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Kumar, Comments on Bajpai
et al
. 2005 www.PalArch.nl, vertebrate palaeontology, 1, 2, (2006)
Comments on ‘Early Eocene land mammals from Vastan Lignite Mine, District Surat (Gujarat), western
India’ by Bajpai, S. et al. published in Journal of the Palaeontological Society of India 50, 1: 101-113, 2005
K. Kumar
Wadia Institute of Himalayan Geology,
33, General Mahadeo Singh Road,
Dehradun 248 001
India
kumark@wihg.res.in
ISSN 1567-2158
Abstract
Bajpai et al.’s recent paper (2005a) describing an important new Early Eocene mammal fauna from the Cambay
Shale of Vastan Lignite Mine, Gujarat, India has a number of errors related to identification, naming, definition,
characterisation and description of new taxa, and measurements of dentitions etc. that need to be recorded and
addressed. This contribution discusses and clarifies some of the errors and will be useful for understanding the
real impact of the Vastan fauna in relation to the India-Asia collision, the mammalian palaeobiogeography and
origin of modern placental mammals.
Contents
1. Introduction
2. General comments
3. Specific comments
4. Concluding remarks
5. Cited literature
Abbreviations
H-GSP Howard University-Geological Survey of Pakistan; IITR/SB/VLM, Indian Institute of Technology,
Roorkee/ Sunil Bajpai/ Vastan Lignite Mine
WIF/A Wadia Institute of Himalayan Geology, Dehradun
PalArch Foundation
Kumar, Comments on Bajpai
et al
. 2005 www.PalArch.nl, vertebrate palaeontology, 1, 2, (2006)
1. Introduction
A rich collection of mandibles, jaws and isolated dental remains of a very important diverse new Early
Eocene terrestrial mammal fauna has recently been described from the Cambay Shale of Vastan Lignite Mine in
Gujarat, India (Bajpai et al., 2005a). The material is remarkably diverse and well preserved. The discovery of
pre-middle Eocene land mammals from India was expected for some time particularly after reports of Early
Eocene mammals from the lignite beds of the Ghazij Formation in Quetta, Kach-Harnai and Gandhera areas of
Baluchistan, Pakistan (Gingerich et al., 1997, 1998, 2001; Ginsburg et al., 1999) and more recent reports of
lower vertebrates and associated mammals including some bats (Bajpai & Kapur, 2004; Rana et al., 2004; Sahni
et al., 2004) from the same lignite mine of Vastan. I have seen equally diverse, well preserved and very similar
material recovered from the same Nummulites burdigalensis Zone of the Cambay Shale of Vastan Lignite Mine
in collection of Dr R.S. Rana of HNB Garhwal University, Srinagar (India) and his collaborators. Papers on
some of this material are already published (Rana et al., 2005) and others are in press. The overall diversity (also
including rodents, bats, ?primates and marsupials mentioned in Rana et al. (2005) and Bajpai et al. (2005a, b) and
richness of the Early Eocene mammalian fauna from Vastan is truly amazing and has surpassed our expectations.
The lower vertebrates are also quite diverse (Bajpai & Kapur, 2004; Rana et al., 2004). Most recently an
exceptionally diverse and well preserved amber-embedded biota has also been reported from the Vastan Mine
(Alimohammadian et al., 2005). I am sure that the Vastan land mammal assemblage will change our perspective
and thinking on the Eocene land mammals of the Indian subcontinent in the context of the India-Asia collision
and the mammalian palaeobiogeography. However, for now the new assemblage seems to have raised some
more interesting questions like endemic versus migrant component, ‘out of India’ versus ‘into India’ migration
etc. than answering the prevailing ones, e.g. timing of first dispersal, bilateral migration and India-Asia collision.
I would like to congratulate Bajpai and his team for discovery of such a marvellous collection of
important fossil material and for scientific content of their paper. However, the overall palaeontological
treatment meted out to such significant fossil material is disappointing. There are quite a few lacunae in the
section dealing with the systematic palaeontology. The quality of this otherwise very important paper has been
compromised particularly in terms of clarity in definition and characterisation of new taxa, accuracy of
descriptions, objective comparisons with closely similar taxa, nomenclature and to some extent in identification,
measurements and illustrations (?coloured photos in plates I - II have been of no help) of fossils. Although I have
not yet seen the material or its casts I feel confident enough to write this commentary and clarify some of the
errors that needed to be addressed for better understanding of the Early Eocene mammalian paleobiogeography.
2. General comments
The fossil material treated in Bajpai et al. (2005a) has been described as representing 7 genera and 10
species, all new with 2 new families (Vastanidae and Cambaytheriidae). The diagnoses of several new taxa lack
clarity and no differential diagnoses have been given for any of the new families or genera. Even characters
mentioned under ‘Diagnosis’ are at times inadequate and inconsistent with description (e.g., Vastania) and
poorly defined in a few cases (e.g., Suratilestes), and comparisons with the allied taxa are at best minimal. The
systematic placement of one of the families (Cambaytheriidae) and of a few genera (e.g., Frugivastodon,
Cambaytherium and Cambaya) is debatable. Besides this a few taxa (e.g., Cambaytherium sp. ‘A’ and C. minor)
are founded on inadequate material.
Curiously, the illustrations of representative specimens of fossil taxa in the three plates incorporated in
the article are not arranged in the order in which they have been described in the text. For example the first plate
comprises photos of Cambaytherium even though this taxon is described in the end. Likewise Anthraryctes,
described in the beginning has its photos in the last plate. In case of Cambaytherium bidens both upper and lower
dentitions represented by dentaries and maxillary/premaxillary fragments have been listed under the ‘Referred
material’ yet the authors have illustrated only the lower dentaries designating them as holotype. Upper dentition
has not been included in the type material despite the fact that one of the named diagnostic familial/generic
characters is based on the upper teeth. Synonymy has not been given for the new genus Gujaratia even though it
has been proposed to include all material previously referred to Diacodexis pakistanensis. No measurements are
given for the new taxon Gujaratia indica even though it has been differentiated from previously known
dichobunids based only on size. All this is unconventional and somewhat confusing.
In naming new taxa, authors have derived names of some genera/species from incomplete name of a
character, location or person. For example Frugivastodon named after Vastan where it was found should have
been named as ‘Frugivastanodon’ and the species Cambaytherium thewissi named after Dr J.G.M. Thewissen
should have been named C. ‘thewisseni’. Similarly, C. bidens so named because it has two incisors in its lower
jaw should have been named C. ‘bidenta’, C. ‘bidentus’ or C. ‘bidentatus’. Also it would have been very
appropriate and logical to name C. thewissi with three incisors in its lower jaw as C. ‘tridenta’, C. ‘tridentus’ or
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C. ‘tridentatus’. As per the systematic lexicon, the correct name for Vastania sahnia named after Dr A. Sahni
would have been V. ‘sahnii’. The suffix ‘therium’ used in naming Cambaytherium is not common only for the
perissodactyls as stated by the authors. It is a Greek word meaning animal, wild animal or wild beast and is
frequently used in naming all kinds of mammals including marsupials.
In systematic palaeontology, description of the designated ‘Type species’ of a genus should precede that
of its other species if any. Accordingly, C. thewissi should have had been described in the text first followed by
C. bidens and or others. Authors have used ‘Mx’/ ‘mx’ at several places under ‘Referred material’ (e.g. on pages
109-110) and elsewhere (e.g. in table 1). I could follow that it means a molar of an uncertain position in the jaw,
but more often than not it is easy to tell the last molar from the anterior ones and therefore, instead of Mx/mx
why not write ‘anterior molar’ or better still make it explicit and write ‘m1 or m2’.
In table 2, dimensions of P4/ of Cambaytherium bidens have been either mixed up – numbers shown
against width probably represent length and vice versa or the tooth has been wrongly identified, and it could
actually be a P3/. Right M/1 of C. bidens holotype has been measured as 120 percent as large as left M/1 of the
same individual – somewhat unusual. In a few cases measurements of teeth given in their description disagree
with illustrations. For example in Suratilestes gingerichi relative width of M/1-M/3 (0.60, 0.86 and 0.93 mm)
given in its description do not seem to match with its figure (3) in plate III.
3. Specific comments
Specific comments are given below genus wise. Familial and ordinal affinities of genera shown in
parenthesis are sensu Bajpai et al. (2005a). References to plates and figures also pertain to plates and figures in
Bajpai et al. (2005a).
Suratilestes (Cimolestidae, Proteutheria)
(Bajpai et al., 2005a: plate III, figures 1-3)
The genus is poorly defined; its diagnosis reads “Cimolestid with relatively long P/4 and anteriorly placed
paraconid on M/1”. The reader is left to guess (i) whether P/4 is longer in respect to adjacent teeth of Suratilestes
or it is longer than P/4 of all other cimolestids or their molars, and (ii) actually how much longer it is. What is so
special if the paraconid is placed anteriorly on M/1? A paraconid will always be anteriorly placed regardless of
tooth position in a jaw, and it can well be seen anteriorly placed in all the three molars of Suratilestes gingerichi
too (plate III, figure 3). The fact is that in M/1 of S. gingerichi it is particularly more anteriorly placed than the
posterior molars and lies almost in the middle in front of protoconid and metaconid (plate III, figure 3).
Frugivastodon (Apatemyidae, Apatotheria)
(Bajpai et al., 2005a: plate III, figures 7-9)
The basis of identification of the lone lower (?anterior) molar as an apatemyid is unclear and apparently
ill-founded; there is no paraconid, which is present in all known apatemyids. This tooth has a strong resemblance
with M/2 of an adapoid primate and lacks typical apatemyid characters. I think it may as well belong to an
?adapoid primate and therefore its comparison with Primates is absolutely necessary. In the past, apatemyids
have been classified with primitive primates as well as insectivorans though presently they are included in their
own order Apatotheria. There is no record of apatemyids outside Europe and North America (McKenna & Bell,
1997; Von Koenigswald et al., 2005). The available evidence is clearly inadequate to announce the presence of
Eocene apatemyids in India. Additional material will be necessary for its accurate taxonomic placement. “P.”
cristatus on page 104 is probably a typo for F. cristatus.
Vastania (Vastanidae, Insectivora)
(Bajpai et al., 2005a: plate III, figures 4-6)
The familial diagnosis for Vastanidae based solely on P/4 morphology is inadequate and inconsistent with
details given under ‘Remarks’. It reads ‘Erinaceomorphs with high protoconid on P/4 and straight and lingual
cristid obliqua, lacking talonid basin’. Vastania sahnia is the only species under the family and authors’ remarks
on it read ‘[…] simple P/4 dominated by a single cusp, and with a talonid with a straight and labial cristid
obliqua’. The genus has been diagnosed by similar-sized M/1 and M/2 with low cuspate paraconid. However, its
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. 2005 www.PalArch.nl, vertebrate palaeontology, 1, 2, (2006)
description giving size of lower molars as 1.11, 1.27 and 1.33 mm long and 0.66, 0.88 and 0.77 mm wide
respectively contradicts this.
Cambaya (Nyctitheriidae, Insectivora)
(Bajpai et al., 2005a: plate III, figures 10-12)
This genus is based on and is known only by an isolated P/4, which shows typical chiropteran characters
like a complete labial cingulum, a small metaconid and a very reduced talonid. I have seen Rana’s collection of
chiropteran dentitions from Vastan some of which is published (Rana et al., 2005) in the same issue of the
journal as this paper of Bajpai et al. (2005a) and I think that the material referred to a new nyctithere Cambaya
by Bajpai et al. (ibidem) may actually belong to a bat. Bajpai et al. (ibidem) have not compared their material
with chiropterans although they have mentioned elsewhere in the text that their collection does contain bats.
Gujaratia (Dichobunidae, Artiodactyla)
(Bajpai et al., 2005a: plate II, figures 4-6)
Authors have excluded Diacodexis pakistanensis known from the Middle Eocene of India and Pakistan
(Thewissen et al., 1983, 1987, 2001; Kumar & Jolly, 1986) from Diacodexis based on the absence of a character
in its upper molars - lingual side slightly extended posteriorly in the form of a rudimentary hypocone, and
renamed it as Gujaratia pakistanensis designating it as the type species for the genus. They have differentiated
the new species Gujaratia indica from G. pakistanensis based only on its larger teeth. However, the size
difference has not been quantified and measurements for G. indica teeth are not given in the paper. A
comparison between metrics of P/4-M/3 in holotypes of D. pakistanensis (HGSP 300 5003, P/4-M/3 length ~ 17
mm) and G. indica (IITR/SB/VLM 511, P/4-M/3 length ~ 18.8 mm) indicates that teeth of the new species are
only about 110-115 percent as large as those for Diacodexis species. For this comparison I have used
measurements of D. pakistanensis given in Thewissen et al. (1983: figure 1b & table 2) and of G. indica from its
scaled figures in Bajpai et al. (2005a: plate II, figures 4-6). I do not think that such a small difference in size of
teeth can qualify to be an independent specific character; but actual dimensions of G. indica may indicate greater
difference. Unless it is established that G. indica is indeed distinct from D. pakistanensis the renaming of D.
pakistanensis as G. pakistanensis cannot be held valid especially because upper molars of G. indica are yet to be
found; so the presumed absence of a rudimentary hypocone in the upper molars of the Vastan species Gujaratia
indica (as implied by Bajpai et al.’s diagnosis for Gujaratia) cannot be established. The available evidence does
not support renaming of Diacodexis pakistanensis and I think it should have had waited at least until the
discovery of upper molars of Gujaratia. It may be mentioned here that out of the material from Kalakot
originally referred to Diacodexis pakistanensis by Kumar & Jolly (1986) one RM2/ (WIF/A 1611) was
subsequently referred to an adapiform primate but rest of the material is still with D. pakistanensis.
Cambaytherium (Cambaytheriidae, Perissodactyla)
(Bajpai et al., 2005a: plate I, figures 1-8; plate II, figures 1-3 & 7-10)
Authors have described Cambaytherium as a perissodactyl under its own new Family Cambaytheriidae.
However, its teeth look extremely similar to those of anthracobunids, which are known by several forms (e.g.
Anthracobune, Jozaria, Nakusia) in India and Pakistan and at least one (Hsanotherium) in Myanmar and are
generally classified under Tethytheria or Proboscidea (West, 1984; McKenna & Bell, 1997; Ginsburg et al.,
1999; Ducrocq et al., 2000). Although some anthracobunids (e.g. Pilgrimella) were referred to Perissodactyla in the
past (Coombs & Coombs, 1979), later they were all clubbed in a separate Family Anthracobunidae under the
Proboscidea (Wells & Gingerich, 1983; Kumar, 1991). Presently Anthracobunidae is referred to Tethytheria
(ancestors for elephants and sea cows) and is unique to India-Pakistan-Myanmar region. While its ordinal
affinity can still be debated, the dentition of Cambaytherium looks closest to anthracobunids and therefore its
comparison with dentition of anthracobunids was absolutely necessary. Surprisingly, Bajpai et al. (2005a) have not
compared their Cambaytherium material with anthracobunids at all. I wonder if they have any associated post-
cranial material to authenticate its affinity with perissodactyls? They do list a proximal astragalus under referred
material of C. bidens but fail to describe, illustrate or even mention it in the text. This is important because
morphology of P/4 of Cambaytherium is quite unlike that of perissodactyls.
In the familial diagnosis of Cambaytheriidae (presently undistinguishable from generic diagnosis of
Cambaytherium), authors write ‘paraconule and metaconule prominent’ even though they have not illustrated
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Kumar, Comments on Bajpai
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. 2005 www.PalArch.nl, vertebrate palaeontology, 1, 2, (2006)
any upper teeth for the genus. Since Cambaytheriidae is presently represented by the type and only genus
Cambaytherium with four species and the upper molar dentition of only C. bidens has been recognised by the
authors, it must have had been included in the type material (either holotype or paratype) of this species.
Otherwise how do we establish the paraconule - metaconule prominence as a generic/familial character?
It is curious to see that authors preferred IIT/SB/VLM 503 containing only the lower dentition (RC-RM/3
and LP/1-LM/3) as holotype for Cambaytherium bidens over IIT/SB/VLM 502, which contains lower as well as
partial upper dentition (LP/1-LP/3, LM/1-LM/3, RP/3-RM/3 as well as RC/-RP1/, RP2/-RP/4, LC/). It has
probably been favoured for the better preservation and less wear, but surely IIT/SB/VLM 502 qualifies to be
designated as a paratype simply because it also contains the upper dentition, which partly forms the basis for
diagnosis of the new family. Cambaytherium bidens is better known species of the genus represented by a large
number of specimens with upper as well as lower teeth, therefore it should have been named as the type species
instead of C. thewissi, which is founded only on the lower dentition though with less worn teeth. In diagnosis of
C. bidens the authors mention presence of two lower incisors, but none of the type or referred specimens has
been shown to possess I/1-I/2 or their alveoli. Incisors cannot be made out from illustrations probably because
the specimens are all black, and they do not find a mention in description either. It is absolutely necessary that all
characters included in the diagnosis of a species must show up either in the holotype or paratype. Likewise
diagnosis of C. thewissi mentions that the species has three lower incisors but again details of specimens given
under ‘Holotype’ do not indicate their presence or of their alveoli. Although in this case description does
mention that the holotype has three alveoli for incisors, the same cannot be seen in the figure. The description of
C. bidens mentions that there is a decrease in tooth size from P/2 to P/4 (table 2); actually it is not the tooth size
but only the tooth length that decreases from P/2 to P/4.
The size difference between C. thewissi and C. bidens seems inadequate for separating them (C. thewissi
has teeth 82-88 percent as large as those of C. bidens) and there is little or no difference in occlusal morphology
of their teeth. However, presence of a diastema between P/1 and P/2 and two lower incisors in C. bidens against
three in C. thewissi do support splitting. I wonder if by any chance C. thewissi dentaries could be of a juvenile, or
the sexual dimorphism discussed by the authors based on canines has anything to do with this. I may well be
wrong but I think it should be looked into because C. thewissi is based on more or less unworn dentaries of a
single individual.
The third species, C. minor represented by two isolated heavily worn partially broken anterior lower
molars has been diagnosed as smaller than other Cambaytherium and approximately 75 percent as large as C.
thewissi. The length of its holotype molar (10.5 mm) is very close to those for M/1s in holotypes of C. thewissi
(11.0-11.3 mm) and C. bidens (10.8-13.0 mm) but the width is much less (only 5.3 mm against 8.0 mm in C.
thewissi and 9.1 mm in C. bidens). So if C. minor has to be separated from other species it can be, but based on
its much narrower tooth (only 66 and 58 percent as wide as M/1s of C. thewissi and C. bidens respectively)
rather than smaller tooth. Its heavily worn and low crown (pl. II, figures 9-10) raises the question if the tooth
could be a deciduous? I think the authors should have waited for additional material before designating it a new
species.
The identification of fourth species (unnamed), Cambaytherium sp. ‘A’ is based only on an isolated
presumed P2/ (IITR/SB/VLM 549). It has been diagnosed as 130 percent as large as C. bidens. A comparison of
its dimensions with those for P2/ of C. bidens given in Bajpai et al.’s (2005a) table 2 indicates that it is 121
percent as long and 102 percent as wide as C. bidens. Therefore, the tooth is more elongated than P2/ of C.
bidens rather than larger. I wonder if this presumed P2/ could actually be a P3/ of C. bidens because often the
main difference between these two teeth is of size. In the absence of a morphological description no further
comments can be made.
4. Concluding remarks
This critique is the result of a keen and careful study of Bajpai et al.’s (2005a) paper. I was prompted to
pen it owing to my deep interest in the Early Tertiary mammalian faunas of India as I myself have been working
on the same though mostly from Himalaya and secondly because I think it is necessary for understanding the real
importance of the Vastan mammal fauna in relation to the India-Asia collision, the mammalian
palaeobiogeography and origin of modern placental mammals. By raising the concern I do not mean to
undermine the scientific value of the paper. My only intention is to highlight the problems so that these could be
redressed in the next communication on Vastan material. I fully understand palaeontologists’ (including me)
keenness in publishing the new material quickly but I am against casual documentation particularly of
occurrences of new taxa just to speed up the publication of an article. With description of as many as 10 new
species the paper in question will be taken as a key contribution on Vastan mammalian fauna so it ought to have
contained maximum and correct information on all the named taxa. Else the authors could have put up a small
note announcing the discovery and then following up with detailed taxonomic treatment of the material.
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It is curious that out of a total of 14 new mammalian species representing 9 new genera recorded from
Vastan (Bajpai et al., 2005a, b), 12 are based on lower dentition, and the upper dentition has not been recognised
for any of them except C. bidens. The two remaining species, viz. Anthraryctes vastanensis and Cambaytherium
sp. A are known only by solitary isolated M3/ and P2/ respectively. So skulls and maxillae of 14 or more species
are waiting to be discovered. As most of the open mines are non-static, enough samples should be taken as soon
as possible. I am sure both Rana’s and Bajpai’s teams will be aware of this and will do all that is required. I
thank the PalArch Foundation in advance for considering publication of my views on an important research
paper.
5. Cited literature
Alimohammadian, H., A. Sahni, R. Patnaik, R.S. Rana & H. Singh. 2005. First record of an exceptionally
diverse and well preserved amber-embedded biota from Lower Eocene (~ 52 Ma) lignites, Vastan,
Gujarat.- Current Science 89, 8: 1328-1330.
Bajpai, S. & V.V. Kapur. 2004. Oldest known gobiids from Vastan Lignite Mine (early Eocene), District Surat,
Gujarat. - Current Science 87, 4: 433-435.
Bajpai, S., V.V. Kapur, D.P. Das, B.N. Tiwari, N. Sarvanan & R. Sharma. 2005a. Early Eocene land
mammals from Vastan Lignite Mine, District Surat (Gujarat), western India. - Journal Palaeontological
Society of India 50, 1: 101-113.
Bajpai, S., V.V. Kapur, J.G.M. Thewissen, B.N. Tiwari & D.P. Das. 2005b. First fossil marsupials from
India: Early Eocene Indodelphis n. gen. and Jaegeria n. gen. from Vastan lignite mine, District Surat,
Gujarat. - Journal of the Palaeontological Society of India 50, 1: 147-151.
Coombs, W.P. & M.C. Coombs. 1979. Pilgrimella, a primitive Asiatic perissodactyl. - Zoological Journal of
Linnean Society 65: 165-192.
Ducrocq, S., A.N. Soe, B. Bo, M. Benammi, Y. Chaimanee, T. Tun, T. Thein & J.J. Jaeger. 2000. First
record of an Anthracobunidae (Mammalia, ?Tethytheria) from the Eocene of Pondaung Formation,
Myanmar. - Comptes Rendus de l’Acadèmie des Sciences de Paris, Earth and Planetary Sciences 330:
725-730.
Gingerich, P.D., S.G. Abbas & M. Arif. 1997. Early Eocene Quettacyon parachai (Condylarthra) from the
Ghazij Formation of Baluchistan (Pakistan): oldest Cenozoic land mammal from South Asia. - Journal of
Vertebrate Paleontology 17, 4: 629-637.
Gingerich, P.D., M. Arif, I.H. Khan & S.G. Abbas. 1998. First Early Eocene land mammals from the Upper
Ghazij Formation of the Sor Range, Baluchistan. In: Ghaznavi, M.I., S.M. Raza & M.T. Hasan. Eds.
1998. Siwaliks of South Asia. - Proceedings 3
rd
GEOSAS Workshop, Islamabad, Geological Survey of
Pakistan: 1-17.
Gingerich, P.D., M. Arif, I.H. Khan, M.-U. Haq, J.I. Bloch, W.C. Clyde & G.F. Gunnell. 2001. Gandhera
quarry, a unique mammalian fossil assemblage from the Early Eocene of Baluchistan (Pakistan). In:
Gunnell, G.F. Ed. 2001. Eocene vertebrates: unusual occurrences and rarely sampled habitats. - New
York, Plenum Press: 251-262.
Ginsburg, L., K.H. Durrani, A.M. Kassi & J.-L. Welcomme. 1999. A new anthracobunid mammal from the
Kach-Harnai lignites of Baluchistan. - Comptes Rendus de l’Acadèmie des Sciences de Paris, Earth and
Planetary Sciences, Series II 328: 23-31.
Koenigswald, Von, W., K. Rose, L. Grande & R. Martin. 2005. Die Lebensweise eozäner Säugetiere
(Pantolestidae und Apatemyidae) aus Messel (Europa) im Vergleich zu neuen Skelettfunden aus dem
Fossil Butte Member von Wyoming (Nordamerika).- Geologisches Jahrbuch Hessen 132: 43-54.
Kumar, K. 1991. Anthracobune aijiensis sp. nov. (Mammalia: Proboscidea) from the Subathu Formation, Eocene
north-western Himalaya, India. - Geobios 24, 2: 221-239.
Kumar, K., M.W. Hamrick & J.G.M. Thewissen. 2002. Middle Eocene prosimian primate from the Subathu
Group of Kalakot, northwestern Himalaya, India.- Current Science 83, 10: 1255-1259.
Kumar, K. & A. Jolly. 1986. Earliest artiodactyl (Diacodexis, Dichobunidae: Mammalia) from the Eocene of
Kalakot, north-western Himalaya, India. - Bulletin of the Indian Society of Geoscientists 2: 20-30.
McKenna, M.C. & S.K. Bell. 1997. Classification of mammals above the species level. – New York, Columbia
University Press.
Rana, R.S., K. Kumar & H. Singh. 2004. Vertebrate fauna from the subsurface Cambay Shale (Lower Eocene)
Vastan Lignite Mine, Gujarat, India.- Current Science 87, 12: 1726-1733.
Rana, R.S., H. Singh, A. Sahni, K.D. Rose & P.K. Saraswati. 2005. Early Eocene Chiropterans from a new
mammalian assemblage (Vastan Lignite Mine, Gujarat, western peninsular margin): oldest known bats
from Asia.- Journal of the Palaeontological Society of India 50, 1: 93-100.
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. 2005 www.PalArch.nl, vertebrate palaeontology, 1, 2, (2006)
Sahni, A., R.S. Rana, R.S. Loyal, P.K. Saraswati, S.K. Mathur, K.D. Rose, S.K.M. Tripathi & R. Garg.
2004. Western margin Palaeocene-Lower Eocene lignites: biostratigraphic and palaeoecological
constraints.- 2
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Submitted: 3 October 2005
Published: 1 January 2006
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