Indraloris

Indraloris is a fossil primate from the Miocene of India he suggested that Sivanasua himalayensis was proba-
and Pakistan in the family Sivaladapidae. Two species bly the same as Indraloris lulli, but left the anities of
are now recognized: I. himalayensis from Haritalyangar, Sivanasua palaeindica open.[3] Tattersall, who also de-
India (about 9 million years old) and I. kamlialensis from scribed additional material of Indraloris, continued to re-
the Pothohar Plateau, Pakistan (15.2 million years old). gard the animal as a lorisid.[4]
Other material from the Potwar Plateau (16.8 and 15.2 Lewis had suggested that Indraloris might derive from
million years old) may represent an additional, unnamed
the Adapidae, a primitive group of primates,[5] and in
species. Body mass estimates range from about 2 kg (4.4 the 1970s some authors provisionally placed Indraloris
lb) for the smaller I. kamlialensis to over 4 kg (8.8 lb) for among the Adapidae.[6] In 1979, American and Indian
the larger I. himalayensis. paleontologists Philip Gingerich and Ashok Sahni re-
Indraloris is known from isolated teeth and fragmentary viewed Indraloris and the Indo-Pakistani "Sivanasua"
lower jaws. The jaw is deep under the last premolars, species. They recognized Sivanasua himalayensis and In-
but becomes shallower towards the front. The lower pre- draloris lulli as representing the same species, Indraloris
molars are elongate. The lower molars are shorter and himalayensis, and created the new genus Sivaladapis
broader than those of Sivaladapis. Indraloris may have for Sivanasua palaeindica and another species that had
been arboreal and at least partly frugivorous. When the been named later, Sivanasua nagrii.[1] Gingerich and
rst Indraloris fossils were discovered in the early 1930s, Sahni considered both Indraloris and Sivaladapis to be
one was misidentied as a carnivoran and the other as adapids.[7]
a loris. The carnivoran identication was corrected in Several other authors suggested similar taxonomic rear-
1968, and in 1979 Indraloris and the related Sivaladapis rangements around the same time. In 1979, Herbert
were identied as late survivors of Adapiformes, an ar- Thomas and Surinder Verma agreed that Indraloris and
chaic primate group.
Sivaladapis were adapids, but placed them in a subfamily
of their own, Sivaladapinae. Also in 1979, Frederick Sza-
lay and Eric Delson placed Indraloris in its own tribe, In-
1 Taxonomy dralorisini, within Adapidae.[8] In 1980, Indian paleon-
tologists S.R.K. Chopra and R.N. Vasishat placed both
Currently, Indraloris is considered to be a valid genus of Pilgrims Sivanasua species in Indraloris and argued
within the family Sivaladapidae, containing two named that Indraloris lulli, Sivanasua himalayensis and Sivana-
species: I. himalayensis from India and I. kamlialensis sua nagrii all represented the same speciesIndraloris
from Pakistan. A third species may be represented in the himalayensis. They listed Sivanasua palaeindica as a sec-
Pakistani material of Indraloris. However, Indraloris has ond Indraloris species, I. palaeindica, [9]
and continued to
had a complicated taxonomic history, and some of the regard Indraloris as a lorisid. Gingerich and Sahni pub-
known material was misidentied as members of other lished in more detail on Sivaladapis in 1984. They then
mammalian groups for decades. placed the two genera in a separate subfamily of Adap-
idae, called Sivaladapinae because that name was pub-
In 1932, British paleontologist Guy Pilgrim described lished two months before Indralorisini.[10] In 1985, Va-
two species from the Miocene of what is now India and sishat continued to classify Indraloris and Sivaladapis in a
Pakistan, Sivanasua palaeindica from Chinji (Pakistan) single genus, and Indraloris himalayensis and Sivaladapis
and Sivanasua himalayensis from Haritalyangar (India). nagrii in a single species, but other authors have not fol-
He attributed both to Sivanasua, a carnivoran genus oth- lowed this classication.[11]
erwise known from Europe.[1] The next year, American
scientist G. Edward Lewis described the new genus and In a 1998 review, primatologist Marc Godinot rec-
species Indraloris lulli from Haritalyangar, which he pro- ognized Sivaladapidae
[12]
as a separate family within the
visionally allocated to the family Lorisidae. The generic Adapiformes, and this classication has been followed
name, Indraloris, combines the name of the god Indra since then. Several genera in addition to Indraloris and
with the generic name Loris, and the specic name, lulli, Sivaladapis are now allocated to Sivaladapidae, which is
honors Richard Swann Lull, at the time director of the known from the Eocene through the Miocene of China,
[13]
[2]
Peabody Museum of Natural History. It was not until Thailand, Myanmar, India, and Pakistan. Sivaladapids
1968 that American anthropologist Ian Tattersall noted are notable for including by far the youngest adapiforms;
that Pilgrims Sivanasua species had been misidentied; members of this group are otherwise known mostly from

1
2 2 DESCRIPTION

the Eocene, but several sivaladapids occurred during the the front and back. A heel is present at the back, part
Miocene.[14] of a small talonid. The tooth has two roots.[17] The p4,
Despite these taxonomic changes, Indraloris remained represented by YGSP 24338, is an elongate, two-rooted
known from only two specimens (the holotypes of In- tooth with a distinct trigonid at the front and talonid at
draloris lulli and Sivanasua palaeindica) until 2005. the back. The protoconid is the highest cusp of the trigo-
Both of those specimensan isolated rst lower molar nid. Two crests descend from it at right angles in a lingual
(m1) and a mandible (lower jaw) fragment with m1, direction (towards the inner side of the tooth): the pro-
respectivelycome from Haritalyangar in the Nagri For- tolophid towards the front, ending at the low paraconid,
and the metalophid towards the back, reaching the elon-
mation.[15] In 2005, however, American paleontologists
Lawrence Flynn and Michle Morgan described ve teeth gate metaconid. The talonid basin is open lingually; on
the labial side, the hypoconid cusp is present. A crest,
of Indraloris from fossil sites in the older Kamlial For-
mation as a second species in the genus, Indraloris kam- the cristid obliqua, reaches from the hypoconid forward
towards the trigonid. No other cusps are visible in the
lialensis. The species was named after the Kamlial
Formation. In addition, they suggested that two lower talonid, but the specimen is worn and poorly preserved;
[16]
the posterolophid, a crest descending from the hypoconid,
jaw fragments from the Kamlial Formation represented a
third, larger species of Indraloris. [17] may end in a small hypoconulid. A weak cingulum is
present on the labial side of the tooth between the proto-
conid and hypoconid.[26] Another tooth, YGSP 32151, is
interpreted as a dp4. It has a more closed trigonid (with
2 Description the protolophid and metalophid making a more acute an-
gle), the protolophid is shorter, and the paraconid is in-
distinct. In the talonid, the hypoconulid and entoconid
Indraloris is known only from isolated teeth and frag- are distinct. The labial cingulum is strong.[17]
ments of the mandible. These show that Indraloris was a
medium-sized sivaladapid, somewhat smaller than Sival- The lower molar of Indraloris is known from four speci-
adapis.[22] In 1982, Gingerich and colleagues estimated mens. GSI D237, [10] an m1 in a piece of jaw, is the holotype
that Indraloris himalayensis may have weighed 3.7 to 4.3 of I. himalayensis. YPM 13802, the holotype of I. lulli
kg (8.2 to 9.5 lb) on the basis of allometric scaling of (= I. himalayensis) was originally identied as an m1, but
tooth size; [23]
Flynn and Morgan estimated a body size Flynn and Morgan suggested in 2005 that it may be an m2
[27]
of about 2 kg (4.4 lb) for I. kamlialensis. [24]
In general, instead. YGSP 44443, the holotype of I. kamlialensis,
the cingula (shelves) on the margins of the cheekteeth are is either m1 or m2, but more likely the former.[28] Part of
[29]
weak in Indraloris.[16] Among the two named species, I. the trigonid is broken o. YGSP 32152, a very worn
kamlialensis is about 20% smaller than I. himalayensis. [17] m1 in a piece of jaw, represents the unnamed large In-
[17]
The unnamed large Indraloris is similar in size to I. hi- draloris. Vasishat suggested in 1985 that these teeth
malayensis. [25] were instead p4s corresponding to molars referable to
Sivaladapis, but this hypothesis has been disproven by the
The mandible is best represented by YGSP 32727, one of discovery of p4s referable to Indraloris.[17]
two specimens of the unnamed large species of Indraloris.
It preserves both the right and left sides of the dentary, Indraloris molars [16]
are short and organized in two main
back to the level of the fourth lower premolars (p4), but is lophs (lobes). They dier from Sivaladapis teeth in
also damaged at the front. The jaw is deep below p4, but being shorter and broader, with a shorter talonid and a
[30]
rapidly becomes shallower further to the front. The roots smaller hypoconulid. In Indraloris himalayensis lower
of two lower incisors and a much larger canine are pre- molars, there are four main cusps (protoconid and meta-
served; the three roots cluster together, with the canine conid in the trigonid, hypoconid and entoconid in the
root above the incisor roots, suggesting that these teeth talonid), which give the crown a rectangular aspect, al-
shared some function. The mental foramen, an opening though the labial cusps (protoconid and hypoconid) are
in the jawbone, is below p4. A root for the deciduous placed somewhat anterior to their lingual counterparts.
second premolar (dp2) is preserved on both the left and In I. kamlialensis, the entoconid is distinct from the
right sides, but the tooth itself is not and it is not possi- hypoconulid, which is large, but the tooth is otherwise
ble to determine whether dp2 had one or two roots. The similar. The cusps are high relative to those of extant
right permanent second premolar (p2) is unerupted, but lorises and approximately equal in height. The cristid
partially visible; it is a blade-shaped cutting tooth. The p3 obliqua, a crest, descends from the hypoconid to a point
bears a single cusp, somewhat anterior to the middle of on the lingual side of the protoconid. On the hypoconid,
the tooth, with crests descending from it towards the front this crest forms a right angle with the posterolophid,
and back, and weak cingula on the inner and outer sides. which runs towards the hypoconulid in the back lingual
It is supported by two roots, which are close together.[17] corner of the tooth. Between the metaconid and ento-
conid, the talonid basin is open. In I. himalayensis at
Isolated lower premolars are known from I. kamlialen- least (the structure is damaged in the only known lower
sis. A p3, YGSP 33157, resembles that of YGSP 32727 molar of I. kamlialensis) there is a well-developed hollow
in possessing a single large cusp connected to crests at
 3

in the trigonid in front of the protoconid and metaconid. [6] Gingerich 1976, p. 95; Gingerich & Sahni 1979, p. 415.
There is a labial cingulum between the protoconid and
hypoconid.[31] YGSP 32152 is so worn that little of its [7] Gingerich & Sahni 1979, p. 416.
structure remains visible. It shows a short trigonid and
a distinct entoconid. A small hypoconulid, close to the [8] Szalay & Delson 1979, p. vii.
entoconid, is suggested by an enamel swelling.[17] This [9] Chopra & Vasishat 1980, p. 132.
specimen is fragmentary enough that it could also repre-
sent a catarrhine primate or a carnivoran.[25] [10] Gingerich & Sahni 1984, table I.
The only known upper tooth of Indraloris is an M3,
[11] Flynn & Morgan 2005, pp. 99100.
YGSP 46009. It is broken at the back labial corner.
The main cusp is protocone; among the other two cusps, [12] Godinot 1998, pp. 241242.
the paracone is higher but the metacone larger. There is
a spur at the back of the protocone, suggesting a rudi- [13] Beard et al. 2007, p. 68.
mentary hypocone. The protocone is connected to the
paracone by a protoloph, which lacks a small cusp (the [14] Beard et al. 2007, p. 68; Godinot 1998, p. 241, g. 10.
paraconule). No crest connects the protocone to the
metacone, but there is a cingulum at the back margin of [15] Gingerich & Sahni 1984, table I; Flynn & Morgan 2005,
the tooth. The tooth bears a strong parastyle (accessory p. 99.
cusp at the front labial corner) and has three roots.[17]
[16] Flynn & Morgan 2005, p. 111.

[17] Flynn & Morgan 2005, p. 114.

3 Distribution and ecology
[18] Gingerich & Sahni 1984, table I; Chopra & Vasishat 1980,
table 2.
Fossils of Indraloris have been found only in the Miocene
Siwalik fossil beds of India and Pakistan. I. himalayen- [19] Lewis 1933, pp. 135, 137; Flynn & Morgan 2005, p. 114.
sis is known only from Haritalyangar,[10] a Late Miocene
site in the Indian state of Himachal Pradesh. This site has [20] Flynn & Morgan 2005, p. 111, table 6.1.
been dated to about 9 million years ago.[32] This site has
also yielded Sivaladapis nagrii.[10] Indraloris kamlialen- [21] Flynn & Morgan 2005, p. 114, table 6.1.
sis is known from two sites in the province of Punjab,
[22] Flynn & Morgan 2005, p. 111; Gingerich & Sahni 1979,
Pakistan, that are both dated to 15.2 million years ago:
p. 415.
Y642 and Y682. Sivaladapis palaendicus has also been
recorded at both sites, and two lorisids are known from [23] Gingerich, Smith & Rosenberg 1982, p. 81, table 5.
Y682. The unnamed large Indraloris is known from
Y642 and an older site, Y801 (16.8 million years old).[33] [24] Flynn & Morgan 2005, p. 116.
All are in the Potwar Plateau region.[34]
[25] Flynn & Morgan 2005, p. 115.
Little is known about sivaladapid ecology. Gingerich and
Sahni suggested that Indraloris was probably arboreal and [26] Flynn & Morgan 2005, pp. 112, 114.
that it may have been more frugivorous (eating fruit) than
Sivaladapis, which they interpreted as a folivore (leaf- [27] Flynn & Morgan 2005, p. 114; Gingerich & Sahni 1984,
eater).[35] Flynn and Morgan interpreted I. kamlialensis table I.
as a mixed feeder.[24] The Late Miocene extinction of
Indian sivaladapids may be related to a decline in forest [28] Flynn & Morgan 2005, pp. 111, 114.
cover in Asia and to competition by immigrating colobine
[29] Flynn & Morgan 2005, p. 112.
monkeys.[35]
[30] Gingerich & Sahni 1984, p. 76; Gingerich & Sahni 1979,
p. 416.
4 References
[31] Lewis 1933, pp. 135136; Flynn & Morgan 2005, pp.
111-112.
[1] Gingerich & Sahni 1979, p. 415.
[32] Pillans et al. 2005.
[2] Lewis 1933, p. 135.

[3] Tattersall 1968, p. 4. [33] Flynn & Morgan 2005, table 6.1.

[4] Tattersall 1968, p. 9. [34] Flynn & Morgan 2005, p. 100.

[5] Lewis 1933, p. 138. [35] Gingerich & Sahni 1984, p. 77.
4 4 REFERENCES

4.1 Literature cited Haritalyangar magnetostratigraphy, Indian Siwaliks:

Beard, K. C.; Marivaux, L.; Tun, S. T.; Soe, A.
N.; Chaimanee, Y.; Htoon, W.; Marandat, B.;
Aung, H. H.; Jaeger, J. -J. (2007). New sival-
adapid primates from the Eocene Pondaung Forma-
tion of Myanmar and the anthropoid status of Am-
phipithecidae. Bulletin of the Carnegie Museum
of Natural History. 39: 67. doi:10.2992/0145-
9058(2007)39[67:NSPFTE]2.0.CO;2.
implications for the age of the Miocene hominids
Indopithecus and Sivapithecus, with a note on a new
hominid tooth. Journal of Human Evolution. 48
(5): 507515. doi:10.1016/j.jhevol.2004.12.003.
PMID 15857653.
Szalay, F.S.; Delson, E. (1979). Evolutionary His-
tory of the Primates. New York: Academic Press.
ISBN 0-12-680150-9.

Chopra, S.; Vasishat, R. (1980). A new Mio- Tattersall, I. (1968). A mandible of Indraloris
Pliocene Indraloris (primate) material with com- (Primates, Lorisidae) form the Miocene of India
ments on the taxonomic status of Sivanasua (Car- (PDF). Postilla. 123: 110.
nivore) from the Sivaliks of the Indian subconti-
nent. Journal of Human Evolution. 9 (2): 129
132. doi:10.1016/0047-2484(80)90069-X.
Flynn, L.J.; Morgan, M.E. (2005). New lower pri-
mates from the Miocene Siwaliks of Pakistan. In
Leiberman, D.E.; Smith, R.J.; Kelley, J. Interpret-
ing the past: Essays on human, primate, and mam-
mal evolution in honor of David Pilbeam. Boston:
Brill Academic Publishers. pp. 81101. ISBN 978-
0391042476.
Gingerich, P.D. (1976). Cranial anatomy and evo-
lution of Early Tertiary Pleasiadapidae (Mammalia,
Primates)". Papers on Paleontology, Museum of
Paleontology, University of Michigan. 15: 1141.
hdl:2027.42/48615.
Gingerich, P.D.; Sahni, A. (1979). "Indraloris
and Sivaladapis: Miocene adapid primates from
the Siwaliks of India and Pakistan. Nature. 279
(5712): 415416. doi:10.1038/279415a0. PMID
16068172.
Gingerich, P.D.; Sahni, A. (1984). Dentition of
Sivaladapis nagrii (Adapidae) from the late Miocene
of India. International Journal of Primatology. 5
(1): 6379. doi:10.1007/BF02735148.
Gingerich, P.D.; Smith, B.H.; Rosenberg, K.
(1982). Allometric scaling in the dentition
of primates and prediction of body weight
from tooth size in fossils. American Journal
of Physical Anthropology. 58 (1): 81100.
doi:10.1002/ajpa.1330580110. PMID 7124918.
Godinot, M. (1998). A summary of adapiform sys-
tematics and phylogeny. Folia Primatologica. 69
(suppl. 1): 218249. doi:10.1159/000052715.
Lewis, G.E. (1933). Preliminary notice of a new
genus of lemuroid from the Siwaliks. Ameri-
can Journal of Science. 5. 26 (152): 134138.
doi:10.2475/ajs.s5-26.152.134.
Pillans, B.; Williams, M.; Cameron, D.; Patnaik,
R.; Hogarth, J.; Sahni, A.; Sharma, J.; Williams,
F.; Bernor, R. (2005). Revised correlation of the
 5

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