tmp1FA6 TMP

C. R.
Palevol 15 (2016) 359378
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Human palaeontology and prehistory
The faunal assemblage of the paleonto-archeological

localities of the Late Pliocene Quranwala Zone, Masol
Formation, Siwalik Range, NW India
Lassemblage faunique des localits palonto-archologiques de la zone
Quranwala, Pliocne nal, formation de Masol, chane frontale des
Siwaliks, Nord-Ouest de lInde
Anne-Marie Moigne a, , Anne Dambricourt Malass a , Mukesh Singh b ,
Amandeep Kaur b , Claire Gaillard a , Baldev Karir b , Surinder Pal b ,
Vipnesh Bhardwaj b , Salah Abdessadok a , Ccile Chapon Sao a ,
Julien Gargani c , Alina Tudryn c
a
b
c
Histoire naturelle de lhomme prhistorique (HNHP, UMR 7194 CNRS), Tautavel, France
Society for Archaeological and Anthropological Research, Chandigarh, India
Gosciences Paris-Sud (GEOPS, UMR 8148 CNRS), universit Paris-Sud, Paris, France
a r t i c l e
i n f o
Article history:
Received 23 June 2015
Accepted after revision 17 September 2015
Available online 18 January 2016
Handled by Anne Dambricourt Malass
Keywords:
Siwalik Frontal Range
Sub-Himalayan oodplain
Late Pliocene
Tatrot faunal assemblage
a b s t r a c t
The Indo-French Program of Research Siwaliks carried out investigations in the Quranwala
zone of the Masol Formation (Tatrot), Chandigarh Siwalik Range, known since the 1960s
for its transitional fauna. This new paleontological study was implemented following the
discovery of bones with cut marks near choppers and akes in quartzite collected on the
outcrops. Nine eldwork seasons (20082015) on 50 hectares of ravines and a small plateau
recovered lithic tools and fossil assemblages in 12 localities with approximately 1500 fossils. Their study shows that the most abundant mammal species are the Proboscideans with
Stegodon insignis. The transition with the Pleistocene fauna is evidenced by Elephas hysudricus, Hipparion antelopinum and Equus sivalensis. The freshwater mammal is also well
illustrated with Hexaprotodon sivalensis. Bovids present the greatest variety with six tribes
from the smallest to the largest. Two types of cervids are observed; Sivatherium giganteum
is visible in several localities and Merycopotamus dissimilis in one. Turtles, with the giant
terrestrial Colossochelys and the freshwater Geoclemys, are abundant. The aquatic predators
are limited (crocodile) and terrestrial carnivores are very scarce (hyena, felid). The faunal
assemblages match the Plio-Pleistocene transitional fauna, also described in the Pabbi Hills
(Pakistan), and mark the beginning of the Equus sivalensis Biostratigraphic interval-Zone,
which extends from 2.6 Ma to 600 ka. The systematic repetition of surveys has, therefore,
allowed the collection of rare taxa, such as Crocuta (2010), Merycopotamus dissimilis (2014)
and a large felid (2015). These latest ndings are signicant for the discovery of Homininae
in Siwaliks.
2015 Acadmie des sciences. Published by Elsevier Masson SAS. This is an open access
article under the CC-BY-NC-ND license (http://creativecommons.org/licenses/
by-nc-nd/4.0/).
Corresponding author. UMR 7194 CNRS/MNHN/EPCC, Tautavel, avenue Lon-Jean-Grgory, 66720 Tautavel, France.
E-mail address: moigne@mnhn.fr (A.-M. Moigne).
http://dx.doi.org/10.1016/j.crpv.2015.09.016
1631-0683/ 2015 Acadmie des sciences. Published by Elsevier Masson SAS. This is an open access article under the CC-BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
360
A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378
r s u m
Mots cls :
Chane frontale des Siwaliks
Plaine alluviale sous-himalayenne
Pliocne nal
Assemblage faunique de Tatrot
Le programme de recherche franco-indien Siwaliks poursuit ses investigations dans la

zone Quranwala de la formation de Masol, connue depuis les annes 1960 pour sa faune
de la n du Pliocne, galement nomme faune de transition Plio-Plistocne. Notre tude
palontologique sest impose la suite de la dcouverte de traces de dcoupe sur des fossiles proches de choppers et dclats en quartzite. Neuf campagnes de terrain (20082015)
sur 50 hectares de paysage particulirement accident (ravins et petits plateaux) ont permis didentier 12 localits o au moins un chopper tait ml des fossiles, lesquels
totalisent 1500 spcimens. Leur tude montre que les mammifres les plus abondants sont
les Proboscidiens, avec Stegodon insignis et Elephas en bien moins grand nombre. Les bovids
prsentent la plus grande varit, avec six tribus de la plus petite la plus grande taille.
Hexaprotodon sivalensis vient en troisime position. Sivatherium giganteum est visible dans
plusieurs localits, Merycopotamus dissimilis dans une seule. Deux types de cervids au
moins ont t observs. La faune de transition est atteste par la co-existence de Stegodon
et Elephas hysudricus, dHipparion antelopinum et dEquus sivalensis. Les fossiles de tortues
sont abondants avec la forme terrestre gante Colossochelys, et celle deau douce, Geoclemys.
En comparaison, les prdateurs sont peu reprsents (crocodile), voire rares (hyne, flin),
tandis que les primates (Procynocephalus) nont pas encore t observs. Lassemblage faunique correspond bien aux faunes de transition Plio-Plistocne galement dcrites dans les
Pabbi Hills (Pakistan) ; elle caractrise le dbut de la biostratigraphie de la zone intervalle
Equus sivalensis, qui stend de 2,6 Ma 600 ka. La rptition systmatique des prospections
a donc permis la collecte de taxons rares comme Crocuta (2010), Merycopotamus dissimilis
(2014) et un grand flin (2015). Ces derniers rsultats sont signicatifs pour la dcouverte
dHomininae dans les Siwaliks.
2015 Acadmie des sciences. Publi par Elsevier Masson SAS. Cet article est publi en
Open Access sous licence CC-BY-NC-ND (http://creativecommons.org/licenses/
by-nc-nd/4.0/).
1. Introduction
This new paleontological study of the Tatrot faunas in
the Chandigarh anticline (Punjab) of the Siwalik Frontal
Range is the necessary consequence of an unexpected discovery during a eld season of the Indo-French Program of
Research Siwaliks (Fig. 1) (Dambricourt Malass, 2016). In
2009, a shaft of bovid tibia was collected in the Quranwala
zone of the Masol Formation, near choppers and akes,
and showed marks on its cortical surface, which resembled butchery activities. At least two other bones with
evidence of suspected hominin scavenging activities were
recorded, and their hominin origins have been demonstrated by 3D topomiscroscopy and experimental protocols
(Dambricourt Malass et al., 2016a, b).
Eight years of surveys in the Masol inlier (20082015)
recorded fossils and artifacts in twelve paleontologicalarcheological localities (Fig. 2). The fossils were documented with very precise geological and geomorphological
data for each one of the localities in order to understand the
local faunal associations, as well as the sedimentary context of their fossilization (silts, sands, gravels, riverbanks,
channels, swamps, and fresh water).
Hence, this research completes the multidisciplinary
analysis of the Masol inlier (Abdessadok et al., 2016;
Chapon Sao et al., 2016a, b; Dambricourt Malass et al.,
2016a, b; Gaillard et al., 2016; Gargani et al., 2016; Tudryn
et al., 2016). The study of the vertebrate fauna allows us
to date these layers and identify the association of vertebrate (mammals and reptiles) in their environmental
condition of fossilization and estimate the potential of nding hominins specimens.
2. General background
The Siwalik Group in the Himalayan foreland basin and
foothills provides fossils from the Miocene (Murree Formation) up to the Middle Pleistocene (Boulder Conglomerate).
They have been described for almost two centuries by
world-renowned paleontologists who succeeded in the
eld (the Potwar Plateau in Pakistan, the Kashmir Valley,
Lesser Himalayan foothills and the Siwalik Frontal Range
in India). They produced the rst important collections
stored at the Geological Survey of India, Kolkata, in London (Falconer, 1868; Falconer and Cautley, 1846; Lydekker,
1879, 1883; Pilgrim, 1910, 1939), and in USA at the American Museum of Natural History (Brown et al., 1924; Colbert,
1935; Gregory et al., 1938) and the Yale University Museum
(de Terra and Teilhard de Chardin, 1936). Among new
species, the discovery of the rst great ape found in mainland Asia gave support to the Central Asian hypothesis of
the origin of the genus Homo, which was the prominent
paradigm during the 1920s. This theory was developed
by numerous paleontologists, such as Matthew, Osborn,
Andrews and Gregory (American Museum of Natural History), Black (1925) (Union Medical College, Beijing) and, to
a lesser extent, Teilhard de Chardin (National Museum of
Natural History, Paris), former student of Marcellin Boule.
After Ernest Hamy (chair of Anthropology of the National
Museum of National History, 1870), human paleontology
became an institutional science in the French Institution
with Boule; as for other geologists, the Tertiary Man, or
the Early Villafranchian Man (Late Pliocene), was inconceivable (Dambricourt Malass et al., 2016a).
361
Fig. 1. A. Location of the Chandigarhs Siwalik Frontal Range, North India (arrow). B. Location of Masol site (1), north of Chandigarh (3), upstream of Patiali
Rao (2). C. View on the Masol anticline and the dome with paleonto-archeological localities, on the dome Masol 1 with two sub-localites and Masol 2,
Masol 3 at a pass, Masol 5 in the small watershed of Pichhli Choe; A and B according to Google Earth, C A. (map data Google 2015). C: A. Dambricourt
Malass.
Fig. 1. A. Localisation de la Chane Frontale des Siwaliks de Chandigarh. B. Localisation du site de Masol (1), au nord de Chandigarh (3), en amont du Patiali
Rao (2). C. Vue sur lanticlinal de Masol avec les localits palonto-archologiques, sur le dme Masol 1 avec deux sous-localits et Masol 2, Masol 3 un
col, Masol 5 dans le petit bassin versant du Pichhli Choe. A et B daprs Google Earth (donnes de carte Google 2015) ; C : A. Dambricourt Malass.
Fig. 2. Geomorphology of the Masol inlier with paleonto-archeological localities, M1, M2 on the anticline dome, M3 at a pass, M4, M5 and M6 in the Pichhli
Choe, M7 to M13, on a circus, on the eastern rank of the Patiali Rao (Google Earth, map data Google 2015).
Fig. 2. Gomorphologie de la boutonnire Masol avec les localits palonto-archologiques, M1 et M2 sur le dme de lanticlinal, M3 un col, M4, M5 et
M6 dans le Pichhli Choe, M7 M13, dans un cirque, sur la rive orientale du Patiali Rao (daprs Google Earth, donnes de carte Google 2015).
362
Nevertheless, in 1926, Black did not hesitate to publish

an isolated tooth from the surface of the Zhoukoudian karst
that he attributed to Sinanthropus pekinensis, the rst suspected Tertiary Man discovered in mainland Asia (Black,
1926). Later, Teilhard de Chardin and Piveteau (1930) compared its faunal assemblage with those of the Nihewan
Basin and dated the S. pekinensis to the Lower Pleistocene.
In 1936, when de Terra and Teilhard de Chardin surveyed
the Potwar Plateau and collected stone tools, the PlioPleistocene boundary was still unclear; the Tatrot and
Pinjor Formations belong to the Upper Siwalik sub-group
(Pilgrim, 1910, 1944), but Tatrot was considered Lower
Pleistocene rather than Late Pliocene as it is today (de
Terra and Teilhard de Chardin, 1936). Thus, the discussions about the Siwalik Group (Lower, Middle and Upper
sub-groups) mainly concerned the equivalence of their Biostratigraphic succession with the European biochronology
(Late PlioceneLower Pleistocene Villafranchian faunae).
The faunal associations of the Potwar Plateau did not
correspond exactly to the European one, and generated
debates between paleontologists over the presence of
Archidiskodon in the Tatrot type locality (Sahni and Khan,
1968), the rst appearance of Equus at the base of the Pinjor Formation (de Terra and Teilhard de Chardin, 1936) and
Leptobos in the Pinjor Formation. Numerous paleontologists proposed to compare this biostratigraphy with the
sequences observed in the South of China and in the Southeast Asian archipelago by discussing the correlations of the
Tatrot Formation with the Kaliglagah and Tjidjulang beds,
and the Pinjor Formation with the Djetis or Puchangan beds
(e.g., Bergh et al., 2001; Colbert, 1935; Hooijer and Colbert,
1951; Koenigswald, 1956; Mishra et al., 2010; Pilgrim,
1939). The Mio-Pliocene biochronology of the Siwaliks
became very important when the same taxa were found
in the Mio-Pliocene sequences of East and South Africa
(Harris, 1976), and recently with Antilopini of the Middle
Awash (Ethiopia) (Bibi, 2011), as well as with the scarce
fossiliferous areas of the Baynunah Formation in the United
Arab Emirates (Bibi et al., 2013; Gilbert et al., 2014).
The Plio-Pleistocene boundary was suspected at the

uppermost level of the Quranwala zone, a fossiliferous
horizon of the Tatrot Formation as dened by Sahni and
Kahn (1964, 1968). Later, Nanda proposed that the Quranwala zone assemblage corresponded to the transitional
fauna between Pliocene and Pleistocene, composed of genera such as Hipparion and Stegodon, which will extinct
during the Pleistocene and new genera as Equus and Elephas, which will develop, and called it pre-Pinjor (Nanda,
1981, 1994, 19961997, 2002). In the same way, the Tatrot
Formation was renamed the Saketi Formation, in a fossiliferous Pliocene sector located 50 km southeast of Masol
in the Himachal Pradesh foothills. The paleomagnetism
was measured in the 1990s along the Patiali Rao, a seasonal river owing from the Masol inlier to the Indus
plain, giving a precise description of the different magnetic polarity reversals (Patnaik, 2012; Patnaik and Nanda,
2010; Ranga Rao, 1993; Ranga Rao et al., 1995; details
in Chapon Sao et al., 2016a). The Gauss/Matuyama reversal is now identied in the Chandigarh anticline between
the Tatrot and Pinjor Formations, and paleontologists have
admitted that the Stegodon/Elephas association characterizes the Latest Pliocene period and is visible below this
paleomagnetic reversal, which actually corresponds to the
Plio-Pleistocene boundary (Kumaravel et al., 2005).
The majority of the fossils collected in the Chandigarh
anticline comes from the Quaranwala zone, dated to Latest
Pliocene, and is particularly mineralized. The fossils of the
Pinjor Formation are less numerous and bones are less mineralized. The fossils also correspond to the very beginning
of the Pleistocene and they have been recovered in pockets by Sahni and Khan on short sedimentary sequences,
the thickness of which has been estimated only at 15 feet
(5 m) above the uppermost limit of the Tatrot Formation
(Sahni and Khan, 1964). Thus, all these fossil assemblages
can be regarded as Plio-Pleistocene transitional species, the
majority between 80100 m below this limit (Chapon Sao
et al., 2016b; Sahni and Khan, 1964, 1968).
4. The fossil sample: stratigraphic origin and
description
3. The Chandigarh anticline and the Quranwala

zone
The Chandigarh anticline is a geological structure of the
Siwalik Frontal Range (or Siwalik Range), 200 km southeast
of the Potwar Plateau and just at the boundary between the
Indus and the Ganga Basins, near the township of Pinjor
(Pilgrim, 1913). Its fossiliferous deposits have been studied
since the beginning of the twentieth century, but the most
important investigations date back to the 195060s with
the geologist M.R. Sahni and the creation of the Geology
Department of the Panjab University in Chandigarh (Union
Territory). Badam (2000) published a historical review for
the 18th commemoration dedicated to Sahni. Today the
geological and fossiliferous sequences of the Chandigarh
anticline are well known; they are composed of only the
Upper Siwalik: (i) the top of the Tatrot Formation, the
complete sequence of the Pinjor Formation and irregular
Boulder Conglomerate Formation (Sahni and Khan, 1968)
(Fig. 3).
Fossils from the Chandigarh anticline are stored and

exhibited in the Centre of Advanced Study Museum,
Panjab University, Chandigarh. The fossils of the IndoFrench program Siwaliks have been stored, classied and
preserved at the Society for Archaeological and Anthropological Research, Chandigarh; they have been restored
when necessary, then determined according to the previous works of the University of Punjab (e.g., Badam, 1973,
1979; Gaur, 1987; Nanda, 1981). The projected surface
of the Masol inlier covers around 80 hectares, but the
core of the anticline is devoid of fossil and the paleontological sector covers a surface of 50 hectares. Up to
now, 1500 fossils have been stored, and 1469 have been
described: 869 herbivores, 37 crocodilians and ve carnivores. They come (1) from fortuitous collecting in the
Quranwala zone (referenced Masol in the list), which
belongs to the collections of the Society for Archaeological and Anthropological Research (SAAR, Chandigarh),
(2) two trial trenches excavated in 2011 in the Masol 2
363
Fig. 3. Geological map of the Chandigarh Siwalik Frontal Range (Abdessadok, Fig. 1A in Chapon Sao et al., 2016, Sahni and Khan, 1968, modied).
Fig. 3. Carte gologique de la chane frontale des Siwaliks de Chandigarh (Abdessadok, Fig. 1A in Chapon Sao et al., 2016, daprs Sahni et Khan, 1968)
locality and (3) (the majority) during the eight eld seasons
(referenced Masol 1 to Masol 13 in the list) (Dambricourt
Malass et al., 2016a). The whole of the collection makes
it possible to establish a good diagnosis of the faunal
assemblages from the various localities, and the number of specimens is signicant and allows comparisons
(Table 1).
4.1. Stratigraphic origin
The base of the fossiliferous Quranwala zone has been
dened at Masol 1 (Abdessadok et al., 2016; Chapon Sao
et al., 2016b). In the stratigraphic log (Fig. 4), the c unit
refers to clays/silts layers and the s unit refers to sandstones. The fossiliferous sequence begins with pink and
orange silts (members ED of c3 unit), covered by ne grey
sandstone and micaceous sandstone (s3 unit), overlaid by
thick coarse sandstone with crossing beddings (s4 unit).
The oldest paleontological-archeological localities are in
units c3 (members ED), s3 and s4, at Masol 1, Masol 2,

Masol 5 and Masol 7; Masol 12 is composed of c3 (members ED) and s3, which yielded fossils but is devoid of tools
and has been selected because of a quartzite cobble layer
in the stratigraphy between two silt deposits. One bovid
tibia with cut marks has been recorded from a Masol 1 sublocality at the top of the dome, unearthed from the unit s3
(Fig. 1D). One bovid metacarpal with cut marks has been
collected in the small T2 terrace of the Pichhli Choe where
this seasonal torrent cuts the c3 unit (members ED), s3 and
s4 below the Masol 6 locality. The other localities belong to
the middle sequence of the log composed of the units c3 to
c6 and s5 to s6, the top of the Masol 2 sequence, Masol 3, 4
and 6 on the western rank of the Patiali Rao. On the eastern
rank, the localities are Masol 8, 9, 10, 11 and 13 in a geological circus forming a large surface of brown silts. This silty
surface (unit c5) is sporadically covered by pebbles/cobbles
in quartzite and dismantled sandstones at the base of M6,
M8 and M13. A bovid bone splinter with cut marks has
364
Table 1
Faunal list of Masol localities (NISP, number of identiable specimens present) (Total includes the collection of the Society for Archaeological and Anthropological Research (SAAR)).
Tableau 1
Liste de la faune des diffrentes localits de Masol (NR nombre de restes) (le total inclut la collection de la Society for Archaeological and Anthropological Research (SAAR)).
Masol 1
Masol 3
1
1
25
12
26
26
55
2
1
15
6
3
1
1
16
1
7
2
2
2
29
5
20
10
2
10
Masol 6
1
90
1
22
20
20
32
2
1
1
14
4
4
1
1
3
3
2
2
6
1
5
8
1
4
10
10
1
1
12
1
1
1
21
10
3
4
22
22
34
1
14
5
4
4
4
13
2
1
3
1
2
2
2
12
1
9
2
19
4
Masol 7
Masol 8
1
1
7
7
12
150
40
27
60
9
Masol 5
5
3
1
2
2
1
76
21
5
46
1
3
Masol 9
Masol 10
Masol 11
Masol 12
Masol 13
1
1
5
12
35
4
4
16
16
22
2
2
11
1
1
1
1
1
12
3
3
3
1
1
4
1
1
1
19
19
24
4
1
1
1
10
3
4
2
2
4
4
1
1
10
1
8
2
3
4
3
1
1
1
2
3
1
2
7
2
4
11
1
9
1
30
4
17
4
4
17
15
5
2
8
Masol A
1
1
13
1
1
65
36
4
387
10
14
15
153
17
23
3
98
1
8
14
5
88
35
53
16
360
12
20
24
21
162
1
1
5
44
2
3
40
2
1
2
106
Masol
total
6
5
1
252
16
77
1
1
120
120
224
9
3
8
21
21
3
3
27
28
2
2
65
6
33
10
1
8
10
3
5
315
82
57
136
9
1
2
28
1
2
162
170
71
1467
CARNIVORA
Crocuta sp.
Panthera
ELEPHANTIDAE
Elephas planifrons
Stegodon insignis
ANTHRA COTERIDAE
Merycopotamus dissimilis
HIPPOPOTAMIDAE
Hexaprotodon sivalensis
BOVIDAE
Bubalus sp.
Duboisia sp.
Hemibos sp.
Hippotragus sp.
Sivacapra sp.
CAMELIDAE
Camelus sivalensis
GIRAFFIDAE
Sivatherium giganteum
TRAGULIDAE
Dorcatherium nagrii
CERVIDAE
Axis like cervid
Cervus punjabiensis
SUIDAE
Propotamocheorus sp.
Sus brachygnatus
EQUIDAE
Equus sivalensis
Hipparion antilopinum
REPTILA
Colossochelys
Geoclemys
Turtles
Varanus
Lacertilia
Gavialis
Crocodylus punjabensis
Molusca
Pesces
PH
GH
TGH
Total
Masol 2
365
Fig. 4. Log of the Quranwala Zone of Masol Formation with paleontological-archeological localities (in Chapon Sao et al., 2016b).
Fig. 4. Log de la zone Quranwala de la formation de Masol avec les localits palonto-archologiques (in Chapon Sao et al., 2016b).
been recorded at Masol 13. Masol 9 is a small circus formed

in eroded sandstones (unit s5) and silts (unit c61), and
rounded by relic cliffs of the sequences s5c6, Masol 10 is
just above M9. Then starting from the sandstones s6 unit,
fossiliferous deposits disappear as on the western rank.
The last sequence of the log corresponds to the uppermost
part of Masol 6 cliffs, which extend from s6 to s13 and c7

to c13 until the crest lining the Pichhli Choe watershed.
In 2015, rare fossils have been collected at Masol 6, just
below the crest in sandstones unit s13, in conglomerate
unit s14 and on the crest in silt unit c17. They have not been
studied.
366
Fig. 5. Masol R10181, Stegodon insignis, right hemimandible with D4-M1 (scale 10 cm), SAAR fossil collection (picture A. Dambricourt Malass).
Fig. 5. Masol R10181, Stegodon insignis, hmimandibule droite avec D4-M1 (chelle 10 cm), collection de fossiles de la SAAR (photo A. Dambricourt
Malass).
4.2. Mammals
The most common mammalian families of the collection
are Elephantidae, Hippopotamidae, and Bovidae.
Proboscidea: Proboscids are very common (252 fossils),
represented by tusks, molars, mandibles, skulls, scapulae, ribs, long bones and foot bones. The Proboscideans
represented in the various lists of the Chandigarh anticline comprise numerous species (Badam, 1979; Gaur,
1987; Gaur and Chopra, 1984). For many years, it was
admitted that the association of Stegodon, Mammuthus
and Elephas could be characteristic of the Latest Pliocene
and appeared below the Gauss-Matuyama paleomagnetic
reversal (Agarwal et al., 1993; Hussain et al., 1992). In
this collection, the challenge was to determine whether
it was possible to recognize several Proboscideans among
Stegodon insignis, S. bombifrons, S. ganesa, E. planifrons, E.
hysudricus, E. platycephalus, Pentalophodon sivalensis and
Stegolophodon stegonoides (according to the nomenclature
of Shoshani and Tassy, 2005).
S. insignis has been identied from 77 fossils, such
as isolated teeth, lower jaws and skull fragments. The
morphology of the lateral view of the maxillary corresponds to S. insignis (Falconer and Cautley, 1846;
Hooijer, 1955). The teeth formula has few thick blades
and the crown is low: lower unworn M1, Masol 1
R10181, 8 blades, height = 50 mm, length = 160 mm and
width = 75 mm, lamina thickness = 19.6 mm, enamel thickness = 4.7 mm (Fig. 5). The dental ranges of S. insignis
are larger and narrower than S. bombifrons, the lamina is narrower and the enamel is ner. A complete
talus presents a Stegodon-like morphology (Falconer
and Cautley, 1846), Masol 6 R10584: height = 137 mm,
width = 116 mm, Dap = 65 mm).
E. hysudricus: only one species (16 fossils) was identied (Falconer and Cautley, 1846). The material is composed
of one skull with the fourth molar, or the rst true molar,
broken on the level of the roots. The isolated teeth are
not very high, with quite separate plates. The lamellate
frequency is seven and the enamel thickness is around
3 mm and folded. The lower unworn last molar Masol
R11082 presents 11+ plates with a laminar frequency
of ve (number of plates per 10 cm of anteroposterior

length) (Fig. 6): length = 230 mm, greatest width = 75 mm,
height = 103 mm. On the occlusal surface, characteristic
rhombuses are present in the center of the plate (thickness = 10.4 mm).
Hexaprotodon sivalensis is well-documented (120 fossils). The hippos are the most abundant taxon at the
different localities; the fossils include several skulls,
mandibles, isolated teeth and postcranial remains, including small bones and the axial skeleton. The cranial elements
(10 remains) from Masol 1, Masol 2 and Masol 6 conrm its
taxonomic status. A cranial fragment at Masol 2 presents
the roots of the last molar, the zygomatic apophysis and
the base of the orbital pit. Occipital and parietal parts are
also signicant of the taxonomic attribution. The Masol 1
R10482 jawbone was discovered in a block of sandy concretion revealing its stratigraphic origin (S3), but most of
the isolated teeth and mandibles were collected on the silts
of Masol 1 and Masol 2. It bears D3, D4 and M1 still in bud;
M1 is 1/3 larger than H. s. koenigswaldi from the Lower
Pleistocene formation of Sangiran (Indonesia).
The mandibular symphyses are broad on the labial
massif and the six incisors are placed on a single range.
Fig. 6. Last lower molar Masol R10182, Elephas hysudricus, SAAR collections, SAAR fossil collection (picture A.-M. Moigne).
Fig. 6. Dernire molaire infrieure, Masol R10182, Elephas hysudricus,
collection de fossiles de la SAAR (photo A.-M. Moigne).

Table 2
Hexaprotodon sivalensis dental measurements (mm) per locality, isolated
teeth (R) and teeth on lower or upper jaw. Note the great variation due to
dimorphism and use-wear stage (germs and used teeth are not measured).
Masol without reference corresponds to the SAAR collection without location.
Tableau 2
Hexaprotodon sivalensis; dimensions dentaires (mm) par localit, dents
isoles (R) et dents sur les mandibules et maxillaire (n). Noter la grande
variation en raison du dimorphisme sexuel et du stade dusure (les germes
et les dents uses ne sont pas mesurs). Masol sans rfrence correspond
aux collections de la SAAR sans localisation.
Locality
Specimen
Type
DMD minmax
DVL minmax
Masol 1
Masol 1
Masol 1
Masol 1
Masol 6
Masol 2
Masol
Masol 1
Masol 8
Masol
R10174
R11345
R10022
n=4
n=6
n=2
R11080
R10135
n=6
n=3
I 1 inf
I2 inf
I3 inf
M2 inf
M3 inf
C sup
I3 sup
Ml sup
M2 sup
M3sup
3026
2420
2820
5436.6
7861
4947.5
20.5
46.6
5246.8
4955.4
34
2218
24.324
39.425.6
4027.5
47.535.3
22.5
38.8
5143.6
5145.7
The I1 and I2 present almost the same section size (Masol

2 R11014, R10188), and this feature could be considered
primitive (Hooijer, 1950). Lower jaw bones are high, thick
and the base of the ramus is at. Six mandibles (from Masol
1, Masol 6 and, Masol 8) bear the last molar; its measurements correspond to a very large Hexaprotodon sivalensis
(Hooijer, 1950) (Table 2).
The isolated higher canine (Masol R10117) is deeply
grooved along the posterior surface and testies to a strong
individual, probably a male. As with the postcranial bones
(ulna, pelvis, and metapodial bones), the H. sivalensis tibia
measurements are within the size range of Hippopotamus:
the transverse distal diameter of Masol 13 R10255 are
82 mm and 73 mm for H. s. koenigswaldi.
Bovidae are represented by many subfamilies in Asia
(e.g., Patnaik and Nanda, 2010). Here, there are 239
remains, including 62 teeth and ankle fragments. The
Bovidea have been separated into six tribes: Bovini,
Alcelaphini, Boselaphini, Hippotragini, Reduncini and Antilopini. Unfortunately, no complete skull has been recorded,
but the taxa were identied based on of ankle morphology
and dental features, such as the lower P4 morphology, the
presence or absence of the basal pillar, the basal cingulum
and the hypsodonty index (Dennell et al., 2006; Gaur, 1983,
1987; Gentry, 1978b; Pilgrim, 1939).
Bovini are the largest bovids, referred to as Hemibos or
Leptobos, as well as Bubalus. Bovini is a large type with
hypsodont and selenodont teeth. The ankles are either
oval shaped or already more attened with a clear former
hull. Hemibos and Bubalus present cavernicolous osseous
ankles, with quadrangular or triangular sections with a hull
well dened on the higher face. The horns develop rst,
slightly backwards, and then go down lower than the top
of the frontal bone (Gaur and Chopra, 1984; Hooijer, 1958;
Pilgrim, 1939). Bubalus sp.: the enamel is thick, rugose
without cingulum, the basal pillars are more or less developed, and the styles are clear. The largest teeth present
the Bubalus fold on both anterior and posterior surfaces.
Hemibos triquetricornis: the teeth of large individuals are
367
lled with cement, the styles are thick and parallel and the
lower molars developed additional styles (Gromova, 1968).
Postcranial bones are often reduced to articulations but can
be separated into two main groups. The maximal weight
estimated according to the distal humerus width (Masol 2
R10184) and distal metacarpal width (Masol 6 R10285) is
close to 1000 kg.
Hippotragini: twenty-one remains are identied with
a partial skull including the frontal and the two straight,
divergent, backwards cylindrical ankles. Many cylindrical
fragments have been recovered on the Masol localities,
which can be confused with highly mineralized antlers.
Teeth are hypsodont, lower molars with the Capra fold,
middle size with a rather short premolar raw. The lower
P4 have a broad bulbous metaconid. The last lower
molars, incomplete, without cement, have straight and
thin styles, and are vertical and parallel. Alveolar crumpling of Capra type and coast punts correspond to the
model B or open habitat adapted ruminants (Khler,
1993). These fossils are assigned to the subfamily Hippotragini (Sivatragus brevicornis (Pilgrim, 1939) is known in
the Tatrot Formation). Postcranial bones are identied in
Masol 1.
Boselaphini (three teeth) are represented by characteristic upper teeth in Masols 1 and 5, but the complete
study of the dental collection requires more evidence.
This kind appears frequently in the faunas of this period.
Dental remains approach the well-known Duboisia in the
Pleistocene sites of Southeast Asia with open teeth, no
cement, discrete basal cingulum and smooth enamel, and
are less hypsodont than are other Bovidae. The size is bigger than Duboisia kroenesenii and resembles the biggest
species, D. saatensis (Geraads, 1979) (Fig. 7A, Masol 5
R11267: maximal length = 24.6 mm, basal width = 20 mm).
Dental occlusion (cutters) does not allow for their classication among specialized animals (Khler, 1993; Rozzi
et al., 2013).
Reduncini tribe (18 remains): two mandibles are close
to Sivacapra sp.; the teeth are fairly hypsodont, small
with unmolarised P4 and a strong Capra fold, which
conrms their dimensions and assignation to the subfamily Reduncini (Fig. 5B). They were compared with
the mandible type from the Centre of Advanced Study
Museum, Panjab University, (UPM Chandigarh) (Gaur,
1987). Postcranial remains bring some indications: the
morphology of the metacarpal bones rather resemble the
A3 Gazella type (Khler, 1993), without a sagittal sulcus
(previously described in the Pinjor Formation as Gazella and
Indoredunca).
Merycopotamus dissimilis: anthracotheres were rst
described in the Upper Siwalik (Falconer and Cautley,
1846) but they are scarce. One large upper molar
(Masol 5 R11286: length = 28.5 mm, width = 28.5 mm,
height = 15 mm) was determined as M. dissimilis (Fig. 7C).
The tooth is an upper M3, unworn and very well preserved,
the crown is selenodont, brachyodont with rugose enamel
teeth, as a typical browser animal. Until now, no postcranial
bones have been identied.
Equidae: (n = 10 fossils) The equid fossils are few and
only the teeth or some long bones could be attributed to
a genus (n = 8). Horses were also discovered in situ during
368
Fig. 7. A. Duboisia sp. Masol 5 R11267, last upper molar. B. Sivacapra sp. Masol 2 R10109, left lower jaw with M1 and M2. C. Merycopotamus dissimilis.
Masol 5 R11286, left upper last true molar. D. Hipparion, premolar, occlusal surface (picture A.-M Moigne).
Fig. 7. A. Duboisia sp. Masol 5 R11267, M3 suprieure. B Sivacapra sp. Masol 2 R10109. hmimandibule droite avec M1 et M2. C. Merycopotamus dissimilis,
Masol 5 R11286, M3 suprieure gauche. D. Hipparion, prmolaire, vue occlusale (photo A.-M. Moigne).
Table 3
Hipparion antelopinum; comparative measurements (mm) of the forth
Upper Premolar * after Gaur, 1987.
Tableau 3
Hipparion antelopinum; dimensions compares de la P4 suprieure
(en mm), * daprs Gaur, 1987.
Locality
Specimen
Length
Width
Index
L protocone
Masol B
Colbert, 1936
Badam, 1973
E. sivalensis*
R10500
S/3
22.5
23.6
24
30
23.5
21.8
25
34.5
0.9
1.08
0.9
1.15
9.6
the trench excavation at Masol 2. Two genera are distinguished: Hipparion and Equus.
Hipparion antilopinum: the upper premolar Masol 3
R10050 is small, the occlusal surface is damaged but on
transverse breakage, the protocon is clear, isolated and oval
shaped (Table 3), which is characteristic of the Hipparion
type as well as the true second premolar (Masol R10471:
length = 40 mm, width = 24 mm). Hipparion presents the
morphological diagnosis of H. antelopinum: crenulated fossette, tendancy of the protocone to be round, complexity of
the enamel ridge, open hypoconal groove complexity of pli
caballin fold (Gromova, 1968). The measurements correspond with H. antelopinum (Badam, 1973; Colbert, 1935)
(Fig. 7 D).
E. sivalensis: the isolated lower M2 (Masol ind. R11459)

is related to E. sivalensis according to Falconer and Cautley
(1846) and Colbert (1935). This specimen is larger than PUA
135B (Centre of Advanced Study Museum, Panjab University, Gaur, 1987). One tooth has been collected in trench B
at Masol 2 (the base of the Quranwala zone).
Tragulidae: In reviews of Siwalik fossils, the Tragulid
genus Dorcatherium is commonly determined with four
species from the Miocene to the Pleistocene. The extended
genera of Tragulid Hyemoschus or Tragulus are very similar to Dorcatherium (Gentry, 1978a). Dorcatherium nagrii
would appear to be the most advanced among the Dorcathers of the Siwaliks, and is probably the ancestor of the
extant species (Gaur, 1992). The attribution to D. nagrii
is mostly established on the size criteria: lower teeth are
selenodont, not hypsodont, narrow, very small, not specialized and characteristic of no ruminant artiodactyls. The
fragmented mandible from Masol 6 R10264 with the last
molar (length = 11 mm) is in the maximum range of this
species.
Girafdae: the fossils (n = 27) have been collected from
Masol 3, 6, 8, 11 and 13, but only rarely at Masol 2 and 5:
one ossicone, eight teeth and one postcranial skeleton were
assigned to the very large Girafd, Sivatherium giganteum.
The gigantic male Pleistocene girafd with four ossicones
was rst described from the Upper Siwalik (Falconer and
Cautley, 1836). The anterior ossicones are conical and the
369
Table 4
Sivatherium giganteum; dental measurements (mm) of Masol specimens compared with those of Kenyan *KNMER/Sivatherium cf. maurusium (Harris, 1976).
Tableau 4
Sivatherium giganteum ; dimensions dentaires (mm) compares celles de lespce kenyanne *KNMER/Sivatherium cf. maurusium (Harris, 1976).
LocaIity
Specimen
Type
Length
DVD max
DVD anterior
DVD posterior
DVD third lobe
Height
Masol3
Masol
*KNMER
UPM
Masol
*KNM-ER
UPM
UPM
Masol
Masol
*KNMER
UPM
UPM
UPM
*KNMER
Masol 13
R10122
R10213
777
A/811
R10213
780
A/542
A/542
R10213
R10213
n=5
A/542
A/542
A/542
797
R10301
P4inf
P4inf
P4inf
M1inf
M1inf
M1inf
M1inf
M2inf
M2inf
M3inf
M3inf
D2inf
D3inf
D4inf
P3inf
P3sup
50.1
40
41.7
55
43
45.4
55
57
50
63
60.466.1
21.3
33
63
34.5
40
35
30
29.5
34
23.2
30
29
30
30
2733
13.2
22
29
42.5
33
29.6
35
30
31.8
31
26
34
28.8
32
30
26
17
29
50
30
35
22
34
35
12
16
22
50
posterior ossicones are palmate. One posterior straight

portion was found, which presents rare humps and a
discrete groove (Masol 2 R10479, diameter 85 80 cm).
According to the diagnosis, the teeth (Table 4) are thick
with rough enamel (e = 3 mm) and very strong styles (Arif
and Vos, 1989; Churcher, 1978; Gaur, 1987; Harris, 1976).
Two mandibles were assigned to adult individuals (Masol
6 R10787 and R10213): the ramus height under M1 is
between 70 and 75 mm, with a thickness between 40 and
47 mm. The molars (raw length = 160 mm) are more slender than the jawbone A/795 from the Centre of Advanced
Study Museum (Panjab University), which bears the cheek
deciduous teeth and the rst true molar (Badam, 1973).
The body skeleton and limbs are heavy (Colbert, 1935). The
postcranial skeleton bones are a distal humerus (Masol 6
R10715), an ulna and a proximal tibia, a talus and a navicular (Masol 13 R10303); on the latest, the articulation
is oblique compared to the bone axis. The medial crest is
very low and forms a continuous, oblique line; the calcaneus facet is broad, as are the two facets for the metatarsal
(Masol 13 R11455).
Camelidae: (three fossils) Camelus sivalensis was rst
described in the Tatrot Formation from Pakistan (Gaur,
1981) and in the Pinjor Formation (Gaur et al., 1984).
The collection is limited with two teeth and one phalanx.
The fourth upper premolars from Masol 1 are complete and worn at. The rst phalanx is broken (Masol
ind. R11439) close to the Falconer observation, proximal
width = 28.4 mm (3.94 inch), length = 19.6 mm (1.61.9
inch) (measurements in inches, Falconer and Cautley,
1846).
Cervidae: At least three genera of cervids are present
in the Siwalik: Cervus punjabiensis (Badam, 1979) Axis or
Dama-like (Dennell et al., 2006) and Rucervus or Muntjac
like (Gaur, 1981). Many fragments of antlers, dental and
postcranial remains were found. The antler from Masol 1
was associated with the bovid striated diaphysis. The beam
and the brow tine are a retting of several antler fragments
and the distal part of the beam shows a at tine (Masol 1
R10020, diameter = 35 mm, height of rst tine is 70 mm,
diameter: 20 mm). This middle size antler, ornamented
by subparallel grooves, can be related to C. punjabiensis

(Badam, 1973).
A shed antler (Masol 1 R10483) and a massacre antler
enriched the collection as an axis morphology with a
very open angle between the brow tine and the thin
and smooth beam without ornaments. This morphology is also observed in the Pabbi Hills (Dennell et al.,
2006). Teeth are often broken and postcranial skeleton
is abundant. Waiting for new discoveries, the nomenclature is still open, but these two kinds lived in the
sub-Himalayan oodplain before 2.6 Ma. Several long
bones are well preserved, in particular, a distal tibia (Masol
3 R10059, distal width = 32.3 mm, distal anteroposterior
diameter = 30.2 mm). The metapodial bones are all broken.
Suidae: Suids are rather rare (six remains) from Masols,
1, 5, 6 and 8, although they have been already indexed
in different studies since Falconer, who described most of
the species. More discussions were conducted on the comparison between Potomochoerus and Prototamochoerus, an
Asian genus. Three hemimandibles, an upper jaw, isolated
teeth and postcranial bones, help us recognize two genera
in Masol.
Propotamochoerus: The hemimandible from Masol 3
R10266 bears three premolars and three molars (Fig. 8). The
diastema is broken, but the inner large section of the canine
root is visible and corresponds to a male. (Masol 3 R10266:
length of cheek teeth is 110 mm, molar row is 67.6 mm).
The fourth premolar is large and square. The third molar is
very short and simple with ve cusps as Propotamochoerus.
The horizontal ramus is thin (height = 45 mm, thickness
under M1 = 28.5 mm).
Sus sp.: the second hemimandible, Masol 5 R10476 is
a verrucosus type. The lower row bears four premolars
and three molars (Fig. 8). The last molar is absent and the
diastema is broken. It is a female without a canine root.
The molar size is similar to Prototamochoerus, but the premolars are less developed than on the rst specimen. The
rst premolar is located near the canine tooth. The horizontal ramus is low and thick (thickness under M1 = 42 mm)
with the typical bulges on the mesial and lateral side.
The measurements of the lower second molar are close to
370
Fig. 8. Sus (up) and Propotamochoerus (below), left hemimandible (scale

10 cm) (picture A. Dambricourt Malass).
Fig. 8. Sus (en haut) et Propotamochoerus (en bas), hmimandibule gauche
(chelle 10 cm) (photo A. Dambricourt Malass).
Javanese S. brachygnatus (Bouteaux et al., 2007) or Chinese

S. lydekkeri (Echassoux et al., 2008). Without the lower M3,
it is difcult to report it as S. falconeri. Postcranial bones,
such as the tibia and talus, are robust.
Carnivora: In the Tatrot Formation, the carnivores are
said to be rare (Gaur, 1983) but many taxa were described
by different authors: Canis pinjorensis, Crocuta crocuta,
Pachycrocuta brevirostris, Felis, cf. Panthera uncia, Megantereon cultridens, cf. Canis cautleyi, and a herpestid, etc.
Carnivore remains in the collection are limited; there are
ve remains: four Hyenids and one Panthera sp. Their presence is in accordance with tooth marks observed on the
long bones of large bovids or hippos. One mineralized
coprolite was also discovered in Masol 9 (R11001).
Hyenidae: one retro-articular process of the temporal
bone with the temporo-mandibular joint (Masol 1 R10243),
one vertebra axis (Masol 5 R10289) and a canine tooth fragment (Masol 5 R10290) correspond to the morphology and
the size of Pachycrocuta (Fig. 9).
Panthera sp.: a left hemimandible (Masol 6 R10542)
is composed of three pieces collected in 2013 and 2015
(Fig. 10). The mandibular corpus is complete until the symphysis, but is broken at the beginning of the masseteric
part of the vertical ramus. The premolar crown is broken,
and the large canine crown is partially damaged. The size
of the dental alveoli corresponds to an adult animal. A
long wear facet due to the upper canine tooth is visible
on the distal part of the canine tooth that extends from the
apex toward the neck of the crown. The mandibular ramus
presents two premolars and the rst molar (carnassial). It
is thick: higher after the canine and lower and thicker at
the retromolar level with a at base. The short diastema
is visible between the canine tooth and the rst premolar P3. Three foramens are visible below the canine. The
edge of the symphysis is slightly convex. The alveolar shape
appears distinctly divergent in its canine arch: height of
the horizontal ramus = 30 mm, maximum length of lower
raw = 54 mm (Fig. 10).
Fig. 9. Hyena, basal view of a right TMJ, size of Pachycrocuta (picture A.

Dambricourt Malass).
Fig. 9. Hyena, vue basale de los temporal droit, avec lATM, taille de
Pachycrocuta (photo A. Dambricourt Malass).
4.3. Reptiles
More than four types of turtles could be described in
these localities and include the large freshwater turtle Geoclemys and the very large terrestrial Colossochelys. This
diversity will be analyzed in a future research program.
The crocodilians have been described in the Quranwala
zone (Sahni and Khan, 1968). We have collected 29 fossils,
teeth, mandibles and dermic plates of Crocodylus penjabensis, according to the upper jaw Masol 6 R11032 (Fig. 11) and
one Gavialis tooth at Masol 1.
5. Expected species
Rodentia: fossil specimens are known in Tatrot
(Patnaik, 2001, 2012). They have been evidenced in the
collection by tooth marks on nine bones of large bovids
at Masols 3, 6 and 13, which can be assigned to Rhizomys,
but no fossils have yet been recorded.
Primates:
Cercopithecidae: the Procynocephaplus was rst
recorded in 1848 in the Pinjor beds near Dehra Dun,
120 km southeast of Chandigarh, and was called P. subhimalayanus. A well-preserved mandible has been recovered
in the 60th, 10 km south of the limit of the Quranwala
zone near Bunga (Verma, 1969), in the Pinjor Formation
close to Khetpurali section were Tatrot is exposed. Patnaik
and Nanda (2010) suspect a Tatrot origin. The fossil was
assigned to P. pinjaury. The dental microwear was recently
analyzed to evaluate the dietary proclivities (Williams and
Holmes, 2012). Verma did not exclude Procynocephalus
from the Latest Pliocene of the Siwaliks due to the presence
371
Fig. 10. Panthera, left hemimandible. A. Occlusal view. B. External lateral view. C. Inner lateral view. (picture A. Dambricourt Malass).
Fig. 10. Panthera, hmimandibule gauche. A. Vue occlusale. B. Vue latrale externe. C. Vue latrale interne (photo A. Dambricourt Malass)
Fig. 11. Right hemimandible Masol 6 R11032, Crocodylus penjabensis (scale 10 cm) (picture A. Dambricourt Malass).
Fig. 11. Hmimandibule droite Masol 6 R11032, Crocodylus penjabensis (chelle 10 cm) (photo A. Dambricourt Malass).
of P. wimani in China within the Hipparion assemblage of

the Honan Province dated to Late Pliocene (Howells and
Tsuchitani, 1977). Procynocephalus were also associated
with cut marks and an enigmatic hominoid fragmented
jaw with two worn molars in Longgupo deposits (Wushan
County, Gansu Province, central China). This assemblage
has been recently dated to 2.48 Ma (Han et al., 2015).
Hominin: the only Hominin known in Asia is the genus
Homo, whose presence is now expected at least 2.48 Ma
in Longgupo cave, central China (Han et al., 2015). Strong
teeth from Longgudong cave, near Longgupo in Jianshi
County, Hubei Province, Central China and dated around
2 Ma (Matuyama period, Hou and Zhao, 2010), have been
assigned to a hypothetical Meganthropus (Zhang et al.,
2004). No fossil has been recovered in Masol, but, as for
the Rodents, their marks have been identied indubitably
on three bovid bones made by akes and/or choppers in
quartzite with intentional gestures (Dambricourt Malass
et al., 2016a, b).
6. Discussion
The vertebrate taxa of the different localities are
grouped according to the lithostratigraphy (Abdessadok
et al., 2016; Chapon Sao et al., 2016b; Tudryn et al.,
2016) and then compared with other Late Pliocene localities in Himachal Pradesh (Nanda, 2002) and in the Pabbi
Hills, Pakistan (Dennell et al., 2006) (Table 5). The Elephantidae and Bovidae are always abundant, but pygmy
hippos (30%) associated with crocodilians and freshwater
turtles, conrm the main uvial environment as lithostratigraphic (Badam, 1987; Gaur and Chopra, 1984; Nanda,
2002; Patnaik, 2012; Pilgrim, 1910; Sahni and Khan,
1968).
Boselaphini, Propotamochoerus, Camelus, Equus and
Merycopotamus have been collected in the basal sequence.
Panthera, Dorcatherium and Hipparion (teeth and/or
mandibles) identied in the middle sequence are not necessarily excluded from the basal one.
372
Table 5
Distribution of genera/species (% NISP) from Masol, the Tatrot Formation from Chandigarh anticline (Gaur and Chopra, 1984), the Saketi/Tatrot Formation
(Nanda, 2002) and the Plio-Pleistocene beds from Pabbi Hills (Dennell et al., 2006). Masol localities are grouped by stratigraphic units (Chapon Sao et al.,
2016b).
Tableau 5
Distribution des espces (% NISP) Masol, dans le Tatrot de lanticlinal de Chandigarh (Gaur et Chopra, 1984), Saketi/Tatrot (Nanda, 2002) et les couches
plio-plistocnes des Pabbi Hills, Pakistan (Dennell et al., 2006). Les localits de Masol sont groupes selon la rpartition stratigraphique (Chapon Sao et al.,
2016b).
Taxon
Masol
Basal
sequence
Masol Middle
sequence base
Masol Middle
sequence up
Tatrot
Gaur
Chopra
Saketi/Tatrot
Nanda
Pabbi Hills
Dennell
Elephas hysudricus
Stegodon insignis
Anancus perimensis
Pentalophodon khetpuraliensis
Hippohypus tatroti
Propotamochoerus sp.
Potamochoerus sp.
Sus brachygnatus/verucoid
Hexaprotodon sivalensis
Hemibos triqueticornis
Leptobos
Bubalus sp.
Proamphibos kashmiricus
Boselaphini/Duboisia
Reduncini/Sivacapra
Gazella
Hippotragini/Hippotragus sp.
Damalops palaeindicus
Dorcatherium nagrii
Camelus sivalensis
Sivatherium giganteum
Rucervus simplicidens
Axis Dama-like
Cervus penjabensis
Anthracothere
Equus sivalensis
Cormohipparion theobaldi
Hipparion antilopinum
Canid
Pachycrocuta sp.
Panthera
2.8
18.6
1.7
31.1
17.5
1.7
2.3
3.4
7.3
0.6
1.7
1.1
7.9
0.6
0.07
0.07
1.1
6.5
8.1
1.6
35.5
21
6.5
1.6
12.9
1.6
3.2
1.6
2.7
20
0.9
1.8
21.8
15.5
1.8
4.5
3.6
0.9
11.8
0.9
8.2
0.14
0.07
0.9
0
X
X
X
X
X
X
X
X
suid
X
X
X
X
X
X
X
X
X
Stegodon and Elephas are represented in a proportion of 20/3. This co-occurrence is signicant from a
biochronological point of view. Indeed the rst appearance
of Elephas hysudricus (and Cervus) has been dated to c. 2.7
Ma in the Mangla-Samwal area in Jhelum basin, Pakistan
(Hussain et al., 1992). According to the Biostratigraphic
zones based on magnetostratigraphy and fauna, Elephas
hysudricus is referred to as the Equus sivalensis Biostratigraphic interval-zone beginning around 2.6 Ma (Nanda,
2002). The Quranwala zone has been dated just below
the Gauss-Matuyama magnetic reversal in the Patiali Rao
(Ranga Rao, 1993; Ranga Rao et al., 1995; see Chapon Sao
et al., 2016a). These results conrm the former proposition
of Colbert (1951) and Sahni and Khan (1964, 1968), who
dated the Tatrot Formation to the Late Pliocene. Stegodon
is well known in the Siwaliks since 4 Ma (Middle Siwalik),
and was associated with the Elephas planifrons Biostratigraphic interval-zone since 3.6 Ma, but it is also associated
with E. hysudricus during the Middle Pleistocene in the
Post-Siwalik Narmada beds (Central India).
The anthracotheres mostly refer to the Middle Siwalik, but Sahni and Khan (1968) collected fossils in the
Quranwala zone and then at the very beginning of the Pinjor Formation (Early Pleistocene). Later, a maxillary from
X
X
X
X
the Mangla-Samwal anticline, Pakistan, dated between 2.4

and 2.65 Ma, conrmed the presence of Merycopotamus
until the end of the Pliocene in the sub-Himalayan oodplain (Hussain et al., 1992). Anthracotheridae was also
found in the Pabbi Hills pre-1.8 Ma (Dennell, 2008). Finally,
Merycopotamus is listed in the Equus sivalensis Biostratigraphic interval-zone where the last appearance is dated close
to 2.5 Ma (Nanda, 2002). This taxon is associated with the
faunal group of the uppermost part of the Tatrot Formation (called the Saketi Formation since the studies of this
fossiliferous zone in Himachal Pradesh, Nanda, 2013).
The association of Hipparion and Equus in the Quranwala
zone is coherent with the Upper Siwalik biozone: Equus
sivalensis appears before 2.6 Ma and Hipparion antilopinum
disappears there around 1.5 Ma. In Jammu and Potwar, the
horses are denitely associated with Hipparion (Barry et al.,
1982), and the end of the Saketi (or Tatrot) Formation is
characterized by the scarcity of the Hipparionines (Nanda,
2002).
Hexaprotodon sivalensis is well known since the Lower
Siwalik, but then disappeared in many localities in the
early Pinjor Formation (Dennell, 2005; Shoshani and Tassy,
2005). It disappeared in the Pakistani Siwalik group around
3.5 Ma (Dennell et al., 2006), but was still present in the
Pabbi Hills due to the favorable water environments with

deep rivers (Dennell, 2005), and in the South of the subHimalayan oodplain around 2.6 Ma as evidenced in the
Masol Formation.
Suidae like Propotamochoerus appears 4 Ma ago in the
Middle Siwalik (Barry, 1995); P. hysudricus is observed in
the Jammu region (Agarwal et al., 1993) and sometimes is
mentioned in the Pinjor Faunal list. In China, this last one is
still present at the end of the Lower Pleistocene (Echassoux
et al., 2008). The faunal list of the Equus sivalensis Biostratigraphic interval-zone includes a great number of suid
species; nevertheless, the Sus genus cannot be identied
without the lower last molar. Within the Masol assemblage,
the Suidae are scarce (less than 2%); however, a complete
hemimandible with three premolars, a large lower P4 and
a simple M3 can be assigned to Propotamochoerus without
problem, as this genus is characteristic of Tatrot.
Camelus sivalensis rst appears in the Middle Siwalik in Pakistan (Barry et al., 1982) and at the top of the
Saketi Formation (Nanda, 2002). Camelus and Hexaprotodon
are attributed to the Elephas planifrons Biostratigraphic
interval-zone.
S. giganteum appears in the Siwalik Group after 2.9 Ma
(Barry, 1995). This is considered a common taxon in the
Elephas planifrons Biostratigraphic interval-zone from the
uppermost part of this group and developed during the
next Equus sivalensis Biostratigraphic interval-zone.
Bovidae are abundant but the determination is not easy
without crania and horncores. The taxonomic keys used
for Masol collection allow the assignation to the tribes
level based on the dental material. The connection between
these observations and the historical nomenclature needs
more data and will be addressed in future work. The presence of Proamphibos or Probison is not yet possible in this
collection. Nevertheless, the large bovids can be assigned
to Hemibos related to the Equus sivalensis Biostratigraphic
interval-zone. The mid-sized bovid is close to the Hippotragini. On the other hand, the Boselaphini is very similar
to the specimen described in Southeast Asia and Sivacapra,
and is also well known in the Pinjor Formation. As these
mid-size bovids are not yet connected to the Tatrot Formation, the Masol assemblage can testify that these bovids
were clearly associated with the faunas of their uppermost
deposits and appear around 2.6 Ma.
Tragulidae are known from the Miocene. Dorcatherium
nagrii is the most advanced species of this family and very
similar to the current genera Tragulus, which was already
described in the Tatrot Formation (Gaur, 1992) and can be
added to the reference list of the Equus sivalensis Biostratigraphic interval-zone.
Cervidae are not completely described; in particular, due
to the postcranial remains, but with typical antlers, we can
assume two genera in the Masol localities: a mid-sized
Cervus punjabiensis and an Axis or Dama-like small cervid.
The latter may be compared with Rucervus simplicidens,
which is described in the literature. The rst appearance of
Cervus is described in the Mangla-Samwal anticline (Pakistan), with Equus sivalensis and Elephas hysudricus (Hussain
et al., 1992).
The localities of the Masol anticline present many similarities with the Mangla-Samwal anticline and can be
373
correlated to the Equus sivalensis Biostratigraphic intervalzone. This well dated association gives a good indication of
the appearance of many taxa and could link the ManglaSamwal area, Jammu area and the uppermost part of the
Tatrot Formation (the Saketi Formation in Himachal and
the Siwalik Frontal Range).
The carnivores are rare in the fauna of Masol; primate
specimens, such as Procynocephalus pinjaury, have not been
yielded in this sector of the Siwalik Frontal Range and the
lack of Rhinocerotidea is noted. This fact is observed in the
Tatrot of Pabbi Hills 300400 km northwest of Chandigarh
in Pakistan (Dennell et al., 2006; Jablonski, 2004). Nevertheless, those data cannot be seen as denitively signicant
of a biochronological frame because rhinos and carnivores
are abundant in the Pinjor Formation; they had necessary
ancestors in the Asian Late Pliocene. Yet, 300 km northeast
of the Chandigarh anticline, in the uppermost Himalayan
basin of the Sutlej River, the Zanda Basin (Tibet) provided
Coelodonta thibetana dated to 3.7 Ma, a rhino genus that
adapted to climate cooler than that in the sub-Himalayan
oodplain (Deng et al., 2011).
This new fossiliferous collection of the Tatrot Formation
coming from the Masol sector has been recorded because
of some cut marks and many stone tools associated with
the same outcrops. The cut marks show, without any doubt,
the presence of a very old Homo-like Hominin (Dambricourt
Malass et al., 2016b). This presence in a sub-Himalayan
Equus sivalensis Biostratigraphic interval-zone opens new
horizons for the origins of taxa engaged in an evolutionary dynamic of appearance-disappearance. This dynamic is
the prelude of the consequences due to the rst Quaternary
glaciation closely linked to the Tibetan Plateau uplifting. For
the prehistoric approach, one critical point is the examination of the faunal association among which this Homo-like
Hominin evolved, the origins of all these species since the
Miocene and the direction of their migrations. Genera, such
as Elephas, Stegodon, Hexaprotodon and Hipparion, Equus,
or families, such as Cervidae, derived from Asian Miocene
ancestors having no exchange with Africa, have been noted
before the Pliocene (Wang et al., 2013). For instance, Equus
appears in East Africa around 2.3 Ma and Elephas around
1.9 Ma, whereas Stegodon, Hexaprotodon and Cervidea
migrated toward South Asia. Many widespread taxa, such
as Sivatherium, Bovidae, Tragulidea and Suidea, are known
during the Miocene or Pliocene in Asia, but also in Africa
and in Europe. For the bovids, except Boselaphini, the genera Bovini, Hippotragini, Alcelaphini, Reduncini and Antilipini
can be compared between Asia, South Africa, East Africa,
and mostly Bovini with Europa. The fossil bovid record provides evidence for greater biological continuity between Africa
and Eurasia in the late Miocene and earliest Pliocene than
is found later in time (Bibi, 2011). Thus, African antelopes
were the ancestors of sub-Himalayan Hippotragini dated
to 3 Ma (Martinez-Navarro, 2010). Later Tragulidae and
Sus migrated toward South Asia. Large bovids, such as Bos,
are probably the descents of the Late Pliocene rather than
the Early Pleistocene species from Upper Siwalik. Indeed,
they are observed as early as the Lower Pleistocene on
the main islands. In the same way, rare carnivores developed later and colonized the greatly expanded Sunda as
Elephas, Bubalus, Epileptobos and Boselaphini (Bergh et al.,
374
2001; De Vos, 1996; De Vos and Long, 2001; Mishra et al.,

2010).
7. Taphonomy of the Herbivora
No bones have been recorded in anatomical connection, as all are mineralized. Differential preservation of
864 fossils (Table 6) shows that the axial and appendicular skeletons are represented approximatively in the
same proportions: respectively, 47.2% and 52.8%, with the
greatest frequency for the shaft, the ribs, the teeth and
the mandibles. For the appendicular skeleton without the
extremities, the anterior (49%) and posterior (51%) skeletons are also in the same proportion. The bones of the
extremities are often small and represent 9.2% of the skeleton, a good representation rather rare. These percentages
reect the equitable preservation of the different parts of
the skeletons despite the lack of connection. This could be
interpreted as the result of successive processes of conservation since the dead of the animal, taking into account
the lithostratigraphic context characteristic of the subHimalayan oodplain (Abdessadok et al., 2016; Tudryn
et al., 2016). This one was regularly exposed to the seasonal monsoon and the changes of the hydraulic energy
from the Himalayan rivers. The layers of quartzite cobbles
interstratied within silts illustrate these changes in the
hydraulic regime. They could correspond to fast and powerful ooding, taking away herds of herbivores from all sizes,
with their carcasses being accessible for the rare scavengers
after the water receded.
The alteration of the fossils is heterogeneous, even
within a single locality; often some of the bones have
been fractured, especially the long bones. They present
important cracking due to geological pressure and tectonic
activity; this is the case for the very large, well-preserved
herbivores, such as Proboscideans and Hippopotamidae
Fig. 12. Fracturing of a mineralized bone from a very large Herbivora

caused by tectonic pressures of the Chandigarh anticline (picture A.
Fig. 12. Fracturation dun os minralis de trs grand herbivore provoque par les pressions tectoniques de lanticlinal de Chandigarh (photo A.
(Fig. 12). In contrast, the bones of cervids or small bovids

are rather intact. The surface is frequently scaled according
to the areas considered. Some are also affected by several
phases of landll and can be covered by very hard sandstone or gravel (Fig. 13). In some localities, the layers rich in
manganese and iron have colored the bony tissues as brown
or black (Fig. 13C). In other areas, the fossils are clear (beigepink) and, in the medullar cavity, they are full of calcite
crystals. This diversity of fossilization allows researchers to
recognize when the carcasses were buried quickly, as very
few have moved since their depository, and why no complete skeletons have been recovered. Finally, some bones
show different types of traces on their surfaces, which can
be attributed to biological alterations, teeth marks and possible trampling. One isolated trace on a big diaphysis of
Proboscidean resembles a long cut mark with two parallel
Table 6
Proportional abundance of various skeletal elements in the Masol fossil assemblage. Small herbivora: Cervidae, Duboisia, Sivacapra, Suidae, Tragulidae,
Large Herbivora: Bovidae, Equidae, Camelus, Very Large Herbivora: Proboscidean, Hippopotamidae, Sivatherium, Anthracoteriidae.
Tableau 6
Abondance relative des diffrents lments squelettiques de lassemblage fossile de Masol. Petits herbivores : Cervidae, Duboisia, Sivacapra, Suidae,
Tragulidae, grands herbivores : Bovidae, Equidae, Camelus, trs grands herbivores : Proboscidiens, Hippopotamidae, Sivatherium, Anthracoteriidae.
Maxillary
Mandible
Teeth
Skull
Vertebrae
Ribs
Scapula
Humeri
Radius/ulnae
Pelvis
Femora
Tibia
Astragali
Calcanei
Metapodial
Metacarpal
Metatarsal
Phalanges
Shaft
Total
Little herbivora
Large herbivora
Very large herbivora
Total
2
14
12
18
2
1
1
3
4
1
6
5
2
1
3
1
5
81
2
19
50
17
21
34
3
16
13
5
11
14
12
6
11
6
11
6
127
384
10
24
103
15
24
41
10
11
18
15
17
6
6
3
7
88
399
14
57
165
50
47
75
14
28
34
24
29
26
23
6
14
7
17
14
220
864
1.6
6.6
19.1
5.8
5.4
8.7
1.6
3.2
3.9
2.8
3.3
3.0
2.7
0.7
1.6
0.8
2.0
1.6
25.4
99.8
375
Fig. 13. Different types of fossilisation. A. Masol 2 East, scapula of Proboscidean partially covered by gravels (circle). B. Masol 2 West, cervid antler in
sandysilt. C. Masol 2 South West, skull of Sivatherium strongly colored by manganese (Photo A. Dambricourt Malass).
Fig. 13. Diffrents types de fossilisation. A. Masol 2 est, omoplate de proboscidien partiellement recouverte de graviers (cercle). B. Masol 2 ouest, bois de
cervid dans des limons sableux. C. Masol 2 sud-ouest, crne de Sivatherium fortement teint par le manganse (Photo A. Dambricourt Malass).
Fig. 14. Fossilized marks. A. Masol 3 R11154, rib with numerous damages with random striations. B. Masol 8 R10893, ulna, Hexaprotodon, marks made by
the pressure of a canine tooth (arrows). C. Masol 8 Proboscidean diaphysis, long mark with two microgrooves (arrow). D. Enlargement of the trace in the
white rectangle Fig. 14C (Photo A. Dambricourt Malass).
Fig. 14. Marques fossilises. A. Masol 3 R11154, cte avec de nombreuses striations alatoires. B. Masol 8 R10893, ulna, Hexaprotodon, marques faites
par la pression dune canine de carnivore (ches). C. Masol 8 grande diaphyse de Proboscidien avec une longue marque forme de deux micro-rainures
parallles. D. Agrandissement de la trace dans le rectangle blanc de la Fig. 14C (Photo A. Dambricourt Malass).
376
microgrooves on a minimum length of 3 cm. Nevertheless,

the cortical bone is destroyed, with typical carried features
(Fig. 14C and D). However, at least three fossils of Bovidae,
a tibia shaft (Masol 1 R10084) about the size of a Leptobos, a
distal metacarpal (Pichhli Choe R10286) and a large splinter (Masol 13 R10298) present a set of traces that cannot be
confused with trampling (Dominguez-Rodrigo et al., 2010).
These marks have been rigorously identied as intentional
cut marks made by a sharp edge of a quartzite cobble (or
ake) through an experimental protocol, and have been
compared with collections of animal marks (Dambricourt
Malass et al., 2016b).
8. Conclusion
The faunal list of the Masol Formation based on the collections from the 12 localities matches the results of former
studies on the Tatrot Formation (The Late Pliocene of the
Upper Siwalik sub-group). This faunal association is classic
in this sector of the Siwalik Frontal Range and is directly
linked with water bodies, as witnessed by the strong presence of H. sivalensis in all the paleontological-archeological
localities, which becomes rare and then absent in the Pinjor
Formation. Elephantidae correspond to 40% of determinate
bones, with the domination of the S. insignis, while Elephas is rare (2% of determinate teeth). Bovids represent
25% of mammals and correspond to at least ve taxa; the
most frequent is Bubalus, and the group Leptobos-Hemibos
is also well represented by many mid-sized bovids (Hippotragini, Reduncini, and Boselaphini). The giant girafd S.
giganteum is observed in many localities (skull, ossicone,
teeth and limb bones), as is Camelus. Cervids and suids
are also observed with a Dama-like mid-sized cervid and a
Verrucosus-like suid. The scarce anthracothere M. dissimilis
has been found again (one molar), along with a rare carnivore with a Panthera (one hemimandible). Until now, no
Rhinoceros or Procynocephalus have been described in the
collections of the 12 localities. The exceptional occurrence
of Hipparion with Equus is characteristic of this period in the
sub-Himalayan oodplains. This corresponds to the lower
part of the E. sivalensis Biostratigraphic interval-zone, following Nandas framework (Nanda, 2002). A preliminary
overview of the biochronology of the so-called transitional fauna showed the origins of herbivores; most
originated from Miocene Asian species (Hipparion, Equus,
Hexaprotodon, Elephas, Stegodon, Cervidae), and the migrations from the sub-Himalayan oodplain occurred toward
South Asia. The presence of a Homo-like Hominin evidenced
by uncontestable cut marks on bovid bones from the
Quranwala zone opens new horizon for the understanding of the oldest Asian sites, such as Riwat (at least 2
Ma, Pakistan, stone tools associated with fauna), Longgupo
(cut marks, stone tools, unknown Hominoid, 2.48 Ma) and
Longgudong (unknown Hominin assigned to Meganthropus
Matuyama chron), both located in central China long before
the Olduvai paleomagnetic event (1.8 Ma).
Acknowledgments
The Indo-French Research Program, Siwaliks, is under
the patronage of Professor Yves Coppens, College of France
and Academy of Sciences, since 2012, it has been supported

by the French Ministry of Foreign Affairs during three years
(201220132014); in 2011 by the ATM grant (Transversal
Action of the Museum) of the National Museum of Natural History (Department of Earth Sciences), in 2006, 2007
and 2011 by the Prehistory Department of the National
Museum of Natural History, Paris; we are thankful to the
Archaeological Survey of India and to the Department of
Tourism, Cultural Affairs, Archaeology and Museums of
Punjab Government for survey permit, to the Embassy of
India in Paris and to the Embassy of France in New Delhi,
for their administrative support. We thank Pr. R.S. Loyal,
Chairman of the Geology Department, Panjab University, Chandigarh, for welcoming us to the Paleontological
Gallery. We are grateful to the Sarpanch of Masol village for
his hospitality. We especially thank B.L. Badam for his comments on the Indo-French cooperation and Robin Dennell
for his precise revision and encouragements to pursue this
eldwork. We pay a special tribute to Jean-Francois Jarrige
(19402014), Former Director of the Guimet Museum, the
French National Museum of Asian Arts, and General Secretary of the Excavations Commission of the French Ministry
of Foreign Affairs.
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C. R.

Palevol 15 (2016) 359378

Contents lists available at ScienceDirect

Comptes Rendus Palevol

Human palaeontology and prehistory

The faunal assemblage of the paleonto-archeological

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

Le programme de recherche franco-indien Siwaliks poursuit ses investigations dans la

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

Nevertheless, in 1926, Black did not hesitate to publish

The Plio-Pleistocene boundary was suspected at the

3. The Chandigarh anticline and the Quranwala

Fossils from the Chandigarh anticline are stored and

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

units c3 (members ED), s3 and s4, at Masol 1, Masol 2,

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

been recorded at Masol 13. Masol 9 is a small circus formed

part of Masol 6 cliffs, which extend from s6 to s13 and c7

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

of ve (number of plates per 10 cm of anteroposterior

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

The I1 and I2 present almost the same section size (Masol

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

E. sivalensis: the isolated lower M2 (Masol ind. R11459)

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

DVD third lobe

posterior ossicones are palmate. One posterior straight

by subparallel grooves, can be related to C. punjabiensis

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

Fig. 8. Sus (up) and Propotamochoerus (below), left hemimandible (scale

Javanese S. brachygnatus (Bouteaux et al., 2007) or Chinese

Fig. 9. Hyena, basal view of a right TMJ, size of Pachycrocuta (picture A.

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

of P. wimani in China within the Hipparion assemblage of

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

the Mangla-Samwal anticline, Pakistan, dated between 2.4

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

Pabbi Hills due to the favorable water environments with

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

2001; De Vos, 1996; De Vos and Long, 2001; Mishra et al.,

Fig. 12. Fracturing of a mineralized bone from a very large Herbivora

(Fig. 12). In contrast, the bones of cervids or small bovids

Very large herbivora

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

microgrooves on a minimum length of 3 cm. Nevertheless,

and Academy of Sciences, since 2012, it has been supported

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

2016. Geology and geomorphology of Masol paleonto-archeological

A.-M. Moigne et al. / C. R. Palevol 15 (2016) 359378

Lydekker, R., 1879. Further notices of Siwalik mammals. Records of the

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