bovid skull and horn-core fossils from the Irrawaddy beds, examined in this study.
Caprinae gen. et sp. indet. Pachyportax latidens

No. Material Area Locality No. Material Area Locality
IR 3138 R. horn core Chaingzauk MKT NM-YNG 004 R. horn core Magway? —
ONG 444 neurocranium no data —
Sivaportax dolabella ONG 588 neurocranium no data —
No. Material Area Locality ONG 604 neurocranium no data —
IR 1272 skull cap Chaingzauk MKT1
IR 1684 R. horn core Chaingzauk MKT Pachyportax giganteus
IR 1729 (3) R. horn core Chaingzauk MKT5 No. Material Area Locality
IR 1755 (2) R. horn core Chaingzauk MKT8 IR 1840 frontal fragment Chaingzauk MKT1
IR 1824 L. horn core Chaingzauk MKT8 IR 3690 L. horn core Chaingzauk MKT
IR 1832 L. horn core Chaingzauk MKT IR 3691 L. horn core Chaingzauk MKT
IR 2136 L. horn core Chaingzauk MKT1 IR 3900 R. horn core Chaingzauk MKT
IR 2766 horn-core tip Chaingzauk MKT8 IR 3908 L. horn core Chaingzauk MKT
IR 2855 L. horn core Chaingzauk MKT neurocranium with R. horn
MGW 063 Magway? —
IR 3533 R. horn core Chaingzauk MKT7 core
IR 3740 L. horn core (base part) Chaingzauk MKT7 NMY no number neurocranium no data —
IR 1413 R. horn core Gwebin Gbn1 NPT 2015/01/2356 L. horn core no data —
IR 1576 L. horn core Gwebin TBG
IR 1766 (3) L. horn core Gwebin TBG1 Proleptobos birmanicus
IR 1897 R. horn core Gwebin TBG1 No. Material Area Locality
IR 1898 horn-core tip Gwebin TBG1 IR 0204 horn-core tip Chaingzauk CHZ15
IR 1899 horn-core tip Gwebin TBG1 IR 0212 horn-core fragment Chaingzauk CHZ15
IR 2091 horn-core fragment Gwebin TBG1 IR 0539 (2) horn-core fragments Chaingzauk CHZ181
IR 3161 horn-core fragment Gwebin TBG1 IR 0716 frontal with L horn core Chaingzauk CHZ27
IR 3313 L. horn core Gwebin Gbn7 IR 1079 L. horn core Chaingzauk MKT8
IR 1080 L. horn core Chaingzauk MKT8
Helicoportax sp. IR 1081 horn-core tip Chaingzauk MKT8
No. Material Area Locality IR 1249 horn-core fragment Chaingzauk MKT2
IR 4067 L. horn core Yenangyaung NYL2 IR 1705 (1) frontal with R. horn core Chaingzauk CHZ11
IR 4508 R. horn core Yenangyaung NYL2 IR 2156 horn-core fragment Chaingzauk MKT2
IR 4494 L. horn core Yenangyaung NYD2 IR 2293 horn-core tip Chaingzauk MKT
IR 4502 L. horn core Yenangyaung NYD2 IR 2305 horn core (middle part) Chaingzauk MKT
IR 2645 R. horn core (base part) Chaingzauk MKT2
Tragoportax sp. IR 2646 horn-core fragment Chaingzauk MKT2
No. Material Area Locality IR 2647 horn-core fragment Chaingzauk MKT2
IR 3593 L. horn core Chaingzauk PAL2 IR 2765 R. horn core (base part) Chaingzauk MKT8
IR 2796 horn-core tip Chaingzauk MKT7
Duboisia sp. IR 2844 frontal with L. horn core Chaingzauk MKT
No. Material Area Locality IR 2845 R. horn core Chaingzauk MKT
neurocranium with L.R. horn Bagan IR 2846 horn-core fragment Chaingzauk MKT
BAM 558/99 —
cores IR 2847 horn core (base part) Chaingzauk MKT
IR 2848 horn core (base part) Chaingzauk MKT
Selenoportax vexillarius IR 2849 horn-core fragment Chaingzauk MKT
No. Material Area Locality IR 2850 frontal with R. horn core Chaingzauk MKT
IR 3693 frontal with L.R. horn cores Chaingzauk MKT IR 2851 horn-core fragments Chaingzauk MKT
IR 3898 R. horn core Chaingzauk MKT IR 2853 horn-core tip Chaingzauk MKT
IR 3901 R. horn core Chaingzauk MKT IR 3044 horn-core tip Chaingzauk MKT1/2
IR 3903 L. horn core Chaingzauk MKT IR 3430 parieto-supraoccipital Chaingzauk CHZ15
IR 3906 L. horn core Chaingzauk MKT IR 3770 frontal with R. horn core Chaingzauk CHZ15
IR 3907 L. horn core Chaingzauk MKT IR 1456 skull with teeth Gwebin TBG1
IR 4515 R. horn core Yenangyaung NYD2 IR 3162 horn-core fragment Gwebin TBG1
ONG 439 frontal with horn pedicels no data — IR 3185 horn-core fragment Gwebin Gbn
IR 3312 frontal with L. horn core Gwebin Gbn7
Selenoportax falconeri PKK 17 skull with horn cores and teeth (Pakokku) —
No. Material Area Locality BAM 555/99 neurocranium no data —
IR 2306 neurocranium Chaingzauk MKT CCZ 447 skull with horn cores and teeth no data —
IR 3665 L. horn core Chaingzauk MKT CCZ 448 skull with horn cores and teeth no data —
IR 3905 R. horn core Chaingzauk MKT CCZ 449 skull with horn cores and teeth no data —
neurocranium with L.R. horn neurocranium with L.R. horn
CCZ 450 no data — PRY 205 no data —
cores cores
skull with L.R. horn cores and no data
PRY 204 —
teeth
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Appendix 2. List of bovid dental fossils from the Irrawaddy beds, examined in this study. Asterisk indicates specimens used for
measurements in Tables 7– 8 and 10 –12.
Bovidae gen. et sp. indet. 1 (small-sized) IR 3079 L. M3 Chaingzauk MKT7
No. Material Area Locality IR 3086 R. P3/P4 Chaingzauk MKT7
* IR 0525 L. mandible with P4–M2 Chaingzauk CHZ27 IR 3087 L. M3 Chaingzauk MKT7
IR 0624 R. M3 Chaingzauk CHZ27 IR 3104 L. M1/M2 Chaingzauk MKT10
* IR 0629 R. M3 Chaingzauk CHZ28 IR 3105 L. M1/M2 Chaingzauk MKT10
IR 0686 R. M3 Chaingzauk CHZ28 IR 3106 R. mandible with P4–M1 Chaingzauk MKT10
* IR 0819 R. mandible with M2–M3 Chaingzauk CHZ IR 3107 R. M3 Chaingzauk MKT10
* IR 0866 L. mandible with M2–M3 Chaingzauk CHZ33 IR 3109 R. mandible with M1–M3 Chaingzauk MKT10
* IR 0889 R. mandible with M1–M2 Chaingzauk CHZ28 IR 3468 R. upper molar Chaingzauk MKT8
* IR 0909 (1) L. mandible with M2–M3 Chaingzauk MKT4 IR 3469 upper molar Chaingzauk MKT8
* IR 0909 (2) L. M1 Chaingzauk MKT IR 3470 upper molar fragment Chaingzauk MKT8
* IR 0930 R. mandible with M1–M2 Chaingzauk MKT1 IR 3472 lower molar fragmet Chaingzauk MKT8
* IR 0936 R. mandible with P3, M2–M3 Chaingzauk MKT1 IR 3474 L. M3 Chaingzauk MKT8
* IR 0980 (1) L. M3 Chaingzauk MKT1 IR 3601 L. M1/M2 Chaingzauk CHZ22
IR 1007 L. M1/M2 Chaingzauk MKT1 * IR-3721 R. M3 Chaingzauk MKT7
* IR 1061 L. mandible with M1 Chaingzauk MKT1 * IR-3749 L. mandible with M2–M3 Chaingzauk MKT8
* IR 1096 R. mandible with M1–M3 Chaingzauk MKT8 IR 3753 L. M1 Chaingzauk MKT8
* IR 1115 (18) M2/M3 Chaingzauk MKT8 * IR-3759 R. M2 Chaingzauk MKT8
* IR 1118 R. M3 fragment Chaingzauk MKT8 * IR-3843 R. M2 Chaingzauk MKT7
IR 1729 (1) L. mandible with M1–M2 Chaingzauk MKT5 IR 3844 R. P2 Chaingzauk MKT7
* IR 1729 (13) R. mandible with P4–M3 Chaingzauk MKT5 * IR-3856 R. M2 Chaingzauk MKT7
* IR 1751 R. mandible with P4–M2 Chaingzauk MKT7 * IR-3876 L. mandible with P4–M2 Chaingzauk MKT
* IR 1757 (2) M2/M3 Chaingzauk MKT8
IR 1801 L. mandible with P4–M3 Chaingzauk MKT Bovidae gen. et sp. indet. 2 (small-sized)
* IR 1817 (1) R. M1 Chaingzauk MKT1 No. Material Area Locality
* IR 1817 (2) M2/M3 Chaingzauk MKT1 * IR 0063 (1) L. M3 Gwebin Gbn1
IR 1818 L. P2 Chaingzauk MKT1 * IR 0063 (2) R. maxilla with P4–M1 Gwebin Gbn1
IR 1819 P3 Chaingzauk MKT1 * IR 0063 (5) R. P2 Gwebin Gbn1
IR 2399 L. mandible with P4–M1 Chaingzauk MKT * IR 1354 L. M2/M3 Gwebin Gbn1
IR 2400 R. M1/M2 Chaingzauk MKT * IR 1356 R. M3 Gwebin Gbn1
IR 2401 R. M1/M2 Chaingzauk MKT * IR 1461 L. M3 Gwebin TBG1
IR 2402 L. M3 Chaingzauk MKT * IR 1464 L. M3 Gwebin TBG1
IR 2403 L. mandible Chaingzauk MKT * IR 1465 L. M3 Gwebin TBG1
IR 2404 R. mandible Chaingzauk MKT * IR 1467 R. M3 Gwebin TBG1
IR 2682 L. M1/M2 Chaingzauk MKT2 * IR 1471 L. mandible with M2 Gwebin TBG1
IR 2683 R. mandible with P2 and P4–M3 Chaingzauk MKT2 * IR 1571 (4) M1 Gwebin TBG1
IR 2684 R. P4 Chaingzauk MKT2 * IR 1572 (1) R. maxilla with P3–M3 Gwebin TBG1
IR 2685 L. M1/M2 Chaingzauk MKT2 * IR 1592 (3) R. P4 Gwebin TBG1 (South)
IR 2686 L. mandible with P4–M1 Chaingzauk MKT2 * IR 1596 (2) R. M2/M3 Gwebin TBG2
IR 2687 L. mandible with M1–M3 Chaingzauk MKT1 * IR 1615 M1 Gwebin Gbn1
IR 2688 R. P4 Chaingzauk MKT1 IR 1625 L. M1/M2 Gwebin Gbn1
IR 2689 L. M1/M2 Chaingzauk MKT1 * IR 1629 (1) L. M3 Gwebin Gbn1
IR 2690 L. M1/M2 Chaingzauk MKT1 * IR 1841 L. M3 Gwebin TBG1
IR 2691 L. M1/M2 Chaingzauk MKT1 * IR 1842 L. M3 Gwebin TBG1
IR 2692 L. M3 Chaingzauk MKT2 * IR 1845 L. M2/M3 Gwebin TBG1
IR 2693 L. M3 Chaingzauk MKT1 * IR 1849 L. M2/M3 Gwebin TBG1
IR 2694 R. M3 Chaingzauk MKT1 IR 1903 R. mandible Gwebin TBG1
IR 2695 L. mandible with M3 Chaingzauk MKT1 * IR 1909 R. M2/M3 Gwebin Psw1
IR 2737 L. M1/M2 Chaingzauk CHZ30 * IR 1912 R. M3 Gwebin Psw2
IR 2785 L. M1/M2 Chaingzauk MKT7 * IR 1948 R. M1 Gwebin Gbn1 (Mg1)
IR 2786 R. M3 Chaingzauk MKT7 * IR 1949 R. M2 Gwebin Gbn1 (Mg1)
IR 2788 L. M1/M2 Chaingzauk MKT8 * IR 1975 L. P4 Gwebin Gbn6
IR 2862 L. mandible with P3–M3 Chaingzauk MKT * IR 1976 L. P4 Gwebin Gbn6
IR 2867 R. mandible with M3 Chaingzauk MKT IR 1977 R. P4 Gwebin Gbn1 (Mg2)
IR 2983 L. M3 Chaingzauk CHZ (east1) * IR 2021 R. M3 Gwebin Gbn1
IR 3048 L. M2 Chaingzauk MKT5 IR 2022 R. M3 Gwebin Gbn1
IR 3049 R. M2 Chaingzauk MKT5 * IR 2034 M2/M3 Gwebin TBG4
IR 3050 R. M2 Chaingzauk MKT5 IR 2043 R. M2 Gwebin TBG5
IR 3053 L. M3 Chaingzauk MKT5 IR 2045 R. M3 Gwebin TBG5
IR 3059 L. mandible with P4–M1 Chaingzauk MKT5 IR 2046 R. M3 Gwebin TBG5
IR 3078 L. M2 Chaingzauk MKT7 * IR 2080 L. M1 Gwebin Psw1
eschweizerbart_xxx

IR 2457 M1/M2 Gwebin Gbn5 IR 0491 (2) L. M3 Chaingzauk CHZ180

* IR 2465 L. M3 Gwebin TBG1 * IR 0496 L. mandible with M2–M3 Chaingzauk CHZ11
IR 2466 R. M3 Gwebin TBG1 * IR 0527 L. mandible with P3–M3 Chaingzauk CHZ27
IR 2467 L. M3 Gwebin TBG1 * IR 0531 R. mandible with M2–M3 Chaingzauk CHZ15
IR 2565 L. M3 Gwebin TBG1 * IR 0535 L. mandible with P4–M2 Chaingzauk CHZ15
IR 2567 R. maxilla with M2–M3 Gwebin Gbn5 IR 0562 R. mandible with M1–M2 Chaingzauk CHZ15
IR 2570 R. M2 Gwebin Psw1 * IR 0574 R. mandible with M2–M3 Chaingzauk CHZ22/23
IR 2571 R. upper molar Gwebin Psw1 IR 0611 L. M1 Chaingzauk CHZ
IR 2572 L. M1/M2 Gwebin Psw1 * IR 0629 (1) M1/M2 Chaingzauk CHZ28
IR 2590 L. M3 Gwebin Psw1 * IR 0631 R. M3 Chaingzauk CHZ27
IR 2595 L. M3 Gwebin Gbn1 IR 0670 L. M2 Chaingzauk CHZ
IR 2597 L. M3 Gwebin Gbn1 * IR 0672 L. mandible with M3 Chaingzauk CHZ28
IR 3023 R. M1/M2 Gwebin Gbn1 * IR 0678 (1) M3 Chaingzauk CHZ29
IR 3024 L. M3 Gwebin Mg1 (Gbn1) * IR 0685 L. mandible with P4–M3 Chaingzauk CHZ29
IR 3025 L. P2 Gwebin Mg1 (Gbn1) IR 0700 R. P4 Chaingzauk CHZ
IR 3204 R. M2 Gwebin TBG4 * IR 0727 L. mandible with M1–M3 Chaingzauk CHZ29
IR 3205 L. M3 Gwebin TBG4 * IR 0729 L. mandible with M3 Chaingzauk CHZ15
* IR 3206 L. M3 Gwebin TBG4 * IR 0735 R. mandible with M2–M3 Chaingzauk CHZ29
* IR 3207 R. M1 Gwebin TBG4 IR 0742 M3 Chaingzauk CHZ22/23
IR 3208 R. upper molar Gwebin TBG4 * IR 0743 L. M1/M2 Chaingzauk CHZ29
IR 3209 L. M3 Gwebin TBG4 * IR 0748 (1) M1/M2 Chaingzauk CHZ27
IR 3213 R. M1/M2 Gwebin TBG4 * IR 0748 (2) M1/M2 Chaingzauk CHZ27
IR 3214 L. M3 Gwebin TBG4 * IR 0770 M1/M2 Chaingzauk CHZ22/23
IR 3215 L. M1 Gwebin TBG4 * IR 0808 R. mandible with P4–M2 Chaingzauk CHZ
IR 3216 L. M1 Gwebin TBG4 * IR 0809 R. mandible with M2–M3 Chaingzauk CHZ
IR 3217 L. M1/M2 Gwebin TBG4 * IR 0811 L. M3 Chaingzauk CHZ
IR 3218 R. M3 Gwebin TBG4 * IR 0817 L. mandible with M1–M3 Chaingzauk CHZ
IR 3220 L. mandible with M3 Gwebin TBG4 * IR 0818 L. mandible with M1–M3 Chaingzauk CHZ
* IR 3221 L. M3 Gwebin TBG4 * IR 0823 (1) R. mandible with P4–M3 Chaingzauk CHZ
IR 3222 L. M3 Gwebin TBG4 * IR 0823 (2) mandible with P4–M3 Chaingzauk CHZ
IR 3320 L. M1/M2 Gwebin TBG4 * IR 0824 L. mandible with M3 Chaingzauk CHZ
IR 3323 R. P3/P4 Gwebin TBG4 * IR 0825 R. mandible with P3–P4 Chaingzauk CHZ
IR 3325 L. M1–M2 Gwebin Psw1 * IR 0826 L. mandible with M2–M3 Chaingzauk CHZ
IR 3327 L. M3 Gwebin Psw1 * IR 0827 L. mandible with M1–M2 Chaingzauk CHZ
* IR 3753 M2/M3 Gwebin MKT8 * IR 0828 R. mandible with M2–M3 Chaingzauk CHZ
IR 3930 R. upper molar Gwebin TBG1 * IR 0829 L. mandible with P4–M1 Chaingzauk CHZ
IR 3963 L. M3 Gwebin MT1 (TBG4) * IR 0830 R. mandible with M3 Chaingzauk CHZ
IR 3964 R. M3 Gwebin MT1 (TBG4) * IR 0833 mandible with dP4–M1 Chaingzauk CHZ31
IR 3966 R. M1 Gwebin MT1 (TBG4) * IR 0842 L. mandible with M2 Chaingzauk CHZ31
IR 3979 R. M1/M2 Gwebin TBG4 * IR 0856 R. mandible with M3 Chaingzauk CHZ23
IR 3980 L. M3 Gwebin TBG4 * IR 0864 R. mandible with M1–M3 Chaingzauk CHZ23
IR 3993 R. mandible with P4–M2 Gwebin Psw3 * IR 0867 R. mandible with M2 Chaingzauk CHZ23
IR 3994 L. M2 Gwebin Psw3 * IR 0873 R. mandible with P4–M1 Chaingzauk CHZ33
IR 3995 L. M2 Gwebin Psw3 * IR 0874 L. mandible with M1 Chaingzauk CHZ33
IR 3996 R. M2 Gwebin Psw3 * IR 0880 L. mandible with M1–M2 Chaingzauk CHZ33
IR 3997 R. M3 Gwebin Psw3 * IR 0892 R. M1/M2 Chaingzauk CHZ33
IR 3998 L. M1/M2 Gwebin Psw3 * IR 0912 L. mandible with M1 Chaingzauk MKT6
IR 3999 L. M3 Gwebin Psw3 * IR 0918 L. M3 Chaingzauk MKT1
IR 4039 R. M3 Gwebin Psw1 * IR 0925 L. mandible with M2–M3 Chaingzauk MKT1
IR 4040 R. lower premolar Gwebin Psw1 * IR 0926 R. mandible with M2–M3 Chaingzauk MKT1
* IR 0928 R. mandible with P4–M2 Chaingzauk MKT1
Bovidae gen. et sp. indet. 3 (medium-sized) * IR 0929 R. mandible with M2–M3 Chaingzauk MKT1
No. Material Area Locality * IR 0931 L. mandible with dP4–M2 Chaingzauk MKT1
IR 4053 R. M3 Yenangyaung SAL1 * IR 0932 L. mandible with M2–M3 Chaingzauk MKT1
IR 4054 L. M3 fragment Yenangyaung SAL1 * IR 0933 R. mandible with M2–M3 Chaingzauk MKT1
IR 4078 R. M1/M2 Yenangyaung NYD2 * IR 0938 R. mandible with M2–M3 Chaingzauk MKT1
IR 4079 R. P3/P4 Yenangyaung NYD2 * IR 0967 R. M3 Chaingzauk MKT1
IR 4091 L. upper molar Yenangyaung NYD2 * IR 1423 L. maxilla with M1–M3 Chaingzauk CHZ15
IR 4092 R. mandible with P3–P4 Yenangyaung NYD2 * IR 1851 L. mandible with M2–M3 Chaingzauk CHZ15
IR 4546 L. mandible with M2 Yenangyaung NYD3 * IR 2129 M1/M2 Chaingzauk MKT1
* IR 2130 M1/M2 Chaingzauk MKT1
Bovidae gen. et sp. indet. 4 (medium-sized) * IR 2141 M1/M2 Chaingzauk MKT2
No. Material Area Locality * IR 2174 M1/M2 Chaingzauk MKT2
IR 0195 R. maxilla with M2–M3 Chaingzauk CHZ17 * IR 2176 M1/M2 Chaingzauk MKT2
eschweizerbart_xxx

* IR 2184 R. mandible with P Chaingzauk MKT IR 2368 L. mandible with M2–M3 Chaingzauk MKT
* IR 2209 M1/M2 Chaingzauk MKT1 IR 2369 L. mandible with molar Chaingzauk MKT
* IR 2223 M1/M2 Chaingzauk MKT1 IR 2370 L. mandible with M2–M3 Chaingzauk MKT
* IR 2240 M1/M2 Chaingzauk MKT1 IR 2371 L. mandible with M2–M3 Chaingzauk MKT
* IR 2288 M1/M2 Chaingzauk MKT IR 2372 R. M3 Chaingzauk MKT
IR 2308 R. mandible with M2 Chaingzauk MKT IR 2373 L. mandible with M3 Chaingzauk MKT
IR 2310 R. mandible with M2–M3 Chaingzauk MKT IR 2374 L. mandible with M1–M3 Chaingzauk MKT
IR 2311 R. mandible with M2–M3 Chaingzauk MKT * IR 2375 L. mandible with M2 Chaingzauk MKT
IR 2312 R. mandible with M1 Chaingzauk MKT IR 2376 L. mandiblewith P4–M2 Chaingzauk MKT
IR 2313 R. mandible with M2 Chaingzauk MKT IR 2377 R. mandible with M1 Chaingzauk MKT
IR 2314 R. mandible with M2–M3 Chaingzauk MKT IR 2378 R. mandible with M3 Chaingzauk MKT
IR 2315 M2? Chaingzauk MKT IR 2379 L. mandible with dP4, M1–M2 Chaingzauk MKT
IR 2316 L. mandible with M2 Chaingzauk MKT IR 2380 R. mandible with M3 Chaingzauk MKT
IR 2317 L. mandible with M2–M3 Chaingzauk MKT IR 2381 R. mandible with P3 –M3 Chaingzauk MKT
IR 2318 R. mandible with M2–M3 Chaingzauk MKT * IR 2382 L. mandible with M1–M3 Chaingzauk MKT
IR 2319 L. mandible with M3 Chaingzauk MKT IR 2383 L. mandible with M2–M3 Chaingzauk MKT
IR 2320 M3 Chaingzauk MKT IR 2384 L. mandible with M2–M3 Chaingzauk MKT
IR 2321 L. mandible with M3 Chaingzauk MKT IR 2385 R. mandible with M3 Chaingzauk MKT
IR 2322 R. mandible with M3 Chaingzauk MKT * IR 2386 R. mandible with M2–M3 Chaingzauk MKT
IR 2323 R. M2 Chaingzauk MKT IR 2387 L. mandible with P3–M3 Chaingzauk MKT
IR 2324 L. mandible with P4–M1 Chaingzauk MKT IR 2388 L. mandible with M3 Chaingzauk MKT
IR 2325 L. M1/M2 Chaingzauk MKT IR 2389 L. mandible with M3 Chaingzauk MKT
* IR 2326 L. M1/M2 Chaingzauk MKT IR 2390 L. mandible Chaingzauk MKT
* IR 2327 R. M1/M2 Chaingzauk MKT IR 2391 L. mandible with M2–M3 Chaingzauk MKT
* IR 2328 L. M1/M2 Chaingzauk MKT * IR 2392 L. mandible with P4–M1 Chaingzauk MKT
IR 2329 L. mandible with M1 Chaingzauk MKT IR 2393 R. mandible with M2–M3 Chaingzauk MKT
IR 2330 R. mandible with M3 Chaingzauk MKT * IR 2394 L. mandible with M2–M3 Chaingzauk MKT
* IR 2331 R. mandible with P4–M3 Chaingzauk MKT IR 2395 R. mandible with M2–M3 Chaingzauk MKT
* IR 2332 L. mandible with P4–M2 Chaingzauk MKT IR 2396 R. mandible with M2–M3 Chaingzauk MKT
* IR 2333 L. mandible with M1–M3 Chaingzauk MKT IR 2397 L. mandible with P4–M1 Chaingzauk MKT
IR 2334 R. mandible with M3 Chaingzauk MKT * IR 2398 R. mandible with M1–M3 Chaingzauk MKT
IR 2335 L. mandible with M3 Chaingzauk MKT L.R. maxillae with P2, dP3, P3,
* IR 2649 Chaingzauk MKT1
IR 2336 L. mandible with M2–M3 Chaingzauk MKT dP4, M1–M3
IR 2337 R. mandible with M1 Chaingzauk MKT IR 2650 R. P3 Chaingzauk MKT1
IR 2338 R. mandible with dP4–M1 Chaingzauk MKT IR 2651 L. P3 Chaingzauk MKT1
IR 2339 L. mandible with M2–M3 Chaingzauk MKT IR 2652 R. M2 Chaingzauk MKT1
IR 2340 L. mandible with M2–M3 Chaingzauk MKT IR 2653 L. M1 Chaingzauk MKT1
IR 2341 L. M2 Chaingzauk MKT IR 2654 L. M3 Chaingzauk MKT1
IR 2342 L. mandible with M1–M3 Chaingzauk MKT IR 2655 L. dP4 Chaingzauk MKT1
IR 2343 L. mandible with M2–M3 Chaingzauk MKT IR 2656 R. P4 Chaingzauk MKT1
IR 2344 R. mandible with M2–M3 Chaingzauk MKT IR 2657 M1 Chaingzauk MKT1
IR 2345 L. mandible with M2–M3 Chaingzauk MKT IR 2658 R. mandible with M1 Chaingzauk MKT1
IR 2346 L. mandible with M3 Chaingzauk MKT IR 2659 R. M1 Chaingzauk MKT1
IR 2347 R. mandible with M1–M2 Chaingzauk MKT IR 2660 R. M1 Chaingzauk MKT1
IR 2348 R. mandible with M3 Chaingzauk MKT IR 2661 L. M3 Chaingzauk MKT1
* IR 2350 L. mandible with M2–M3 Chaingzauk MKT IR 2663 R. M3 Chaingzauk MKT1
* IR 2353 L. mandible with M2–M3 Chaingzauk MKT IR 2666 L. M1 Chaingzauk MKT2
IR 2356 R. mandible with P4 Chaingzauk MKT IR 2669 L. M2 Chaingzauk MKT2
IR 2357 L. mandible with M2 Chaingzauk MKT IR 2670 L. M3 Chaingzauk MKT2
IR 2359 L. M2 Chaingzauk MKT IR 2672 R. P4 Chaingzauk MKT2
IR 2360 R. mandible with M3 Chaingzauk MKT IR 2673 R. P4 Chaingzauk MKT2
IR 2361 L. M3 Chaingzauk MKT IR 2674 L. M1 Chaingzauk MKT2
IR 2363 L. mandible with M2 Chaingzauk MKT IR 2676 L. M1/M2 Chaingzauk MKT2
IR 2364 L. mandible with M2 Chaingzauk MKT IR 2677 R. M1/M2 Chaingzauk MKT2
IR 2365 lower molar fragment Chaingzauk MKT IR 2678 L. M2–M3 Chaingzauk MKT2
IR 2366 L. mandible with P4 Chaingzauk MKT IR 2679 R. M3 Chaingzauk MKT2
eschweizerbart_xxx

IR 2681 L. mandible with M1–M3 Chaingzauk MKT2 IR 2990 L. P4 Chaingzauk CHZ15

IR 2696 L. M3 Chaingzauk CHZ15 * IR 2991 R. M1/M2 Chaingzauk CHZ15
IR 2697 L. M3 Chaingzauk CHZ30 IR 2992 R. mandible with M3 Chaingzauk CHZ
IR 2723 R. M2 Chaingzauk CHZ30 IR 2993 R. M1/M2 Chaingzauk CHZ15
IR 2724 L. P3/P4 Chaingzauk CHZ30 IR 2994 R. P3/P4 Chaingzauk CHZ15
IR 2730 R. M2 Chaingzauk CHZ15 IR 3046 R. M2 Chaingzauk MKT5
IR 2731 L. M3 Chaingzauk CHZ30 IR 3047 L. M1 Chaingzauk MKT5
IR 2732 R. M3 Chaingzauk CHZ15 IR 3051 L. M1 Chaingzauk MKT5
IR 2733 L. mandible with M1 Chaingzauk CHZ30 IR 3052 L. M2 Chaingzauk MKT5
IR 2734 L. mandible with M3 Chaingzauk CHZ30 IR 3054 R. M3 Chaingzauk MKT5
IR 2735 L. M3 Chaingzauk CHZ30 IR 3055 R. M3 Chaingzauk MKT5
IR 2736 L. M3 Chaingzauk CHZ30 IR 3056 L. M3 Chaingzauk MKT5
IR 2767 L. M2 Chaingzauk MKT8 IR 3057 L. mandible with M3 Chaingzauk MKT5
IR 2768 R. M1/M2 Chaingzauk MKT8 IR 3058 L. mandible with M3 Chaingzauk MKT5
IR 2769 R. P3/P4 Chaingzauk MKT8 IR 3063 L. M1 Chaingzauk MKT1/2
IR 2770 L. M1 Chaingzauk MKT8 IR 3064 L. M1 Chaingzauk MKT1/2
IR 2771 R. P2 Chaingzauk MKT8 IR 3065 R. M1/M2 Chaingzauk MKT1/2
IR 2772 L. M1/M2 Chaingzauk MKT8 IR 3066 L. M2 Chaingzauk MKT1/2
IR 2773 L. mandible with molar Chaingzauk MKT8 IR 3080 R. M1/M2 Chaingzauk MKT7
IR 2774 molar fragment Chaingzauk MKT8 IR 3081 R. M3 Chaingzauk MKT7
IR 2775 L. M1/M2 Chaingzauk MKT8 * IR 3082 R. M3 Chaingzauk MKT7
IR 2776 R. M1/M2 Chaingzauk MKT8 * IR 3083 R. M1/M2 Chaingzauk MKT7
IR 2777 R. molar fragment Chaingzauk MKT8 * IR 3084 R. M3 Chaingzauk MKT7
IR 2781 R. M1/M2 Chaingzauk MKT7 IR 3085 R. upper molar Chaingzauk MKT7
IR 2782 L. M1/M2 Chaingzauk MKT7 IR 3088 R. M1/M2 Chaingzauk MKT7
IR 2783 R. M3 Chaingzauk MKT7 IR 3089 R. M3 Chaingzauk MKT7
IR 2784 L. M3 Chaingzauk MKT7 IR 3090 L. M3 Chaingzauk MKT7
IR 2801 R. M3 Chaingzauk CHZ15 IR 3091 L. mandible with M1–M2 Chaingzauk MKT7
IR 2802 R. M3 Chaingzauk CHZ15 IR 3092 R. mandible with M2–M3 Chaingzauk MKT7
IR 2803 R. P4 Chaingzauk CHZ15 IR 3093 L. mandible fragment Chaingzauk MKT7
IR 2864 R. mandible with M2–M3 Chaingzauk MKT IR 3108 R. manidble with P4–M1 Chaingzauk MKT10
IR 2865 L. mandible with M2–M3 Chaingzauk MKT IR 3111 L. M1/M2 Chaingzauk MKT10
IR 2868 R. mandible with P3–M1 Chaingzauk MKT IR 3113 R. M1/M2 Chaingzauk MKT10
IR 2869 L. M3 Chaingzauk MKT IR 3135 L. mandible with M1–M2 Chaingzauk MKT
IR 2870 L. mandible with P2–P3 Chaingzauk MKT * IR 3136 R. mandible with M1–M2 Chaingzauk MKT
IR 2871 L. mandible withP4–M3 Chaingzauk MKT IR 3137 L. mandible with P4–M2 Chaingzauk MKT
IR 2872 L. mandible with M2 Chaingzauk MKT IR 3145 R. mandible with M1/M2 Chaingzauk CHZ15
IR 2873 R. mandible with M2–M3 Chaingzauk MKT IR 3146 L. M3 Chaingzauk CHZ15
IR 2874 L. mandible with M2–M3 Chaingzauk MKT IR 3467 R. upper molar Chaingzauk MKT8
IR 2875 L. mandible with M1–M3 Chaingzauk MKT IR 3477 L. P3 Chaingzauk MKT8
IR 2876 L. mandible with P4–M1 Chaingzauk MKT IR 3478 R. P2 Chaingzauk MKT8
IR 2877 L. mandible with M2–M3 Chaingzauk MKT IR 3482 R. mandible with M2–M3 Chaingzauk CHZ
IR 2878 R. mandible with M3 Chaingzauk MKT * IR 3569 mandible with P2–P3, M1–M3 Chaingzauk MKT8
IR 2879 L. mandible with M2 Chaingzauk MKT IR 3598 L. mandible with M1/M2 Chaingzauk CHZ22
IR 2880 L. mandible with P4–M2 Chaingzauk MKT IR 3599 L. mandible with M1–M2 Chaingzauk CHZ22
IR 2881 L. M1/M2 Chaingzauk MKT IR 3600 L. M1/M2 Chaingzauk CHZ22
IR 2882 L. M1/M2 Chaingzauk MKT IR 3615 R. mandible with M1–M2 Chaingzauk CHZ
IR 2883 L. M1/M2 Chaingzauk MKT IR 3616 L. mandible with M1/M2 Chaingzauk CHZ
IR 2884 R. M1/M2 Chaingzauk MKT IR 3644 L. mandible with P4–M3 Chaingzauk MKT
IR 2885 L. M1/M2 Chaingzauk MKT IR 3645 L. mandible with P4–M3 Chaingzauk MKT
IR 2886 L. M3 Chaingzauk MKT IR 3646 L. mandible with M2–M3 Chaingzauk MKT
IR 2887 L. mandible Chaingzauk MKT IR 3647 L. mandible with M1–M3 Chaingzauk MKT
IR 2888 L. mandible Chaingzauk MKT IR 3648 R. mandible with M2–M3 Chaingzauk MKT
IR 2968 R. M3 Chaingzauk MKT1/2 * IR 3712 L. M3 Chaingzauk MKT7
IR 2969 L. M3 Chaingzauk MKT1/2 * IR 3713 L. M3 Chaingzauk MKT7
IR 2984 R. M2 Chaingzauk CHZ (east1) * IR 3717 R. M3 Chaingzauk MKT7
IR 2985 L. M2 Chaingzauk CHZ (east1) * IR 3718 L. M2 Chaingzauk MKT7
IR 2986 L. P4 Chaingzauk CHZ15 IR 3719 R. M3 Chaingzauk MKT7
IR 2988 L. P4 Chaingzauk CHZ15 * IR 3720 R. M3 Chaingzauk MKT7
IR 2989 L. M3 Chaingzauk CHZ15 * IR 3722 L. M1 Chaingzauk MKT7
eschweizerbart_xxx

* IR 3723 R. M3 Chaingzauk MKT7 * IR 1463 (1) maxilla with M2 Gwebin TBG1

* IR 3725 R. M2 Chaingzauk MKT7 * IR 1469 L. mandible with M3 Gwebin TBG1
* IR 3748 R. M3 Chaingzauk MKT8 * IR 1470 maxilla with M2 Gwebin TBG1
IR 3750 R. mandible with M3 Chaingzauk MKT8 * IR 1571 (1) M2 Gwebin TBG
* IR 3752 L. mandible with P4–M3 Chaingzauk MKT8 * IR 1571 (2) M3 Gwebin TBG
* IR 3753 L. M1 Chaingzauk MKT8 * IR 1571 (3) P4 Gwebin TBG
* IR 3754 R. M3 Chaingzauk MKT8 * IR 1571 (6) M3 Gwebin TBG
IR 3756 R. M3 Chaingzauk MKT8 * IR 1572 (2) M1/M2 Gwebin TBG
* IR 3757 R. mandible with M1–M3 Chaingzauk MKT8 * IR 1573 R. mandible with M2–M3 Gwebin TBG
* IR 3767 R. M3 Chaingzauk CHZ15 * IR 1574 M3 Gwebin TBG
* IR 3825 R. mandible with M1–M3 Chaingzauk PAL * IR 1592 (1) M1/M2 Gwebin TBG
* IR 3826 R. mandible with M2–M3 Chaingzauk PAL * IR 1592 (2) M1/M2 Gwebin TBG
* IR 3828 R. mandible with M3 Chaingzauk PAL * IR 1596 (1) M3 Gwebin TBG
* IR 3834 R. M3 Chaingzauk MKT7 * IR 1623 (1) M1/M2 Gwebin TBG
IR 3835 L. M3 Chaingzauk MKT7 * IR 1623 (2) M1/M2 Gwebin TBG
IR 3836 R. M3 Chaingzauk MKT7 * IR 1629 (2) M2 Gwebin TBG
* IR 3837 L. M2 Chaingzauk MKT7 * IR 1843 L. M3 Gwebin TBG
* IR 3838 R. M2 Chaingzauk MKT7 * IR 1844 L. M2 Gwebin TBG
* IR 3839 L. M1 Chaingzauk MKT7 * IR 1847 R. M3 Gwebin TBG
* IR 3840 R. M2 Chaingzauk MKT7 * IR 1858 R. M2 Gwebin TBG
* IR 3841 L. M2 Chaingzauk MKT7 * IR 1859 R. maxilla with P4–M1 Gwebin TBG
* IR 3842 R. M1 Chaingzauk MKT7 * IR 1860 L. M1/M2 Gwebin TBG
* IR 3846 R. M3 Chaingzauk MKT7 * IR 1861 L. M2 Gwebin TBG
* IR 3847 R. M3 Chaingzauk MKT7 IR 1902 R. mandible with M2 Gwebin TBG1
IR 3848 L. M3 Chaingzauk MKT7 * IR 1905 P3 Gwebin Psw1
* IR 3851 R. M3 Chaingzauk MKT7 * IR 1906 L. M2 Gwebin TBG1
IR 3854 R. M3 Chaingzauk MKT7 * IR 1907 L. M2 Gwebin TBG1
* IR 3855 R. M1 Chaingzauk MKT7 IR 1910 L. M3 Gwebin TBG1
* IR 3858 L. M2 Chaingzauk MKT7 * IR 1913 R. P4 Gwebin Psw1
* IR 3859 R. M3 Chaingzauk MKT7 * IR 2018 P3 Gwebin Gbn1 (Mg2)
* IR 3860 L. M3 Chaingzauk MKT7 IR 2020 L. maxilla with P4–M1 Gwebin Gbn1
* IR 3861 L. M1/M2 Chaingzauk MKT7 * IR 2025 M1/M2 Gwebin Gbn
* IR 3862 R. M2 Chaingzauk MKT7 IR 2030 R. P4 Gwebin TBG4
* IR 3868 R. mandible with M2–M3 Chaingzauk MKT * IR 2031 P4 Gwebin TBG4
* IR 3869 R. mandible with P3, M1–M2 Chaingzauk MKT * IR 2032 P3 Gwebin TBG4
* IR 3870 L. mandible with M2–M3 Chaingzauk MKT * IR 2033 R. M1 Gwebin TBG4
* IR 3871 L. mandible with M2–M3 Chaingzauk MKT IR 2034 R. M1 Gwebin TBG4
* IR 3872 R. mandible with M2–M3 Chaingzauk MKT * IR 2036 L. M3 Gwebin TBG4
* IR 3873 L. mandible with M1–M3 Chaingzauk MKT * IR 2037 L. M3 Gwebin TBG4
* IR 3874 R. mandible with P4–M2 Chaingzauk MKT * IR 2038 mandible with P2–P3, M1–M3 Gwebin TBG4
IR 3875 R. mandible with M1–M3 Chaingzauk MKT * IR 2042 mandible with P3–P4 Gwebin TBG5
* IR 3877 R. mandible with M2–M3 Chaingzauk MKT * IR 2043 M2 Gwebin TBG5
* IR 3878 R. mandible with M2–M3 Chaingzauk MKT * IR 2045 R. M3 Gwebin TBG5
* IR 3887 L. mandible with P4–M2 Chaingzauk MKT * IR 2046 M3 Gwebin TBG5
* IR 3888 R. mandible with M1–M2 Chaingzauk MKT * IR 2048 R. mandible with P4–M3 Gwebin TBG5
MGW 005 R. mandible with P4–M3 — — * IR 2088 M3 Gwebin TBG1
* ONG 577 L. mandible with P4–M3 — — * IR 2089 P4 Gwebin TBG1
IR 2458 L. M3 Gwebin Gbn1
Bovidae gen. et sp. indet. 5 (medium-sized) * IR 2459 L. M3 Gwebin Gbn1
No. Material Area Locality * IR 2564 L. maxilla with P3–P4 Gwebin TBG1
* IR 0003 (10) R. M3 Gwebin Gbn1 IR 2580 L. mandible Gwebin TBG1
IR 0003 (11) R. M3 Gwebin Gbn1 IR 2591 L. M3 Gwebin Gbn1
* IR-0063 P3 Gwebin Gbn1 IR 2592 L. M1 Gwebin Gbn5
* IR 0063 (3) P4 Gwebin Gbn1 IR 2601 L. M3 Gwebin TBG4
* IR 0063 (7) M1/M2 Gwebin Gbn1 IR 3153 L. M3 Gwebin TBG1
* IR 0063 (8) M1/M2 Gwebin Gbn1 IR 3154 L. M1/M2 Gwebin TBG1
* IR 1342 (1) P4 Gwebin Gbn1 IR 3155 L. M1 Gwebin TBG1
* IR 1351 R. mandible with P2–P3 Gwebin Gbn1 IR 3156 P4 Gwebin TBG1
* IR 1353 P4 Gwebin Gbn1 IR 3168 R. M1/M2 Gwebin Psw1
* IR 1406 L. mandible with M2–M3 Gwebin Gbn1 IR 3172 L. M1/M2 Gwebin Gbn1
* IR 1407 (1) R. M3 Gwebin Gbn1 IR 3173 R. lower molar Gwebin Gbn1
* IR 1407 (2) R. M3 Gwebin Gbn1 IR 3200 P4 Gwebin TBG4
* IR 1408 (2) M2 Gwebin Gbn1 IR 3201 R. M2 Gwebin TBG4
* IR 1461 M1/M2 Gwebin TBG1 IR 3202 R. M3 Gwebin TBG4
eschweizerbart_xxx

IR 3203 L. molar fragment Gwebin TBG4 IR 1729 (8) L. M3 Chaingzauk MKT5

IR 3211 L. P4 Gwebin TBG4 IR 1729 (12) R. mandible with M2–M3 Chaingzauk MKT5
IR 3212 L. P4 Gwebin TBG4 * IR 2300 M2 Chaingzauk MKT
IR 3219 R. M3 Gwebin TBG4 * IR 2405 R. M3 Chaingzauk MKT
IR 3223 L. M3 Gwebin TBG4 * IR 2406 R. M2 Chaingzauk MKT
IR 3224 L. M3 Gwebin TBG4 * IR 2407 R. M2 Chaingzauk MKT
IR 3315 L. upper molar Gwebin TBG1 (north) * IR 2408 L. M2 Chaingzauk MKT
IR 3316 R. upper molar Gwebin Mg1 IR 2409 L. P4 Chaingzauk MKT
IR 3317 M1 Gwebin Mg1 IR 2410 L. mandible with P2–P4 Chaingzauk MKT
IR 3318 L. M1/M2 Gwebin Gbn1 IR 2411 L. mandible with M1–M2 Chaingzauk MKT
IR 3319 R. M1/M2 Gwebin TBG4 * IR 2412 L. mandible with M3 Chaingzauk MKT
IR 3324 R. M2–M3 Gwebin TBG4 IR 2779 L. M1/M2 Chaingzauk MKT7
IR 3326 L. M1/M2 Gwebin Gbn1 IR 2780 L. M1/M2 Chaingzauk MKT7
IR 3965 L. M3 Gwebin TBG4 IR 2861 R. mandible with P3–M1 Chaingzauk MKT
IR 3974 R. P3 Gwebin TBG4 * IR-2967 L. M2 Chaingzauk MKT1/2
IR 3975 L. P4 Gwebin TBG4 IR 3110 L. M3 Chaingzauk MKT10
IR 3978 R. M2 Gwebin TBG4 IR 3869 R. mandible with P3 and M1–M2 Chaingzauk MKT
IR 3992 R. I2 Gwebin Psw3 * IR 3891 L. mandible with M1–M2 Chaingzauk MKT
* IR 4027 L. M1/M2 Gwebin Psw3
Bovidae gen. et sp. indet. 7 (large-sized)
Bovidae gen. et sp. indet. 6 (large-sized) No. Material Area Locality
No. Material Area Locality IR 0003 (2) R. M1/M2 Gwebin Gbn1
* IR 0131 L. M1/M2 Chaingzauk CHZ * IR 0003 (4) L. M3 Gwebin Gbn1
* IR 0610 L. M3 Chaingzauk CHZ * IR 0003 (5) R. M2 Gwebin Gbn1
IR 0731 R. M2 Chaingzauk CHZ29 IR 0003 (12) L. M2? Gwebin Gbn1
* IR 0749 R. P2 Chaingzauk CHZ29 IR 0016 L. upper molar Gwebin Gbn2
IR 0868 L. M3 fragment Chaingzauk CHZ33 * IR 1463 (2) R. P4 Gwebin TBG1
* IR 0881 L. mandible with dP4–M2 Chaingzauk CHZ29 IR 1574 R. mandible with M3 Gwebin TBG
* IR 0934 L. mandible with dP3–M2 Chaingzauk MKT1 * IR 1601 R. P3 fragment Gwebin TBG3
* IR 1086 R. mandible with P3–M3 Chaingzauk MKT8 * IR 1846 L. M3 Gwebin TBG
IR 1116 (11) L. M3 Chaingzauk MKT8 * IR 1848 R. M3 Gwebin TBG
IR 1193 R. mandible with dP4, M1–M2 Chaingzauk MKT * IR 1850 R. M2 Gwebin TBG
* IR 1265 L. M1/M2 Chaingzauk MKT2 * IR 1857 R. M3 Gwebin TBG
IR 1654 (1) maxilla with M1–M2 Chaingzauk MKT * IR 1862 L. M3 Gwebin TBG
IR 1654 (2) mandible with P3–M1 Chaingzauk MKT * IR 1934 R. maxilla with M1–M2 Gwebin Gbn1 (west)
IR 1654 (3) mandible with M1–M3 Chaingzauk MKT IR 2025 R. M2 Gwebin Gbn village
IR 1654 (4) L. mandible with M1–M3 Chaingzauk MKT * IR 2026 L. M2/M3 Gwebin TBG1
IR 1654 (5) mandible with M2–M3 Chaingzauk MKT IR 2031 R. P4 Gwebin TBG4
IR 1654 (6) mandible with M1–M3 Chaingzauk MKT IR 2035 L. M2 Gwebin TBG4
IR 1654 (7) L. mandible with M1–M3 Chaingzauk MKT IR 2047 L. M3 Gwebin TBG5
IR 1654 (8) L. mandible with P4–M3 Chaingzauk MKT * IR 2094 R. P2 Gwebin Gbn5
IR 1654 (9) mandible with P4–M3 Chaingzauk MKT * IR 2460 R. M2 Gwebin Gbn1
IR 1654 (10) R. mandible with M1–M3 Chaingzauk MKT IR 3167 L. P4 Gwebin Psw1
* IR 1680 R. P3 Chaingzauk MKT2 * IR 3314 L. P2 Gwebin TBG1 (north)
IR 1729 (2) R. mandible with M2–M3 Chaingzauk MKT5 IR 3939 L. P4 Gwebin TBG1
IR 1729 (4) L. M3 Chaingzauk MKT5 IR 3976 R. M1/M2 Gwebin TBG4
IR 1729 (6) L. M3 Chaingzauk MKT5 IR 3977 upper molar fragment Gwebin TBG4
IR 1729 (7) R. M3 Chaingzauk MKT5
eschweizerbart_xxx

Nishioka+ (2019) Myanmar Bovids

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Nishioka+ (2019) Myanmar Bovids

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Palaeontographica, Abt. A: Palaeozoology – Stratigraphy Article

Bovidae (Mammalia, Artiodactyla) from the Neogene

Keywords: Artiodactyla, Mammalia, Miocene, Pliocene, Southeast Asia

© 2019 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

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1. Introduction Miocene Chinji Formation, which ranges from 14.2

Zin-Maung-Maung-Thein et al. (2011) Chavasseau et al. (2013) This study

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rawaddy fossils sold in Thailand may have come from 3.3 Abbreviations

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Table 2. Measurements of skulls of Sivaportax dolabella and Duboisia sp.

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Duboisia sp. terior keel may be explained by the intraspecific vari-

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Bovidae gen. et sp. indet. 2 (small-sized)

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Table 10. Measurements of cheek teeth of the medium-sized

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Text-fig. 7. Occlusal morphotypes of lower

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Siwaliks, namely the latest Miocene or early Pliocene.

Moreover, the fossil assemblage from the Chaingzauk

area suggest that the three species (S. vexillarius, S. fal-

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Explanation of the plates

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Palaeontographica, Abt. A: Palaeozoology – Stratigraphy Article

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