Miocene hyaenids (Hyaenidae; Carnivora) from Siwaliks of Pakistan
Introduction
Carnivore remains are very rare in the Siwaliks, and are found in different environments compared to herbivores (Dennell et al., 2006). The carnivorans from the Siwalik deposits of Pakistan have always been a vital source to understand Eurasian carnivore evolution and present-day distribution. The fossil carnivorans from the Middle Miocene to Late Pleistocene of the Siwaliks are of great value for understanding the evolution of the lineages of various taxa, and their distribution and extinction. Despite the scarcity of carnivore remains, a variety of carnivores belonging to families including Mustelidae, Viverridae, Felidae, and Hyaenidae are the components of the Siwalik faunas that are important paleontologically (Pilgrim, 1932; Barry, 1983; Pickford, 2007; Ghaffar et al., 2019; Jiangzuo et al., in press). Hyaenids represent dominantly in the carnivore remains of the Siwalik Group (Pilgrim, 1932, Colbert, 1935, Ghaffar et al., 2019). At least eight species of Hyaenidae have been recognized from the Siwaliks, predominantly the Pakistani Siwaliks (Werdelin and Solounias, 1991).
New carnivoran material belonging to the family Hyaenidae from the Middle Miocene to Early Pliocene localities of the Pakistani Siwaliks is presented here. The studied carnivoran material belongs to Thalassictis cf. T. proava, Ictitherine indet. and Lepthayena sivalensis and contributes to knowledge of the Siwalik Carnivora. Although still rare, the new material represents a new stratigraphic occurrence of T. proava. Future collection will surely shed more light on this poorly known fauna.
The described specimens have been collected from the outcrops near the Kanhatti, Nagri and Hasnot villages in Potwar Plateau of Pakistan (Fig. 1). The outcrops of the Kanhatti village (Lat. 32°41′1.7″N, Long. 72°25′02.5″E) represents the Middle Miocene of the Chinji Formation (Heissig, 1972, Shah, 2009, Khan et al., 2013) and is situated in the Khushab district, Punjab, Pakistan (Fig. 1). The outcrops of the Chinji Formation are predominantly clays (mud- and siltstone) of brick red color (70%) and ash grey or brownish grey, fine to medium grained sandstone (30 %) (Pilbeam et al., 1977, Pilbeam et al., 1979, Shah, 2009). Sethi-Nagri (Lat. 32°46′28.5″N, Long. 72°29′43.4″E), located in the Potwar Plateau, is the type locality of the Nagri Formation; it is a famous expedition locality of the Yale University, USA-Geological Survey of Pakistan (Y-311). The Nagri Formation consists of sandstone with subordinate clay and conglomerate. The sandstone is greenish-grey medium to coarse-grained, cross-bedded and massive. The fossiliferous level of the Nagri type section was radiometrically bracketed between 10.1 and 11.2 Ma (Pilbeam et al., 1997, Barry et al., 2002).
The Hasnot outcrops (Lat. 32°50′44.7″N, Long. 73°17′50.1″E) have been assigned to the Dhok Pathan Formation (Late Miocene–Early Pliocene). The Dhok Pathan Formation is typically represented by monotonous cyclic alternations of sandstone and clay beds. The sandstone color varies: it is commonly grey, light grey, gleaming white or reddish brown. The clay is orange, brown, dull red or reddish brown and occasionally rusty orange, represented by monotonous cyclic alternations of sandstone and clay beds. Interbedded in these layers is conglomerate that is in the form of lenses and this layer is an essential character of the upper part (Shah, 2009). A detailed account of the lithology, stratigraphy and magnetostratigraphy have been provided in Pilbeam et al. (1979), Johnson et al. (1982), and Barry et al. (2002).
AK5, Akkasdag I, 2000–2001 excavations; AMNH, American Museum of Natural History, United States; GSI, Geological Survey of India; GSP, Geological Survey of Pakistan; NHMUK, Natural History Museum, London; PUPC, Punjab University Paleontological Collection, Lahore, Pakistan.
Section snippets
Material and methods
The material described in this article consists of a maxillary fragment, an isolated lower premolar, a molar and two mandible fragments, collected from the Siwaliks of Potwar Plateau, Pakistan, and is housed in the Dr. Abu Bakr Fossil Display and Research Centre, University of the Punjab, Lahore, Pakistan. Crown terminology and measurements follow those of Werdelin and Solounias (1991) and Borths and Seiffert (2017). The measurements were taken by digital calipers and are expressed in mm. The
Systematic palaeontology
Order Carnivora Bowdich
Family Hyaenidae Gray
Genus Thalassictis Gervais ex. Nordmann
Thalassictis cf. T. proava Pilgrim, 1910
(Fig. 2A–I)
Holotype: GSI-D 126, a left mandibular ramus containing p4 and m1 (Pilgrim, 1910).
Type locality: Chinji, Salt Range (Pilgrim, 1910).
Stratigraphic range: Chinji Formation (Pilgrim, 1932), and Nagri–Dhok Pathan Formations (reference herein); Middle–Late Miocene of the Siwaliks.
New material: PUPC 17/272, left p3 (Hasnot); PUPC 18/217, left mandible fragment with m1
Discussion
The Siwalik carnivorans have been comparatively neglected relative to the other Siwalik mammalian groups. However, considerable work was done by Baker (1835), Falconer and Cautley (1846), Falconer (1868), Bose (1880), Lydekker (1884), Matthew (1929), Pilgrim (1932), Colbert (1935), Prasad (1968), Bakr (1986), Ghaffar (2005), Pickford (2007), and Ghaffar et al. (2019). Surprisingly, carnivores are rare in the fossil record of the Siwalik Group and include few fossil specimens of identified
Conclusions
Three carnivoran taxa, Thalassictis cf. T. proava, Ictitherine indet. and Lepthayaena sivalensis, have been recorded from Pakistan in the Middle to Late Miocene of the Siwalik Group. The identification is based on morphometric studies. Thalassictis proava and L. sivalensis have a long list of synonymies with a complex history. The newly described material indicates the presence of T. proava for the first time in the Late Miocene–Early Pliocene localities of the Siwalik Group, extending its
Acknowledgements
We are thankful to Mr. Anaar Khan for his hosting and guidance during the field work. Grateful to two anonymous reviewers for their critical comments to improve the manuscript. This study is supported by University of the Punjab, Lahore, Pakistan.
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