New hyaenodonta (Mammalia) from the middle Eocene of Myanmar

Please cite this article in press as: de Bonis, L., et al., New hyaenodonta (Mammalia) from the middle Eocene of Myanmar.
C. R. Palevol (2017), https://doi.org/10.1016/j.crpv.2017.12.003
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C. R. Palevol xxx (2017) xxx–xxx
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General Paleontology, Systematics, and Evolution
New hyaenodonta (Mammalia) from the middle Eocene
of Myanmar
Nouveaux hyaenodontes (Mammalia) de l’Éocène moyen de Birmanie
Louis de Bonisa,∗
, Floreal Soléb
, Yaowalak Chaimaneea
, Aung Naing Soec
,
Chit Seind
, Vincent Lazzaria
, Olivier Chavasseaua
, Jean-Jacques Jaegera
a
IPHEP: Institut de paléoprimatologie et de paléontologie humaine, évolution et paléoenvironnements, UMR CNRS 7262, Université de
Poitiers, 6, rue Michel-Brunet, 86073 Poitiers cedex 9, France
b
Royal Belgian Institute of Natural Sciences (RBINS), Operational Directorate Earth and History of Life, 29, rue Vautier, 1000 Bruxelles,
Belgium
c
Department of Higher Education Ministry of Education, Naypyitaw, Myanmar
d
Department of Geology, West Yangon University, 11393 Yangon, Myanmar
a r t i c l e i n f o
Article history:
Received 13 November 2017
Accepted after revision 16 December 2017
Available online xxx
Handled by Lorenzo Rook
Keywords:
Hyaenodonta
Hyainailouridae
Hyaenodontidae
Pondaung Formation
Nova species
a b s t r a c t
The middle Eocene Pondaung Formation in Myanmar has yielded a rich mammalian fauna
including several Primate taxa. Hyaenodonta are known by the genera Kyawdawia, Yarshea,
Orienspterodon, and two other indeterminate taxa. We describe here new material of Kyaw-
dawia, including some morphological details, a new species of the hypercarnivorous genus
Propterodon and an indeterminate species, different from those described earlier in Myan-
mar, and characterized by a reduction of m3 and would belong to a third lineage with the
same evolutionary trend as Galecyon and the Limnocyoninae. The hyainailourines (Orien-
spterodon) and hyaenodontines (Propterodon) are recorded for the first time in Southeast
Asia and these subfamilies appeared in quasi the same time in Europe illustrating probably
a profound change in the carnivorous fauna among Laurasia.
© 2018 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved.
Mots clés :
Hyaenodonta
Hyainailouridae
Hyaenodontidae
Formation de Pondaung
Nova species
r é s u m é
La formation de l’Éocène moyen de Pondaung, en Myanmar, a livré une riche faune mam-
malienne, dans laquelle on a signalé plusieurs taxons de Primates. Parmi les autres ordres
de mammifères, celui des Hyaenodonta était connu jusqu’à présent par les genres Kyaw-
dawia, Yarshea, Orienspterodon et deux autres taxons indéterminés. Nous décrivons dans cet
article du nouveau matériel de Kyawdawia, qui apporte des précisions sur certains détails
morphologiques, une nouvelle espèce du genre hypercarnivore Propterodon ainsi qu’une
forme indéterminée, qui diffère de toutes les espèces décrites à ce jour en Birmanie. Elle se
∗ Corresponding author.
E-mail address: louis.debonis@univ-poitiers.fr (L. de Bonis).
https://doi.org/10.1016/j.crpv.2017.12.003
1631-0683/© 2018 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved.

2 L. de Bonis et al. / C. R. Palevol xxx (2017) xxx–xxx
caractérise, en particulier, par la réduction de m3, et peut représenter une troisième lignée
montrant la même tendance évolutive que les Limnocyoninae et Galecyon. Les hyainailour-
inés (Orienspterodon) et les hyaenodontinés (Propterodon) sont les plus anciens connus
dans le Sud-Est asiatique. Ces sous-familles apparaissent quasiment à la même époque en
Europe, et la fin de l’Éocène moyen pourrait correspondre à un changement majeur de la
faune de carnassiers en Eurasie.
© 2018 Académie des sciences. Publié par Elsevier Masson SAS. Tous droits réservés.
1. Introduction
The middle Eocene Pondaung Formation, exposed in the
northwestern part of Central Myanmar, is mainly com-
posed of fluviatile sediments, and has produced a rich
vertebrate fauna. Since the first fossils reported by Pilgrim
and Cotter (1916), about 50 mammalian species have
been discovered such as artiodactyls (Ducrocq et al., 2000;
Holroyd and Ciochon, 1995; Métais, 2006; Métais et al.,
2000; Pilgrim, 1928; Tsubamoto et al., 2003, 2011, 2012,
2013), perissodactyls (Colbert, 1938; Holroyd et al., 2006;
Métais et al., 2006; Rémy et al., 2005; Tsubamoto et al.,
2003, 2005), rodents (Dawson et al., 2003; Marivaux et al.,
2005), and primates (Ba Maw et al., 1979; Beard et al.,
2005, 2007, 2009; Chaimanee et al., 2000, 2012; Ciochon,
1985; Colbert, 1938; Gebo et al., 2002; Gunnel et al., 2002;
Jaeger et al., 1998, 1999; Marivaux et al., 2008a, 2008b;
Shigehara et al., 2002; Takai et al., 2001). The age of the
Pondaung Formation has been estimated to be late mid-
dle Eocene (ca. *37 Ma) by magnetostratigraphy (Benammi
et al., 2002) and fission track dating (Tsubamoto et al.,
2002). However, recent radiometric analyses (U–Pb age)
on zircons obtained from a tuffacaceous bed in Pondaung
Formation indicate an age range between 40.22–40.31 Ma
(Khin Zaw et al., 2014), which is slightly older than that
obtained from the other methods. Almost 30 fossiliferous
localities have been recognized in the Pondaung Formation
and its stratigraphic study, conducted by Aung Naing Soe
et al. (2002), has demonstrated that sediments that yielded
fossil mammal remains were deposited in the same short
time interval and are issued from the upper part of the
Pondaung Formation. Geochemical, sedimentological and
paleoenvironmental data indicate a tropical, humid envi-
ronment under strong monsoon influences (Licht et al.,
2013a, 2013b, 2014). Most studies on the Pondaung fos-
sil mammals have focused on anthracotheriid artiodactyls
and primates because they are the most abundant taxa that
have been collected since the beginning of the twentieth
century. Here we report new discoveries that increase our
understanding of the Hyaenodonta of that formation.
The first hyaenodonts recovered in Myanmar were
briefly described (Egi and Tsubamoto, 2000) from speci-
mens coming from the Pondaung area as two indetermi-
nate species in an overall review of the fossil mammalian
fauna of Myanmar. Few years later (Egi et al., 2004),
one of these species was published as Yarshea cruenta
on fragments of the same individual (NMMP-KU 0045;
field number Bhn-31, right mandibular fragment with m1;
NMMP-KU 0046, right m3 trigonid; NMMP-KU 0214, right
m2 and m3 talonids, metastylar blade and protocone of left
M1 or M2, a tooth root fragment). The material was com-
ing from the “Upper Member” of the Pondaung Formation,
eastern side of Pondaung Range, central Myanmar in the
Bahin basin (Egi et al., 2004). Later the other species was
also described (Egi et al., 2005) as Kyawdawia lupina, with
cranial and post-cranial elements associated with the holo-
type. The dentition was quite complete but, belonging to
an old individual, the teeth are worn. In the same article,
a fragment of mandible bearing p4 talonid and m1 coming
from the same area and level is figured as Proviverrinae
indet. Smaller than Yarshea and Kyawdawia, this mandible
differs also by characters which seem to be more African
proviverrine-like. Other remains were recorded from the
same Pondaung Formation (Peigné et al., 2007). Some of
them were reported as Kyawdawia lupina but a M3 fig-
ured as Hyaenodontidae sp. indet. is too large to belong to
the other known Pondaung species at that time. The same
year another genus, Orienspterodon Egi et al., 2007, was
created for very large specimens from Pondaung and the
Chinese species Pterodon dahkoensis Chow Minchen, 1975
was assigned to this genus.
The new material which is described in the present arti-
cle consists in mandibular fragments and isolated teeth.
These new finds both increase the known diversity of the
hyaenodont fauna in Pondaung and provide new morpho-
logical details for previously described taxa.
2. Systematics
Hyaenodonta Van Valen, 1967
Hyainailouroidea Pilgrim, 1932
Hyainailouridae Pilgrim, 1932
Indohyaenodontinae Solé et al., 2013
Kyawdawia Egi et al., 2005
Kyawdawia lupina Egi et al., 2005
This species is represented in the new material by an
isolated p3, a fragment of mandible with p4–m2, a P4, 2
upper molars and some fragments of teeth.
2.1. Description of the lower teeth
The isolated p3 (NMMP-PK4-5-12-10a) is identified as
a lower one, but an upper one cannot be excluded; it is
little worn, smaller than p4 (Table 1), with a triangular
almost symmetrical lateral shape. There is a thin paracristid
descending from the apex of the protoconid to meet a small
paraconid and, distally, a better developed talonid.
The dentary (NMMP-PK2-2-11-13) is broken off at mid-
height and we cannot estimate the depth (Fig. 1). The
mandible would have been quite robust being 10.6 mm
breadth at the level of m1–m2 (Fig. 1A–C). The p4 is higher
than m1 (Table 1), robust and without any trace of wear.
The surface of enamel is slightly granular. The crown is

L. de Bonis et al. / C. R. Palevol xxx (2017) xxx–xxx 3
Table 1
Measurements (mm) of the lower teeth of Kyawdakia lupina.
Tableau 1
Mesures (mm) des dents inférieures de Kyawdakia lupina.
p3 p4 m1 m2 m3 References
L w h L w h L TrL trh trw talw L trL trh trw talw L trL trh trw tlw
NMMP-PK2-11-2-2017 9.2 5.8 8.8
NMMP-PK2-02-11-2013 13.3 6.2 10.8 11.9 6.5 7.7 6 5.9 15.9 9.9 11.5 9.5 7.5
NM11 Bhn 2000 13.6 8.2 7.4 6 Peigné et al., 2007
NPL-12-2010 7.5 15 8.4 8.9 Peigné et al., 2007
PKg 1051 14.2 8 7.7 7.7 Peigné et al., 2007
NMMP-TMC-2016 12.1 6.3 9.9 5.3 4.1
NMMP-KU 42 5.2 14.2 6.3 12.3 17.6 9.6 8.6 7.5 18.4 10.3 9.1 7.2 Egi et al., 2005
NMMP-KU1288 9.4 6.4 9.3 15.2 7 13.6 13.4 6.9 8.5 6.4 6.2 20.3 12.5 16.1 11 8.3 Egi et al., 2005
NMMP-MGW 0022 11 9.7 5.6 4.7 5 Egi et al., 2005
L: length; w: width; h: height; TrL: trigonid length; trh: trigonid height; trw: trigonid width; talw: talonid width.
Fig. 1. Kyawdawia lupina (PK2-2-11-13). Fragment of dentary with p4–m2. A. Lingual view. B. Buccal view. C. Occlusal view (stereo). Isolated P4. D. Lingual
view. E. Buccal view. Scale bar = 10 mm.
Fig. 1. Kyawdawia lupina (PK2-2-11-13). Fragment de mandibule avec p4–m2. A. Vue linguale. B. Vue buccale. C. Vue occlusale (stéréo). P4 isolée. D. Vue
linguale. E. Vue buccale. Échelle = 10 mm.
bucco-lingually flattened, the mesial half being narrower
than the distal one with a sharp paralophid running from
the top to the base of the crown. There is another smaller
crest distally from mid-height to the base. There is a well-
developed talonid (3.3 mm length) without a clear cristid
obliqua but with a high and powerful hypoconid and a
narrow elongated lingual flat surface replacing the weak
cingulid surrounding the tooth. The m1, slightly shorter
than p4 and far smaller than m2 (Table 1), is almost com-
plete except the disto-lingual corner of the talonid. The
trigonid is complete with a swollen metaconid smaller than
the paraconid but nevertheless well developed and bearing
a small vertical mesio-lingual ridge. There are wear facets
on the top of the three cuspids of the trigonid and we can-
not know the original height of the cuspids. The wear facet
is rounded and horizontal in the protoconid with appear-
ance of the dentine in the middle, elongated and slightly
buccally inclined with traces of dentine in the paraconid
and it is oval and horizontal in the metaconid with also
a pit of dentine. Another wear facet extends in the dis-
tal surface of the protoconid and metaconid. The talonid
is slightly narrower than the trigonid (talw/trw = 98). The
lingual half of the talonid is broken off. The cristid obli-
qua, whose a mesial part is missing, is perpendicular to the
protoconid and joins the conical hypoconid, a wear facet
running along the crest and the top of the hypoconid. The
ratio width of trigonid/width of talonid (98.3) is slightly
higher than those (77.3 to 96.8) of published m1 of K. lupina
(Egi et al., 2005; Peigné et al., 2007). The m2 is very well
preserved. The trigonid, overlooked by the protoconid, is
high without significant traces of wear in the top of the cus-
pids except a very tiny one, concerning only the enamel, in
the apex of the protoconid. Like in specimen NMMP-Pkg-
1051, the protoconid height surpasses the trigonid length.

The paraconid is very oblique, nearly perpendicular to the
axis of the tooth raw and slightly separated from the meta-
conid. The latter is well developed but smaller and slightly
lower than the former. A large vertical shearing wear facet
with vertical striations extends on the buccal surface of the
protoconid and partly the paraconid. There is not a clear
cristid obliqua in the talonid but a large hypoconid elon-
gated with a slightly convex buccal face and a lingual one
gently sloping to the deep and wide talonid basin. The ento-
conid, separated from the metaconid by a deep notch, is as
high as the hypoconid but thinner; it constitutes a crest
separated from the hypoconulid by a small notch, the lat-
ter being separated from the hypoconid by a larger one. A
moderate cingulid runs along the buccal base of the tooth
except at the level of the disto-buccal part of the proto-
conid. The talonid breadth is smaller than the trigonid one
and the ratio (79) is slightly smaller than those (87.2–100)
previously published (Egi et al., 2005; Peigné et al., 2007).
2.2. Upper teeth
P4 (Fig. 1C, D) is three rooted but a root is broken off.
It is fresh except a small pit of wear in the top of the main
cusp and a small part of the mesio-buccal corner broken
off. Its measurements are: width = 8.4 mm, height = 9.7 mm.
We cannot take the length. The main cusp is triangular in
lateral view; distally a thin vertical crest runs down to an
elongated and trenchant talon. In the distal wall, there is
a small horse-shoe shaped contact facet for M1. Another
smaller mesial accessory cusp is present. Lingually a true
protocone does exist as a basin limited by the cingulum in
which a small transverse crest extends from the main cusp
to a sharp small cusp, dividing the basin in two parts. The
height of the crown is similar to the length of the teeth
but the latter is may be underestimating because a missing
fragment of enamel. The newly found upper molars do not
differ from those described in previous publications.
3. Discussion
This is the third publication in which Kyawdawia lupina
is reported, the ﬁrst being Egi et al. (2005) and the second
Peigné et al. (2007). Because of its relative abundance,
this species was probably the most common carnivorous
mammal in the Pondaung area during the end of the
middle Eocene. It was probably an opportunistic feeder
like are some of the extent canids. The morphology of the
lower molars displays some cutting areas along the buccal
face of the trigonid as well as large talonids dedicated to a
crushing function. The new material provides new details
of characters of the lower molars and principally those of
the upper P4 which was quite unknown until now.
Hyaenodontoidea Leidy, 1869
Hyaenodontidae Leidy, 1869
Hyaenodontinae Leidy, 1869
Propterodon Martin, 1906
Propterodon panganensis nov. sp.
Holotype: dentary with p4–m1, alveoli of m2 and m3
(NMMP-Pg1-2011-29).
Etymology: from the name of the locality where the
specimen was recorded. Diagnosis: small sized Propterodon
with a low and trenchant p4 having a trenchant small
talonid. Mesiodistally ﬂattened trigonid of m1 without
metaconid and short trenchant talonid with a small nar-
row postfossid. Presence of individualized hypoconid in
m1. Complete absence of cingulid.
Differential diagnosis: differs from Hyaenodon and other
hyaenodontids by the shape of p4, and the relative large
size of m1. Propterodon morrisi (= P. irdinensis) differs by
its larger size, the larger talonid of m1 and the presence of
a minute metaconid in molars. P. pishigouensis differs by
its larger size, with a relatively larger and powerful p4 of
which the main cusp is distally tilted with very developed
accessory mesial and distal cuspids. P. tongi differs by its
larger size, more robust dentary, and, following Tong and
Lei, 1986, a m1 similar to that of P. irdinensis (with a minute
metaconid?).
Type locality: Pangan 1, Myanmar.
Geological age: upper part of the Pondaung Formation,
40–37 Ma, almost equivalent to the European Bartonian or
MP 15–MP 16 (European Land Mammal Age).
Distribution: Pondaung formation (Myanmar) only.
Measurements in mm: p4 length 5.5, width 2.15, height
5.06. m1L 5.1, trigL 4.16, trigw 2.73, trigh 4.2, talw 1.6.
m2L (6.5), m3L (7.5) Height and width of the dentary: level
p3–p4 10.3 × 5.3; level p4–m1 10.6 × 4.9; level m1–m2
11 × 4.6; level m2–m3 11.5 × 7.7. Height of the coronoid
process: 21.7 mm. () = measured from alveoli.
4. Description
This taxon is represented by an incomplete dentary
bearing p4–m1 (Fig. 2A–C) and part of the mesial alve-
olus and the distal alveolus of p3, alveoli of m2 and m3
with pieces of the tooth roots and the coronoid process
(NMMP-Pg1-2011-29). Beyond its far smaller size, it dif-
fers dramatically from Kyadawia lupina by its morphology
and probably by its ecology and diet. It belongs clearly to
a different genus undescribed in Myanmar until now. The
dentary is slender, shallow with the lower border slightly
concave and slightly tapering forward. The mesial border of
the coronoid process is very oblique with an angle of 134◦
relatively to the horizontal line of the tooth row. Part of the
distal surface and the angular area are missing off, but it
seems that it was ventrally pointed. The masseteric fossa
is wide but not especially deep. It extends beneath m3 and
is dorsally limited by a robust ridge along the mesial bor-
der of the coronoid process. Lingually, a shallow depression
mesially limited by a moderate ridge corresponds to the
temporal muscle. The position of the mandibular condyle is
visible: it is located below the tooth row. The p4 is complete
except a fragment of enamel missing from the bucco-distal
corner. It is almost as high as m1 with a triangular lat-
eral outline. A weak paralophid ends at the base of a small
mesio-lingual paraconid. Distally there is a small but clear
talonid with a hypoconid of which the top is obliterated
by a small wear facet and, lingually a narrow and shallow
basin limited by a low entocristid. There is no trace of a

Fig. 2. Propterodon paganensis nov. sp. (Pg1-2011-29). Fragment of dentary with p4–m1. A. Lingual view. B. Buccal view. C. Occlusal view (stereo). Scale
bar = 10 mm.
Fig. 2. Propterodon paganensis nov. sp. (Pg1-2011-29). Fragment de mandibule avec p4–m1. A. Vue linguale. B. Vue buccale. C. Vue occlusale (stéréo).
Échelle = 10 mm.
cingulid. The paraconid of m1 is lower than the protoconid
and it is elongate mesially and slightly lingually directed.
The protoconid is transversally ﬂattened. The protoconid
is shifted distally. A wide quasi vertical wear facet with
vertical striations extends in the buccal surface of the para-
conid and large part of the protoconid. The metaconid is
absent. A short talonid composed by a conical hypoconid is
aligned on the protoconid and separated from it by a buc-
cal depression. A very short entocristid is present but does
not reach the metaconid, leaving the small talonid basin
lingually open. Like in p4, there is no cingulid.
5. Comparisons and discussion
The fossil from Pondaung fauna shares with the Hyaen-
odon species the distally shifting of the protoconid on
m1, the absence of metaconid, the very oblique coronoid
process, the coronoid process longer than high, the ven-
trally curved angular process, and the wide masseteric
fossa. These features distinguish the Pondaung taxon from
hyainailourines (e.g., Pterodon) and teratodontines (e.g.,
Dissopsalis), two hyaenodont subfamilies that convergently
acquired hypercarnivorous dentition (Borths et al., 2016).
The morphology of the m1 (long and distally shifting of the
protoconid) distinguishes the Pondaung taxon from hyper-
carnivorous proviverrines such as Oxyaenoides – the latter
genus actually shares mandibular features with Hyaenodon
(Solé et al., 2015).
We estimated the index (Lm1/Lp4) × 100 for some of
the oldest Hyaenodon species. It equals 74 in the North
American H. venturae (this species possesses the relatively
longest m1 among Hyaenodon (Lavrov and Emry, 1998)),
69 in H. minor, 68 in H. brachyrhynchus, H. requieni, and
H. eminus. This index actually equals 93 in NMMP-Pg1-
2011-29. Therefore, the Pondaung fossil differs from the
Hyaenodon species by the relative size of the m1: the lat-
ter is indeed longer than in Hyaenodon. This feature is
primitive among hyaenodontids. Moreover, the species of
Hyaenodon have ﬂattened and shearing lower molars with-
out metaconid but p4 is normally more robust relative to
NMMP-Pg1-2011-29, far higher than m1 and more conical.
Consequently, NMMP-Pg1-2011-29 does not represent a
Hyaenodon species despite the presence of similar dental
and mandibular features.
Propterodon is the closest relative to Hyaenodon (Borths
et al., 2016). Propterodon morrisi, Matthew and Granger,
1924 (= P. irdinensis Matthew and Granger, 1925) from the
Eocene of Mongolia is founded on a dentary with p3, m1,
m3 and the alveoli of p1, p2, p4 and m2. It is close by the
size to NMMP-Pg1 2011-29 (Matthew and Granger, 1925,
Fig. 3). Unfortunately, the absence of p4 does not allow any
comparison of that tooth but m1 is different “with rather
large, high-crested heel nearly as broad as the body of the
tooth, a minute metaconid, paraconid considerably smaller
and lower than metaconid” (Matthew and Granger, 1925,
p. 4). In NMMP-Pg1-2011-29, the paraconid cannot be
lower than the metaconid which is absent, the protoconid

Fig. 3. Hyaenodonta indet. sp. (PK5 6-12-10). Fragment of dentary with talonid of m2 and the complete m3. A. Lingual view. B. Buccal view. C. Occlusal
view (stereo). D. Radiography showing the roots of m2 and m3. Scale bar = 10 mm.
Fig. 3. Hyaenodonta indet. sp. (PK5 6-12-10). Fragment de mandibule avec le talonide de m2 et m3 complète. A. Vue linguale. B. Vue buccale. C. Vue
occlusale (stéreo). D. Radiographie montrant les racines de la m2 et la m3. Échelle = 10 mm.
is less high relative to the paraconid and the talonid is
relatively smaller than that of P. irdinensis and it is nar-
rower than the trigonid. Another species, from the Eocene
of China, is attributed to the genus with a question mark?
Propterodon pishigouensis Tong and Lei, 1986. This taxon is
known from a piece of dentary with p4–m1. It differs by the
large size of p4, the protoconid of which being very high
and distally tilted, with relatively larger mesial and distal
accessory cuspids, the m1 with a large and high talonid
(Tong and Lei, 1986, Fig. 2). Another dentary bearing
p1–m3 is the type specimen of Propterodon tongi Liu and
Huang, 2002. That mandible differs from that of Pondaung
by its larger size (height of the dentary at p4–m1 14.5
against 10.6, at m1–m3 15.4 against 11.5: and length of
p4 5.5 against 8, Lm1 6.5 for 5.1). The ventral border of the
dentary is going up rapidly and is staying in a horizontal
line in NMMP-Pg1-2011-29 (Liu and Huang, 2002, Fig. 1).
The teeth are also different, p4 of P. tongi bears a clear
paraconid and a larger talonid, while m1 of P. tongi is worn
out and we just can see that it is smaller relatively to p4.
Regarding the relative sizes of the p4 and m1, the index
(Lm1/Lp4) × 100 equals 81 in P. tongi and 80 in P. iridensis
(unpublished specimen). The species? P. pishigouensis has
an index close to that of the Pondaung taxa: 95 vs 93.
To conclude, we consider that Pg1-2011-29 represents
a new species of Propterodon.
Family indet.
Hyaenodonta indet. sp.
NMMP-PK5 6-12-10 is a fragment of dentary (Fig. 3A–D)
with the mesial root and the talonid of m2 and the com-
plete m3. Its tooth measurements (mm) are: L m2 = (5.43);
talw m2 = 1.71; Lm3 = 3.76; Ltrg m3 = 2.12: trh m3 = 2.22;
trw m3 = 2; tlw m3 = 1.2.
The dentary is deep relatively to its width [index
(w/h)100 = 45.5]. The distal root of m1 is robust. The mesial
root of m2 is large, in the talonid, a high and relatively tren-
chant hypoconid has a convex buccal surface and a slightly
convex, almost ﬂat, vertical lingual surface while the thin
very low entocristid limits lingually an almost ﬂat talonid
basin. The talonid is slightly tapering distally. There is no

hypoconulid but a small notch separates hypoconid and
entocristid. In m3, the enamel is slightly weathered. The
protoconid overlooks the high trigonid with a lingual face
only slightly convex. The postparacristid is very oblique,
almost perpendicular, relatively to the tooth row axis. The
base of the paraconid contacts that of the metaconid. The
latter is less voluminous but as high as the paraconid and, in
occlusal view, the trigonid is V shaped in isosceles triangle.
The metaconid is slightly more distal than the protoconid.
The talonid is narrow and high because the size and shape
of the hypoconid. There is no talonid basin but a surface
sloping gently toward the lingual border without clear
entocristid nor entoconid. It is difficult to distinguish the
wear facets but it seems that there is one shearing in front
of the protoconid and, partly, the paraconid and another
one on the wall constituted by the distal surface of the pro-
toconid and metaconid which are both in a same plane. Like
around the talonid of m2, there is no trace of any cingulid
around the tooth.
6. Comparisons
The comparisons will deal with the hyaenodonts of
which m3 is reduced relative to m2: our comparison thus
will focus on Galecyon and on Limnocyoninae.
The genus Galecyon has been described in North Amer-
ica (Gingerich and Deutsch, 1989; Zack, 2011) and Europe
(Smith and Smith, 2001). The relative reduction of m3 is
mentioned in the original diagnosis as one of the major
features of this hyaenodont genus (Zack, 2011).
We can measure the reduction of the last molar rela-
tively to m2 (index (Lm3/Lm2) × 100). This index is 69 in
the Pondaung fossil, 97 in Galecyon morloi, 77 in Galecyon
peregrinus, 68 in G. mordax and 64 in G. chronius (means of
several specimens for the latter), higher for the two former
species but close to that of Pondaung in the latter one.
In NMMP-PK5 6-12-10, the dentary is relatively deep
like in Galecyon but the trigonid of the m3 is more
open and the lingual cuspids clearly separated, the pro-
toconid is lower relative to the paraconid, the metaconid
is slightly more distal, and the paraconid is lower with
a postparacristid more elongate and directed more disto-
mesially and not more conical as in Galecyon. The talonid of
Galecyon has a robust hypoconid and a crista oblica directed
toward the protoconid like in NMMP-PK5 6-12-10 but the
basin is deeper with an entocristid slightly higher. Despite
the common presence of a reduced m3, the differences
between the m3 of the Asian fossil and the Galecyon species
– especially the opening of the trigonid and length of the
paracristid – imply that it is hard to refer the Pondaung
taxon to Galecyon.
The Limnocyoninae are first recorded in the late
Palaeocene of Asia (Meng et al., 1998), but they mainly
radiated in the early and middle Eocene of North Amer-
ica (Gunnel, 1998). The whole subfamily Limnocyoninae is
characterized by m3 reduced or absent (Morlo and Gunnell,
2003, 2005; Polly, 1996). The m3 is absent in Limno-
cyon Marsh, 1872a, Thinocyon Marsh, 1872b, Oxyaenodon
Wortman, 1899, and Iridodon Morlo and Gunnell, 2003.
In Prolimnocyon Matthew and Granger, 1915, m3 is dra-
matically reduced. The index m3/m2 is very low (63) in
P. chowi Meng et al., 1998 – the oldest limnocyonine – and
in P. haematus Gingerich and Deutsch, 1989 (means of sev-
eral specimens for the latter), (43) in P. atavus Matthew and
Granger, 1915, and the m3 is knob-like in P. antiquus (Morlo
and Gunnell, 2003, fig. 18) so far more reduced than that of
NMMP-PK5 6-12-10. The m3 of P. chowi recalls that of the
Pondaung taxon in having a V shaped trigonid (in occlusal
view) and a metaconid slightly more distal than the proto-
conid. However, the m3 of P. chowi differs from that of the
Pondaung taxon in having a postparacristid clearly shorter
and more transversally aligned and large hypoconulid and
hypoconid. The m3 of P. atavus is more similar in having a
longer and more oblique postparacristid, but, as mentioned
above, the m3 is much more reduced compared to the m2
– the elongation and modification of the position of the
postparacristid could correspond to a convergence towards
more secant teeth. Therefore, based on these comparisons,
the fossil from Pondaung could represent a taxon different
from the Limnocyoninae.
Following the observations of Smith and Smith (2001)
based on the hyaenodonts from Dormaal, it has been pro-
posed that m3 was slightly smaller than m2 in the earliest
hyaenodonts (Solé, 2013; Solé et al., 2013). Based on this
postulate, one hypothesized that (1) the relative size of m3
increased in several groups of Hyaenodonta (e.g., Proviver-
rinae, Hyaenodontinae), whereas (2) the relative size of m3
decreased in the Limnocyoninae and Galecyon. Regarding
this hypothesis, the Pondaung taxon could represent a third
lineage that experienced such reduction.
7. Overall conclusion
The Pondaung Formation had produced at least three
different taxa of Hyaenodonta, Kyawdawia lupina, Yarshea
cruenta and Orienspterodon dahkoensis. Two indeterminate
species, were also described, Proviverrinae indet. for a frag-
ment of mandible in Egi et al. (2005) and Hyaenodontidae
sp. indet. in Peigné et al. (2007) for a large M3, none of
them matching the new material. Thus, the two new car-
nivorous significantly augment the list of the Pondaung
predators. The larger species, O. dahkoensis, would occupy
the place of a large hypercarnivore, the two medium-sized
Yarshea and Kyawdawia acted like canids, the former being
probably more carnivorous than the latter, Propterodon was
a small hypercarnivore similar to the extant Martes and
the smaller species was preying upon small vertebrates or
invertebrates. These five species, together with the other
indeterminate species, constituted a quite complete guild
of predators.
Concerning the paleo-biogeography, we note that all the
genera are recorded only in Myanmar (Yarshea and Kyaw-
dawia) or in Myanmar, Mongolia or China (Propterodon
and Orienspterodon). The relationships of these carnivorous
with those of other continents are thus actually unknown.
However, one can note that the hyainailourines (Orien-
spterodon) and hyaenodontines (Propterodon) recorded in
the Pondaung Formation, are the first ones recorded in
the Southeast Asia. It is interesting to note that these sub-
families appeared almost contemporaneously in Europe:
hyaenailourines appeared around MP16 reference level (ca.
39 Ma) (Solé et al., 2013) and the hyaenodontines around

MP17a (ca. 38 Ma) (Lange-Badré, 1979). The hyaenodonts
of the Pondaung Formation thus might illustrate a profound
modification of the carnivorous fauna among Laurasia.
Acknowledgements
We thank the villagers of Bahin, Paukkaung, Nyaung-
pinle, and Maggyigan of Pondaung area and their
authorities for their help, kindness and enthusiasm which
greatly facilitated our fieldwork. We are grateful to Sabine
Riffaut (IPHEP) for her work on the photos by LdeB, Arnaud
Mazurier (Univ. Poitiers) for the CT-scan sections. We also
thank Xavier Valentin and Mana Rubgumrung for their
participation to the fieldwork and their technical assis-
tance and Christopher Beard for his participation to the
fieldwork. This work has been supported by the ANR-09-
BLAN-0238-02 Program, CNRS UMR 7262, the University
of Poitiers, the Ministry of Culture of the Republic of
the Union of Myanmar, a 2015 grant from the Leakey
foundation and National Geographic grant number NG-
W344-14. F.S. was financially supported by the Federal
Science Policy Office of Belgium (Belspo Brain Pioneer
project BR/175/PI/CARNAGES).
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ပုံေတာင္ဂ်ီးယားမ်ဳိးစုပ႐ိုင္းမိတ္၏ ေက်ာက္ျဖစ္႐ုပ္ၾကြင္း ေတြ႕ရွိ
10 March 2018 12:05(+0630) (လြန္ခဲ့ေသာ 23 နာရီ ခန္႔က)
ၿမိဳင္ၿမိဳ႕နယ္ ၾကာကန္က်စ္ေခ်ာင္ႏွင့္ သန္းဥေတာက်စ္ေခ်ာင္မွ ပုံေတာင္ဂ်ီးယားမ်ဳိးစုပ႐ိုင္းမိတ္၏ ေ
က်ာက္ျဖစ္႐ုပ္ၾကြင္း ေတြ႕ရွိ
ေနျပည္ေတာ္ မတ္ ၉
ပုံေတာင္ဂ်ီးယားမ်ိဳးစုပ႐ိုင္းမိတ္၏ ေက်ာက္ျဖစ္႐ုပ္ၾကြင္းကို မေကြးတိုင္း ေဒသႀကီး ၿမိဳင္ၿမိဳ႕နယ္ ေပါ
က္ေခါင္းေက်းရြာအနီး ၾကာကန္က်စ္ေခ်ာင္ႏွင့္ သန္းဥေတာက်စ္ေခ်ာင္တို႔မွ ပထမဆုံးအႀကိမ္ ေတြ႕ရွိ
ခဲ့ေၾကာင္း သိရသည္။
ျပင္သစ္ႏိုင္ငံ Poitier University မွ ပါေမာကၡJean-Jacques Jaegerဦးေဆာင္ေသာပညာရွင္အဖြဲ႕ႏွ
င့္ ျမန္မာႏိုင္ငံသာသနာေရးႏွင့္ ယဥ္ေက်းမႈ ဝန္ႀကီးဌာန ေရွးေဟာင္းသုေတသနႏွင့္ အမ်ဳိးသားျပတို
က္ဦးစီးဌာနႏွင့္ ပညာေရးဝန္ႀကီးဌာနမွ ပညာရွင္မ်ားပူးေပါင္း၍ ေက်ာက္ျဖစ္႐ုပ္ၾကြင္းသုေတသန လု
ပ္ငန္းမ်ားကို ေဖေဖာ္ဝါရီ ၈ ရက္မွ ၂၈ ရက္ အထိ ပူးေပါင္းေဆာင္ရြက္ ခဲ့ျခင္းျဖစ္သည္။
အဆိုပါအဖဲြ႕သည္ ေဖေဖာ္ဝါရီ ၁၅ ရက္တြင္ မေကြးတိုင္းေဒသႀကီး ၿမိဳင္ၿမိဳ႕နယ္ ေပါက္ေခါင္းေက်းရြာ
အနီး ၾကာကန္က်စ္ေခ်ာင္၌ ပုံေတာင္ဂ်ီးယား ပ႐ိုင္းမိတ္၏ အံသြား အမွတ္(၁)ႏွင့္ (၂)ပါရွိေသာ ဘ
ယ္ဘက္ေအာက္ေမး႐ိုးပါ သြားႏွစ္ေခ်ာင္းကို လည္းေကာင္း၊ ေဖေဖာ္ဝါရီ ၁၈ ရက္တြင္ ယာရွည္က်
စ္ေခ်ာင္မွ ပုံေတာင္ဂ်ီးယားညာဘက္ေအာက္ေမး႐ိုးပါ သြားႏွစ္ေခ်ာင္းႏွင့္ သြားအလြတ္ အေသးတစ္ေ
ခ်ာင္းကို လည္းေကာင္း ေတြ႕ခဲ့ျခင္းျဖစ္သည္။
ေက်ာက္ျဖစ္႐ုပ္ၾကြင္းသုေတသနလုပ္ငန္းမ်ားကို ဆက္လက္ေဆာင္ရြက္ ခဲ့ရာတြင္ ေဖေဖာ္ဝါရီ ၂၂ ရ
က္တြင္ သန္းဥေတာက်စ္ေခ်ာင္မွ ပုံေတာင္ဂ်ီးယား၏ ဘယ္ဘက္ေမး႐ိုးပါ အံႀကိဳသြားအမွတ္(၃)ႏွင့္
အံသြားအမွတ္(၃)ပါ တစ္ေခ်ာင္းကို လည္းေကာင္း၊ ယာရွည္က်စ္ေခ်ာင္မွ ပုံေတာင္ဂ်ီးယား၏ ညာဘ
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အသစ္ တင္သမွ် သတင္းမ်ား အလို အေလွ်ာက္ပုိ ့
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Home ျပည္တြင္းသတင္း ႏိုင္ငံတကာ သတင္း ေဆာင္းပါး TV on Livestream မႈခင္း Voice of facebook အျခားက႑

3/11/2018 ပံေ◌တာင် ီးယားမ် ဳ ိးစပ႐ိင္◌းမိတ္၏ ေ◌က်ာက္◌ျဖစ္႐ပ ကင္◌း ေ◌ တ႕ရွိ - သစ း လင္ (ေ◌န႔စဥတင္◌း)
http://www.thithtoolwin.com/2018/03/blog-post_554.html?utm_source=feedburner&utm_medium=email&utm_campaign=Feed%3A+thithtoolwin%2FyI… 2/4
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က္ပိုင္း အံသြားအမွတ္(၁)ႏွင့္ အမွတ္(၃)ပါသြားတစ္ေခ်ာင္း၊ အႀကီးသြားအလြတ္ ႏွစ္ေခ်ာင္းကို လ
ည္းေကာင္း၊ ေပါက္ေခါင္းက်စ္ေခ်ာင္ အမွတ္(၂) ေျမသားေရး ေဆးဆန္ခါခ်ရာမွ ပုံေတာင္ဂ်ီးယား၏
ညာဘက္ေမး႐ိုးပါ အံသြားအမွတ္(၁)ႏွင့္ အမွတ္(၃)ပါ တစ္ေခ်ာင္းတို႔ကို လည္းေကာင္း ေတြ႕ရွိခဲ့သ
ည္။ ထိုသို႔ ေတြ႕ရွိ ရေသာ ပုံေတာင္ဂ်ီးယားမ်ဳိးစု၏အံသြားမွာ ယခင္ေတြ႕ရွိခဲ့ဖူးေသာ ပုံေတာင္ ဂ်ီး
ယားမ်ဳိးစု၏ အံသြားကဲ့သို႔ ေၾကြလႊာမ်ားတြန္႔ေခါက္မႈနည္းပါးကာ ေခ်ာမြတ္ေန သည္ကို ေတြ႕ရွိရၿပီး
အရြယ္အစားမွာလည္း အနည္းငယ္ေသးငယ္ေနသည္ကို ေတြ႕ရွိရေၾကာင္း၊ ပုံေတာင္ဂ်ီးယားမ်ဳိးစုပ႐ို
င္းမိတ္၏ ေက်ာက္ျဖစ္႐ုပ္ၾကြင္းကို ၾကာကန္က်စ္ေခ်ာင္ႏွင့္ သန္းဥေတာက်စ္ေခ်ာင္မွ ယခုႏွစ္တြင္
ပထမဆုံး ေတြ႕ရွိ ခဲ့ျခင္းျဖစ္ေၾကာင္း သိရသည္။
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အမာၿမိဳ႕နယ္ခြဲ ေဒၚညိမ္း ေက်းရြာအုပ္စု ခါးျပတ္ေ
က်းရြာေန ဖခင္ျဖစ္သူ ဦးတင့္ က်ပန္းလုပ္ က...
ေျမာင္းျမၿမိဳ႕နယ္တြင္၊ ဆင္လိမၼာတစ္ေကာင္က ေဗ
ဒင္ဆရာတစ္ဦးကုိ ႏွာေမာင္းျဖင့္ ညွစ္၍ အစြယ္ျဖင့္
ထုိးသတ္မႈ ျဖစ္ပြား ၊ ဆင္ဦးစီးကုိ လူေသမႈျဖင့္ အမႈ
ဖြင့္
PHOTO - YWM (ဆင္ထီး ဂိုးပလိပ္ေမာင္ ကုိ ျမင္ေ
တြ႕ရစဥ္) ေျမာင္းျမ၊ မတ္ ၈။ ဧရာ၀တီတိုင္း၊ ေျမာင္းျ
မၿမိဳ႕နယ္တြင္ ဆင္လိမၼာတစ္ေကာင္က ေဗဒင္ဆရာ
တ...
မူလတန္းအရြယ္မွစ လိင္ပညာေပးျခင္းသည္ သက္
ငယ္မုဒိမ္းကို ေလ်ာ့က်ေစမည့္ နည္းလမ္းတစ္ခု
“ကၽြန္မက အဲဒီလိုကိုင္တာေတြကို မႀကိဳက္ဘူးလို႕
အေဒၚ့ကို ျပန္ေျပာျပတယ္။ ဒါေပမယ့္ အေဒၚက ကို
ယ့္ဦးေလးေတြပဲ ကိုင္ပါေစ၊ ဘာျဖစ္လဲဆုိၿပီး ျပန္ေျ
ပာ...
ေပါက္ေတာျမိဳ႕တြင္ ဆယ္တန္းေျဖမည့္ ေက်ာင္း
သားတစ္ဦး ဓါးထိုးခံရ၍ ေသဆံုး
နိရဥၥရာ၊ မတ္ ၅ ရခိုင္ျပည္နယ္ ေပါက္ေတာျမိဳ႕တြင္
ေဆြမ်ိဳးခ်င္း ရန္စရွိရာမွ ဦးေလးေတာ္စပ္သူ တစ္ဦး
က ဓါးျဖင့္ထိုးသျဖင့္ အသက္ (၁၆) ႏွစ္အရြယ္...
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3/11/2018 Myanmar fossil find turns human history on its head - our earliest ancestors came from Asia, not Africa | Daily Mail Online
http://www.dailymail.co.uk/sciencetech/article-2154753/Myanmar-fossil-turns-human-history-head--earliest-ancestors-came-Asia-Africa.html 1/47
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Myanmar fossil ind turns human
history on its head - our earliest
ancestors came from Asia, not Africa
Tooth from Myanmar is similar to tooth from Libya from 37 million years ago
Pre-human ancestors migrated between continents
Asia, not Africa, is the birthplace of our anthropoid ancestors
By ROB WAUGH
PUBLISHED: 08:43 GMT, 5 June 2012 | UPDATED: 08:43 GMT, 5 June 2012
The birthplace of the human race is Asia - our earliest ancestors came to Asia in a
huge migration 37-38 million years ago, before they evolved into present-day apes
and humans.
A team of palaeontologists in Myanmar has found the tooth of a pre-human ancestor
- afrasia djijidae, so-called because it forms a missing link between Africa and Asia -
that is very similar another early ancestor found in Libya.
Four similar teeth were found after six years of sifting through sediment - a ind that
helps seal Asia as the starting point for our species.
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A team of palaeontologists in Myanmar has found the tooth of a pre-human ancestor - afrasia
djijidae, so-called because it forms a missing link between Africa and Asia - that is very similar
another early ancestor found in Libya
The 37-million-year-old tooth proves that early human ancestors lived in Asia, and only moved
to Africa fairly late in the process of evolution - the latest in a series of discoveries that have
'sealed' Asia as the starting point for our species
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The human family tree: A team of palaeontologists in Myanmar has found the tooth of a pre-
human ancestor - afrasia djijidae, so-called because it forms a missing link between Africa and
Asia. It is very similar another early ancestor found in Libya
‘Not only does Afrasia help seal the case that anthropoids irst evolved in Asia, it also
tells us when our anthropoid ancestors irst made their way to Africa, where they
continued to evolve into apes and humans,’ says Chris Beard, Carnegie Museum of
Natural History palaontologist.
He worked with an international team that included scientists from the University of
Poitiers.
‘Afrasia is a game-changer because for the irst time it signals when our distant
ancestors initially colonized Africa. If this ancient migration had never taken place,
we wouldn’t be here talking about it.’
Paleontologists have been divided over exactly how and when early Asian
anthropoids made their way from Asia to Africa.
The trip could not have been easy, because a more extensive version of the modern
Mediterranean Sea called the Tethys Sea separated Africa from Eurasia at that time.
While the discovery of Afrasia does not solve the exact route early anthropoids
followed in reaching Africa, it does suggest that the colonization event occurred
relatively recently, only shortly before the irst anthropoid fossils are found in the
African fossil record.
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¿Not only does Afrasia help seal the case that anthropoids irst evolved in Asia, it also tells us
when our anthropoid ancestors irst made their way to Africa, where they continued to evolve
into apes and humans,¿ says Chris Beard, Carnegie Museum of Natural History palaontologist
¿Not only does Afrasia help seal the case that anthropoids irst evolved in Asia, it also tells us
when our anthropoid ancestors irst made their way to Africa, where they continued to evolve
into apes and humans,¿ says Chris Beard, Carnegie Museum of Natural History palaontologist
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Myanmar’s 37-million-year-old Afrasia is remarkable in that its teeth closely resemble
those of Afrotarsius libycus, a North African primate dating to about the same time.
The four known teeth of Afrasia were recovered after six years of sifting through tons
of sediment near Nyaungpinle in central Myanmar.
Details of tooth shape in the Asian Afrasia and the North African Afrotarsius fossils
indicate that these animals probably ate insects.
The size of their teeth suggests that in life these animals weighed around 3.5 ounces,
roughly the size of a modern tarsier.
‘For years we thought the African fossil record was simply bad,’ says Professor Jean-
Jacques Jaeger of the University of Poitiers in France, the team leader and a
Carnegie Museum research associate. ‘The fact that such similar anthropoids lived at
the same time in Myanmar and Libya suggests that the gap in early African
anthropoid evolution is actually real. Anthropoids didn’t arrive in Africa until right
before we ind their fossils in Libya.’
The search for the origin of early anthropoids—and, by extension, early human
ancestors—is a focal point of modern paleoanthropology.
The discovery of Afrasia shows that one lineage of early anthropoids colonized
Africa around 37–38 million years ago, but the diversity of early anthropoids known
from the Libyan site that produced Afrotarsius libycus hints that the true picture was
more complicated.
These other Libyan fossil anthropoids may be the descendants of one or more
additional Asian colonists, because they don’t appear to be specially related to
Afrasia and Afrotarsius. Fossil evidence of evolutionary divergence—when a species
divides to create new lineages—is critical data for researchers in evolution.
The groundbreaking discovery of the relationship between Asia’s Afrasia and North
Africa’s Afrotarsius is an important benchmark for pinpointing the date at which
Asian anthropoids colonized Africa.
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