taxonID	type	description	language	source
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	description	(Fig. 2 A 1 - A 4)	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	materials_examined	HOLOTYPE. — BMNH / NHM 4847, an isolated left P 4 figured by Matthew (1929: 472, text-fig. 11) and by Pilgrim (1931: pl. 2, figs 3, 3 a). TYPE LOCALITY. — Indeterminate / unknown locality in the Siwaliks Hills (Falconer 1868; Pilgrim 1931).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	diagnosis	DIAGNOSIS. — Small species of Enhydriodon intermediate in dimensions between Sivaonyx bathygnathus and Enhydriodon sivalensis, with voluminous conical hypocone in P 4 lacking ridge directed into central valley; two post-protocone cusps oriented linguodistally, the most lingual one not blocking the lingual outlet of the central valley. Parastyle large but low. “ Lutrine crest ” well developed and swollen, but not forming a separate cusp. Lingual opening of m 1 trigonid narrow U-shaped (Pickford 2007).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	materials_examined	STRATIGRAPHIC RANGE. — Unknown, probably Tatrot to Pinjor formations (Matthew 1929; Pilgrim 1932; Pickford 2007); Pinjor Formation (in this study). REFERRED MATERIAL. — PUPC 83 / 01, right mandibular fragment with m 1 and root of m 2.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	description	DESCRIPTION The corpus is robust but not fully preserved (Fig. 2 A 1 - A 4). A deep mylohyoid line is present which subsequently joins the partially preserved pterygoid shelf at the lingual side while the preserved buccal side is bulged. Its preserved length is 40.1 mm. It bears an m 1 and root of m 2. The metaconid is partially broken in m 1, rest of the tooth is well-preserved, and it is moderately worn. In m 1, the paraconid is thick and eye-shaped as a result of moderate wear. It connects with the protoconid through a crest and forms a pronounced carnassial notch. The trigonid basin is small and lingually blocked by a portion of a long and strong mesial cingulid which covers the base of the tooth completely. The protoconid is thick and has the postprotocristid equipped with a well-developed cuspid. A large gap separates this cuspid from the buccal cusp of the talonid, i. e., the hypoconid and the closely attached hypoconulid. The entoconid is also well-developed. It connects with the meta- and hypoconulid through its crests and completely closes the distal and lingual side of the talonid basin. The talonid basin is much larger than the trigonid basin and partially open at its buccal side. There is a small diastema between m 1 and m 2. Only the alveolus with root of m 2 is preserved. It was single-rooted and situated much higher than m 1.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FF1EFCDAAB17252B.taxon	discussion	REMARKS The wide paraconid and postprotocristid equipped with a well-developed cuspid, large talonid basin, more or less equally developed para-, proto-, and metaconid, and a large cingulid (Fig. 2 A 1 - A 4) associate the specimen with the genus Enhydriodon. There are two species of Enhydriodon known from the Siwaliks, E. falconeri and E. sivalensis (Pilgrim 1932; Verma & Gupta 1992; Willemsen 1992, 1999; Pickford 2007). Enhydriodon falconeri is known from the latest Pliocene Tatrot Formation, while E. sivalensis has been recorded from the Middle to Upper Pleistocene Pinjor Formation (Pickford 2007). The present specimen has been collected from Pleistocene outcrops of Bhimber, Azad Jammu Kashmir (AJK), that are equivalent to the Pinjor Formation. This further justifies its association with the genus Enhydriodon rather than Sivaonyx. However, the studied specimen is smaller than the known specimens of E. sivalensis, and morphologically PUPC 83 / 01 is extremely similar to GSI-D 161, described and figured by Pilgrim (1932) as E. cf. falconeri (see Fig. 2 A 4, A 5) which is why we also attribute it to E. cf. falconeri. Nevertheless, it has been previously noted by Pilgrim (1932: 87) that “ in respect of the structure of its ml, Enhydriodon falconeri closely resembles Sivaonyx. In fact, except by its larger size and the relatively slightly broader talonid it does not seem that the genera can be separated on the characters of m 1 alone ”.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA914FC38FA3AADFF2667.taxon	type_taxon	TYPE SPECIES. — Lutra bathygnatha Lydekker, 1884 by original designation.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	description	(Fig. 2 B 1 - B 3) urn: lsid: zoobank. org: act: 123 FFCDF-CE 50 - 4 AE 6 - AB 31 - 1 F 563001 A 874	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	etymology	DERIVATION OF NAME. — After Dr Muhammad Sarwar, the second vertebrate palaeontologist of the Punjab University and who have done extensive field work during the 1963 - 1977 throughout Pakistan, that resulted in the identification and exploration of new localities and became a source of guidance for the field work for the future palaeontologists of Pakistan.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	materials_examined	HOLOTYPE. — PUPC 17 / 64, lP 4. TYPE LOCALITY. — Hasnot, Jhelum, Punjab, Pakistan.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	diagnosis	DIAGNOSIS. — A small Sivaonyx species having P 4 with round borders / external walls, basal lingual cingulum is straight at mesial border, postparacrista short and having less buccal concavity, lack a cusplet between proto- and hypocone, small mesial valley hence, less pinched mesial border, and very long posthypocrista, the crest that joins hypocone with metastyle / metacone. DIFFERENTIAL DIAGNOSIS. — Differs from all Sivaonyx species being small in size, having round outline, basal lingual cingulum is straight at mesial border, and small mesial valley hence, less pinched mesial border. It differs from Sivaonyx bathygnathus in having short postparacrista, less spaced proto- and hypocone, and having no cusplet between proto- and hypocone; from Sivaonyx gandakasensis in short postparacrista, long posthypocrista, and more buccal concavity of postparacrista; from Sivaonyx hessicus Lydekker, 1890 in having weak buccal concavity of postparacrista, no cusplet between proto- and hypocone, very long posthypocrista; from Sivaonyx hendeyi Morales, Pickford & Soria, 2005 in lacking notch between paracone and metastyle, having very long posthypocrista, and weak buccal cingulum; from Sivaonyx ekecaman Werdelin, 2003 in having longer pre- and postparacrista, slightly distally positioned protocone, lack of notch between paracone and metastyle, weak buccal concavity of postparacrista, no cusplet between proto- and hypocone, large median valley, and very long posthypocrista; and from Sivaonyx soriae in having slightly distal position of protocone, weak buccal concavity of postparacrista, no cusplet between proto- and hypocone, very long posthypocrista. STRATIGRAPHIC RANGE. — Dhok Pathan Formation of the Middle Siwaliks (Late Miocene).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	description	DESCRIPTION PUPC 17 / 64 is nearly rectangular with round outline, well-preserved and slightly worn (Fig. 2 B 1 - B 3). All the cusps are well-developed and prominent. The paracone is the highest, massive, with a rounded external wall and a prominent parastyle. The parastyle is low in height and it is separated from paracone by a small notch. Three crests originate from the paracone, mesial crest (preparacrista) connects the paracone to the parastyle, distal (postparacrista) to the metacone and the metacrista, and the third (lingual crest) connects to the protocone through crista obliqua. The metacrista is small and the metastyle / metacone is bulbous. The paracone is connected to the metacone. A prominent slanting cingulum is also present between the paracone and metacone and is connected to metastyle. The coupled protocone and hypocone are well-developed and distinct, separated by only a small notch. The protocone is slightly higher than the hypocone. The hypocrista is extremely long and becomes slightly round distally, rendering tooth round outline. The median fossette is large, oval in shape and separates trigon and talon. A thick cingulum fully covers the base of the tooth that has three roots (Fig. 2 B 1 - B 3).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBEA916FC03F994AFB8278F.taxon	discussion	COMPARISON The presence of a protocone far from the paracone and a large cingulum that covers the base of the tooth in PUPC 17 / 64 differentiate it from Paludolutra, and presence of a less developed hypocone, a protocone far from the paracone, a more sectorial paracone-metastyle, and that it is longer than wide in addition to its small size differentiate it from Enhydriodon. Vishnuonyx and Sivaonyx are the other two “ enhydriodontine ” genera present in the Siwaliks. PUPC 17 / 64 differs from the P 4 of all Vishnuonyx species in having round outline, robust parastyle, short metacrista, very long posthypocrista, and wider central basin in addition to incomplete lingual shelf. Even the space between the protocone and hypocone is smaller in PUPC 17 / 64 than in Vishnuonyx while the space between the parastyle and paracone is wider in Vishnuonyx. Sivaonyx is represented by two species, a small, Sivaonyx gandakasensis and a large, S. bathygnathus (Pickford 2007; Grohé et al. 2013; Ghaffar & Akhtar 2016). PUPC 17 / 64 is smaller than any known P 4 of both of these species (Table 2). However, it shares more morphological similarity with the P 4, GSI-D 157, of Sivaonyx bathygnathus first described and figured by Pilgrim (1932: pl. 2, fig. 16) and later figured by Pickford (2007: fig. 9 A). These similarities include the large paracone with a round lingual wall, low but heavy parastyle, a large and bulbous metastyle / metacone, a large and wide median fossette, and a very long posthypocrista (Fig. 2 B 1 - B 3). Based on these similarities, we are allocating PUPC 17 / 64 to Sivaonyx. However, due to its minute size, its rounded outline, the short metacrista, small space between protocone and hypocone, the more developed cingulum, less pinched preprotocrista or anterotransverse wall of the tooth, we do not assign it to one of the known species but designate it as a new species, Sivaonyx sarwari n. sp. It differs from other Sivaonyx species as given in differential diagnosis.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBCA916FC62FF7BAAB92669.taxon	materials_examined	HOLOTYPE. — GSI D 33, left mandible with p 4 - m 1 and alveoli of c 1 - p 3 (Lydekker 1884). TYPE LOCALITY. — Hasnot, Punjab, Pakistan (Lydekker 1884).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBCA916FC62FF7BAAB92669.taxon	diagnosis	DIAGNOSIS. — Large species of the genus Sivaonyx with slightly sectorial aspect to P 4 and m 1, but showing a talonid basin broader than the trigonid (Pickford 2007).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBCA916FC62FF7BAAB92669.taxon	materials_examined	STRATIGRAPHIC RANGE AND GEOGRAPHIC DISTRIBUTION. — This species well-known from the Late Miocene, Dhok Pathan Formation of Siwaliks (Pilgrim 1932; Pickford 2007) and Southern China (Zong 1997; Qi et al. 2006). REFERRED MATERIAL. — PUPC 16 / 29, lp 4.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBCA916FC62FF7BAAB92669.taxon	description	DESCRIPTION PUPC 16 / 29 is a left p 4. The base of tooth is partially broken postero-lingually. It is monocuspid, slightly worn and subtriangular in shape. It has a thick and strong cingulum at the base labio-lingually. The protoconid is large, pyramidal, slightly compressed buccally and highest. The distal cuspid is small, slightly buccally positioned, and separated from the protoconid by a small notch. A wide basal platform is present in the front of the distal cuspid lingually. It is double-rooted, and the distal root is more preserved (Fig. 2 C 1 - C 3).	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
FD0C1313FFBCA916FC62FF7BAAB92669.taxon	discussion	COMPARISON The described p 4, PUPC 16 / 29, is broad distally, with a strong, and a broad cingulid, no mesial accessory cusp, outwardly situated distal cusp; and these are the characters of p 4 given in the diagnosis of Sivaonyx bathygnathus given by Pilgrim (1932). Further, while describing p 4 (GSI-D 244), Pilgrim (1932) writes: “ distal accessory cusp is almost as strong and probably little inferior in height to the main cusp and it lies well on the outside of the median axis ” and the same morphology is described by Pickford (2007), i. e., presence of a distal accessory cusp, a cingulid that covers the base of whole tooth, and a broadly expanded distal platform. In this regard, morphology and dimensions of the specimen (Fig. 2 C 1 - C 3; Table 2) are close to S. bathygnathus, hence, we referred it to Sivaonyx bathygnathus.	en	Mahmood, Khalid, Morlo, Michael, Abbas, Sayyed Ghyour, Babar, Muhammad Adeeb, Khan, Muhammad Akbar (2025): The Lutrinae (Mustelidae, Carnivora, Mammalia) from the Upper Miocene to the Lower Pleistocene deposits of Pakistan. Geodiversitas 47 (5): 301-311, DOI: 10.5252/geodiversitas2025v47a5, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/geodiversitas2025v47a5.pdf
