[Papers in Palaeontology, 2019, pp.

1–24]

AN ENIGMATIC NEW UNGULATE-LIKE MAMMAL

FROM THE EARLY EOCENE OF INDIA
by SHAWN P. ZACK 1 , 2 , KENNETH D. ROSE 3 , LUKE T. HOLBROOK 4 ,
KISHOR KUMAR 5 , RAJENDRA S. RANA 6 and THIERRY SMITH 7
1
Department of Basic Medical Sciences, University of Arizona College of Medicine-Phoenix, Phoenix, AZ 85004, USA; zack@email.arizona.edu
2
School of Human Evolution & Social Change, Arizona State University, Tempe, AZ 85287, USA
3
Center for Functional Anatomy & Evolution, Johns Hopkins University School of Medicine, Baltimore, MD 21205, USA; kdrose@jhmi.edu
4
Department of Biological Sciences, Rowan University, Glassboro, NJ 08028, USA; holbrook@rowan.edu
5
Wadia Institute of Himalayan Geology, 33 General Mahadeo Singh Road, Dehradun, Uttarakhand 248001, India; kumark@wihg.res.in
6
Department of Geology, H.N.B. Garhwal University, Srinagar, Uttarakhand 246175, India; Rajendra.Rana1@gmail.com
7
Directorate Earth & History of Life, Royal Belgian Institute of Natural Sciences, Rue Vautier 29, B-1000, Brussels, Belgium; thierry.smith@naturalsciences.be

Typescript received 8 May 2019; accepted in revised form 26 June 2019

Abstract: We report a new genus and species of herbivo-
rous mammal, Pahelia mysteriosa, from the early Eocene
Cambay Shale Formation, Tadkeshwar Lignite Mine, Gujarat,
India. The new taxon, approximately the size of a small
phenacodontid (e.g. Ectocion parvus), is represented by three
mandibular fragments, the most complete of which docu-
ments nearly the entire symphysis and mandibular body plus
P3–M3. Pahelia has incipiently selenolophodont molars with
strong exodaenodonty, absent paraconids, weak but distinct
entolophids, and prominent ectostylids. Molar size increases
distally, but M3 does not develop a prominent third lobe.
Premolars are simple, with prominent protoconids and short
talonids but little development of other trigonid cusps. The
mandibular symphysis is strongly fused, and there is an
enlarged alveolus for an anterior tooth. The combination of
features present in the new taxon does not closely match that
of any known mammal, but there are some similarities to a
diversity of ungulates from Africa, Asia, Europe and North
America. Preserved morphology is insufficient to assess the
affinities of the new taxon with confidence, but a link to
Quettacyonidae, also endemic to the Indian subcontinent, is
morphologically and biogeographically plausible. If this sce-
nario is correct, it suggests that P. mysteriosa could be a part
of the endemic mammalian fauna of India prior to its initial
faunal contact with Asia.
Key words: Mammalia, Eutheria, ungulate, Eocene, Cam-
bay Shale Formation, India.

T H E Cambay Shale Formation has produced the richest,
most diverse terrestrial vertebrate fauna from the early
Eocene of India. Mammals are particularly well docu-
mented and include the earliest representatives of Artio-
dactyla, Perissodactyla, Primates, Rodentia, Lagomorpha,
Chiroptera, Lipotyphla, Hyaenodonta and Tillodontia
known from the Indian subcontinent (Bajpai et al.
2005a–c, 2006, 2007, 2008, 2009; Rose et al. 2006, 2007,
2008, 2009a, b, 2013, 2014; Smith et al. 2007, 2015, 2016;
Rana et al. 2008, 2015; Kumar et al. 2010; Kapur & Baj-
pai 2015; Dunn et al. 2016; Kapur et al. 2017a, b).
Despite India’s lengthy isolation from both Africa and
Eurasia, the level of endemism of the Cambay Shale
fauna, to date, has been relatively modest. While some
groups are distinctive at the family level, particularly cam-
baytheriids, which are endemic to the Indian subconti-
nent (Ginsburg et al. 1999; Bajpai et al. 2005a, 2006;
Rose et al. 2006, 2014), most groups show evidence for a
relatively close relationship to taxa from North America,
Europe, or mainland Asia, particularly the first two land-
masses (Smith et al. 2007; Rana et al. 2008; Kumar et al.
2010; Rose et al. 2013). Even cambaytheres, although dis-
tinct and endemic, show clear affinities to early Holarctic
perissodactyls (Rose et al. 2014).
Until recently, Cambay Shale vertebrates were primarily
known from the Vastan Lignite Mine, Surat District,
Gujarat State, India. Smith et al. (2016) recently described
material from two horizons at a second locality, the Tad-
keshwar Lignite Mine. The Tadkeshwar mammal assem-
blage is broadly similar to the Vastan fauna, sharing
several species in common; differences include the pres-
ence of at least one pantodont and a small species of the
cambaythere Cambaytherium (Smith et al. 2016).
The present contribution reports a new genus and spe-
cies of a mammal from both horizons at Tadkeshwar
(lower TAD-1, upper TAD-2). It is represented by three
tooth-bearing lower jaw fragments, the morphology of
which is strongly distinct from that of any previously

© The Palaeontological Association doi: 10.1002/spp2.1288 1

2 PAPERS IN PALAEONTOLOGY

described mammal, living or extinct. While it would be

desirable to have additional specimens, particularly the
upper dentition, three trips to Tadkeshwar Mine subse-
quent to the discovery of the first two specimens from
TAD-2 produced only a single mandible from TAD-1,
and no new exposures at the level of TAD-2 have been
located. The new form is broadly similar to early ‘ungu-
lates’, a diverse, almost certainly polyphyletic assemblage
of taxa that are generally both terrestrial and herbivorous.
There are intriguing similarities to a number of previ-
ously known ‘ungulate’ groups, including forms from
North America, Europe and Africa, as well as taxa known
from the Indian subcontinent. In all instances, there are
also substantial contrasts that suggest that any relation-
ship is likely to be distant. The new form is best regarded
as a new genus of uncertain higher-level affinities. Discov-
ery of such a distinctive form in the Cambay Shale assem-
blage has implications for the biogeography and early FIG. 1. Left M3 of Pahelia mysteriosa gen. et sp. nov. (GU/
evolution of Indian mammalian faunas, although these RSR/TAD 9203), in occlusal view illustrating dental terminology
implications are tempered by phylogenetic uncertainty. used in this study. Abbreviations: cro, cristid obliqua; el, entolo-
phid; end, entoconid; enm, endometacristid; es, ectostylid; hcr,
hypocristid; hyd, hypoconid; hyld, hypoconulid; mc, mesial cin-
MATERIAL AND METHOD gulid; med, metaconid; pcr, paracristid; pec, preentocristid;
pom, postmetacristid; pop, postprotocristid; prd, protoconid;
prm, premetacristid.
The new fossils are catalogued in the collection of the
Department of Geology, H.N.B. Garhwal University, Sri-
nagar, Uttarakhand, India. The specimens were scanned TABLE 1. Pahelia mysteriosa gen. et sp. nov. dental and
on ultra-high resolution computed tomography (CT) at mandibular measurements (mm).
the Duke University Shared Materials Instrumentation GU/RSR/TAD GU/RSR/TAD GU/RSR/
Facility (SMIF) using a Nikon XT H 225 ST and at the 9201 (type) 9203 TAD 9608
Royal Belgian Institute of Natural Sciences using a RX-
Solution EasyTom. Minor digital preparation was done to Side Left Left Left
P3 L 5.6
the most complete of the three specimens (GU/RSR/TAD
P3 W 3.9
9201) to remove sediment filling anterior alveoli and to
P4 L 5.1
GU/RSR/TAD 9608 to reconstruct the back of the den- P4 W 4.1
tary. Specimens were manipulated and digitally imaged in M1L >5.3
MeshLab. The three specimens were further prepared, M1TrW 3.9
moulded and cast at the fossil microvertebrate prepara- M1TaW
tion laboratory, Royal Belgian Institute of Natural M2L 6.6 5.9
Sciences, Brussels. M2TrW 5.1 4.2
Dental terminology used in this work largely follows M2TaW 4.8 3.7
Szalay (1969) and is illustrated in Figure 1. Nomenclature M3L 7.5 7.8 7.5
for trigonid crests follows Orliac & Ducrocq (2012). Use M3TrW 5.4 5.4 4.9
M3TaW 4.7 4.4 4.0
of hypocristid and postentocristid for the buccal and lin-
P4MD 11.7 9.9
gual portions of the postcristid follows Kay (1977). Use
M3MD 15.8 c. 15.4 12.8
of entolophid for a transverse entoconid crest directed
towards the hypoconid follows Cifelli et al. (1989). L, maximum length; MD, mandibular depth (measured lin-
gually); TaW, maximum talonid width; TrW, maximum trigonid
width; W, maximum width.
Comparative material examined
Anthracotheriidae. Anthracotherium spp. (sensu Tsub-
The following specimens (or casts thereof) and references amoto et al. 2002): Colbert (1938); Tsubamoto et al.
were used to make detailed morphological comparisons (2002); Soe (2008). Siamotherium krabiense: Suteethorn
to the new taxon (see Comparisons, below) (Table 1). et al. (1988).
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
Apheliscidae. Aletodon gunnelli: UM 65059, 66301; Aphe-
liscus insidiosus: USGS 7131, 12608; Gingerichia geoteretes:
UM 83932, 84535; Haplaletes disceptatrix: AMNH FM
35897, USNM 9410, YPM-PU 14829, 17547, 19835; Hap-
laletes pelicatus: USNM 21008; Haplomylus speirianus:
USNM 521645; Litomylus dissentaneus: AMNH FM 35922,
YPM-PU 16849; Phenacodaptes sabulosus: YPM-PU 14372,
14398, 17591; Utemylus serior: UW 1078.
Cambaytheriidae. Cambaytherium spp.: Bajpai et al.
(2005a, 2006); Rose et al. (2006, 2014); Smith et al.
(2016).
Hyracoidea. Dimaitherium patnaiki: Barrow et al. (2010).
Microhyrax lavocati: Tabuce et al. (2001, 2007). Seggeurius
amourensis: UO-K212; Court & Mahboubi (1993); Benoit
et al. (2016).
Louisinidae. Louisina spp.: Hooker & Russell (2012). Pro-
louisina atavella: Wa-356; Hooker & Russell (2012). Teil-
hardimys brisswalteri: MNHN CR-1180, CR-1264, CR-
234, CR-236; Hooker & Russell (2012). Teilhardimys mus-
culus: Tabuce et al. (2006); Hooker & Russell (2012).
Teilhardimys reisi: Tabuce et al. (2006).
Ocepeia. Ocepeia daouiensis: MNHN.F PM41, PM49;
Gheerbrant et al. (2001, 2014) and Gheerbrant (2010).
Ocepeia grandis: MNHN.F PM34, PM37; Gheerbrant et al.
(2014).
Periptychidae. Conacodon cophater: USNM 404950. Cona-
codon entoconus: USNM 404965. Goleroconus alfi:
McKenna et al. (2008). Haploconus angustus: USNM
15378, 408061, 408064. Hemithlaeus kowalevskianus:
USNM 15452, 405093, 405152, 464050. Mithrandir gillia-
nus: NMMNH P-3083, USNM 15439, 372299.
Quettacyonidae. Machocyon abbasi: GPS-UM 4208, Gin-
gerich et al. (1999). Quettacyon parachi: GPS-UM 4000,
Gingerich et al. (1997, 1999). Sororocyon usmanii: GPS-
UM 4007, Gingerich et al. (1998).
Institutional abbreviations. AMNH FM, Division of Fossil
Mammals, American Museum of Natural History, New York,
USA; GSP-UM, Geological Survey of Pakistan-University of
Michigan collection, Quetta, Pakistan; GU/RSR/TAD, Tadkesh-
war collection, Department of Geology, H.N.B. Garhwal Univer-
sity, Srinagar, Uttarakhand, India; MNHN, Mus
eum National
d’Histoire Naturelle, Paris, France; NMMNH, New Mexico
Museum of Natural History and Science, Albuquerque, New
Mexico, USA; OCP DEK/GE, Collections of the Office Ch
erifien
des Phosphates, Khouribga, Morocco; UM, Museum of Paleon-
tology, University of Michigan, Ann Arbor, Michigan, USA; UO,
3
Universit
e d’Oran, Algeria; USGS, United States Geological Sur-
vey, Denver registry, Denver, Colorado, USA (transferred to
USNM); USNM, Department of Paleobiology, United States
National Museum of Natural History, Smithsonian Institution,
Washington D.C., USA; UW, University of Wyoming Geological
Museum, University of Wyoming, Laramie, Wyoming, USA;
YPM-PU, Princeton University Collection, Yale Peabody
Museum, Yale University, New Haven, Connecticut, USA.
SYSTEMATIC PALAEONTOLOGY
Class MAMMALIA Huxley, 1880
Subclass THERIA Parker & Haswell, 1897
Infraclass EUTHERIA Gill, 1872
Order & Family INCERTAE SEDIS
Genus PAHELIA nov.
LSID. urn:lsid:zoobank.org:act:1E2C3170-16B3-48F7-B47E-
CFBDA127D2AD
Derivation of name. Hindi paheli, puzzle or riddle, in ref-
erence to the uncertain affinities of the new taxon.
Type and only known species. Pahelia mysteriosa sp. nov.
Diagnosis. As for the type species.
Age and distribution. As for the type species.
Pahelia mysteriosa sp. nov.
Figures 1–6
LSID. urn:lsid:zoobank.org:act:9A7A2A91-5890-419C-BEDB-
5F2AA452381E
Derivation of name. Latin mysterium, secret or mystery.
Holotype. GU/RSR/TAD 9201, mandible including sym-
physis and left mandibular body with P3–M3 (Fig. 2,
restored mandible shown in Fig. 3).
Hypodigm. Holotype; GU/RSR/TAD 9203, mandibular
fragment with left M3 (Fig. 4); GU/RSR/TAD 9608,
mandibular fragment with left M2–3 (Fig. 5).
Diagnosis. An eutherian mammal differing from all
known mammals in possessing the following combination
of features: mandibular symphysis fused, relatively hori-
zontal; sharp laterally projecting crest at anterior border
of masseteric fossa; enlarged proclivous anterior tooth
4 PAPERS IN PALAEONTOLOGY

FIG. 2. GU/RSR/TAD 9201, mandible with left dentary and P3–M3, holotype of Pahelia mysteriosa gen. et sp. nov. in: A, buccal;
B, lingual; C, occlusal; D, oblique occlusobuccal; E, oblique anterobuccal view. F–H, M2–3 showing details of molar morphology, in:
F, buccal; G, lingual; H, occlusal view. The oblique occlusobuccal view (D) is included to show crown relief, while the oblique anterobuccal
view (E) is included to show the size of the enlarged anterior tooth alveolus. Abbreviations: ccr, coronoid crest; cro, cristid obliqua; hyd,
hypoconid; med, metaconid; mf, mental foramen; tc, second P4 talonid cusp. Scale bars represent 10 mm (A–E); 5 mm (F–H).

(sensu Walsh 1998); premolars simple, premolariform,
with no paraconid on P3–4, no metaconid on P3 and a
rudimentary metaconid on P4, and short, simple talonids;
M1 < M2 < M3; molar talonids almost as tall as trigonids
but distinctly narrower; molars exodaenodont; paraconid
absent on all lower molars, paracristid ends at mid-width;
molar metaconid directly lingual to protoconid; molar
protocristid with a deep trigonid notch; cristid obliqua
on M2–3 contacting protocristid at trigonid notch; hypo-
conulid absent on M2; entoconid with incipient
entolophid on M2–3; M3 hypoconulid present but not
forming complete third lobe; strong ectostylid present on
lower molars.
Description. The mandibular symphysis of the holotype is
fused and extends posteriorly to a point opposite P3. The
symphysis is relatively shallow and almost horizontal. The
mandibular body is deep but relatively narrow trans-
versely. The ventral margin of the horizontal ramus is dis-
tinctly convex and is deepest beneath M3. Two mental
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA 5

FIG. 3. Restored mandible of GU/RSR/TAD 9201 in: A, occlusal; B, oblique right anterobuccal view. The mandible was restored by
mirroring the left side of GU/RSR/TAD 9201. Scale bar represents 10 mm.

foramina are preserved, one beneath the mesial root of P2
and one beneath the centre of P4.
Posteriorly, a sharp, roughly vertical coronoid crest
projects laterally at the anterior margin of the lateral sur-
face of the ascending ramus. More posteriorly on GU/
RSR/TAD 9608, running anteroventrally from the
mandibular condyle, is a blunt, curving condyloid crest,
which subdivides the masseteric fossa into anterodorsal
and posteroventral portions. The crest fades rapidly ante-
riorly. The fossa itself appears to have extended ventrally
to the level of the alveolar margin, leaving a relatively
deep ramus ventral to the fossa. In GU/RSR/TAD 9203
and 9608, but apparently not in the holotype, there is a
small foramen near the front of the masseteric fossa,
immediately posterior to the coronoid crest and at the
vertical level of the alveolar margin of M3 (Figs 4, 5).
This buccal mandibular foramen, which appears to com-
municate with the medullary cavity of the mandible, is in
the same position as the ‘labial mandibular foramen’
described in some Mesozoic eutherians (e.g. Maelestes:
Wible et al. 2009). The content of the latter foramen is
unclear. Davis (2012) suggested a vascular, nutrient func-
tion in peramurans. Transmission of a branch of the infe-
rior alveolar nerve is also possible based on the broad
similarity to the retromolar foramen sometimes encoun-
tered in humans (e.g. Lizio et al. 2013).
The mandibular condyle is positioned well dorsal to
the level of the tooth row. The condyle is poorly pre-
served, but its base is flush with the back of the ascending
ramus and the condyle does not appear to have projected
laterally. Although the coronoid process is not preserved,
it appears to have ended somewhat anterior to the
6 PAPERS IN PALAEONTOLOGY

FIG. 4. GU/RSR/TAD 9203, left dentary fragment with M3, referred to Pahelia mysteriosa gen. et sp. nov. in: A, buccal; B, lingual; C,
occlusal; D, oblique occlusobuccal view. The oblique occlusobuccal view (D) is included to show crown relief. Abbreviation: bmf, buc-
cal mandibular foramen (labial mandibular foramen of Wible et al. 2009). Scale bar represents 10 mm.

mandibular condyle. Its development and height cannot
be assessed.
Although the tip is broken, the base of a large,
dorsoventrally deep angular process is preserved in GU/
RSR/TAD 9608. Beginning at the posterior margin of the
ascending ramus, the process appears to have projected
posteriorly, with no evidence of a ventral or medial
inflection, although there is a weak, blunt crest on the
ventral margin of the medial side of the angular process.
Dorsal to this crest, a shallow depression, presumably for
attachment of m. pterygoideus medialis, can be identified.
More dorsally on the medial side of the ascending ramus,
the opening of the mandibular foramen is positioned on
the ascending ramus, well posterior and ventral to the
dentition. The foramen opens posteriorly.
On the left side of the holotype are preserved the
crowns of P3–M3 plus roots of P2 and single alveoli of
two more anterior teeth. On the right side, the same alve-
oli up to the mesial alveolus of P3 are preserved, but only
the mesial root of P2 remains.
The most anterior complete alveolus is relatively large,
much larger than the succeeding premolars, and single-
rooted, with the root being compressed mesiodistally but
deep buccolingually. The root is oriented parallel to the
long axis of the symphysis (Fig. 6), implying a relatively
proclivous crown. The left and right alveoli are separated
by a distinct gap on the midline of the mandibular sym-
physis. Such a gap is atypical in instances in which I1 is
enlarged, suggesting that the large tooth probably repre-
sents a more posterior locus. There are faint depressions
on either side of the midline that suggest that an addi-
tional, small alveolus was present anterior to the large
alveolus, further evidence that the enlarged tooth is not
I1. However, it is not possible to determine if the
enlarged tooth represents I2, I3, or even an enlarged
canine.
The single alveolus of the next tooth is small, circular,
and set immediately posterior to the large alveolus, with
no intervening diastema. Based on the maximum dimen-
sions of the alveolus, the root of this tooth appears to
have been wider than the succeeding P2. Given that P1 is
generally smaller than P2, this alveolus probably housed
either I3 or C1, with P1 likely to be lost. The two roots of
P2 are positioned immediately posterior to the second
alveolus. Both roots of P2 are small and circular, with the
mesial root being slightly larger than the distal. While it
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA 7

F I G . 5 . GU/RSR/TAD 9608, left dentary with M2–3, referred to Pahelia mysteriosa gen. et sp. nov. in: A, buccal; B, lingual; C, occlusal
view. Abbreviations: ap, angular process; bmf, buccal mandibular foramen; cc, condyloid crest; ccr, coronoid crest; maf, mandibular
canal; mc, mandibular condyle; mf, mental foramen. Scale bar represents 10 mm.
8 PAPERS IN PALAEONTOLOGY
FIG. 6. Anteriorly inclined transverse microCT section through the mandible of GU/RSR/TAD 9201 to show alveoli, particularly the
alveoli of the enlarged anterior locus. View is from below. Note that while the alveolus of the second anterior tooth appears smaller
than the mesial root of P2, this reflects the angulation of the section. Abbreviations: a1, first anterior tooth; a2, second anterior tooth.
is possible that these roots actually pertain to two single-
rooted teeth, this is unlikely given their close appression
and the larger size of the more mesial root, which would
contrast with the typical progression of increasing premo-
lar size distally.
P3 is again set immediately posterior to P2 with no
intervening diastema and appears to be modestly larger
than the preceding tooth. The base of the crown is com-
plete, but the apex of the protoconid is broken. The
crown is dominated by the large protoconid with no
development of other trigonid cusps and only a rudimen-
tary talonid distally. The protoconid is relatively circular,
but bears distinct mesial and distal carinae. The mesial
carina makes a direct descent of the mesial margin of the
protoconid, intersecting with a transverse mesial cingulid
basally. The latter forms a narrow shelf lingually. Both
portions of the cingulid terminate distally against the base
of the protoconid. The distal protoconid carina also
makes a direct descent and has a diffuse contact with the
distal cingulid basally. The distal cingulid defines two
asymmetric shelves, a smaller buccal shelf and a narrow,
lingually sloping talonid lingually. There is no develop-
ment of differentiated talonid cusps. Instead, the distal
cingulid curves mesiolingually to form a low entocristid,
terminating against the base of the protoconid.
P4 is slightly shorter than P3 and similar in morphol-
ogy. The trigonid is also dominated by a tall, conical pro-
toconid (damaged buccally). There is no development of
a paraconid mesially, but there is also no lingual shelf
and the mesial cingulid is generally weaker than on P3.
Also unlike P3, there is a small metaconid, positioned low
on the lingual surface of the protoconid, just distal to the
midpoint of the cusp. The metaconid takes the form of a
small swelling, connected to the protoconid by a crest
that curves mesially and then buccally. In addition, the
distal protoconid carina is thickened at its terminal
end, forming an incipient cuspule distobuccal to the
metaconid.
The talonid of P4 is broader and better differentiated
than on P3, but remains very short mesiodistally. The
short cristid obliqua is directed mesiolingually from the
poorly defined hypoconid, contacting the base of the pro-
toconid. Distally, a notch separates the hypoconid from a
second, more lingual cusp, presumably an entoconid or
hypoconulid. The latter is continuous with a low ento-
cristid that, as on P3, terminates against the lingual base
of the protoconid.
M1 is slightly longer than P4, but probably lower. The
crown of M1 is worn and damaged lingually, with the
metaconid damaged and the lingual two-thirds of the
talonid missing. Given the damage, the description of
molar morphology will begin with M2. M1 morphology
appears to be nearly identical to that of M2, to the extent
that it can be determined.
M2 of the type is substantially larger than M1. In the
type, the crown of M2 is strongly exodaenodont, extending
more basally on the buccal than the lingual side, and com-
prises a trigonid and talonid that are subequal in length
(M2 of GU/RSR/TAD 9608 is heavily worn and provides
no useful morphologic information). The trigonid is bicus-
pid, comprising a buccal protoconid and a smaller meta-
conid, with no paraconid. The protoconid apex is worn,
but the cusp appears to have been somewhat compressed
transversely, with a flattened lingual surface and a convex
buccal surface. Mesially, a distinct paracristid descends the
mesial margin of the protoconid, running basally and
slightly lingually. Distally, the protoconid portion of the
protocristid appears to have had a similar basolingual ori-
entation, but its development is obscured by wear. Both
crests terminate relatively high on the crown.
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
The metaconid is smaller than the protoconid, but
appears to have been subequal in height. It is positioned
directly lingual to the protoconid. The metaconid bears
two weak and poorly defined mesial crests: a premetacris-
tid directed mesiobuccally towards the paracristid, and an
endometacristid extending directly buccally towards the
protoconid (terminology sensu Orliac & Ducrocq 2012).
Distally, the metaconid portion of the protocristid (post-
metacristid) is better defined. It is directed basally and
distobuccally, joining the protoconid portion of the pro-
tocristid in a deep trigonid notch. At the mesial margin
of the trigonid, well below the paracristid and premeta-
cristid, is a weak mesial cingulid that runs transversely
and rises slightly towards the lingual side of the crown.
Buccally, between the trigonid and talonid, the hypo-
flexid is filled by a well-developed ectostylid cusp.
Although much lower on the crown, this cusp is posi-
tioned just mesial and buccal to the apex of the hypoco-
nid. A distinct crest descends along the ectostylid
mesially, ending against the base of the protoconid.
The talonid is distinctly narrower than the trigonid and
somewhat lower. As with the trigonid, it consists of two
well-developed cusps, a slightly larger buccal hypoconid
and a smaller but slightly taller entoconid situated directly
lingual to it. A distinct hypoconulid is absent, although it
may be represented by a thickening of the distal end of
the hypocristid. As with the protoconid, the hypoconid is
somewhat transversely compressed. The cristid obliqua
extends nearly directly mesially, joining the protocristid at
the trigonid notch. On the distal side of the hypoconid,
the hypocristid also runs nearly directly distally, with only
a small lingual component. In the absence of a defined
hypoconulid, it terminates at the distal margin of the
crown and does not meet the entoconid.
Lingually, the entoconid is somewhat compressed
mesiodistally. The preentocristid is weak and is only
developed basally, forming a low ridge at the lingual mar-
gin of the talonid and terminating against the base of the
metaconid. Running transversely from the entoconid apex
towards the hypoconid is a weak crest that forms an
incipient entolophid. There is no crest directed towards
the expected position of the hypoconulid (posten-
tocristid), leaving a distinct distolingual notch between
the entoconid and hypocristid.
M3 of the holotype is larger and wider than M2 and
nearly unworn. The trigonid morphology is similar to
that of M2 (and is somewhat clearer due to little wear),
with a somewhat transversely compressed protoconid
bearing well-developed mesial and distal crests, no para-
conid, and a metaconid that is smaller but subequal in
height and positioned directly lingual to the protoconid.
As on M2, two mesial crests descend the metaconid. On
M3, however, the endometacristid is clearly stronger than
the premetacristid. Both crests are weaker than the
9
metaconid portion of the protocristid. The protocristid
morphology is clearer than on M2 due to the lack of
wear. The protoconid and metaconid portions of the pro-
tocristid converge on a deep, V-shaped notch that also
forms the termination of the cristid obliqua.
As on M2, there is a distinct ectostylid in the M3 hypo-
flexid. It is better developed in GU/RSR/TAD 9203 than
in the holotype, and is even stronger in GU/RSR/TAD
9608, where it is incipiently divided into a more mesial
and a more distal cusp. The holotype has a weak ectocin-
gulid around the base of the trigonid between the ectosty-
lid and mesial cingulid that is absent in GU/RSR/TAD
9203 and 9608. The latter specimen has a short, weak cin-
gulid descending the buccal surface of the hypoconulid.
The talonid of M3 of GU/RSR/TAD 9201 lacks a
defined hypoconid. Instead, the cristid obliqua and
hypocristid form a continuous crest that is concave
mesially and convex distally, terminating on a well-devel-
oped, transversely oriented hypoconulid that forms a
small third lobe on M3. Although the latter cusp is pre-
sent, it is isolated from the entoconid. As on M2, the
entoconid is mesiodistally compressed and its best-devel-
oped crest runs nearly transversely, although on M3 it
runs slightly mesially as well as buccally. Just distal to this
crest is a second, weaker transverse crest. The preen-
tocristid, in contrast, is essentially absent, although there
is a small entoconulid at the base of the entoconid. Due
to the absence of the hypoconid, the entoconid is the tal-
lest cusp on the M3 talonid of GU/RSR/TAD 9201.
The talonids of M3 of GU/RSR/TAD 9203 and 9608 are
more similar to the talonid of M2 of GU/RSR/TAD 9201.
Unlike M3 of the holotype, both have well-developed
hypoconids and better developed preentocristids, especially
GU/RSR/TAD 9608. Hypoconulid development in GU/
RSR/TAD 9608 is similar to that of the holotype, although
the hypocristid is slightly lower. In GU/RSR/TAD 9203,
the hypoconulid is smaller and isolated from the hypocris-
tid, forming an enlarged cingular shelf that does not con-
nect with either the hypoconid or the entoconid.
Type locality and horizon. Locality TAD-2, 2–3 m below
Lignite 1, Cambay Shale Formation, Tadkeshwar Lignite
Mine, Surat District, Gujarat, India, Ypresian, lower Eocene.
Age and distribution. In addition to the holotype, GU/
RSR/TAD 9203 is from the type locality. GU/RSR/TAD
9608 is from locality TAD-1, 10–16 m below Lignite 1.
COMPARISONS
The three specimens of Pahelia mysteriosa document a
distinctive new mammal that does not show obvious
affinities to any known group. At a most basic level, the
10 PAPERS IN PALAEONTOLOGY
dental morphology and formula of P. mysteriosa indicate
eutherian affinities. The morphology of the new taxon
includes numerous features indicative of herbivorous
adaptation, particularly the incipient lophodonty or selen-
odonty of the molar series. Among Paleocene and Eocene
eutherian mammals, the dental morphology of Pahelia is
most consistent with a relationship to the diverse taxa
broadly classed as ungulates. Early ungulates, however,
are both extraordinarily diverse and almost certainly poly-
phyletic, encompassing at least two unrelated radiations
of primarily herbivorous and terrestrial eutherian mam-
mals, one within Afrotheria and the other within
Laurasiatheria (Madsen et al. 2001; Murphy et al. 2001;
Amrine-Madsen et al. 2003; Prasad et al. 2008; Meredith
et al. 2011; O’Leary et al. 2013). In this sense, identifica-
tion of Pahelia as an ungulate does little to clarify its
affinities. A clearer understanding of the affinities of
Pahelia is required to confidently interpret the implica-
tions of the new taxon for the evolution of the Indian
mammalian fauna.
Among the broad sweep of ungulate diversity, individ-
ual features of P. mysteriosa are shared with numerous
taxa, but a smaller number of groups show sufficient sim-
ilarity to warrant more detailed comparisons. These
include four families of condylarths (European Louisini-
dae, North American Apheliscidae and Periptychidae, and
Indo-Pakistani Quettacyonidae), one family of basal artio-
dactyls (Anthracotheriidae), a family closely related to
perissodactyls (Cambaytheriidae), and two taxa known
from the early Cenozoic of Africa (Ocepeia and Hyra-
coidea). The following section compares P. mysteriosa
with each of these groups, with the goal of narrowing
down the possible affinities of Pahelia.
Louisinidae
Pahelia shares a number of similarities with the advanced
louisinid Teilhardimys and its immediate relatives (Gigar-
ton, Prolouisina, Louisina) (Russell 1964; Tabuce et al.
2006; Hooker & Russell 2012). Advanced louisinids and
Pahelia share simple premolars dominated by a single
trigonid cusp. Both show substantial molar exodaen-
odonty and a tendency to develop an ectostylid in the
hypoflexid. At least on M2–3, molar talonids are narrower
than trigonids in Teilhardimys and its relatives. As in
Pahelia, there is a tendency to reduce or lose the para-
conid and hypoconulid on lower molars. Some specimens
of Teilhardimys even show a very weakly defined entolo-
phid (e.g. Hooker & Russell 2012, fig. 27J1).
Differences include the much smaller size of known
louisinids. The mandibular symphysis is unfused in
known louisinids and, where known from alveoli, anterior
teeth are small. While premolar morphology is simple,
particularly in Teilhardimys, the talonid is proportionally
much larger on P4 than in Pahelia and the premolars are
more transversely compressed in louisinids. Molar cusps
are more basally inflated in morphologically similar loui-
sinids (e.g. Teilhardimys) than in Pahelia. Louisinids have
a mesially directed premetacristid that joins with the
paracristid to enclose the trigonid mesially. In contrast,
the accessory metaconid crest in Pahelia is directed buc-
cally, towards the protoconid. The cristid obliqua in loui-
sinids generally contacts the trigonid below the trigonid
notch, and may be indistinct or buccal to the midpoint
of the protocristid. The morphology of some specimens
of Teilhardimys reisi is more similar to Pahelia, but the
protocristid itself is poorly defined in T. reisi (Tabuce
et al. 2006). No louisinid has an entolophid as strongly
developed as in Pahelia, and the hypoconulid, while
small, is typically connected to both the hypoconid and
entoconid in louisinids. In Pahelia, even on M3, which
has a strong hypoconulid, the entoconid is isolated from
the hypoconulid. Finally, M3 is substantially smaller in
morphologically similar louisinids and the hypoconulid is
effectively absent in the most derived species, contrasting
with the enlarged M3 and hypoconulid lobe in Pahelia.
A further consideration is that the greatest similarity
between Pahelia and louisinids is with the temporally late,
morphologically advanced Teilhardimys. Earlier and/or
more conservative taxa (e.g. Paschatherium, Walbeckodon)
show additional differences from Pahelia (more complex
premolars; larger paraconids and hypoconulids; lack of an
ectostylid) (Denys & Russell 1981; L
opez-Mart
ınez et al.
2006; Hooker & Russell 2012). Therefore, unless Pahelia
is deeply nested within Louisinidae, the similarities to
advanced louisinids are likely to be convergent.
Apheliscidae
Members of the North American condylarth family Aphe-
liscidae show some similarity to Pahelia, particularly Hap-
laletes and its likely descendant Utemylus as well as
members of Apheliscinae (Gazin 1956, 1959; Gingerich
1983; Zack et al. 2005a). Haplaletes and Utemylus resem-
ble Pahelia in having strongly exodaenodont molars and
lingual molar cusps that approach or even exceed the
height of corresponding buccal cusps. These taxa also
share with Pahelia reduced molar paraconids, a morphol-
ogy also present in the apheliscine Apheliscus, resulting in
short trigonids. As a group, apheliscids also have molar
metaconids positioned directly lingual to the protoconid.
Aside from the presence of a tall entoconid in Haplaletes,
apheliscid molar talonids resemble those of Pahelia in
having strongly reduced hypoconulids on M1–2. Most
apheliscids also have a narrow M3 talonid with a mod-
estly enlarged hypoconulid, again similar to the
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
morphology present in P. mysteriosa. More anteriorly,
apheliscid premolars are relatively unmolarized, except in
Haplomylus; and some taxa, including Aletodon and mem-
bers of Apheliscinae, have simple, premolariform premo-
lars very similar to those of Pahelia.
Despite these similarities, apheliscids also show signifi-
cant contrasts with Pahelia. Anteriorly, where known,
apheliscids have an unfused mandibular symphysis. There
is no evidence for substantial incisor or canine enlarge-
ment in any apheliscid, although the incisor region is
poorly known in many taxa. The anterior dentition of
apheliscids is also unreduced, where known, contrasting
with P. mysteriosa.
While premolar complexity may be similar to Pahelia,
apheliscid lower premolars, particularly P3–4, are derived
in being relatively elongate, reflecting elongation of the
rostrum. This contrasts with Pahelia, in which both the
rostrum and P3–4 are comparatively short. Turning to
the molars, no known apheliscid has an ectostylid,
although a buccal cingulid is variably developed in
Haplaletes.
If trigonid morphology is relatively similar, aside from
hypoconulid reduction, apheliscid talonids differ substan-
tially. In all apheliscids, talonids of M1–2 are either sub-
equal in width to or wider than trigonids, and overall
talonid size is larger. In apheliscids, the cristid obliqua
contacts the back of the trigonid below or buccal to the
trigonid notch, rather than contacting the protocristid as
in P. mysteriosa. While small, the hypoconulid in aphelis-
cids is connected to both the hypoconid and entoconid,
and there is no development of a transverse entolophid in
any apheliscid. Finally, apheliscid molar sizes contrast
with Pahelia. In many apheliscid taxa, M2 is slightly larger
than M1 rather than substantially larger, as in Pahelia.
With the exception of Apheliscus, where they are sube-
qual, M3 is smaller than M2 in apheliscids, contrasting
with the enlarged M3 present in Pahelia.
Periptychidae
Pahelia is superficially similar to members of another
North American condylarth family, Periptychidae, partic-
ularly to relatively small-bodied forms in the subfamilies
Anisonchinae and Conacodontinae, as well as the early
periptychine Hemithlaeus (Matthew 1937; Archibald et al.
1983a, b; McKenna et al. 2008). Like Pahelia, periptychids
tend to have relatively compact, simple premolars lacking
well-developed metaconids or talonids. Small periptychids
also share relatively tall molar cusps and comparatively
small talonids. Anisonchines tend to reduce or lose the
molar paraconid (e.g. Goleroconus, Haploconus), making
them particularly similar to Pahelia (Matthew 1937;
McKenna et al. 2008).
11
If the features just mentioned combine to produce a
superficial similarity between periptychids and Pahelia,
there are also numerous differences in detail that indicate
that a close relationship is unlikely. Known periptychids
have an unfused mandibular symphysis and relatively
small anterior teeth. Periptychids show no development
of exodaenodonty. If anything, the lingual sides of the
lower molars of Haploconus project further basally than
the buccal sides. On the molar trigonids, periptychids
have an oblique protocristid with the metaconid distolin-
gual to the protoconid rather than the transverse pro-
tocristid of Pahelia. Additionally, the protoconid and
metaconid are subequal in size and height in periptychids,
whereas the metaconid is smaller than the protoconid in
Pahelia. Although periptychid molar talonids are relatively
small, they are generally subequal to or wider than the
trigonids, rather than distinctly narrower. The cristid
obliqua contacts the back of the trigonid more basally,
and because of this it fails to join the protocristid. More
distally on the talonid, the hypoconulid is distinct, and
the three major talonid cusps are joined by a continuous
postcristid (hypocristid plus postentocristid) at the distal
margin of the talonid. In contrast, Pahelia lacks posten-
tocristids on M2–3 and a distinct hypoconulid on M2, and
the hypocristid is directed distally rather than distolin-
gually on both molars. Periptychids lack the incipient
entolophid that is present in Pahelia. The entoconid of
periptychids further differs in joining a well-developed
entocristid that closes off the talonid basin lingually.
Additionally, the entoconid itself projects lingual to the
trigonid in periptychids, while its apex is directly distal to
the metaconid apex in Pahelia. Cingular development is
stronger in periptychids; most taxa have a distinctive
mesiolingual cingulid at the base of the trigonid that is
absent in Pahelia. Buccally, there is no development of an
ectostylid in any periptychid taxon. In contrast to the dis-
tal increase in molar size in Pahelia, molar sizes are sube-
qual in periptychids, particularly M2–3. M1 is slightly
smaller than M2–3 in some anisonchines (e.g. Mithrandir),
but in other periptychids it is subequal to M2–3. Taken
together, the substantial number of differences suggests
that the similarities between Pahelia and periptychids rep-
resent convergence or parallelism.
Quettacyonidae
Another possible relative of Pahelia among condylarths is
Quettacyonidae, a group of relatively large-bodied condy-
larths endemic to the early Eocene of Pakistan (Gingerich
et al. 1997, 1998, 1999). Quettacyonids are similar to
Pahelia in having simple premolars dominated by the
protoconid, with limited development of other trigonid
cusps and a small talonid. Like Pahelia, quettacyonid
12 PAPERS IN PALAEONTOLOGY
lower molars frequently lack a paraconid and have the
cristid obliqua connected to the midpoint of the pro-
tocristid. Development of the back of M3 is also similar,
with both taxa having an M3 hypoconulid that is larger
than M1–2 but does not form a full third lobe.
There are, however, numerous distinctions between the
dentitions of Pahelia and members of Quettacyonidae.
Quettacyonids are distinctly larger. Anteriorly, Machocyon
has an unfused mandibular symphysis (the symphysis is
unknown in other quettacyonids). Premolar protoconids
are erected in Pahelia versus retroflexed in quettacyonids,
where they have an inclined mesial and vertical distal
margin. Molar cusps are much lower crowned in quetta-
cyonids and show no development of exodaenodonty.
Molar buccal cingulids are stronger in quettacyonids and
there is no tendency to develop an ectostylid. Quettacy-
onid trigonids are more compressed mesiodistally than in
Pahelia and, unlike quettacyonids, Pahelia does not have
a trigonid fovea. Talonid width is subequal to trigonid
width in quettacyonids and there is no development of an
entolophid, with the exception of Sororocyon usmanii
(Gingerich et al. 1998). The hypoconulid is well devel-
oped on all three molars of quettacyonids and is twinned
with the entoconid, resulting in a longer, more transverse
hypocristid than in Pahelia. Finally, M2 is the largest
lower molar in quettacyonids, rather than M3 in Pahelia.
Cambaytheriidae
The perissodactyl relative Cambaytherium (also from the
early Eocene Cambay Shale Formation) and Pahelia share
a fused mandibular symphysis and the presence of an
enlarged anterior tooth (C1 in the case of Cam-
baytherium) (Bajpai et al. 2005a, 2006; Rose et al. 2014).
Although there is substantial variation, some individuals
of Cambaytherium have simple premolars (e.g. GU 7004).
Both taxa lack molar paraconids, have distally directed
hypocristids, and lack a distinct postentocristid. In both
genera, molar size increases posteriorly.
Beyond these similarities, there are substantial differ-
ences in molar morphology between Cambaytherium and
Pahelia. At a basic level, the molars of Cambaytherium
are much more bunodont than those of Pahelia. While
both genera lack molar paraconids, Cambaytherium has a
complete paracristid running from the protoconid to the
metaconid to enclose a trigonid fovea. In contrast, the
paracristid of Pahelia is short and terminates in the valley
between the protoconid and metaconid. While Pahelia
does have a mesial crest emerging from the metaconid: it
runs directly buccally and does not contact the paracris-
tid. In Cambaytherium, the metaconid is subequal in size
to the protoconid and positioned distolingual to that
cusp, contrasting with Pahelia in which it is distinctly
smaller than the protoconid and directly lingual. Addi-
tionally, the metaconid is twinned in Cambaytherium, fur-
ther extending the trigonid distolingually.
Molar talonid widths are subequal to trigonid widths
in Cambaytherium, while talonids are distinctly narrower
than trigonids in Pahelia. The cristid obliqua contacts the
trigonid below the protoconid in Cambaytherium, rather
than joining with the protocristid at the trigonid notch as
in Pahelia. In Cambaytherium, the hypoconulid is promi-
nent on all molars, contrasting with the absence of this
cusp on M2 (and presumably M1) of Pahelia. The ento-
conid is distolingual to the hypoconid in Cambaytherium
rather than directly lingual. While both genera lack a con-
nection between the entoconid and the area of the hypo-
conulid, Cambaytherium differs from Pahelia in lacking
an incipient entolophid. On M3, Cambaytherium consis-
tently has multiple hypoconulid cusps (two to four), pro-
ducing a well-developed third lobe. While there is
variation in the structure of M3 in Pahelia, no specimen
has any accessory cusp beyond a prominent hypoconulid.
Finally, like most compared taxa, Cambaytherium lacks
molar exodaenodonty and ectostylids.
Anthracotheriidae
While Pahelia differs substantially from known Vastan
artiodactyls (Diacodexis indicus and D. parvus: Bajpai et al.
2005a; Kumar et al. 2010), which have typical dichobu-
noid features, including retention of the paraconid on
lower molars and a broad talonid on which the hypoconu-
lid is shifted lingually towards the entoconid, there are
similarities to somewhat younger Asian artiodactyls, par-
ticularly early representatives of Anthracotheriidae (Col-
bert 1938; Suteethorn et al. 1988; Tsubamoto et al. 2002;
Soe 2008). Some early anthracotheres have a fused
mandibular symphysis (Colbert 1938), in common with
Pahelia. Premolar morphology in early artiodactyls is gen-
erally simple, with anthracotheres being no exception, and
the most basal known anthracothere, Siamotherium, has
premolars that are proportionately similar to those of
Pahelia (Suteethorn et al. 1988). Unlike Diacodexis,
anthracotheres strongly reduce or lose the paraconid, and
some develop a transverse crest from the metaconid
towards the protoconid, both features shared with Pahelia.
Similarly the cristid obliqua joins the protocristid in some
early anthracotheres (e.g. Anthracokeryx birmanicum: Soe
2008). The hypoconulid is retained on M1–2 of anthra-
cotheres, but it resembles Pahelia in being isolated from
the entoconid. Instead, there is variable development of an
entolophid, as in the case of Pahelia. Finally, M3 is the lar-
gest molar in anthracotheres, while M1 is the smallest.
Differences are also significant. Aside from Siamoth-
erium, early anthracotheres have longer, narrower
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
premolars than Pahelia, and a stronger metaconid is gen-
erally present on P4. Siamotherium itself lacks several fea-
tures characteristic of Pahelia. It has an unfused
symphysis, lacks a transverse metaconid crest, and the
cristid obliqua terminates below the protocristid. More
generally, early anthracotheres are more bunodont and
lack exodaenodont molars and an ectostylid. Anthra-
cotheres have broader talonids than trigonids, and the
height discrepancy between the trigonid and talonid is
less than in Pahelia. As noted above, the hypoconulid is
stronger on all molars in anthracotheres. This is particu-
larly true of M3, where the hypoconulid is relatively larger
than in Pahelia and forms a distinct third lobe. The fact
that the most basal anthracothere lacks several features
that suggest a link between Pahelia and other anthra-
cotheres is particularly compelling evidence that the simi-
larities that are present are probably convergent.
Ocepeia
Among early African ungulates, the possible stem paenun-
gulate Ocepeia (Gheerbrant et al. 2001, 2014; Gheerbrant
2010) shows some similarities to Pahelia. Pahelia and
Ocepeia share simple, premolariform premolars, including
P4. Ocepeia grandis and Pahelia also have similar develop-
ment of the P4 metaconid as a small, curved ridge low on
the lingual side of the protoconid (observed on a cast of
MNHN.F PM37), although a metaconid is lacking entirely
in O. daouiensis. Molars share the distinctive presence of
an incipient entolophid. A postentocristid connecting the
entoconid and hypoconulid is absent in O. daouiensis and
P. mysteriosa but present in O. grandis.
Despite these limited similarities, there are numerous
substantial differences that make a close relationship appear
unlikely. Ocepeia has a mandibular symphysis described as
‘partly fused’ (Gheerbrant et al. 2014) and a single-rooted
P2, while Pahelia has an extensively fused symphysis and a
probably double-rooted P2, assuming our interpretation of
the alveoli mesial to P3 in Pahelia is correct. While both
taxa have simple premolars, Ocepeia has a very poorly
developed talonid on P4, with the distal cristid of the proto-
conid extending to the distal margin of the crown. In con-
trast, Pahelia has a small but distinct talonid on P4.
Molar morphology shows more dramatic distinctions.
At a basic level, the molar morphology of Ocepeia is
much more bunodont and less lophodont than in Pahelia.
Turning to specific features, Ocepeia has molar talonids
that are equal to or wider than trigonids, contrasting with
the narrow talonids present in Pahelia. Unlike Pahelia,
Ocepeia retains well-developed molar paraconids that are
placed at the lingual margin of the crown and connected
to the protoconids by strong paracristids. Perhaps as a
result, molar metaconids are positioned distolingual to
13
the protoconids in Ocepeia, compared with directly lin-
gual in Pahelia. Ocepeia also has a distinct distal crest on
the metaconid that functions to extend the protocristid
distolingually. Pahelia lacks such a crest. Talonids of Oce-
peia have a strongly developed mesoconid that interrupts
the elongate cristid obliqua, while the latter crest is short
and unbroken in Pahelia. The cristid obliqua contacts the
trigonid beneath the trigonid notch in Ocepeia, rather
than joining the protocristid. Ocepeia also retains well-
developed hypoconulids on M1–2, while this cusp is lack-
ing at these loci in Pahelia. The hypoconulid is lingually
positioned in Ocepeia, resulting in the presence of elon-
gate hypocristids on M1–2. In Pahelia, the hypocristids on
M1–2 are short and directed almost directly distally. On
M3, Ocepeia has a much more dramatically expanded
hypoconulid lobe. Molar sizes also differ, with Ocepeia
having M1–3 of subequal size, aside from the expanded
hypoconulid lobe of M3. This contrasts with the progres-
sive distal size increase seen in Pahelia. Finally, the most
distinctive features of the Pahelia dentition (strong exo-
daenodonty, prominent ectostylids) are absent in Ocepeia.
Brief comment on another basal paenungulate,
Abdounodus hamdii, is also warranted. Pahelia and
Abdounodus share simple premolars, with an essentially
unmolarized P4. One M3 of A. hamdii (OCP DEK/GE
308) has a small ectostylid, although this does not appear
to be a consistently developed feature. Beyond this, there
is little to suggest a relationship. Abdounodus has an
unfused symphysis, single-rooted P2, strongly bunodont
molars lacking well-developed crests, talonids subequal or
wider than trigonids, prominent paraconids, low but dis-
tinct (and twinned) hypoconulids, and subequal M1–3,
with the possible exception of the number of P2 roots, all
substantial contrasts with Pahelia. Unlike other early Afri-
can ungulates, A. hamdii lacks a transverse entolophid,
and the entoconid is isolated from other talonid cusps.
There is nothing in the morphology of A. hamdii to sug-
gest a particularly close relationship to Pahelia.
Hyracoidea
Similarities between Pahelia and early members of Hyra-
coidea (Court & Mahboubi 1993; Rasmussen & Simons
2000; Tabuce et al. 2001; Barrow et al. 2010) are some-
what stronger than with Ocepeia. Pahelia and early hyra-
coids share a fused mandibular symphysis. Preserved
evidence of the anterior dentition is very suggestive, given
that both Pahelia and hyracoids have multiple proclivous
anterior teeth set at a distinct angle to the remainder of
the dentition, including an enlarged, non-median anterior
tooth (Fig. 6). In the cheek dentition, both early hyra-
coids and Pahelia show reduction of the molar paraconids
and the hypoconid on M2 (and probably M1), lack of a
14 PAPERS IN PALAEONTOLOGY
connection between the entoconid and hypoconulid on
M2, general molar bunolophodonty, and a progressive
increase in molar size from M1 to M3.
Significant differences include a more elongate rostrum
with accompanying elongate premolars in hyracoids,
more molarized P3–4 in hyracoids (including the oldest
known hyracoid, early Eocene Seggeurius: Benoit et al.
2016), which have well-developed metaconids on P4 and
large, relatively complex talonids on P3–4. In the molars,
the size gradient between M1 and M3 is more pronounced
in Pahelia and the M2 hypoconulid is more reduced. In
Pahelia, M1–2 trigonids are substantially wider than their
respective talonids, but they are subequal in hyracoids.
Pahelia has stronger protocristids and entolophids than
Seggeurius, although slightly more advanced taxa (e.g.
Dimaitherium) show similar (or stronger) development of
both crests. Pahelia lacks a metastylid or any development
of a twinned metaconid (well-developed features in even
the earliest hyracoids). Finally, hyracoids lack an ectosty-
lid or the strongly asymmetric enamel present in Pahelia.
PHYLOGENETIC ANALYSIS
None of the taxa compared with Pahelia shows clear and
convincing evidence of affinity. The resemblance of Pahe-
lia to a few taxa, particularly periptychids, is limited and
superficial, and a close phylogenetic link appears unlikely.
In most instances, however, at least some seemingly com-
pelling similarities are present but they are balanced by
equally significant contrasts. With evidence limited to the
lower cheek teeth and mandible, it is difficult to deter-
mine which (if any) features are providing a real signal of
relationships and which reflect convergence or parallelism.
It is for precisely this purpose that phylogenetic analysis
was introduced: to objectively compare conflicting
hypotheses of relationships with differing suites of sup-
porting character evidence.
One difficulty in testing the relationships of Pahelia is
the broad range of possible affinities for the new genus.
The taxa to which Pahelia shows meaningful similarities
are themselves phylogenetically diverse. Hyracoids and
probably Ocepeia are paenungulate afrotheres (Gheerbrant
et al. 2014, 2016), while apheliscids and louisinids may be
non-paenungulate afrotheres (Zack et al. 2005b; Penkrot
et al. 2008; Hooker & Russell 2012). Cambaytheres,
anthracotheriids and probably periptychids are laura-
siatheres, only distantly related to afrotheres. Quettacy-
onids are themselves of undetermined affinities. No
existing character–taxon matrix includes more than a few
of these taxa. Quettacyonids have never been included in
a published phylogenetic analysis.
To investigate the affinities of P. mysteriosa, the new
taxon was initially added to three existing matrices that
included one or more of the taxa that share similarities
with Pahelia. Specifically, Pahelia was added to the char-
acter–taxon matrices found in Tabuce et al. (2011), Mui-
zon et al. (2015) and Gheerbrant et al. (2016). A
representative quettacyonid, Machocyon abbasi was also
added to the first mentioned matrix. Results were highly
variable and contradictory, with Pahelia recovered in a
variety of mutually incompatible positions, failing to
meaningfully clarify the relationships of the new genus.
Methods and results of these initial analyses are presented
in Zack et al. (2019a, appendix S1).
None of the initial matrices included all, or even most,
taxa potentially relevant to the relationships of Pahelia.
To remedy this situation, an additional character–taxon
matrix was constructed consisting of 92 dental characters
(10 ordered) scored for 71 ingroup and two outgroup
taxa. Characters and scorings are modified from the char-
acter–taxon matrix developed as part of the senior
author’s unpublished dissertation and subsequently used
as one of two primary data sources in the recent analysis
of eutherian phylogeny by Halliday et al. (2015). In addi-
tion to Pahelia and all taxa showing significant similarity
to Pahelia, taxa sampled for this present analysis comprise
a diversity of early Cenozoic ungulates, including early
representatives of Artiodactyla, Perissodactyla, Hyra-
coidea, Proboscidea, Macroscelidea, Litopterna and
Notoungulata, as well as a variety of taxa that have, at
one time or another, been classified in the wastebasket
taxon Condylarthra. In addition to Pahelia, three quetta-
cyonid taxa (Machocyon abbasi, Quettacyon parachai and
Sororocyon usmanii) were scored. However, scorings for
Q. abbasi were less complete but otherwise identical to
those of M. abbasi, so Q. abbasi was excluded from the
analysis following the principle of safe taxonomic reduc-
tion (Wilkinson 1995). Along with Pahelia and represen-
tative of quettacyonids, the Late Cretaceous Indian
ungulate Kharmerungulatum vanvaleni was also included
to test for a potential affinity of Pahelia to other early
Indian ungulate taxa. The matrix is available in Zack
et al. (2019b), along with the three initial matrices
described above. A list of specimens used to score the
new matrix is available in Zack et al. (2019a, appendix
S2) and a character list in Zack et al. (2019a, appendix
S3).
The matrix was analysed in TNT version 1.5 (Goloboff
& Catalano 2016). The matrix was initially analysed using
the Ratchet algorithm, with the analysis rerun until mini-
mum length trees were recovered one hundred times.
Shortest trees (most parsimonious trees; MPTs) recovered
by this algorithm were submitted for TBR branch swap-
ping to fully sample minimum length trees. Dummy
characters were used to construct constraint analyses and
to enforce ingroup monophyly. The dummy characters
were deactivated prior to calculation of tree statistics.
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
Because results of the analysis are poorly resolved, at least
with regard to the relationships of Pahelia (see below), no
attempt was made to calculate node support statistics
(e.g. bootstrap or Bremer support). Character state distri-
butions were examined using WinClada 1.00.08 (Nixon
2002).
Results
Analysis of the new matrix produced 700 shortest trees
(length, 444; consistency index, 0.60; retention index,
0.24). The strict consensus is poorly resolved, with most
taxa, including Pahelia, collapsed into a large polytomy at
the base of the ingroup. Most groups that are recovered
by the strict consensus are distinctive, widely recognized
clades (e.g. Hyracoidea), although there are a few sur-
prises such as the grouping of the aberrant arctocyonid
Deltatherium with the primitive periptychids Maiorana
and Mimatuta. The majority rules consensus is somewhat
better resolved (Fig. 7), but Pahelia remains part of a
large polytomy at the base of the ingroup. Examination
of the constituent trees demonstrates that Pahelia itself is
responsible for much of the poor resolution.
In all trees, quettacyonids and proboscideans plus basal
African ungulates diverge near the base of the ingroup
while apheliscids, louisinids, and macroscelideans are
nested deeply within the ingroup. In exactly 50% of trees
(350 out of 700), Pahelia is recovered as the sister taxon of
the apheliscid Haplaletes disceptatrix (Fig. 8), widely sepa-
rated from quettacyonids and proboscideans plus basal
African ungulates. In the remaining 50% of trees, Pahelia
diverges basally, close to quettacyonids and proboscideans
plus basal African ungulates (Fig. 9A) and widely separated
from apheliscids and their allies. The strict and majority
rules consensus trees collapse all of the nodes between these
two positions, resulting in a poorly resolved tree. In the sec-
ond group of trees, Pahelia, quettacyonids and pro-
boscideans/basal African ungulates can have three basic
relationships (Fig. 9B–D). First, Pahelia can be nested
within Quettacyonidae, with this clade forming the sister
taxon to the proboscidean/basal African ungulate clade at
or near the base of the ingroup (Fig. 9B). Second, the same
clade of Pahelia and quettacyonids can diverge just above
the proboscidean/basal African ungulate clade (Fig. 9C).
Third, Pahelia and quettacyonids can form a pectinate ser-
ies at the base of the proboscidean/basal African ungulate
clade (Fig. 9D). The remaining Indian taxon, Kharmerun-
gulatum vanvaleni, is not recovered close to other Indian
taxa in any tree. Instead, it emerges as the sister taxon of
either the basal mesonychian Oxyprimus cuspidatus or the
mioclaenine hyopsodontid Mioclaenus turgidus.
To assess how much less parsimonious other potential
relationships are, the analysis was rerun with Pahelia
15
constrained to monophyly with various groups not sup-
ported by the MPTs. Unsurprisingly, given that the two
taxa were closely allied in some unconstrained trees, con-
straining Pahelia and Ocepeia to monophyly produces trees
only a single step longer than the MPTs. Constraining
Pahelia and Louisinidae to monophyly adds five steps, but
when Chambius is excluded from the constraint definition
the added length is reduced to one additional step. In con-
trast, constraining Pahelia to monophyly with either Perip-
tychidae or Artiodactyla adds six steps, while an exclusive
relationship to hyracoids adds four. Finally, constraining
early Indian endemic ungulates (Pahelia, Machocyon,
Kharmerungulatum) to monophyly adds two steps, as does
an exclusive constraint of Pahelia plus Kharmerungulatum.
BIOGEOGRAPHICAL IMPLICATIONS
Because it supports two very different relationships for
Pahelia, the phylogenetic analysis leaves considerable
uncertainty concerning Pahelia’s biogeographical affinities.
The new genus may be allied with North American Aphe-
liscidae and, more distantly, with European Louisinidae
and African Macroscelididae. Alternatively, Pahelia may
be most closely allied to another Indian endemic clade,
Quettacyonidae, with both groups potentially linked to
Proboscidea. The latter relationship would support a
broader relationship for both Pahelia and Quettacyonidae
with Afrotheria, a clade whose Paleocene–Eocene record
is primarily restricted to Africa, but with significant
potential records known from Holarctica, most notably in
the form of apheliscids and louisinids (Zack et al. 2005b;
Penkrot et al. 2008; Hooker & Russell, 2012). In sum, the
presence of Pahelia in the early Eocene of India could
indicate early Palaeogene faunal exchange either with
Holarctic continents or with Africa. At the same time, the
potential link between Pahelia and Quettacyonidae would
point to an extensive endemic history on the Indian sub-
continent, regardless of broader biogeographical links.
Pahelia and Holarctic faunas
A link between Pahelia and either Apheliscidae or Louisi-
nidae would align with the pattern shown by other com-
ponents of the Cambay Shale mammalian fauna in
linking the new taxon to North American and European
Paleocene and Eocene mammals. Artiodactyls, euprimates,
rodents, bats and tillodonts from the Cambay Shale For-
mation all have close relatives in Europe (Smith et al.
2007; Rana et al. 2008; Rose et al. 2009a, b; Kumar et al.
2010). In the case of tillodonts and artiodactyls, links to
early Eocene North American mammals are also evident
(Rose et al. 2009b; Kumar et al. 2010).
16 PAPERS IN PALAEONTOLOGY

FIG. 7. Majority rules consensus of 700 most parsimonious trees derived from analysis of the character taxon matrix described in
the text. Numbers below branches indicate percentage support, where less than 100%.
ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA 17

FIG. 8. One of two basic topologies among the most parsimo-

nious trees recovered by the phylogenetic analysis, the consensus
of which is illustrated in Figure 7. Strict consensus of 350 trees
in which Pahelia mysteriosa and Haplaletes disceptatrix are recov-
ered as sister taxa.

One notable contrast between Pahelia and other Cam-

bay Shale taxa with potential European and/or North
American affinities concerns the likely time depth of a
connection. Despite their evident similarities, Pahelia dif-
fers substantially from any known apheliscid or louisinid,
implying a relatively distant connection. In contrast,
other Cambay Shale mammals showing links to Europe
and North America indicate more immediate faunal
exchange. In the case of artiodactyls and rodents, species
from India and Europe are congeneric (Diacodexis and
Meldimys, respectively) (Rana et al. 2008; Kumar et al.
2010), as are several bats (Archaeonycteris, Hassianycteris,
Icaronycteris) (Smith et al. 2007). Diacodexis and
Icaronycteris are also shared at the generic level with
North American faunas. Individual euprimate and til-
lodont genera are not shared between India and either
Europe or North America, but Ni et al. (2016) found the
European euprimate species Donrussellia gallica to be
nested within a clade of Cambay Shale primates (along
with D. provincialis in some analyses). These strong simi-
larities among European, North American and Indian
taxa indicate immediate contacts, a conclusion strength-
ened by the fact that artiodactyls, euprimates and til-
lodonts are unknown from the European Paleocene, nor
are artiodactyls or euprimates known from the North
American Paleocene. Given its more distinct morphology,
it is unlikely that Pahelia arrived in India in the same
dispersal event or events that brought artiodactyls, bats,
euprimates, rodents and tillodonts to the subcontinent
(or to Europe if the direction of dispersal is reversed).
Instead, Pahelia would suggest either a much earlier dis-
persal event or, less likely, a sustained faunal exchange
spanning much of the early Palaeogene.

Pahelia and African faunas

The potential link between Pahelia and Afrotheria identi-

fied by the phylogenetic analysis, combined with the pri-
marily African distribution of early Cenozoic afrotheres,
raises the possibility that Pahelia documents faunal
exchange between Africa and India. Although some
palaeobiogeographical models suggest the potential for
early Palaeogene faunal exchange between Africa and
India (e.g. Ali & Aitchison 2008; Chatterjee & Scotese
18 PAPERS IN PALAEONTOLOGY
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA 19

FIG. 9. The second basic topology among the most parsimonious trees recovered by the phylogenetic analysis, the consensus of
which is illustrated in Figure 7. A, strict consensus of 350 trees in which Pahelia mysteriosa is associated with quettacyonids. B–D,
detail of the basal relationships to show different sub-topologies: B, strict consensus of 118 trees in which P. mysteriosa is nested within
Quettacyonidae as the sister taxon of basal African ungulates plus Proboscidea; C, strict consensus of 114 trees in which P. mysteriosa
is nested within Quettacyonidae, with this group positioned above basal African ungulates plus Proboscidea; D, strict consensus of 118
trees in which P. mysteriosa and Quettacyonidae form a pectinate series basal to basal African ungulates plus Proboscidea.

2010; Chatterjee et al. 2013), to date, faunal evidence for
an African (or more generally Gondwanan) affinity of the
Cambay Shale fauna has come primarily from the her-
petofauna (Smith et al. 2016). Aside from Pahelia, the
only mammalian taxon with potential African affinities is
the possible adapisoriculid Bharatlestes kalamensis (=Indo-
lestes kalamensis and Bharatlestes kalami) (Kapur et al.
2017a, b: the specific name introduced in Kapur et al.
2017b, kalami, appears to be an unjustified emendation of
kalamensis). Most of the potential herpetological links
involve semiaquatic taxa for which over-water dispersal is
plausible. In the case of madtsoiid snakes, the apparent
link between Cambay Shale and African taxa may reflect
limited sampling of Gondwanan snakes rather than evi-
dence of a specific connection between Africa and India
(Rio & Mannion 2017). With regard to Bharatlestes, in
the phylogenetic analysis presented by Kapur et al.
(2017a, b), Bharatlestes was recovered as the sister taxon
to a European clade of adapisoriculids (Bustylus and
Adapisoriculus), rather than to African taxa (e.g. Afrodon
chleuhi), making a biogeographical link to Europe more
plausible than a link to Africa.
An additional difficulty with linking Pahelia and quet-
tacyonids specifically to African mammals (as opposed to
Afrotheria as a clade) concerns the timing of potential
faunal exchange between Africa and the Indian subconti-
nent. While the Indian subcontinent was linked to Africa
via Madagascar until approximately 85 Ma (Masters et al.
2006), it is highly unlikely that similarities shared by
Pahelia and quettacyonids with stem proboscideans are a
relic of biogeographical links dating back to the Creta-
ceous. Molecular divergence estimates, which tend to be
substantially older than estimates based on the fossil
record, place the initial radiation of Paenungulata at less
than 80 Ma (Meredith et al. 2011), after the separation of
India and Africa. Any direct phylogenetic link between
Indian taxa and Proboscidea must reflect dispersals after
the separation of India (and Madagascar) from Africa
(e.g. Chatterjee et al. 2017). Chatterjee & Scotese (2010)
raise the possibility of dispersal along an island arc link-
ing India and the Arabian Peninsula at the eastern edge
of the African continent (see also Chatterjee et al. 2017),
but the limited evidence for African links in the Cambay
Shale fauna as a whole and limited options for faunal
exchange between Africa and India make a Cenozoic Afri-
can origin for Pahelia appear unlikely.
Pahelia and Indian faunas
Discussion of both a Holarctic and an African affinity for
Pahelia have presupposed some sort of early Cenozoic
faunal exchange with India. The close affinity between
Pahelia and Quettacyonidae in 50% of trees raises the
possibility of more substantial endemism on the Indian
subcontinent. While Pahelia shares intriguing dental simi-
larities with quettacyonids, there are substantial differ-
ences outlined above, most notably in the much more
bunodont morphology of known quettacyonids, and any
relationship was probably somewhat distant. The sister
taxon relationship between P. mysteriosa and Sororocyon
usmanii in many trees is probably an artefact of the lim-
ited differentiation between scorings of S. usmanii and
Machocyon abbasi, which differ in a single character state.
Given the uniform morphology of known quettacyonids,
it is unlikely that they are paraphyletic with regard to the
contemporaneous but very distinct Pahelia. If Pahelia and
Quettacyonidae are sister taxa, the restriction of both to
the Indian subcontinent would suggest that they form
part of an endemic radiation of herbivorous mammals.
The degree of divergence between Pahelia and quettacy-
onids would further indicate that the time depth of such
a radiation must be sought early in the Paleocene, if not
earlier. A relatively ancient divergence for Pahelia would
be consistent with the apparently primitive presence of a
buccal mandibular foramen, a feature recorded primarily
in Mesozoic mammals (Davis 2012).
The possibility of an endemic Indian clade comprising
Pahelia and Quettacyonidae should not be surprising in
view of what is known of the palaeogeography of India
prior to the early Eocene. With the partial exception of
Madagascar, India appears to have been isolated from
other continental land masses for much of the Cretaceous
and Paleocene (Masters et al. 2006; Chatterjee et al.
2017), an interval that includes the Cretaceous–Palaeo-
gene (K/P) boundary mass extinction. The origin of Pahe-
lia and quettacyonids might ultimately be from taxa that
dispersed via island arcs (e.g. Chatterjee & Scotese 2010;
Rose et al. 2014; Kapur et al. 2017a, b) (or some other
mechanism) early in the Paleocene or from endemic sur-
vivors of the K/P event. Such early dispersers or endemic
holdovers from the Cretaceous would have formed the
basis of an endemic mammalian radiation in India occur-
ring in parallel with Paleocene mammalian radiations on
20 PAPERS IN PALAEONTOLOGY
other continents, one that was subsequently decimated by
the arrival of taxa from northern continents close to the
Paleocene–Eocene boundary.
The Late Cretaceous Indian mammal Kharmerungula-
tum vanvaleni, known from a single worn lower molar,
shows evidence for incipient herbivorous adaptation (Pra-
sad et al. 2007) and could potentially underlie a post-K/P
herbivorous radiation. While K. vanvaleni was not linked
to Pahelia or Machocyon in the MPTs, only two addi-
tional steps are required to recover trees in which these
taxa form a clade, a minimal increase of less than one
half of 1% of total tree length. Unfortunately, K. van-
valeni is known only from a single, worn lower molar,
which does not provide a wealth of phylogenetically
informative character data, making any assessment of its
relationships to better-known taxa tenuous. Until addi-
tional material becomes available, the best that can be
said is that a relationship between Pahelia and Quettacy-
onidae is not substantially unparsimonious.
With regard to the more completely documented Quet-
tacyonidae, the disparity between Pahelia and quettacy-
onids is within the range of morphological disparity seen
in Paleocene–Eocene ungulate radiations on other conti-
nents. In North America, the morphological contrast
between the selenolophodont phenacodontids (e.g. Menis-
cotherium) and bunodont arctocyonids (e.g. Anacodon)
exceeds the disparity between Pahelia and quettacyonids.
The same is true in South America, where the Itabora
ı
fauna, probably similar in age to the Cambay Shale
(Woodburne et al. 2014), includes lophodont notoungu-
lates (e.g. Colbertia) alongside bunodont didolodontids
(e.g. Lamegoia) (Paula Couto 1952a, b; Cifelli 1983).
At present, the hypothesis that Pahelia and Quettacy-
onidae are members of an endemic Indian radiation of
ungulate-like herbivorous mammals appears to best fit
the morphological and biogeographical evidence. Linking
Pahelia with North American Apheliscidae is equally par-
simonious from a morphological perspective, but implies
a deep time connection to Holarctic faunas that does not
align with the broader biogeographical pattern of the
Cambay Shale mammalian fauna. Linking Pahelia (and
Quettacyonidae) to African proboscideans and their rela-
tives also contrasts with the broader biogeographical
affinities of the Cambay Shale mammalian fauna, which
lacks taxa with clear African links. The time depth of an
endemic radiation of Indian ungulates is uncertain. A link
between Pahelia, Quettacyonidae and the Late Cretaceous
Indian mammal Kharmerungulatum is not supported but
remains plausible in view of the very fragmentary record
of this genus. Additional Cretaceous–Eocene mammalian
records from the Indian Subcontinent will be required
both to test the hypothesis that Pahelia and Quettacy-
onidae belong to an endemic radiation and to understand
its origins and morphological scope.
CONCLUSIONS
The fossils described above document a distinctive new
herbivorous mammal, Pahelia mysteriosa, from the early
Eocene Cambay Shale Formation. The new taxon combi-
nes incipiently selenolophodont molars with simple,
crowded premolars, an enlarged, proclivous anterior
tooth, and an elongate, fused mandibular symphysis: a
combination of features that sets P. mysteriosa strongly
apart from other taxa known from the Cambay Shale
assemblage, or any other early Eocene mammal. Other
features, including the presence of strong molar exodaen-
odonty and prominent ectostylids further emphasize the
distinctiveness of the new form.
With only the morphology of the mandible and lower
cheek dentition known, there is not enough information
to confidently reconstruct the relationships of Pahelia.
Phylogenetic analysis supports two divergent affinities: a
relationship to Apheliscidae and, more distantly, Louisini-
dae; or a relationship to Quettacyonidae, potentially with
a more distant affinity to Proboscidea. More material,
particularly the upper dentition, is required to discrimi-
nate between these possibilities. Taking biogeographical
considerations into account favours a relationship to
endemic Indo-Pakistani quettacyonids, potentially as part
of a larger endemic radiation of herbivorous mammals in
the Paleocene of the Indian subcontinent.
Acknowledgements. We thank N. Egi, E. Gheerbrant, P.D. Gin-
gerich and R. Tabuce for providing casts of comparative taxa.
We are grateful to staff at the Duke University SMIF facility
for preparing the microCTs of GU/RSR/TAD 9201 and 9203
and to Ulysse Lefevre for scanning GU/RSR/TAD 9608.
Jonathan Perry provided insights into the potential significance
of features of the mandible of GU/RSR/TAD 9608. We thank
Cecilia Cousin, Rachel Dunn, Annelise Folie, Ross MacPhee,
Waqaz Mirsa, Corentin Noiret, Raman Patel, Abhishek Singh,
and Thomas Steeman for field assistance. Nathan Vall
ee-Gill-
ette prepared the Pahelia specimens. This manuscript was
greatly improved through the input of two anonymous
reviewers. The Gujarat Mineral Development Corporation
facilitated excavations at Tadkeshwar mine. TnT is made avail-
able with the sponsorship of the Willi Hennig Society. Field-
work and research were supported by grants from the Leakey
Foundation to KDR and TS and a grant from the National
Science Foundation (NSF DEB 1456826) to LTH. Additional
support from the National Geographic Society, Wadia Institute
of Himalayan Geology, and Belgian Science Policy Office (Bel-
spo BRAIN project BR/121/A3/PalEurAfrica) is gratefully
acknowledged.
DATA ARCHIVING STATEMENT
The data matrices for this study are available in MorphoBank:
http://morphobank.org/permalink/?P2798.
 ZACK ET AL.: NEW EOCENE UNGULATE FROM INDIA
Appendices S1–S3 are available in the Dryad Digital Repository:
https://doi.org/10.5061/dryad.kn4f23q.
Digital models of all specimens are available in MorphoSource:
https://www.morphosource.org/Detail/ProjectDetail/Show/project_id/
425.
This published work and the nomenclatural acts it contains have
been registered with Zoobank: http://zoobank.org/References/
FEC7FC08-8905-4F65-8331-13CD23CE88A2.
Editor. Hannah O’Regan
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