Open Journal of Geology, 2011, 1, 37-44
http://dx.doi.org/10.4236/ojg.2011.13004 Published Online October 2011 (http://www.SciRP.org/journal/ojg)
A New Species of Fossil Mus (Muridae, Mammalia)
from the Late Quaternary Deposits of Narmada
Valley, Central India
Bahadur Singh Kotlia1, M ou li sh re e Jos hi 2, Lalit Mohan Joshi1
1Department of Geology, Kumaun University, Nainital, India
2Faculty of Petroleum and Renewable Energy Engineering, Universiti Teknologi Malaysia (UTM),
Johor Bahru, Malaysia
Received April 18, 2011; revised July 21, 2011; accepted August 29, 2011
Abstract
A new species of fossil Mus (Muridae, Rodentia) is described from the Pleistocene fluviatile deposits of the
Narmada valley (Central India). The species, Mus narmadaensis sp. Nov., has a comparatively smaller lower
molar which is characterized by a narrow molar with well connected cusps, small anterior expansion of lin-
gual anteroconid, protoconid and metaconid, reduced posterior cingulum in addition to hypoconid and ento-
conid nearly at the same level. The large M3 has centrally placed bulbous hypoconid. Among the extant spe-
cies, the present one is closest to M. shortridgei in having similarly placed protoconid and metaconid in M1
and a well developed hypoconid in M3.
Keywords: Fossil Mus, Late Quaternary, Narmada basin, Central India
1. Introduction
Considered to be the most successful groups of living
mammals, the murid rodents were originated in the In-
dian sub- continent about 14 ma ago. At present, they are
found all over the world with ability to adapt themselves
to varied environmental conditions and show marked
species diversity. Today more than 70% of the murid
species are found in the Indo-Australian region, whereas,
26% murid taxa are found in Africa [1]. The oldest
known fossil murid, Antemus chinjiensis was evolved
from a cricetid Potwarmus primitivus and was recovered
from the Chinji Formation (Siwalik sub-group) in the
Potwar Plateau [2]. The study of fossil murids in the In-
dian subcontinent was initiated by [3-6] and followed by
[2] who made significant contribution to the study of
Pakistan Siwalik by describing various murid taxa. Sub-
sequently, a sizeable work on the Afghanistan murids
was done by [7-11]. As far as the Indian murids are con-
cerned, a number of researches, e.g. [12-26] have shown
that the murids were widespread in the country from the
Pliocene onwards.
Among the murids, the genus Mus has been reported
from various parts of India, e.g., from Kurnool caves
[27], Saketi [19], Kashmir basin [22], Narmada valley
[21,25,28], Upper Pleistocene of Bhimtal [23,24], and
Dulam [25]. The great diversity of Mus both in terms of
number and taxa indicates that the probable place of its
origin was the Indian subcontinent. However, an early
stock migration to the African continent during Mio-
Pliocene time has been suggested [29]. Elsewhere in
Asia, Mus has been described from China [30], Crete
[31], former USSR [32], Hungary [33], Japan [34] and
Thailand [35].
We report here the lower molars of a new species of
Mus from the Devakachar section of the Hirdepur For-
mation of the Narmada deposits.
2. Area of Study, Litho-Chronology and
Fossil Material
Narmada, the largest river in the Central India, originates
at the plateau of Amarkantak (22º40’N; 81º40’E) and
after traversing across the middle of the Indian sub conti-
nent, it joins the Gulf of Cambay near Baroda. The cour-
se of the river is controlled by the east-west lineament.
Between Bhedaghat (23º8’N; 79º48’E) and Hoshan-
gabad (22º45’N; 77º45’E), the river forms a trough in

B. S. KOTLIA ET AL. 38
which about 50m thick Quaternary fluviatile deposits are
preserved. Though the deposits are much thicker in the
south, the fossiliferous deposits are exposed in the north-
ern fringe in the sections exposed along river Narmada
and its tributaries.
The Narmada deposits have been divided into seven
lithostratigraphic Formations [36]. The present study
area forms a part of the flood plain facies of the Hirdepur
Formation (Figure 1(a)), comprising greyish homoge-
nous calcareous silt, interlayered with coarse sand, grav el
and conglomerate with high degree of calcification. We
studied a 17m thick profile at Devakachar (23º23’N; 79º
07’E), exposed by the Sher River (see Figures 1(a) and
(b)). It consists of sand, silt and cemented conglomerate
including a fossil bearing horizon. The fossiliferous layer
is 0.5 m in thickness and is composed of medium to
coarse grained brownish coloured sand. It is about 9 m
above the base of the profile (Figure 1(b)).
(a)
(b)
(c)
Figure 1. (a) Geological map of the Narmada Valley showing the study sites; modified after [36]; (b) Lithology of the
Devakachar sections (present work) and Hirdepur Formation (stratotype section of the Hirdepur Formation is taken from
[36]; 1(c) Chronology around Homo erectus locality in the Narmada valley after [26].

B. S. KOTLIA ET AL. 39
The basin is very well known for a large number of
vertebrate fossils including Elephas, Equus, Bos and
several others [37,38]. Lately, a discovery of the skull-
cap of Homo erectus [39] and additional Homo material
[40] has made Narmada valley an important site for pa-
laeontological studies. However, the microvertebrates
have only been mentioned in a handful of reports, e.g.
[21,41,42]. The detailed magnetic stratigraphy of the
Surajkund and Hirdepur Formations [36,43] and absolute
date of the Toba volcanic ash found in the sediments [44]
suggest that the boundary of both the formations lies at
74 ka BP and the top of the Narmada sequence is Holo-
cene [40,42]. Several lithics recovered from the Dhansi
Formation (see Figure 1(c)) may represent the first un-
equivocal evidence for an early Pleistocene hominin pre-
sence in India [45]. The Homo erectus horizon is only
slightly older than the present fossil horizon.
We recovered a large number of microvertebrate re-
mains, such as, murid rodents, lizards and fish from the
Devakachar section. The murids are represented by lower
molars and incisors. The lizards consist of dentaries,
whereas, the cyprinid and channid fishes have teeth and
spines. Her e, we report only the murid material.
3. Systematic Palaeontology
Order: Rodentia
Family: Muridae
Genus: Mus
Mus narmadaensis sp. nov.
Type locality: Devakachar, 120 km southwest of Ja-
balpur (Madhya Pradesh).
Horizon and age: The horizon, a medium to coarse
grained sand, is Middle to Upper Pleistocene in age.
Referred material: Two LM1s (NAR/1, NAR/2), One
LM3 (NAR/3). Broken incisors (NAR/I1-NAR/I6 (Fig-
ures 2(a)-(e ) ).
Etymology: The species has been named after the type
area.
Holotype: LM1 (NAR/1, Figure 2(a)).
Paratype: LM1 (NAR/2, Figure 2(b)).
Measurements: See Table 1 for measurements.
3.1. Differential Diagnosis
Smallest Mus ever reported, M1 with highly reduced
posterior cingulum, M3 with a large second chevron;
differing from Mus auctor [2] in having narrower M1 and
centrally placed hypoconid in M3; from Mus sp. [2] in
having a smaller M1 and from Mus sp. [19] in having a
reduced posterior cingulum in M1; from M. flynni [19] in
having a larger hypoconid in M3; from M. jacobsi [22] in
having poorly developed labial cingulum and lack of
(a) (b) (c)
(e)
(d)
Figure 2. Lower molars of Mus narmadaensis sp. nov. (a)
LM1 (NAR/1); (b) LM1 (NAR/2); c LM3 (NAR/3); (d) and
(e), murid incisors (NAR/11 and NAR/16).
Table 1. Length/width measurements of lower molars (mm)
in Mus narmadaensis sp. nov.
Sp.no. tooth type length Width
NAR/1 LM1 1.27 0.73
NAR/2 LM1 1.26 0.75
NAR/3 LM3 0.76 0.64
accessory cusps in M1; and differing from M. dhailai [23 ]
in having a smaller M1 and much larger hypoconid in
M3.
3.2. Description
M1 is a small and narrow cusp. The asymmetrical ‘X’
pattern is formed by the four anterior cusps. The labial
cusps lie posterior to the lingual cusps in the first chev-
ron. The labial anteroconid is smaller than the anteriorly
displaced lingual anteroconid. The cusps are very strongly
connected and the connection between the labial antero-
conid and protoconid is stronger than between the lingual
anteroconid and metaconid. The hypoconid and entoco-
nid are more or less at the same level, the former being
slightly bigger than the later. The posterior cingulum is
small, oval, transversally flattened and highly reduced.
M1 has two roots.
M3 is roughly triangular in outline. The protoconid and
metaconid are at the same level and are more or less of
the same size in the anterior chevron. The hypoconid and
entoconid are merged together to form a bulbous chevron
which is centrally placed. The specimen has one com-
plete root.

B. S. KOTLIA ET AL. 40
3.3. Comparisons
Mus narmadaensis sp. nov. can be differentiated from M.
auctor [2], Mus sp.[19] and M. jacobsi [22] in having the
following characters; smaller M1, marginal anterior dis-
placement of lingual anteroconid relative to the labial
anteroconid, protoconid and metaconid at the same level,
poorly developed labial cingulum and highly reduced
posterior cingulum in M1. However, M3 of the present
species is bigger than that of M. jacobsi and M. auctor.
In the M3 of the present species, the hypoconid is cen-
trally placed, whereas, it is displaced lingually in M.
auctor and labially in M. jacobsi.
The present M1s differ from M. flynni [19] in having a
much smaller M1 with lingual anteroconid sh owing small
anterior displacement relative to labial anteroconid, hy-
poconid and entoconid occupying the same plane and a
highly reduced posterior cingulum. M3 of M. narmadaen-
sis sp. nov. is larg er than that of M. flynni and also ha s a
larger hypoconid. The Narmada species is close to Mus
sp. [2] in the relative position of cusps in the anterior
chevron and a reduced posterior cingulum in M1 but it
has a much smaller size. Also, the connection of cusps is
much stronger in the Narmada species. The present spe-
cies is similar to M. dhailai [23,24] in the relative posi-
tion of the labial and lingual anteroconid, protoconid, me-
taconid and in having a reduced posterior cingulum in M1
and similarly placed hypoconid in M3 but differs from it
in having a much smaller M1 and a bigger M3 with a bet-
ter developed hypoconid (Tables 2 and 3). A compari-
son of various species of Mus is shown in Figure 3.
3.4. Enamel Ultrastructure in Murid Incisor
The rodent enamel microstructure has the highest degree
of complexity among mammals [46-50]. In most rodents,
the incisor enamel is made up of two layers, an inner
portion known as Portio Interna (PI) with intersecting
prisms which appear as Hunter-Schreger Bands (HSB) in
the longitudinal section, and an outer portion known as
Portio Externa (PE) with radial enamel in which the
prisms are oriented parallel to each other. The presence
of these two layers in the rodent incisor enamel is re-
garded as a characteristic feature which distinguishes it
from lagomorphs where only Portio Interna with HSB is
developed [51]. Biomechanically, the HSBs serve as stre-
ngthening device inhibiting crack propagation [49,52-54],
whereas the radial enamel of the Portio Externa helps to
maintain a sharp cutting edge because of its higher resis-
tance to wear [55-57]. The evolution of enamel of the
rodent incisor is independent from that of the molar
enamel [58].
In rodent incisors, three are three basic types of HSBs,
e.g., pauciserial, multiserial and uniserial [46]. Paucise-
rial HSBs are primitive with highly variable band thick-
ness [59,60]. This condition gave rise to the uniserial and
multiserial HSBs. In the multiserial enamel, the HSBs
are 3-6 prisms wide and are inclined to the Enamel Den-
tine Junction (EDJ) [61]; whereas, in the uniserial HSBs,
the band thickness is reduced to a single prism and the
Interprismatic Matrix (IPM) may be parallel or angular
to the prism direction [50-61]. In the highly derived
uniserial HSBs, the IPM runs rectangular to the prism
direction and serves to strength en the enamel in the third
dimension. We studied the enamel ultrastructure of the
incisor in the longitudinal section.
3.5. Lower Incisor of Mus; NAR/I4 (Figures
4(a)-(b))
The longitudinal section reveals a PI with typical unise-
rial HSBs which are two prisms thick and a PE with the
radial enamel. The HSBs are inclined at an angle of 60º
to the Enamel-Dentine Junction (EDJ) (Figure 4(b)). As
Table 2. Comparative length/width measurements of different species of Mus.
measurements (mm)
(M1) (M3)
Name Reference Locality Age
Length width lengthwidth
Mus auctor [2] Dhok Pathan Fm., Upper Si wa li k 5.7 ma 1.472 0.928 0.6800.800
Mus sp. [19] Tatrot Fm., Upper Siwalik 2.5 ma 1.400 0.940 ---- ----
Mus flynni [19] Tatrot Fm., U p pe r Siwalik 2.5 ma 1.687 1.040 0.6170.653
Mus jacobsi [22] Kashmir basin, NW India 2.4 ma 1.550 0.956 0.5600.520
Mus sp. [2] Dhok Pathan Fm., Pakistan SiwalikEarly Pleistocene1.490 0.90 ---- ----
Mus narmadaensis sp. nov. present work Devakachar, Narmada valley Upper Pleistocene1.270 0.730 0.7600.640
Mus dhailai [23] South-central Kumaun Himalaya Upper Pleistocene1.582 0.968 0.6130.645

B. S. KOTLIA ET AL.41
Table 3. Characters and position of various cusps in different species of Mus
sp. Mus auctor Mus sp. Mus flynni Mus jacobsiMus sp. Mus narmadaensis
sp. nov. Mus dhailai
locality Dhok Pathan Fm.,
Pakistan Tatrot Fm., Upper Siwalik Tatrot Fm., Upper Siwalik Kashmir basin, NW India
Dhok Pathan
Fm.
Pakistan
Devakachar,
Narmada
valley
South-central
Kumaun
Himalaya
age Late Miocene 2.5 ma 2.5 ma 2.4 ma Early Pleistocene Late Pleistocene Late Pleistocene
author [2] [19] [19] [22] [2] present study [24]
M1 lingual
antero-conid
twice the size
of labial
anteroconid
twice the size
of labial
anteroconid
thrice the size
of labial
anteroconid
thrice the size
of labial
anteroconid
twice the size
of labial
anteroconid
slightly bigger
than labial
anteroconid
thrice the size
of labial
anteroconid
labial
antero-conid
smaller and
posteriorly
displaced
very small/
posteriorly
displaced
very small/
posteriorly
displaced
very small/
posteriorly
displaced
almost at same level of
lingual
anteroconid
almost at same level
of lingual
anteroconid
very small and
posteriorly
displaced
protoconid posterior
to metaconid posterior to metaconid almost at the level of metaconidposterior to metaconid posterior to metaconid almost at the level of metaconid posterior to metaconid
metaconid smaller than
protoconid smaller than protoconid almost equal to protoconid almost equal to protoconidsmaller than protoconid almost equal to protoconid smaller than protoconid
hypoconid posterior
to entoconid posterior to entoconid almost at same level posterior to entoconidmore or less at same level more or less at same level more or less at same level
posterior
cingulum medium large large large large Medium small medium
M3
hypoconid large, lingually placed ---- medium, centrally placed large, labially placed ---- very large, centrally placed small, lingually placed
(a) (b) (c) (e)
(d) (f) (g) (h)
Figure 3. Comparative morphology of the lower molars in various species of Mus. (a) Mus auctor [2]; (b) Mus sp. [19]; (c) M.
flynni [19]; (d) M. jacobsi [22]; (e) Mus sp. [2]; (f) M. narmadaensis sp. nov. (present study); (g) M. dhailai [23]; (h) M. shor-
tridgei.
the bands move towards the outer enamel, the angle of
inclination gradually decreases from 60º to 30º and the
prisms become parallel to the EDJ. The prisms of alter-
nating bands intersect at an angle of 90º at the PE-PI
junction and the crystallites of the IP run perpendicular
to the long axis of the prisms. The outer and thick
enamel is made up of horizontal interlocking prisms. The
IP makes an angle of about 90º with the longitudinal
prisms of HSB.
The enamel thickness decreases towards the incisal

B. S. KOTLIA ET AL. 42
ab
Figure 4. Longitudinal views of the enamel ultrastructure in the lower incisor in Mus (Bar represents 1 mm).
direction (Figure 4(a)). Near the tip of the incisor, the
HSBs are more closely spaced just below the outer enamel
and the IPM is dense below the PE. The PI is reduced
towards the incisal end. At the tip of th e incisor, only the
radial enamel of the PE is present (Figure 3(a)). The
crystallites of the IPM are rectangular and serve to
strengthen the enamel in a third dimension. This feature
is generally seen in the derived species of murids.
4. Discussion
The reduced posterior cingulum in M1 in the present
specimen points to its affinity with Pahari section of Mus
[13]. In India, Mus is represented by three subgenera,
Mus with M. booduga and M. dunni; Pyromys with M.
saxicola, M. shortridgei and M. platythrix; and Coelomys
with M. mayori, M. pahari and M. crociduroides [14].
Coelomys section includes Asiatic species such as M.
mayori, M. pahari, M. crociduroides and M. shortridgei
[13]. The Narmada Mus resembles M. pahari in general
outline and the placement of cusps in the anterior chev-
ron of M1. However, the second chevron in M3 of M.
pahari is weakly developed and shows a small lingually
placed hypoconid. M. crociduroides has a weak second
chevron in M3 and is therefore different from the present
species. M. mayori differs from M. narmadaensis sp . nov.
in having a posteriorly displaced protoconid in M1 and a
lingually placed and weakly developed hypoconid in M3.
Among the extant species, M. narmadaensis sp. nov.
shows closest resemblance with M. shortridgei in having
similarly placed protoconid and metaconid in M1 and a
well developed hypoconid in M3.
A very small size of the M. narmadaensis sp. nov.
may be attributed to its getting iso lated from the stock at
the onset of glacial age during the Pleistocene period.
Murids are very sensitive to the climatic changes and it is
believed that the onset of cold climatic conditions wiped
out several species of murids while some migrated to
warmer regions [25]. It may be postulated that M. nar-
madaensis sp. nov. was one of those species that mi-
grated towards Central India from the Lesser Himalayan
region at the onset of glaciation. It may have lived there
in isolation for a considerable period due to which it
could not evolve more progressively although its enamel
shows some derived characters as much as in other
Pleistocene species.
5. Acknowledgements
The study was spon sored by the University Grants Com-
mission and Council of Scientific and Industrial Resear-
ch, New Delhi. The laboratory facilities were provided by
the Department of Geology, Kuma un University , Naini tal.
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