Protocetid Cetaceans (Mammalia) from the Eocene of India
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Palaeontologia Electronica palaeo-electronica.org Protocetid cetaceans (Mammalia) from the Eocene of India Sunil Bajpai and J.G.M. Thewissen ABSTRACT Protocetid cetaceans were first described from the Eocene of India in 1975, but many more specimens have been discovered since then and are described here. All specimens are from District Kutch in the State of Gujarat and were recovered in depos- its approximately 42 million years old. Valid species described in the past include Indocetus ramani, Babiacetus indicus and B. mishrai. We here describe new material for Indocetus, including lower teeth and deciduous premolars. We also describe two new genera and species: Kharodacetus sahnii and Dhedacetus hyaeni. Kharodacetus is mostly based on a very well preserved rostrum and mandibles with teeth, and Dhe- dacetus is based on a partial skull with vertebral column. The Kutch protocetid fauna differs from the protocetid fauna of the Pakistani Sulaiman Range, possibly because the latter is partly older, and/or because it samples a different environment, being located on the trailing edge of the Indian Plate, directly exposed to the Indian Ocean. Sunil Bajpai. Department of Earth Sciences, Indian Institute of Technology, Roorkee 247667, Uttarakhand, India. Current address: Birbal Sahni Institute of Palaeobotany, Lucknow 226007, Uttar Pradesh, India. [email protected]; [email protected]. J.G.M. Thewissen (corresponding author). Department of Anatomy and Neurobiology, Northeast Ohio Medical University, Rootstown, Ohio 44272, U.S.A. [email protected]. Keywords: Eocene; Mammalia; Cetacea; India; New species; New genus INTRODUCTION 1998, 2000; Thewissen and Hussain, 1998) and many specimens of the remingtonocetine genera The earliest cetaceans, pakicetids and ambu- Remingtonocetus (Bajpai et al., 2011), Dalanistes locetids are only known from the Eocene of India (Thewissen and Bajpai, 2001) and the andrewsi- and Pakistan (reviewed by Thewissen et al., 2009). phiine genera Andrewsiphius and Kutchicetus These families are followed, in time as well as on (Thewissen and Bajpai, 2009) have been recov- the cladogram, by remingtonocetids and proto- ered in Kutch (Gujarat, India). Remingtonocetids cetids. Unambiguous remingtonocetid fossils are had small eyes and large middle ear cavities. Rem- also only known only from India and Pakistan ingtonocetines and andrewsiphiines are both com- (Sahni and Mishra, 1975; Kumar and Sahni, 1986; mon in localities indicative of murky water, but Gingerich et al., 1995b; Bajpai and Thewissen, http://zoobank.org/6A475A48-F702-49BB-8A8F-1B732123348A PE Article Number: 17.3.34A Copyright: Paleontological Society October 2014 Submission: 31 January 2014. Acceptance: 11 September 2014 Bajpai, Sunil and Thewissen, J.G.M. 2014. Protocetid cetaceans (Mammalia) from the Eocene of India. Palaeontologia Electronica Vol. 17, Issue 3;34A; 19p; palaeo-electronica.org/content/2014/914-whales-from-india BAJPAI AND THEWISSEN: WHALES FROM INDIA Remingtonocetus also lived in more clear water Differential diagnosis. Indocetus is a medium-sized localities (Thewissen and Bajpai, 2009). protocetid, with teeth larger than most protocetids Protocetids are found in many continents, from the Indian continent, but smaller than those of including Africa (Kellogg, 1936; Williams, 1998; Kharodacetus, and only slightly larger than teeth of Bianucci and Gingerich, 2011; Gingerich and Cap- Rodhocetus. The orbit is barely elevated above the petta, 2014), North America (Hulbert et al., 1998; palate (unlike Kharodacetus and Dhedacetus). Geisler et al., 2005; Uhen, 2008, 2014a, 2014b), Indocetus has a flat supraorbital shield with large and South America (Uhen et al., 2011). Apparently orbits, similar to all other protocetids and different then, protocetids were the first cetaceans able to from pakicetids, ambulocetids and reming- cross oceans, and to colonize tropical and subtrop- tonocetids. ical oceans. Discussion. Gingerich et al. (1994) diagnosed Rod- Several genera of protocetids are known from hocetus as distinct from Indocetus based on the the desert of Kutch in western India and the moun- convexity of the posterior surface of the exoccipital, tains of the Sulaiman Range in Pakistan. From the shorter cervical vertebrae, unfused sacral ver- Kutch, Indocetus ramani Sahni and Mishra, 1975, tebrae and the shorter femur. However, the post- Babiacetus indicus Trivedi and Satsangi, 1984, cranial elements on which the Indocetus Babiacetus mishrai Bajpai and Thewissen, 1998, comparisons are based have been reassigned to and Gaviacetus sahnii Bajpai and Thewissen, other taxa, and these elements are not known for 1998, have been reported. Specimens initially Indocetus. This leaves the exoccipital as the only identified as Protocetus (Sahni and Mishra, 1975) diagnostic feature. It is possible that the exoccipital have been reassigned later to Remingtonocetus shape constitutes a consistent and sharp differ- (Kumar and Sahni, 1986). In the Sulaiman Range, ence between Rodhocetus and Indocetus, but it is there may be nine genera with 10 species of also possible that this difference does not merit dis- Eocene whales: Rodhocetus kasranii Gingerich et tinction of separate genera. The m1 crown of the al., 1994 (spelling amended by Gingerich et al., specimen referred to Indocetus below (IITR-SB 2001a); Babiacetus indicus Trivedi and Satsangi, 2986) is 125% as large, in linear dimensions, as 1984 (reported by Gingerich et al., 1995a), the same tooth of Rodhocetus kasrani (GSP-UM Takracetus simus Gingerich et al., 1995b; Gaviace- 3012, Gingerich et al., 1995b). This size difference tus razai Gingerich et al., 1995b; Artiocetus clavis may merit distinction at the specific level, but is Gingerich et al., 2001b; Rodhocetus balochistan- probably not a diagnostic character at the genus ensis Gingerich et al., 2001b; Qaisracetus arifi Gin- level. Based on the limited amount of available evi- gerich et al., 2001a; Makaracetus bidens, dence, it is likely that Rodhocetus is a junior syn- Gingerich et al., 2005, and Maiacetus inuus Gin- onym of Indocetus, however, we stop short of gerich et al., 2009. Specimens of Indocetus synonymizing these taxa, as a more detailed com- reported from the Sulaiman Range by Gingerich et parison of holotypes and a study of the variability of al. (1993) have now been reassigned to Reming- South Asian protocetids are needed. At present, it tonocetus (Gingerich et al., 1995b). is preferable to maintain the status quo. The Kutch deposits with fossil cetaceans were Gingerich et al. (2001b, supplementary infor- dated to 42 million years ago by Ravikant and mation) showed that there are significant differ- Bajpai (2010). The localities in the Sulaiman ences in the rostrum shape of the Indo-Pakistani Range have some species in common with those protocetids. The shape of the rostrum of Indocetus in Kutch, and it appears clear that some of the fau- matches that of Rodhocetus. The rostrum of nal differences are due to environmental differ- Artiocetus, Takracetus and Maiacetus (Gingerich et ences between the two regions. Here we report al., 2009) is broader, whereas that of Gaviacetus new protocetid cetaceans from Kutch and review and Babiacetus (based on its lower jaws) is nar- the already described species from India. rower. In Makaracetus the rostrum is downturned (Gingerich et al., 2005), unlike any other proto- SYSTEMATIC PALAEONTOLOGY cetid. It is likely that rostrum differences reflect dif- ferences in feeding and quantification of these Class MAMMALIA Linnaeus, 1758 differences based on original, undeformed material Order CETACEA Brisson, 1762 may expose subtle ecological differences among Family PROTOCETIDAE Stromer, 1908 these taxa. Genus Indocetus Sahni and Mishra, 1975 Type species. Indocetus ramani Sahni and Mishra, 1975, by original designation. 2 PALAEO-ELECTRONICA.ORG Indocetus ramani Sahni and Mishra, 1975 ventrally by a deep groove that contained the Figure 1.1-4 nerves to the eye and orbit. The face is low, with Holotype. LUVP 11034, fragments of orbits with the orbit placed only slightly above the palate, palate, interorbital region, and ear region with unlike Kharodacetus (see below). crown for one molar. IITR-SB 2986 (Figure 1.1-2) is a well-pre- Other material. IITR-SB 2000-33, left mandible with served mandible for Indocetus ramani. The caudal dp3 and dp4, Babia Hill; IITR-SB 2986, left mandi- end of dp3 is preserved, and it shows that a small ble with dp3 fragment, dp4, m1, and unerupted m2; cusp is present at the end of the postprotocrista. Rato Nala. IITR-SB 4001, palatal/orbital fragment The dp4 has four cusps that are lined up from ros- with crowns for right P3-M1, Rato Nala; VPL 1014, tral to caudal, and are connected by a sharp crest. right tympanic, Bajpai and Thewissen (1998); VPL The rostral cusp is the lowest, and the second cusp 1017, endocast of cranial cavity, palatal, interorbital the highest, the third and fourth cusp decrease in fragment, Bajpai et al. (1996); VPL 1018, endocast size from the second cusp. The lingual cingulum is of cranial cavity, Bajpai et al. (1996). complete, and the labial cingulum is damaged for Type locality. Rato Nala, 1 km north of the village of as far as it can be studied. The m1 bears two Harudi. cusps, a high protoconid on the trigonid and a low Type stratum. Harudi Formation, Eocene, around hypoconid. The rostral face of the protoconid is 42 million years old. concave, bounded by a sharp lingual crest and a Diagnosis. Same as for genus. much weaker labial crest. Both of these crests Description. IITR-SB 4001 shows alveoli for P2 and ascend the protoconid partially. A sharp crest crowns for P3 to M1 (Figure 1.3-4, Table 1). There descends from the protoconid caudally and is con- are two alveoli for P2, followed by a diastema, tinued over the rostral and caudal aspect of the whereas no diastemata occur between P3 and M1. hypoconid. Only the tip of m2 is erupted. P3 and P4 are similar in shape. They lack a proto- The mandible shows a gently ascending coro- cone, but have a large lingual bulge, similar in size noid process, and a strongly dorso-ventrally con- to that of Babiacetus (Gingerich et al., 1995a).