Fossil Study of Pigs (Family Suidae) from Litra-Chaudhwan Formation of Sulaiman Range
Safdar Iqbal1, Muhammad Faizan1*, Muhammad Shehzad1, Misbah Faiz2
and Abdul Ghaffar1*
1Department of Zoology, Cholistan University of Veterinary and Animal Sciences, Bahawalpur
2Department of Zoology, Ghazi University, Dera Ghazi Khan
ABSTRACT
Dharab locality of Sulaiman Range near Sakhi Sarwar Tomb is highly fossiliferous in Litra- Chaudhwan formations, dated from the Late Miocene to the Early Pliocene of Middle Siwalik of Pakistan. Fossils of this locality are reported as bovids, tragulids, rhinocerotids, girafids, suids, and proboscideans. The present study emphasizes the Suidae family and the collected specimen is similar to Propotamochoerus hysudricus. This species is extinct with a moderately sized body and large cheek teeth. The specimen discussed from this locality is reported for the first time in this article and is a virtuous addition to the Paleontological field. This material broadens our findings for the evolution and diversification of Propotamochoerus hysudricus (Suidae) in the Sulaiman Range of Pakistan.
Article Information
Received 23 December 2023
Revised 05 May 2025
Accepted 19 May 2025
Available online 19 December 2025
(early access)
Published 13 May 2026
Authors’ Contribution
SI and MF planned this study. MS and MF executed the experimental work and analyzed the data. SI and MF wrote the article. AG reviewed and proofread the article.
Key words
Sulaiman Range, Sakhi Sarwar, Suidae, Litra formation, Chaudhwan formation
DOI: https://dx.doi.org/10.17582/journal.pjz/20231223165239
* Corresponding author: [email protected], [email protected]
0030-9923/2026/0004-1609 $ 9.00/0
Copyright 2026 by the authors. Licensee Zoological Society of Pakistan.
This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/).
Introduction
Suidae represents one of the most critical taxa from the economic point of view in the order Artiodactyla. Furthermore, it is essential owing to its substantial contribution to the evolutionary linkage of mammals between demand Artiodactyla and Cetacea (Pickford, 1993). It is a diversified group in terms of species and has been studied by numerous scholars (Falconer, 1868; Lydekker, 1883; Pilgrim, 1926; Colbert, 1935; Pickford, 1988; Batool et al., 2015). The Suidae family was reported in Eurasia and Africa in the past with numerous genera (Kostopoulos and Sen, 2016; Pickford and Obada, 2016; Spassov et al., 2018; Mors et al., 2019).
The diversity of vertebrate fauna of the Cenozoic era has been found in the Bugti Hills (Balochistan) (Forster-Cooper, 1913; Metais et al., 2009). The specimens yielded from the Chitarwata formation mostly belong to Oligocene to early Miocene (Welcomme and Ginsburg, 1997; Marivaux et al., 2001, 2005, 2006; Antoine et al., 2003; Welcomme et al., 2001; Metais et al., 2003). The diversity of artiodactyla is very high (Metais et al., 2009) and includes many undescribed suid teeth. Despite their advanced wear and scarcity, the remains of Suidae are of significant attention, as the Paleogene ancient suids needs to be better studied (Ducrocq, 1998; Liu, 2003; Harris and Liu, 2007).
Conversely, certain genera have described very well on origin of their relatively more affluence as others have described in less because of the fixed figure of specimens such like Propotamochoerus hysudricus; consequently, each fresh fossil collection of said species is a novel addition of family Suidae in paleontological study. Fossil remnants for extinct genus Propotamochoerus of family Suidae have recognized from Upper Miocene to lower Pliocene of Asia as well as Europe (Lydekker, 1884; Pilgrim, 1926; Pearson, 1928; Van der Made and Han, 1994). The Propotamochoerus hysudricus species belong to Propotamochoerus genus existing in Siwalik ridges within range of age between 10Ma to 06 (Ma) (Aslam et al., 2019).
In this study, for the first time, we locatde and identified mandibular fragment of Propotamochoerus hysudricus from Sulaiman Range of Dharab locality near Sakhi Sarwar, Dera Ghazi khan, Pakistan having the age of Late Miocene- Early Pliocene from Litra formation to Chaudhwan formation.
Materials and Methods
The specimen under study was gathered from Dharab village of Sulaiman Range near Sakhi Sarwar Tomb. This village is surrounded by freshwater sedimentary rocks deposits which have plenty of vertebrate fossil fauna. Sulaiman Range has complete sequence Late Miocene in the form of Litra Formation.
Specimen was collected from surface and some mandibular parts and p4 were detached from the major part of specimen, which were carefully picked and wrapped in cotton. Specimen was left mandibular fragment having m3-p4 dentition Collected specimen was brought to PC-CUVAS, Bahawalpur for further study. Specimen was cleaned and broken pieces were attached carefully with the help of elfy and glue.
Specimen was catalogued as PC-CUVAS 55/2023. The catalogue numerator represent the serial of collection of the year and denominator represent year of collection. For the necessary measurements in (mm), Digital as well as manual Vernier Caliper was used. Magnifying lens was also used to observe the uncertain and very minute characters for dental morphology. Teeth length of spacemen by anterior-posterior and width as lingual-buccal were measured (Khan et al., 2015). Taxonomy as well as terminology followed (Pickford, 1998).
Systematic Paleontology
Order- Artiodactyla
Sub order- Suiformes
Family-Suidae
Subfamily-Suinae
Genus Propotamochoerus
Species Propotamochoerus hysudricus
Generic diagnosis
Specimen under study PC-CUVAS-55/2023 (Fig. 1) is medium sized mandibular fragment. Specimen contain m3- p4, while anterior part having rest of teeth missing due to broken ramus. Specimen has well preserved dentine and having rugose enamel.
From distal to mesial of mandible is m3 which is simple and well preserved containing four main cusps i.e., entoconid and protoconid as well as hypoconid and paraconid. Protoconid and hypoconid are elongated and more pointed than metaconid and entoconid. Cingulum in moderate quantity is found as mesial cingulum. Above the hypoconid minute ridge is buccal cingulum. Median accessory cusp is also pronounced. A portion of cingulum at termination side of median valley make the basal pillar. Posterior part of cingulum expends posteriorly to give rise to posterior accessory cusp and talonid. The color of enamel is dark brown and smooth in appearance.
Protoconid is some larger than paraconid and is blunt inward. Well-developed valley behind paraconid form buccal cingulum. Metaconid cusp is some pointed than paraconid. Behind metaconid distal cingulum is seen clearly. In given specimen m1 is in front of 3 m2. This tooth is not well preserved and crown is damaged. Outer ridges of protoconid and hypoconid are unclearly marked. Metaconid is diffused and cingulum not marked.
At the end of specimen is p4 which show clearly roots embedded in mandibular ramus, as ramus at the area of p4 is broken. Crown of p4 is well preserved. The central cusp is folded lingual side to form inner cusp. The anterior cingulum is moderately high. There some space exists between m1 and p4 due to damaged area of some part of m1. Dental measurements of studied teeth are listed below in the Table I.
Table I. The comparative dental measurements (in mm) of Propotamochoerus hysudricus from Sulaiman range.
|
Species/specimen |
Nature/position |
Length |
Width |
W/L ratio |
|
Propotamochorous hysudricus |
||||
|
CUVAS 55/23* |
lm3 |
25.2 |
13.34 |
|
|
lm2 |
17.6 |
12.37 |
||
|
lm1 |
12.5 |
9.66 |
||
|
lp4 |
12.5 |
9.8 |
||
|
PUPC 16/103 |
lm3 |
31.9 |
16.4 |
.51 |
|
lm2 |
21 |
15.3 |
.72 |
|
|
PCGU-7/21 |
lm2 |
17.7 |
14 |
|
|
lm3 |
31.07 |
15.52 |
||
|
PCGU-136/21 |
lm3 |
36.45 |
19.50 |
|
|
PUPC 95/15 |
lm3 |
35.6 |
18 |
.50 |
|
lm2 |
23.4 |
16 |
.68 |
|
|
lm1 |
16.4 |
12.5 |
.76 |
|
|
PUPC 97/41 |
lm3 |
30 |
17.3 |
.57 |
|
lm2 |
20.9 |
16.8 |
.80 |
|
|
lm1 |
16.8 |
12.8 |
.76 |
|
|
lp4 |
16.7 |
10.7 |
.64 |
|
|
PUPC 97/40 |
lm3 |
34.9 |
19.5 |
.55 |
|
lm2 |
22 |
16 |
.72 |
|
|
lm1 |
16.8 |
12.7 |
.75 |
|
|
lp4 |
15.9 |
10 |
.62 |
|
|
PUPC16/86 |
lp4 |
18.3 |
12.3 |
.67 |
|
PUPC 16/85 |
lm3 |
29.4 |
15 |
.51 |
|
PUPC94/65 |
lp4 |
14.8 |
9.3 |
.62 |
|
lm1 |
15.5 |
11.8 |
.76 |
|
|
lm2 |
20 |
15.6 |
.78 |
|
|
PUPC 02/69 |
m3 |
27 |
14 |
.51 |
|
m2 |
19.2 |
13.3 |
.69 |
|
|
m1 |
13.7 |
10.7 |
.78 |
|
|
PUPC 16/188 |
m1 |
18.7 |
12.2 |
.65 |
|
PUPC 99/02 |
lp4 |
14.5 |
11 |
.75 |
|
PUPC 94/65 |
p4 |
15.6 |
12.6 |
.80 |
|
GSP 17 |
p4 |
15.5 |
11.6 |
.76 |
|
GSP 2503 |
p4 |
16.7 |
9.2 |
.55 |
|
GSP 10998 |
p4 |
14.4 |
11.3 |
.78 |
|
GSP12343 |
p4 |
16.6 |
11.4 |
.68 |
|
GSP 5823 |
p4 |
16 |
9.40 |
.58 |
|
GSP 12732 |
m1 |
17.6 |
13 |
.74 |
|
PUPC 07/32 |
m1 |
16 |
12 |
.75 |
|
GSP 10998 |
m1 |
16.6 |
12.5 |
.75 |
|
GSP 2807 |
m1 |
18.4 |
11.7 |
.63 |
|
Table continues on next column................ |
||||
|
Species/specimen |
Nature/position |
Length |
Width |
W/L ratio |
|
GSP 6226 |
m1 |
17.5 |
12.4 |
.70 |
|
GSP 4773 |
m3 |
31 |
16 |
.51 |
|
GSP 5112 |
m3 |
30 |
15 |
.50 |
|
GSP 7384 |
m3 |
31.3 |
17 |
.54 |
|
PUPC 15/17 |
m3 |
28.9 |
15 |
.51 |
|
PUPC 15/18 |
m2 |
21.4 |
14.3 |
.66 |
|
PC-GCUF11/162 |
p4 |
12.7 |
14.4 |
1.13 |
|
PC-GCUF11/162 |
p4 |
12.3 |
14.5 |
1.16 |
|
PC-GCUF 12/24 |
p4 |
15.3 |
9.5 |
.62 |
|
B 742 |
p4 |
10.7 |
15.3 |
1.43 |
|
Tetraconodon magnus |
||||
|
PUPC 13/37 |
lm3 |
29 |
55 |
.53 |
|
B 71 |
m3 |
32.5 |
50 |
1.54 |
|
Listriodon pentapotaminae |
||||
|
PC-GCUF155/18 |
p4 |
12.25 |
7.2 |
|
|
K 13/808 |
p4 |
15.30 |
12.30 |
|
|
K 23/721 |
p4 |
16.10 |
12.50 |
|
|
K 16/425 |
p4 |
18.90 |
14 |
|
|
IVAU K 83 |
p4 |
17.2 |
11.50 |
|
|
GSP 7017 |
m1 |
14.60 |
13 |
|
|
HGSP 8427/3664 |
m1 |
18.6 |
13.4 |
|
|
Listriodon guptai |
||||
|
PAK 1386 |
p4 |
16.80 |
11.50 |
|
|
PAK 1385 |
m1 |
19.20 |
13.30 |
|
|
Conohyus sindiensis |
||||
|
K41/836 |
m3 |
21.5 |
12.2 |
|
|
PUPC 94/7 |
m3 |
24.5 |
14.0 |
|
|
PUPC 18/10 |
lm3 |
23.58 |
13.57 |
|
|
Conohyus indicus |
||||
|
K16/482 |
m3 |
29.6 |
22 |
|
|
MO3-101 |
m3 |
19.40 |
16.2 |
|
*The studied specimen Tetraconodon magnus, Listriodon pentapotaminae, Listriodon guptai, Conohyus sindiensis, C. indicus
Discussion
The suids of Siwalik are considered important collection and also important as they form noteworthy connection with other areas of world. Pilgrim (1926) was the first paleontologist who studied massive divergence in suids of Siwalik group. Pilgrim described that Tetraconodon, Lophochoerus, Hipopopotamodon, Hippohyus and Sivahyus are endemic genera. Pilgrim also described that certain common genus like Listriodon, Sivachorus conohyus, Bunolistriodon and Propotamochoerus as well as Sus in Siwalik, moved to Europe. Africa and other parts of the world (Sarwar et al., 2016; Colbert, 1935b; Van der Made, 1996; Mathew, 1929; Pickford, 1988; Babar et al., 2021; Aslam et al., 2015).
The common of Siwalik and Europe are Listrodon, Bunolistrodon, Conohyus, Sus and Propotamochoerus (Pilgrim, 1926; Pickford, 1988; Colbert, 1935; Van der Made, 1996). Suids of Africa evolved from primitive Siwalik Sus (Pickford, 1988). Propotamchoerus in Asia and Europe is well known. Specimen of different ages of genus Hipopotamodon was present from 10-7 million years ago. The range of H. sivalense and P. hysudricus is very closer and seemingly Microstonyx. Though it is the Late Miocene fossil species that disappeared through Pliocene (Pickford, 1988).
Studied specimen has been accumulated from Dharab area of Late Miocene to Early Pliocene locality of Litra and Chaudhwan formation of Sulaiman Range. As Bunodont dentition is evident and more than one accessory conids are found. Also, protoconid and hypoconid are elongated than paraconid and metaconid suggest specimen to include in the Suidae family. The genera of Late Miocene to Pliocene represent comparatively larger sized teeth. As specimen belong to same age, so suggested to belong Late Miocene-Early Pliocene of Sulaiman Range. The Siwalik suids along with other regions of world form assemblages.
All over the world, the species in genus Propotamochoerus are as P. wui, P. provincialis, P. palaeochoerus, P. hyotheriodes, P. hysudricus and MN11-13 form of Europe (Van der Made, 1996; Pickford, 1988). Among all species of genus Propotamochoerus, P. provincialis and Propotamochoerus palaeochoerus have rcognized from the Europe (Fortelius et al., 1996). According to Sarwar et al. (2016), P. hysudricus is originated from Hyotherium near about 11 million years ago. P. hysudricus is distinguished from Hyotherium by large size and shape of p4 and sagittal cusplets. Lower crowned molars along shallow furrows place it in P. hysudricus. As Propotamochoerus hysudricus is well recognized from Potwar Plateau in Pakistan and India (Pilgram, 1926; Colbert, 1935; Lydekker, 1884).
In Indo-Pak sub-continent, all over the world significant the faunal replacement (the change of diet) was identified; the adaptations of the mammals from the wooded area to an open habitat during the middle Miocene range in age (14.5 to 11.8 Ma) and it is very significant age regarding climatic, global vegetation as well as faunal modifications. The technical as well as mathematical data regarding dentition record are selected as the molars with high crown were curiously well-structured in the Cenozoic era that consistently reinforced in taxa having Hypsodont dentition (Eronen et al., 2012; Tapaltsyan et al., 2015). The specimens are also compared with the already published data to correlate the size of molars (Fig. 2).
Conclusion
Propotamochoerus hysudricus was discovered from Nagri and Dhok Pathan Formation (10.2 to 6.8) Ma in Late Miocene (Barry et al., 2002). The Sulaiman Range reveals the most complete sequence of Litra Formation (Late Miocene) group and yield as diversified fossil collection. As Nagri and Dhok Pathan Formation of Siwalik group is similar in age with Litra and Chaudhawn Formation of Sulaiman Range. Similar community of both formations of Potwar Plateau diversified. The p3 of Propotamochoerus hysudricus is relatively narrow as well as oval in shape. Conversely, the p3 of the P. hysudricus of India is constantly shorter. For the first time Propotamchoerus hysudricus species of family suidae is reported from Dharab area of Sulaiman range near Sakhi Sarwar tomb.
Declarations
Funding
This research was self-funded by the authors. No external funding was received for this study.
Generative AI and AI-assisted technology statement
The authors declare that they have not used generative AI or AI-assisted technologies in the writing or editing of this manuscript.
Statement of conflict of interest
The authors have declared no conflict of interest.
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