CTENODACTYLOID RODENT ASSEMBLAGE

F R O M KARGIL FORMATION, L A D A K H M O L A S S E G R O U P :
AGE A N D P A L A E O B I O G E O G R A P H I C
IMPLICATIONS FOR THE INDIAN
S U B C O N T I N E N T IN THE O L I G O - M I O C E N E
AVINASH C. N A N D A & ASHOK SAHNI

NANDA A.C. & SAHNI A. 1998. Ctenodactyloid rodent assemblage from Kargil Formation, Ladakh Molasse Group:
age and palaeobiogeographic implications for the Indian subcontinent in the Oligo-Miocene. [Assemblage de ron-
geurs ct~nodactylo~des de la Formation Kargil, Groupe de la Molasse de Ladakh: fige et pal~og~ographie du sous-
continent indien ~ l'Oligo-Mioc~ne]. GEOBIOS, 31, 4: 533-544. Villeurbanne, le 31.08.1998.

Manuscrit d~pos~ le 18.03.1996; accept~ d~finitivement le 05.06.1997.

ABSTRACT - Two ctenodactyloid rodents, Fallomus r a z a e FLYNN,JACOBS& CHEEMAand E ladakhensis nov. sp., are
being described from the Kargil Formation of the Ladakh Molasse Group of Kargil area. Fallomus (?
Chapattimyidae) is considered an endemic offshoot of the South Asian Eocene ctenodactyloid stock. The new species
is relatively larger and more hypsodont with an ectostylid borne on a separate pillar. At Kargil, Iberomeryx (=
Cryptomeryx) savagei and Lophiomeryx kargilensis were earlier reported and indicate an Upper Oligocene age for
the lower part of the Ladakh Molasse Group. Both these taxa show Eurasian affinity. From the upper part of the
Ladakh Molasse Group i.e. Pashkyum Formation, Hyoboops palaeindicus is known and indicates a Lower Miocene
age for the lower part of the Pashkyum Formation. The fossil vertebrates suggest an age for the Ladakh Molasse
Group extending from Upper Oligocene to Middle Miocene. The incisor enamel ultrastructure of Fallomus is multi-
serial with average inclination of the Hunter Schreger Bands varying between 30 °. 35 °.

KEYWORDS: CTENODACTYLOID RODENTS, OLIGO-MIOCENE, ENAMEL STRUCTURE, LADAKH MOLASSE

GROUP, KARGIL FORMATION.

RI~SUMI~ - Deux esp~ces de rongeurs ct~nodactylo~des, Fallomus r a z a e FLYNN,JACOBS& CHEEMAet F. ladakhensis

nov. sp. sont d~crites dans la Formation Kargil du Groupe de la Molasse Ladakh dans la r~gion de Khargil. Fallomus
(? Chapattimyidae) est consid6r~ comme un descendant end~mique des ct~nodactyloides de l'Eoc~ne sud-asiatique.
La nouvelle esp~ce est relativement plus grande et plus hypsodonte avec un ectostylide sur un pilier s~par~. De
Kongil, Iberomeryx (= Cryptomeryx) savagei et Lophiomeryx kargilensis ont d~j~ ~t~ signal,s indiquant un fige
Oligoc~ne sup~rieur pour la partie inf~rieure du Groupe de la Molasse de Ladakh. Tous ces taxons montrent une
affinit~ europ~enne. Hyoboops palaeindicus connue de la partie sup~rieure du groupe i.e. (Formation Pashkyum)
indique un fige Miocene inf~rieur pour la partie inf~rieure de la Formation Pashkyum. Ainsi les vertebras fossiles
sugg~rent un fige de l'Oligoc~ne sup~rieur au Miocene moyen pour le Groupe de la Molasse de Ladakh.
L'ultrastructure de l'~mail des incisives de Fal.lomus est de type multis~ri~ avec une inclinaison moyenne des bandes
d'Hunter Schreger comprise entre 30 ~ 35 °.

MOTS-CLI~S: RONGEURS CTI~NODACTYLOIDES, OLIGO-MIOCI~NE, STRUCTURE DE L'I~MAIL, GROUPE DE LA

MOLASSE DE LADAKH, FORMATION KARGIL.

INTRODUCTION
A d d i t i o n a l finds of r o d e n t s from t h e Oligo-Miocene
s e q u e n c e s of t h e L a d a k h Molasse G r o u p in the
n o r t h w e s t e r n H i m a l a y a h a v e i n t e r e s t i n g implica-
tions for r o d e n t diversification in S o u t h Asia a n d
m i g r a t o r y r o u t e s in t h e context of a r i s i n g m o u n -
r a i n r a n g e s . T h e L a d a k h Molasse G r o u p occurs as
a n a r r o w a r c u a t e belt, all along t h e s o u t h e r n m a r -
gin of t h e L a d a k h P l u t o n i c Complex, a n d its wes-
t e r n e x t r e m i t y t e r m i n a t e s at K a r g i l t o w n s h i p
(Tewari 1964). It is exposed e a s t w a r d s at Leh,
N y o m a and H a n l e and passes into Tibet w h e r e it is
referred as Xigaze Group by Chinese geologists (fide
S r i k a n t i a & R a z d a n 1985). This g r o u p is best expo-
sed along W a k k a River section, Kargil a n d h a s yiel-
ded from its basal p a r t a diverse a n d significant
f a u n a which h a s been described by N a n d a & S a h n i
(1990). I n addition to this a single Hyoboops tooth
was r e p o r t e d from Baroo Colony of Kargil by Dixit
 534
et al. (1971) and was described in detail by Savage
et al. (1977). Location of Kargil area and position of
fossil localities are shown in the geological map
(Fig. 1). Interestingly from the Trans Himalayan
region vertebrate fossils are known from Hundes
and Kailash region of Tibet and were reported in
the last century by Buckland (1823) and Strachey
(1851). These findings were considered as belon-
ging to the Pleistocene by Savage et al. (1977).
The present fossil locality is the same which have
earlier yielded the artiodactyals, Lophiomeryx
and Iberomeryx (= Cryptomeryx savagei), worn
ctenodactyloid maxillary f r a g m e n t and boid
snakes (Nanda & Sahni 1990). Continued field
work in the area suggests that Kargil locality
exposed along Wakka River is stratigraphically
older than the one producing Hyoboops (Dixit et
al. 1971; Savage et al. 1977). This observation
finds some supports from earlier studies by
Brookfield & A n d r e w s - S p e e d (1984). These
authors followed the biostratigraphic scheme of
Bhandari et a1.(1977), but were able to differen-
tiate six lithounits. Their unit 1 corresponds to
the horizon yielding the present forms (i.e. the
Kargil Formation of Bhandari et al. 1977), while
the Hyoboops find is probably equivalent to their
unit 5 (Pashkyum Formation of Bhandari et al.
1977). Field evidences support this as there is no
strike continuity between the two exposures as
Raiverman & Misra (1975, fig. 4) have shown a
fault in the intervening area. Srikantia and
Razdan (1980, p. 528 ) have also considered that
the "outlier" which yielded Hyoboops at Baroo
represents the upper part and not the basal part
of their "Indus Group" (= Ladakh Molasse Group
of Tewari 1964 and of present work). According to
these workers due to time transgressive phase the
u p p e r p a r t overlaps the L a d a k h Plutonic
Complex. At present on account of the lack of the
diagnostic fossils, this observation can not be vali-
dated and in need of future confirmation. The pre-
sent age evidences are based on plants, charo-
phytes, molluscs, and spores and pollens (Sahni &
Bhatnagar 1962; Tewari & Dixit 1971; Tewari &
Sharma 1972; Bhandari et a1.1977; Mathur 1983).
More evidences based on flora and fauna are
required to established age precisely.
The geology of the Ladakh area is beset with seve-
ral problems regarding multiplicity of stratigra-
phic nomenclature and its usages, and it is there-
fore essential to clarify the terminology used in the
present paper. While excellent summaries of the
regional geology have been provided by Shah et al.
(1976), Frank et al. (1977), Thakur (1981), Sahni et
al. (1984), Srikantia & Razdan (1985), one of the
simplest classification for the Ladakh Molasse
Group of the Kargil area has been given by
Bhandari et al. (1977). These workers gave the
details of these various lithounits and described
the palynological fossils from Wakka Chu section.
One of us (ACN) has mapped the area on a scale of
1:50 000 and was able to demarcate the various
formations of Bhandari et al. (1977) in the vicinity
of Kargil (Fig. 1). In a significant contribution,
Brookfield & Andrews - Speed (1984) carried out
sedimentological studies in the area and recogni-
zed six lithological units for the Ladakh Molasse
Group in Kargil area. The first three units show a
transition from arid, ephemeral stream deposits,
through braided and meandering stream sedimen-
tation and mark the filling in a large basin and
suggest climatic variations from semi-arid to tem-
perate, humid conditions. The three other younger
units show a change from deep lake basin turbi-
dites through semi-arid ephemeral stream deposits
to large braided and meandering streams, followed
by a major change to extreme aridity (Brookfield &
Andrews-Speed 1984, p. 175, Table i).
GEOLOGY
The Ladakh Molasse Group is best exposed along
Wakka River section and has been divided into
three formations - Kargil, Tarumsa and Pashkyum
by Bhandari et al. (1977). A brief description of the
various formations is given below and the position
of the various formations and their geographic
extension in Kargil area is shown in Figure 1.
KARGIL FORMATION
Kargil Formation, the oldest formation of the
Ladakh Molasse Group, overlies unconformably
on the granitoids of the Ladakh Plutonic Complex
and is best exposed along both sides of Wakka
River near Akchamal village. The sequence is cha-
racterized by conglomerates, greyish green fine to
gritty sandstones and purplish red mudstones.
The measured thickness of the Kargil Formation
along Wakka River is about 190 m and passes
transitionally into the overlying T a r u m s a
Formation. About 42 m above the contact of the
molasse and granitoid basement, from the same
purple red mudstone (Fig. 2) which yielded the
vertebrate fauna earlier described by N a n d a &
Sahni (1990), an additional collection of ctenodac-
tyloid rodents was made and is being described
here. Earlier workers reported shells of gastro-
pods and lamellibranchs associated with plant
fossils (Sahni & Bhatnagar 1962; M a t h u r 1983)
and palynological fossils (Bhandari et al. 1977)
from this section. However, they did not report
any vertebrate fossils.
TARUMSA FORMATION
The Tarumsa Formation (700 m thick) is best expo-
sed east of the Tarumsa village and is characterized
 535

FIGURE 1 - Geological m a p of
K a r g i l area. I n s e t geological m a p ~ q 1 Quaternary ~ Ladakh Granite N
Deposits Present Fossil
s i m p l i f i e d a f t e r Sch~irer et al.
(1984). Carte gdographique de la ~, I ' ~ " ~ Pashk u j - ' ~ Dras Voloanics X F' L°oalitY
0 O.,. ll_ __1 ymFm. , .,
rdgion de Kargil, simplifide d'apr@s ~" X Baroo Colony + -+
Schdrer et al. (1984). -r 0 4 ~ Tarumsa Fm. ~ Thrust F, Locality , + T/<7tk-

< // I ~
~ I .r,,argl
. . . Pro. F "-. ,,,,, Fault .
+ t ÷. ÷ + + / ~/~-- ~ ~ -,.r
m ~- Y "" La HamDo[ina
-~ ~ : . ~
oh ~in~
_~

+%z-L-%+ +I
+] >;r'".l ++ ++ ::.7

+\ ~ . . . : : .. : : . ~ , - v
+'~ ~ . . : :~.-. :~'v

+ ~ -- . . . . . . \-_-_-_---/v ,.7 v v
-,,-y-r .,/,. ,'Y y ~.,-~ ~,~ .-? < ~..~-- ----_-- / V '4 0 1 2 3 km
- -~.~...,- .,~ ~ ~- ~- ~- -.,- ? \ - - - / V I I I I
+1~ v ~ - " ~ ~..~v-'~---~ ''v
V,/~v --.c.,.~ .~.:; V V
4- - ' ' + ~ V V
150kml +-..T<argil + ?'ra/?s+~-L.++ + N ~,Kargil
0 ~ : + I ~NDI~:~
" 0 ' , 2

INDEX
" -'xZanskar Sedimen [ [ ~
F~J LadakhMolasseGroup OphioIiticMelange
autochthonous
Indus Formation ~ Lamayuruunit
Parautochthonous
rn-4F'BDrasvolcanics 17"-71Sedimentand
NindamFormation IZZI acidvolcanics

by thickly bedded multistoried sandstone bodies lithe-successions, belonging to the Kargil and the
with mudstone interval and minor conglomeratic Pashkyum Formations show close similarities in
horizons. Fragmentary bone pieces are noticed in Wakka River section. Brookfield & Andrews - Speed
the Tarumsa Formation which is otherwise rich in (1984, Table 1) classified these beds to their lithou-
plant and invertebrate fossils mainly molluscs and nit 5 (i.e. Pashkyum Formation of Bhandari et al.
these are described by Sahni & Bhatnagar (1962) 1977) whereas the Kargil Formation belongs to
and Mathur (1983). Bhandari et al. (1977) observed their lithounit 1. On the basis of the sedimentologi-
spores, pollens, algal remains and highly carboni- cal studies, these authors prefer to put these beds
zed cuticles from the lower part of the formation with unit 5 i.e. the Pashkyum Formation (Brook-
and suggested an Eocene - Miocene age. field & Andrews-Speed 1984, p. 188).

PASHKYUM FORMATION COMMENTS ON R O D E N T S F R O M KARGIL

The Pashkyum Formation (850 m thick) is best A small rodent assemblage found in association
exposed in Wakka Chu River section and east ofvil- with Iberomeryx and Lophiorneryx, provides new
lage Pashkyum. In addition one isolated patch at insight into the evolution and affinities of the
Cheskor village and at least four patches, including Kargil vertebrate assemblage. F r a g m e n t a r y ver-
one at the Baroo colony, along Suru river section tebrates were initially recorded from Wakka Chu
are also mapped (Fig. 1). The sandstones and mud- section by Brookfield & Andrews-Speed (1984)
stones are purple in colour. The four isolated from a fluvial sedimentary sequence disconforma-
patches along Suru River section, west of Tarumsa bly overlying the granitic rocks of the Ladakh
- Pashkyum area, show red mudstone intervals and batholith. Screening and sorting of the Kargil,
seem similar to the Kargil Formation. In fact the sediments representing the lower part of the
 536

In a definitive paper, Flynn et al. (1986) described

T
Early Miocene rodents from the Dera Bugti loca-
lily of Baluchistan (Pakistan). They described four
new genera, Baluchimys, Lindsaya, Lophibaiu-
chia and Hodsahibia, in a new subfamily the
K Baluchimyinae of the redefined family Chapattiy-
A F Conglomerate
• midae of the Superfamily Ctenodactyloidea. Of
~ Sandstone
special interest in the present context is their des-
R cription of new genus and species of Fallomus (?
G ~ Mudstone Chapattimyidae) as this taxon has been reported
from two Pakistan localities, Dera Bugti and
I Current bedding Zinda Fir Dome (de Bruijn et al. 1981; Flynn &
L c=~ Vertebratefossil Cheema 1994), and now this taxon is being repor-
ted from Kargil. A map of the various localities of
Fish remains Fallomus is provided in Figure 7.

Invertebratefossil
One of best studies of Early Miocene vertebrate
assemblages from the Indian subcontinent is the
Plant fossil one at Bugti which has recently been restudied by
Raza & Meyer (1984). The Bugti m a m m a l s as now
known comprise of large forms which exhibit a
F mixture of Asian and African elements (Flynn et
al. 1986). The Bugti Bone Bed has been assigned
O to the Bugti Member (about 100 m) of the
R Iberomeryx savagei, Lophiomeryx Chitarwata Formation (Hemphill & Kidwai 1973).
M kargilensis, Faflomus razae, F, These fossiliferous sediments include near shore
ladakensis, boid snakes etc. invertebrates such as Ostrea along with sharks
A and other bony fishes. Raza & Meyer (1984) have
T suggested that the Bugti m a y be correlatable to
the Orleanian Mammal Age of Europe extending
I I
Tarumsa Formation from 21 to 18 Ma and this is based on their com-
O parisons to the Rusinga Island, Kenya faunas.
However, this correlation requires further docu-
N mentation. Fallomus is also reported from coeval
beds about 100 kms to the northeast of Bugti in
the Zinda Fir Dome area (Flynn & Cheema 1994).
It is important in the context of the Kargil Forma-
lion to understand the age, sedimentation and bio-
tas of the Chitarwata Formation exposed east of
Chitarwata. These sediments comprise mainly
FIGURE 2 - Stratigraphic column of the lower part of the argillaceous facies with red, grey and green clays-
L a d a k h Molasse Group, W a k k a River section. Colonne strati- tones with minor amounts of siltstones and sand-
graphique de la partie infdrieure du Groupe de la Molasse de stones. Hemphill & Kidwai (1973) considered the
Ladakh, coupe de la rivi~re Wakka. Chitarwata Formation to be Late Oligocene and
Lower Miocene in age on the basis of palynoas-
L a d a k h Molasse Group (= Indus Group of semblage from the Toi Nala Section. Recently
Srikantia & Razdan 1980), have produced several Downing et al. (1993) have restudied the Chitar-
isolated and j a w f r a g m e n t s of c h a p p a t y m i d wata Formation at Dalana and consider these
rodents assignable to one new species of Fallo- deposits as estuarine marked by intense bioturba-
mus. The significance of rodents in the biochrono- tion, some subaerial exposures leading to pedoge-
logy of the thick succession of Tertiary sequences nesis. Most of the small mammals can be related to
in India and Pakistan has been constantly stres- those described from Bugti. There are minor diffe-
sed by several workers (Sahni & Khare 1973; rences between the Bugti and the Zinda Fir
Jacobs 1978; (de Nruijn et al. 1981; Flynn et al. rodents and these are mostly related to frequency
1986). In the present context of the Kargil rodent of occurrence rather than any major compositional
assemblage, what is important is the temporal differences in the assemblage. Significantly, the
and spatial distribution of Oligo-Miocene cteno- non-baluchimyinae chapattimyid genus, Fallomus,
dactyloid and chapattiymid rodents of Pakistan. is also reported from here.
 537

Further to the northwest at Kohat, de Bruijn et al.
(1981) have documented a rodent similar to Fallo-
m u s from beds lying just above a regional uncon-
formity dated at 18.3 Ma. The Murree rodents
include some typical Lower Siwalik elements such
as Sayimys, cricetids, scurids, a rhizomid and a
possible thryonomyoid. Kargil rodents described
below are housed in the repository of the Museum
of Wadia Institute of Himalayan Geology, Dehra
Dun, India. These rodents are collected from the
same locality from which Nanda & Sahni (1990)
described the vertebrates fossils. However, they
have quoted the geographical position incorrectly.
The locality lies on the main Kargil-Homboting La-
Batalik Road and is just before the Wakka River
Bridge and is 3.5 km. east southeast of the Kargil
township. Its position is shown in the geological
map and stratigraphic column (Figs 1, 2).
SYSTEMATIC D E S C R I P T I O N S
Superfamily CTENODACTYOLOIDEATu]lberg, 1899
Family ?CHAPATTIMYIDAEHussain, de Bruijn &
Leinders, 1978
Genus F a l l o m u s FLYNN, JACOBS & CHEEMA, 1986

~6
8

~t

11 12

FIGURE 3 - Scanning electron photomicrographs o f F a l l o m u s razae FLYNN,JACOBS& CHEEMA(Figs. 1-9) and F a l l o m u s l a d a k h e n s i s

nov. sp. (Figs. 10-12). Bar represents 1 ram. 1-2. WIMF/A 1701, left lower first molar, 1, occlusal view, 2, labial view. 3-4. WIMF/A
1702, right lower second molar, 3, occlusal view, 4, labial view. 5-6. WIMF/A 1703, left lower second molar, 5, occlusal view, 6, labial
view. 7-8. WIMF/A 1704, left lower second molar, 7, occlusal view, 8, labial view. 9. WIMF/A 1705, upper right second molar, occlu-
sal view. 10-12. WIMF/A 1706, holotype, right lower second molar, 10, occlusal view, 11, lingual view, 12, labial view.
Microphotographies M E B de Fallomus r a z a e FLYNN, JACOBS • CHEEMA (Figs. 1-9) et Fallomus ladakhensis nov.sp. (Figs. 10-12).
Echelle: 1 ram. 1-2, premiere molaire infgrieure gauche, 1, rue occlusale, 2, rue labiale. 3-4, deuxi~me molaire infdrieure droite, 3,
rue occlusale, 4, vue labiale. 5-6, deuxi~me molaire infgrieure gauche, 5, rue labiale. 7-8, deuxi~me molaire infdrieure gauche, 7, rue
occlusale, 8, rue labiale. 9, deuxi~me molaire supdrieure droite en rue occlusale. 10-12, holotype, deuxi~rne molaire infdrieure droi-
te, 10, rue occlusale, 11, rue linguale, 12, rue labiale.
 538
Fallomus r a z a e FLYNN, JACOBS • CHEEMA, 1 9 8 6
Fig. 3.1~9
The material referable to this species in the
Kargil collection consists four lower molars, an
upper molar and numerous fragmentary dental
elements. As the occurrence of this taxon requires
detailed documentation, all the available speci-
mens have been described.
WIMF/A 1701 (Fig. 3.1-2) exhibits about the
same degree of wear as specimen GSP 21236 from
Pakistan and illustrated in Fig. 5H (Flynn et al.
1986). The protoconid is a large, vertically projec-
ting, conical cusp which is linked to the antero-
posteriorly compressed metaconid by a high ridge.
The talonid is not appreciably wider than the tri-
gonid and this suggests that the tooth is a LM1.
The anterior cingulum is not as well developed as
in WIMF/A 1703 but it is nevertheless distinct.
The hypoconulid is fairly characteristic: it is ver-
tically oriented, isolated and cut off by a deep val-
ley from the ridge joining the hypoconid to the
entoconid. It is situated medianly and somewhat
lingually. The well developed hypoconid is linked
by a transverse high ridge to the entoconid. The
ectostylid is a small and feeble cusp in this parti-
cular specimen though it is better developed in
the other molars in the collection. This specimen
shows some tendency towards hypsodonty but it
should be borne in mind that the specimen is rela-
tively unworn (Fig. 3.2). The lingual view shows
the slender nature of the c u s p s .
WIMF/A 1702 (Fig. 3.3-4) is a more worn specimen
and is a RM2. The cusps are bunodont and wear has
allowed the joining of the hypoconulid with the
obliquely trending ridge joining the entoconid and
the hypoconid. The protoconid is a massive cusp
and on its posterior slope at the base is situated a
small poorly developed ectostylid (Fig. 3.4).
FIGURE 4 - Scanning electron photomicrograph o f F a l l o m u s ladakhensis nov. sp. Bar represents 1 mm. WIMF/A 1707, paratype,
right lower last molar. 1. labial view. 2. occlusal view. 3. lingual view. Microphotographies M E B de Fallomus ladakhensis nov. sp.
Echelle: 1 ram. Paratype, derni~re molaire infgrieure droite, 1, vue labiale, 2, vue occlusale, 3, rue linguale.
WIMF/A 1703 is probably a LM2 and is fairly hyp-
sodont (Fig. 3.5-6). This tooth is marked high
ridges and relatively little wear on the trigonid.
The anterior cingulum is well developed. Again,
wear has resulted in the linking of the hypoconu-
lid with the ridge joining the hypoconid and the
entoconid. A prominent ectostylid is present at
the base of the valley separating the trigonid and
the talonid.
WIMF/A 1704 (Fig. 3.7-8) is similar in dental
morphology to the other teeth described. It is fair-
ly bunodont, worn down and possesses a promi-
nent ectostylid. The above described lower molars
are indistinguishable from those described by
Flynn et al. (1986) from the Bugti. In some ins-
tances, as mentioned before there seems to be a
tendency towards hypsodonty.
WLMF/A 1705 (Fig. 3.9) represents the only iden-
tifiable upper right molar in the collection. It is
worn posteriorly and better specimens are proba-
bly needed to confirm the association of this upper
molar with the lowers. The ridge joining the para-
cone to the protocone is lingually inclined and
somewhat oriented in the same manner as GSP
21209 (Flynn et al. 1986, fig. 5C). The posteroloph
is highly worn and exhibits a prominent metaco-
ne situated at the posterior corner of the tooth.
Fal!omus ladakhensis nov. sp.
Figs 3.10-12; 4.1-3
t I o l o t y p e - WIMF/A 1706, right lower second molar.
P a r a t y p e - WIMF/A 1707, right lower last molar.
T y p e L o c a l i t y - Purple red mudstone, about 3.5 km east sou-
theast of Kargil, Wakka Chu section,Ladakh
E t y m o l o g y - Named after the region of Ladakh
S p e c i f i c d i a g n o s i s - A relatively larger, more
hypsodont species with a rather high parapet joi-
 539

P r e s e n t Collection
E razae E razae E ladakhensis
F l y n n et al. 1986
Table 5* Table 4, n ° n ° WIMF/A N ° WIMF/A
21257 21258 21259 21260 1701 1702 1703 1704 1706 1707

M1
Length(L) 1.40-1.68 2.21
Width(W) 1.20-1.46 1.74
W/L 0.86-0.87 0.79

M2
Length(L) 1.52-1.66 2.53 2.49 1.90 2.45
Width(W) 1.44-1.62 2.57 2.53 1.86 2.49
W/L 0.95-0.98 1.02 1.02 0.98 1.02

M3 3.24
Length(L) 1.54 1.46 1.70 1.68 2.53
Width(W ) 1.38 1.34 1.32 1.46 0.78
W/L 0.90 0.92 0.78 0.87

*Observed range

TABLE 1 - C o m p a r a t i v e m e a s u r e m e n t s (in m i l l i m e t r e s ) of Lower t e e t h of v a r i o u s species of Fallomua Mensurations comparatives

(en ram) des dents infdrieures de diverses esp~ces de F a l l o m u s .

ning the metaconid and the protoconid. RM3 hyp-
sodont with well developed and distinct lophids.
Ectostylid borne on a separate pillar.
Description - W I M F / 1 7 0 6 (Fig. 3.10-12 ) is a
well preserved tooth in a fairly advanced stage of
wear. In spite of this it appears to be quite hypso-
dont with deeply incised valleys between the talo-
nid and the trigonid as well as between the meta-
conid and the protoconid. The ectostylid is very
prominent and only somewhat lower than the
hypoconid.The cusps also appear to be relatively
more robust than those of F a l l o m u s razae.
W I M F / A 1707 (Fig. 4) is a r a t h e r elongated,
high crested tooth. The anterior cusps are verti-
cally oriented while the posterior ones are incli-
ned anteriorly. The protoconid is a massive cylin-
drical cusp and is joined to the metaconid along
its anterior margin. The hypoconulid is joined to
the posterolophid. Ectostylid is not differentia-
ted. C o m p a r a t i v e m e a s u r e m e n t s of the lower
molars of t h e species of F a l l o m u s are given in
Table 1.
FALLOMUS: INCISOR ENAMEL
ULTRASTRUCTURE
Ctenodactyloid rodents from the Indian subconti-
nent have also been studied from the perspective
of their dental microstructure using polarizing
light microscopy (Sahni 1980) and scanning elec-
tron microscopy (Hussain et al. 1978, Sahni 1984
& 1986; Martin 1992). Martin (1992) in a mono-
graphic t r e a t m e n t of enamel ultrastructure of
selected genera of the major rodent families firm-
ly established the generalized nature of paucise-
rial enamel and the more derived condition (in
rodents) of the multiserial and uniserial enamel.
Earlier, Wahlert (1968) had stated that the pauci-
serial pattern was the most generalized, a view-
point that was strongly supported by Sahni in all
his papers (Sahni 1980, 1984, 1986, 1989). Sahni
(1986) pointed out that the pauciserial enamel
pattern should be considered as a functional
grade in rodent enamel evolution as several
Eocene rodent families possessed this generalized
pattern. Martin (1992) provided concrete evidence
in the orientation patterns of the inter prismatic
and prismatic crystallites by showing that in most
generalized and temporally older rodent enamels,
the crystallite orientation of the inter prismatic
layers was parallel to that of the prismatic layer.
In the Kargil material, there are a few fragmen-
tary incisors but only one that can be attributed
to F a l l o m u s with some degree of confidence. The
presently studied incisor represents a small por-
tion in the upper jaw close to maxillary alveolus.
All the presently studied enamel ultrastructure
relates to this single incisor fragment. This frag-
ment was sectioned transversely, obliquely and
longitudinally and is described herewith (Figs 5,
6). The enamel thickness of F a l l o m u s is variable
and ranges between 200 to 260 ~m which is
appreciably greater than the thickness of chapat-
timyids and bahichimyines so far reviewed by
Martin (1992). However, the variability m a y also
be on account of the basal sections studied. The
pattern is clearly multiserial with an average
inclination of HSB's about 30-35 ° (Martin 1992).
In longitudinal section the limited observations
 540

FIGURE 5 - 1. Transverse section showing portio externa, Hunter Schreger

Bands and enamel dentine junction, vertically arranged mosaics. Bar
represents 10 l~m. 2. Transverse section, bar represents 1000 gin. 3.
Transverse section, bar represents 100 llm. 4. Longitudinal section showing
portio externa, multiserial Hunter Schreger Bands, 4-5 Prism thick, gently
inclined, bar represents 100 ~lm. Coupe transversale de l'dmail montrant la
portion externe, les bandes d'Hunter Schreger en mosafque jusqu~t la ligne
de fonctions entre l'dmail et la denture. Echelle: 10 ~trn. 2, coupe transversa-
le, ~chelle 1000 ttm. 3, coupe transversale, 4chelle 100 #rn. 4, coupe longitu-
dinale montrant la partie externe, les bandes de Hunter Schreger, faiblement
inclin~es, fortunes de 4 it 5 prismes.

i n d i c a t e a h i g h l y developed portio i n t e r n a and a of the total thickness. The m u l t i s e r i a l H S B ' s cross

r e l a t i v e l y t h i n portio e x t e r n a (PE). Towards the
c o n t a c t w i t h t h e EDJ, t h e H S B ' s h a v e a t e n d e n c y
t o w a r d s bifurcation. The p r i s m s are u s u a l l y 4
p r i s m s wide. I n t h e PE zone, t h e p r i s m s are incli-
n e d s h a r p l y in t h e incisal direction. I n t r a n s v e r s e
section, t h e P E a g a i n is seen to be less t h a n 10%
over at an angle. T o w a r d s t h e base of t h e r o d e n t
root contact, the H S B ' s are seen to f l a t t e n out. At
the base of the P I zone, t h e r e is a d i v e r g e n t angle
b e t w e e n the crystallites of t h e p r i s m a t i c l a y e r a n d
those of t h e i n t e r p r i s m a t i c layer. The p r e s e n t des-
cription gives us only a basic idea of t h e incisor
 541

FIGURE 6 - 1. Transverse section showing multiserial H u n t e r Schreger Bands. Mosaic arranged laterally, bar represents 100 ~m.
2. Transverse section u n d e r high magnification showing multiserial H u n t e r Schreger Bands. Mosaic a r r a n g e d vertically, bar
represents 10 ~m. Coupe transversale montrant la structure multisdride des bandes de Hunter Schreger disposdes en mosa~que.
dchelle: 100 #m. 2, ddtail, ~chelle 10 #m.

structure of Fallomus. More material is needed to ly multiserial and not typical for structures t h a t
make a detailed study. However, the present work have been described for older (Eocene and Oligo-
has established t h a t the incisor pattern is typical- cene) rodent enamels.
 542

DISCUSSION sediments to the south suggests also an Eocene

event for the initiation of the Kargil molassic
By coincidence or tectonic design, all the Fallomus deposits. However, apart from the occurrence of
localities known to date from the Indian subconti- Iberomeryx in the basal Kargil Formation and the
nent lie in Oligo-Miocene sediments near the existence of Hyoboops in the overlying P a s h k y u m
Indus River. The northernmost of these is the pre- sediments there is no evidence that is reliable for
sently described Kargil locality which is situated age determination. Keeping in mind the large
on the west northwest-east southeast trending thickness of the P a s h k y u m Formation and occur-
Indus River close to this region. This zone marks rence of Hyoboops in the lower part, the upper
the suturing of the Indian and Asian Plates, and limit of the Ladakh Molasse may extend at least
the Kargil Formation represents one of the initial in the Middle Miocene.The evidence of the rodents
s e d i m e n t a r y cycles to be established as the described here is not inconsistent w i t h an
Ladakh-Kohistan-Gandese Batholith uplifted and Oligocene age. However, in the Bugti and the
provided a provenance for basins to the south. Zinda Pir Dome localities, the basal Chitarawata
Unlike the localities in Pakistan, the Kargil loca- Formation may be somewhat younger than the
lity is represented by completely freshwater sedi- Late Oligocene though at present there is no hard
ments, and in this respect m a y be close to the evidence for this. The rodents themselves, espe-
Kohat locality described by de Bruijn et al. (1981). cially in the base of the Kargil, are somewhat hyp-
The other two southern localities are more estua- sodont which is basically a derived character for
rine and brackish water in character and reflect the ctenodactyloids. Also, the enamel structure is
the gradual withdrawal of the Tethys along the fairly derived for the incisors studied from Kargil
course of the present river (Sahni et al. 1983). and represents a multiserial pattern a change
Distribution of the various localities of Fallomus that usually took place for ctenodactyloids during
is shown in Figure 7. The age of the Kargil locali- the Oligocene. Though at present the rodents pro-
ty so far had been established on the basis of the vide some circumstantial evidence for assigning a
presence of fairly typical European taxa, Ibero- some what younger than Middle Oligocene age for
meryx in association with Lophiomeryx which are the Kargil Formation, a precise age assignment
in fact quite common in the Lower and Middle must await more detailed work. Fallomus repre-
Oligocene rocks. It should be noted that in a pre- sents the youngest and most widespread member
vious paper, Nanda & Sahni (1990) described of the Family Chapattimyidae sensu Flynn et al.
Iberomeryx by its junior synonym Cryptomeryx. (1986) and belongs to the end line of an evolutio-
This age is not inconsistent with the presently nary tree which represents endemic evolution of
known stratigraphic d a t a which places the primitive ctenodactyloid rodents on the Indian
emplacement of the batholith as principally an subcontinent starting with the Birbalomys - Cha-
Eocene event. The fact that the basal conglomera- pattimys assemblage from the S u b a t h u Forma-
tic beds overlying the Ladakh batholith have tion of India and the Kuldana and coeval forma-
clasts of granitoid boulders of large size and also tions of Pakistan. The material at hand presently
contain clasts of deep water late Cretaceous - from Kargil is not good enough to make any gene-
Palaeocene foraminifera limestone and volcanics ral comments on the evolutionary status of the
derived from the trench and foredeep island arc Kargil chapattimyids. There is no doubt of the

:t
• "'i ....J . f ,.. X - Fallornus
P'.. o,.~.,oj°
I "~ "2"~'~%~ "~'" " - "~ Locality
• • I ." ~ _Leh r"
) AFGHANISTAN cJ /",~ K a m i l " ~ " T"
[. (," Islamabad ~. - k.~ 0 500 km
\ : / -'~. 9..'x---.x
,.-,..' f ,-" t
_.
/ ~i ' " / ~~ • ,~ ~"~_
"~-
t Zinda Pir X j-~ ./ ,~:,-,'~..,.
"~'t.-- . . . . . . . . . . . v l~ /" DELHI l ,. "'v'.
"x. ^ / d ,," • [..~ ~v ~ ' p "--.~ ..
' Dera B u g t i ~ / .... Xl/. "~-.~.._.,,
Y / .... •. . . . . . . ,
• ( ,.';, "" .......
\"/~.-.._.r~ _ )
~ C I N D I A
FIGURE 7 - G e o g r a p h i c a l d i s t r i -
b u t i o n of Fallomus. Rdpartition
gdographique de F a l l o m u s .
 543
p r e s e n c e of Fallomus razae a n d this species can
be d i s t i n g u i s h e d on t h e basis of size, h y p s o d o n t y
a n d cusp m o r p h o l o g y from F. ladakhensis nov. sp.
M i g r a t i o n r o u t e s for the Oligo-Miocene r o d e n t s
are h a r d e r to c h a r t as the geology and s t r a t i g r a -
p h y of t h e Oligo-Miocene (i.e. pre Siwalik) interval
needs to be b e t t e r defined. However, at p r e s e n t it
a p p e a r s t h a t Fallomus was an endemic offshoot of
t h e E o c e n e ctenodactyloid stock w h i c h g r a d u a l l y
s p r e a d a l o n g the coastal deposits t h a t were shif-
t i n g d e p o c e n t r a l l y s o u t h w a r d s . The E u r o p e a n affi-
nities h a v e been s t r e n g t h e n e d by additional m a t e -
rial of Iberomeryx while the relationships to Africa
a n d c e n t r a l Asia m u s t await additional data. The
occurrence of E u r o p e a n a n d s u b c o n t i n e n t forms
s u g g e s t s t h a t the K o h i s t a n - L a d a k h - G a n d e s e
M a g m a t i c Arc h a d s u t u r e d earlier (Cretaceous -
Palaeocene) w i t h the I n d i a n P l a t e a n d also h a d a
connection with the E u r o p e a n l a n d m a s s .
T h e r e is little p a l a e o m a g n e t i c d a t a t h a t can be
u s e d to precisely c o n s t r a i n the spatial distribu-
tion of t h e L a d a k h , S o u t h Tibet, K a r a k o r a m a n d
B a l u c h i s t a n b l o c k s in t h e E a r l y T e r t i a r y .
S e d i m e n t a r y a n d biotic d a t a at p r e s e n t provide us
w i t h only w i t h b r o a d outlines of t h e palaeobiogeo-
g r a p h y detailed b y S a h n i & M i t t r a (1980a,b).
A d d i t i o n a l w o r k in t h e L a t e Oligocene - L o w e r
Miocene s e d i m e n t s is of t h e u t m o s t i m p o r t a n c e as
this w a s t h e t i m e for the initiation of t h e foreland
b a s i n in t h e I n d i a n L e s s e r H i m a l a y a s t h a t was to
form a p r e c u r s o r b a s i n to t h e y o u n g e r Siwalik
b a s i n a n d a t i m e for a g r e a t f a u n a l turnover.
Acknowledgements - The authors are thankful to Dr. V.C.
Thakur, Wadia Institute of Himalayan Geologyfor various faci-
lities. The work was carried out under project (SP/G2/P02/85)
sponsored by Department of Science and Technology, Govt. of
India and was awarded to one of us (ACN). Thanks are due to
Dr. K.R. Gupta for scientific discussion and for arranging gene-
rous financial support. A.C. Nanda is also thankful to his
various co-workers, Dr. N.S. Virdi, Dr. Rohtash Kumar, Dr. D.C.
Sati, Dr. R.K. Sehgal, Mr. Govind Mehra and Mr. Alok Tripathi
for the help in the field and for discussion, and to Dr. Kishor
Kumar for the help in the preparation of the plates. Dr. (Mrs.)
Neera Sahni (SEM/EDX Lab., Panjab University) helped in
electron microscope micrography of the rodent incisor material.
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A.C. NANDA
Wadia Institute of Himalayan Geology
Dehra Dun, 248001, India
t . SAHNI
Centre of Advanced Study in Geology
Panjab University
Chandigarh, 160014, India