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 flakes in quartzite collected on the outcrops. Nine fieldwork seasons (2008–2015) 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 findings are significant for the discovery of Homininae in Siwaliks.
Le programme de recherche franco-indien « Siwaliks » poursuit ses investigations dans la zone Quranwala de la formation de Masol, connue depuis les années 1960 pour sa faune de la fin du Pliocène, également nommée faune de transition Plio-Pléistocène. Notre étude paléontologique s’est imposée à la suite de la découverte de traces de découpe sur des fossiles proches de choppers et d’éclats en quartzite. Neuf campagnes de terrain (2008–2015) sur 50 hectares de paysage particulièrement accidenté (ravins et petits plateaux) ont permis d’identifier 12 localités où au moins un chopper était mêlé à des fossiles, lesquels totalisent 1500 spécimens. Leur étude montre que les mammifères les plus abondants sont les Proboscidiens, avec Stegodon insignis et Elephas en bien moins grand nombre. Les bovidés présentent la plus grande variété, avec six tribus de la plus petite à la plus grande taille. Hexaprotodon sivalensis vient en troisième position. Sivatherium giganteum est visible dans plusieurs localités, Merycopotamus dissimilis dans une seule. Deux types de cervidés au moins ont été observés. La faune de transition est attestée par la co-existence de Stegodon et Elephas hysudricus, d’Hipparion antelopinum et d’Equus sivalensis. Les fossiles de tortues sont abondants avec la forme terrestre géante Colossochelys, et celle d’eau douce, Geoclemys. En comparaison, les prédateurs sont peu représentés (crocodile), voire rares (hyène, félin), tandis que les primates (Procynocephalus) n’ont pas encore été observés. L’assemblage faunique correspond bien aux faunes de transition Plio-Pléistocène également décrites dans les Pabbi Hills (Pakistan) ; elle caractérise le début de la biostratigraphie de la zone intervalle Equus sivalensis, qui s’étend de 2,6 Ma à 600 ka. La répétition systématique des prospections a donc permis la collecte de taxons rares comme Crocuta (2010), Merycopotamus dissimilis (2014) et un grand félin (2015). Ces derniers résultats sont significatifs pour la découverte d’Homininae dans les Siwaliks.
Siwalik Frontal Range, Sub-Himalayan floodplain, Late Pliocene, Tatrot faunal assemblageChaîne frontale des Siwaliks, Plaine alluviale sous-himalayenne, Pliocène final, Assemblage faunique de TatrotpresentedHandled by Anne Dambricourt MalasséIntroduction
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 field season of the Indo-French Program of Research ‘Siwaliks’ (Fig. 1) (Dambricourt Malassé, 2016). In 2009, a shaft of bovid tibia was collected in the Quranwala zone of the Masol Formation, near choppers and flakes, 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 Malassé et al., 2016a and Dambricourt Malassé et al., 2016b).
Eight years of surveys in the Masol inlier (2008–2015) recorded fossils and artifacts in twelve paleontological-archeological 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, Chapon Sao et al., 2016b, Dambricourt Malassé et al., 2016a, Dambricourt Malassé et al., 2016b, Gaillard et al., 2016, Gargani et al., 2016 and 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 finding hominins specimens.
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 field (the Potwar Plateau in Pakistan, the Kashmir Valley, Lesser Himalayan foothills and the Siwalik Frontal Range in India). They produced the first important collections stored at the Geological Survey of India, Kolkata, in London (Falconer and Cautley, 1846, Falconer, 1868, Lydekker, 1879, Lydekker, 1883, Pilgrim, 1910 and Pilgrim, 1939), and in USA at the American Museum of Natural History (Brown et al., 1924, Colbert, 1935 and Gregory et al., 1938) and the Yale University Museum (de Terra and Teilhard de Chardin, 1936). Among new species, the discovery of the first 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 Malassé et al., 2016a).
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 first 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 Plio-Pleistocene boundary was still unclear; the Tatrot and Pinjor Formations belong to the Upper Siwalik sub-group (Pilgrim, 1910 and Pilgrim, 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 Pliocene–Lower 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 first 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 South-east 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 and 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 and Gilbert et al., 2014).
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 1950–60s 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).
The Plio-Pleistocene boundary was suspected at the uppermost level of the Quranwala zone, a fossiliferous horizon of the Tatrot Formation as defined 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, Nanda, 1994, Nanda, 1996 and Nanda, 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 flowing 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 and Ranga Rao et al., 1995; details in Chapon Sao et al., 2016a). The Gauss/Matuyama reversal is now identified 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 80–100 m below this limit (Chapon Sao et al., 2016b, Sahni and Khan, 1964 and Sahni and Khan, 1968).
The fossil sample: stratigraphic origin and description
Fossils from the Chandigarh anticline are stored and exhibited in the Centre of Advanced Study Museum, Panjab University, Chandigarh. The fossils of the Indo-French program ‘Siwaliks’ have been stored, classified 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, Badam, 1979, Gaur, 1987 and 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 five 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 locality and (3) (the majority) during the eight field seasons (referenced Masol 1 to Masol 13 in the list) (Dambricourt Malassé 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 significant and allows comparisons (Table 1).
Stratigraphic origin
The base of the fossiliferous Quranwala zone has been defined at Masol 1 (Abdessadok et al., 2016 and 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 E–D of c3 unit), covered by fine 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 E–D), s3 and s4, at Masol 1, Masol 2, Masol 5 and Masol 7; Masol 12 is composed of c3 (members E–D) 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 sub-locality 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 E–D), 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 been recorded at Masol 13. Masol 9 is a small circus formed in eroded sandstones (unit s5) and silts (unit c6–1), and rounded by relic cliffs of the sequences s5–c6, 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.
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 and 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 and 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 identified 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 and 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 finer. 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 identified (Falconer and Cautley, 1846). The material is composed of one skull with the fourth molar, or the first 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 five (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 confirm 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 significant 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. 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 flat. 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 testifies 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 identified 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, Gaur, 1987, Gentry, 1978b and 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 flattened with a clear former hull. Hemibos and Bubalus present cavernicolous osseous ankles, with quadrangular or triangular sections with a hull well defined on the higher face. The horns develop first, slightly backwards, and then go down lower than the top of the frontal bone (Gaur and Chopra, 1984, Hooijer, 1958 and 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 filled 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 identified 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 (Köhler, 1993). These fossils are assigned to the subfamily Hippotragini (Sivatragus brevicornis (Pilgrim, 1939) is known in the Tatrot Formation). Postcranial bones are identified 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 classification among specialized animals (Köhler, 1993 and 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 confirms 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 (Köhler, 1993), without a sagittal sulcus (previously described in the Pinjor Formation as Gazella and Indoredunca).
Merycopotamus dissimilis: anthracotheres were first 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 identified.
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 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 and 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.
Giraffidae: 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 Giraffid, Sivatherium giganteum. The gigantic male Pleistocene giraffid with four ossicones was first described from the Upper Siwalik (Falconer and Cautley, 1836). The anterior ossicones are conical and the 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 and 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 first 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 first 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 flat. The first phalanx is broken (Masol ind. R11439) close to the Falconer observation, proximal width = 28.4 mm (3.9–4 inch), length = 19.6 mm (1.6–1.9 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 refitting of several antler fragments and the distal part of the beam shows a flat tine (Masol 1 R10020, diameter = 35 mm, height of first 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 floodplain 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 five cusps as Propotamochoerus. The horizontal ramus is thin (height = 45 mm, thickness under M1 = 28.5 mm).
Sussp.: 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 first specimen. The first 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 Javanese S. brachygnatus (Bouteaux et al., 2007) or Chinese S. lydekkeri (Echassoux et al., 2008). Without the lower M3, it is difficult 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 five 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).
Pantherasp.: 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 first molar (carnassial). It is thick: higher after the canine and lower and thicker at the retromolar level with a flat base. The short diastema is visible between the canine tooth and the first 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).
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.
Expected species
Rodentia: fossil specimens are known in Tatrot (Patnaik, 2001 and Patnaik, 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 first 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 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 identified indubitably on three bovid bones made by flakes and/or choppers in quartzite with intentional gestures (Dambricourt Malassé et al., 2016a and Dambricourt Malassé et al., 2016b).
Discussion
The vertebrate taxa of the different localities are grouped according to the lithostratigraphy (Abdessadok et al., 2016, Chapon Sao et al., 2016b and 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, confirm the main fluvial environment as lithostratigraphic (Badam, 1987; Gaur and Chopra, 1984, Nanda, 2002, Patnaik, 2012, Pilgrim, 1910 and 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) identified in the middle sequence are not necessarily excluded from the basal one.
Stegodon and Elephas are represented in a proportion of 20/3. This co-occurrence is significant from a biochronological point of view. Indeed the first 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 and Ranga Rao et al., 1995; see Chapon Sao et al., 2016a). These results confirm the former proposition of Colbert (1951) and Sahni and Khan, 1964 and Sahni and Khan, 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 the Mangla-Samwal anticline, Pakistan, dated between 2.4 and 2.65 Ma, confirmed the presence of Merycopotamus until the end of the Pliocene in the sub-Himalayan floodplain (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 definitely 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 and 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 sub-Himalayan floodplain 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 identified 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 first 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 tribe's 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 first 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 correlated to the Equus sivalensis Biostratigraphic interval-zone. This well dated association gives a good indication of the appearance of many taxa and could link the Mangla-Samwal 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 300–400 km northwest of Chandigarh in Pakistan (Dennell et al., 2006 and Jablonski, 2004). Nevertheless, those data cannot be seen as definitively significant 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 floodplain (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 Malassé 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 first 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., 2001, De Vos, 1996, De Vos and Long, 2001 and Mishra et al., 2010).
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 reflect 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 sub-Himalayan floodplain (Abdessadok et al., 2016 and 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 interstratified within silts illustrate these changes in the hydraulic regime. They could correspond to fast and powerful flooding, 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). 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 landfill 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 (beige-pink) 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 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 identified as intentional cut marks made by a sharp edge of a quartzite cobble (or flake) through an experimental protocol, and have been compared with collections of animal marks (Dambricourt Malassé et al., 2016b).
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 five 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 giraffid 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 floodplains. This corresponds to the lower part of the ‘E. sivalensis Biostratigraphic interval-zone’, following Nanda's 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 floodplain 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 (2012–2013–2014); 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 fieldwork. We pay a special tribute to Jean-François Jarrige (1940–2014), 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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Geological map of the Chandigarh Siwalik Frontal Range (Abdessadok, Fig. 1A in Chapon Sao et al., 2016, Sahni and Khan, 1968, modified).
Carte géologique de la chaîne frontale des Siwaliks de Chandigarh (Abdessadok, Fig. 1A in Chapon Sao et al., 2016, d’après Sahni et Khan, 1968)
Log of the Quranwala Zone of Masol Formation with paleontological-archeological localities (in Chapon Sao et al., 2016b).
Log de la zone Quranwala de la formation de Masol avec les localités paléonto-archéologiques (in Chapon Sao et al., 2016b).
Masol R10181, Stegodon insignis, right hemimandible with D4-M1 (scale 10 cm), SAAR fossil collection (picture A. Dambricourt Malassé).
Masol R10181, Stegodon insignis, hémimandibule droite avec D4-M1 (échelle 10 cm), collection de fossiles de la SAAR (photo A. Dambricourt Malassé).
Last lower molar Masol R10182, Elephas hysudricus, SAAR collections, SAAR fossil collection (picture A.-M. Moigne).
Dernière molaire inférieure, Masol R10182, Elephas hysudricus, collection de fossiles de la SAAR (photo A.-M. Moigne).
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).
A. Duboisia sp. Masol 5 R11267, M3 supérieure. B Sivacapra sp. Masol 2 R10109. hémimandibule droite avec M1 et M2. C. Merycopotamus dissimilis, Masol 5 R11286, M3 supérieure gauche. D. Hipparion, prémolaire, vue occlusale (photo A.-M. Moigne).
Sus (up) and Propotamochoerus (below), left hemimandible (scale 10 cm) (picture A. Dambricourt Malassé).
Sus (en haut) et Propotamochoerus (en bas), hémimandibule gauche (échelle 10 cm) (photo A. Dambricourt Malassé).
Hyena, basal view of a right TMJ, size of Pachycrocuta (picture A. Dambricourt Malassé).
Hyena, vue basale de l’os temporal droit, avec l’ATM, taille de Pachycrocuta (photo A. Dambricourt Malassé).
Panthera, left hemimandible. A. Occlusal view. B. External lateral view. C. Inner lateral view. (picture A. Dambricourt Malassé).
Panthera, hémimandibule gauche. A. Vue occlusale. B. Vue latérale externe. C. Vue latérale interne (photo A. Dambricourt Malassé)
Right hemimandible Masol 6 R11032, Crocodylus penjabensis (scale 10 cm) (picture A. Dambricourt Malassé).
Fracturing of a mineralized bone from a very large Herbivora caused by tectonic pressures of the Chandigarh anticline (picture A. Dambricourt Malassé).
Fracturation d’un os minéralisé de très grand herbivore provoquée par les pressions tectoniques de l’anticlinal de Chandigarh (photo A. Dambricourt Malassé).
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 Malassé).
Différents types de fossilisation. A. Masol 2 est, omoplate de proboscidien partiellement recouverte de graviers (cercle). B. Masol 2 ouest, bois de cervidé dans des limons sableux. C. Masol 2 sud-ouest, crâne de Sivatherium fortement teinté par le manganèse (Photo A. Dambricourt Malassé).
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 Malassé).
Marques fossilisées. A. Masol 3 R11154, côte avec de nombreuses striations aléatoires. B. Masol 8 R10893, ulna, Hexaprotodon, marques faites par la pression d’une canine de carnivore (flèches). C. Masol 8 grande diaphyse de Proboscidien avec une longue marque formée de deux micro-rainures parallèles. D. Agrandissement de la trace dans le rectangle blanc de la Fig. 14C (Photo A. Dambricourt Malassé).
Faunal list of Masol localities (NISP, number of identifiable specimens present) (Total includes the collection of the “Society for Archaeological and Anthropological Research” (SAAR)).
Liste de la faune des différentes localités de Masol (NR nombre de restes) (le total inclut la collection de la « Society for Archaeological and Anthropological Research » (SAAR)).
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.
Hexaprotodon sivalensis ; dimensions dentaires (mm) par localité, dents isolées (R) et dents sur les mandibules et maxillaire (n). Noter la grande variation en raison du dimorphisme sexuel et du stade d’usure (les germes et les dents usées ne sont pas mesurés). Masol sans référence correspond aux collections de la SAAR sans localisation.
Sivatherium giganteum; dental measurements (mm) of Masol specimens compared with those of Kenyan *KNMER/Sivatherium cf. maurusium (Harris, 1976).
Sivatherium giganteum ; dimensions dentaires (mm) comparées à celles de l’espèce kenyanne *KNMER/Sivatherium cf. maurusium (Harris, 1976).
LocaIitySpecimenTypeLengthDVD maxDVD anteriorDVD posteriorDVD third lobeHeightMasol3R10122P4inf50.135––50MasolR10213P4inf403029.6––30*KNMER777P4inf41.729.5––––UPMA/811M1inf55343534–35MasolR10213M1inf4323.23028.8–22*KNM-ER780M1inf45.430––––UPMA/542M1inf5529––––UPMA/542M2inf57-––––MasolR10213M2inf503031.832–34MasolR10213M3inf633031301735*KNMERn = 5M3inf60.4–66.127–33––––UPMA/542D2inf21.313.2–––12UPMA/542D3inf3322–––16UPMA/542D4inf632926262922*KNMER797P3inf34.542.5––––Masol 13R10301P3sup4033–––50
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).
Distribution des espèces (% NISP) à Masol, dans le Tatrot de l’anticlinal de Chandigarh (Gaur et Chopra, 1984), Saketi/Tatrot (Nanda, 2002) et les couches plio-pléistocènes des Pabbi Hills, Pakistan (Dennell et al., 2006). Les localités de Masol sont groupées selon la répartition stratigraphique (Chapon Sao et al., 2016b).
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.
Abondance relative des différents éléments squelettiques de l’assemblage fossile de Masol. Petits herbivores : Cervidae, Duboisia, Sivacapra, Suidae, Tragulidae, grands herbivores : Bovidae, Equidae, Camelus, très grands herbivores : Proboscidiens, Hippopotamidae, Sivatherium, Anthracoteriidae.
Little herbivoraLarge herbivoraVery large herbivoraTotal%Maxillary2210141.6Mandible141924576.6Teeth125010316519.1Skull181715505.8Vertebrae22124475.4Ribs–3441758.7Scapula1310141.6Humeri11611283.2Radius/ulnae31318343.9Pelvis4515242.8Femora11117293.3Tibia6146263.0Astragali5126232.7Calcanei–6–60.7Metapodial2111141.6Metacarpal16–70.8Metatarsal3113172.0Phalanges167141.6Shaft51278822025.4Total8138439986499.8