See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/260450534

Eocene out-of-India dispersal of Asian

dipterocarps
Dataset January 2011

CITATIONS

READS

28

147

7 authors, including:
Monalisa Mallick

Paul F. Greenwood

Halliburton Technology Center, Pune

University of Western Australia

8 PUBLICATIONS 175 CITATIONS

111 PUBLICATIONS 1,645 CITATIONS

SEE PROFILE

SEE PROFILE

Mulagalapalli R Rao

Roger E. Summons

Birbal Sahni Institute of Palaeobotany

Massachusetts Institute of Technology

34 PUBLICATIONS 235 CITATIONS

171 PUBLICATIONS 8,619 CITATIONS

SEE PROFILE

All in-text references underlined in blue are linked to publications on ResearchGate,

letting you access and read them immediately.

SEE PROFILE

Available from: Mulagalapalli R Rao

Retrieved on: 22 September 2016

Review of Palaeobotany and Palynology 166 (2011) 6368

Contents lists available at ScienceDirect

Review of Palaeobotany and Palynology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / r ev p a l b o

Research papers

Eocene out-of-India dispersal of Asian dipterocarps

Suryendu Dutta a,, Suryakant M. Tripathi b, Monalisa Mallick a, Runcie P. Mathews a, Paul F. Greenwood c,
Mulagalapalli R. Rao b, Roger E. Summons d
a

Department of Earth Sciences, Indian Institute of Technology Bombay, Mumbai-400076, India

Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow 226007, India
WA Biogeochemistry Centre, The University of Western Australia, 35 Stirling Hwy, Crawley, WA, 6009, Australia
d
Department of Earth, Atmospheric and Planetary Sciences, Massachusetts Institute of Technology, Cambridge, MA 02139, USA
b
c

a r t i c l e

i n f o

Article history:
Received 16 February 2011
Received in revised form 11 April 2011
Accepted 9 May 2011
Available online 23 May 2011
Keywords:
Early Eocene
Dipterocarpaceae
Biogeography
Resin chemistry
Palynology

a b s t r a c t
The Dipterocarpaceae, a well known and economically important family of trees of the tropical rain forests of
Asia, comprise over 470 species. These angiosperm trees contribute to 30% of the total area in typical lowland
evergreen forests in Southeast Asia. Despite their remarkable diversity and regional ecological dominance, the
origins and phytogeographical evolution of the family are poorly understood. The earliest dipterocarp fossils
recorded in SE Asia come from Oligocene (3423 Ma) sediments of Borneo. Here, we report an occurrence of
Asian dipterocarps from approximately 53 Ma old sediments from western India based on fossil resin
chemistry and palynological data. An important implication of our nding is that Asian dipterocarps must
have originated in Gondwana and dispersed from India into Asia once the land connection between the Indian
and Asian plate was well established during the middle Eocene (4941 Ma). Moreover, the present study
supports the hypothesis which suggests that many angiosperms did not originate in the SE Asian region, but
dispersed into the area from western Gondwanaland.
2011 Elsevier B.V. All rights reserved.

1. Introduction
The Dipterocarpaceae comprise large trees that dominate the
canopy of lowland equatorial forests. They typically contribute to 30%
of the total area in lowland evergreen forests in Southeast Asia (Aiba
and Kitayama 1999) and play a dominant role in Asian rain forest
ecology (Ashton 1982, 1988). There is no equivalent elsewhere in the
tropics of a single, highly diverse, family dominating large-tree
forests over such large areas (Corlett 2007). The Dipterocarpaceae
comprise three subfamilies: the Dipterocarpoideae in Asia, the
Pakaraimoideae in South America and the Monotoideae in Africa.
There are approximately 520 species in 17 genera amongst which the
Asian dipterocarps include 470 species alone. This family is well
known as one of the major sources of valuable commercial hardwood
timber while its resins and leaves contribute signicantly to the rural
economies of SE Asia. Two opposing hypotheses have been proposed
to explain the origin of the Asian dipterocarps. Some hypothesize
that the family originated in Southeast Asia, most probably from
West Malaysia in the late Mesozoic (Lakhanpal 1970; Sasaki 2006)
and migrated into India during the late Cenozoic Era. The occurrence
of bicadinanes diagnostic of Dipterocarpaceae resins in late Cenozoic

Corresponding author. Tel.: + 91 22 2576 7278; fax: + 91 22 2576 7253.

E-mail address: s.dutta@iitb.ac.in (S. Dutta).
0034-6667/$ see front matter 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2011.05.002

uvio-deltaic oils from across SE Asia represented a further

connection with this age (Stout 1995). Others suggest that
Dipterocarpaceae have a Gondwanan origin and reached Asia by
rafting on the Indian plate (Ashton 1982; Ashton and Gunatilleke
1987; Dayanandan et al. 1999; Ducousso et al. 2004). Here, we report
the occurrence of Dipterocarpaceae from more than 53 Ma old
sediments from India based on fossil resin chemistry and palynological data. Our ndings suggest that Asian dipterocarps migrated from
India into Asia as the land connection between the Indian and the
Asian plate was established at ca. 50 Ma ago (Scotese et al. 1988;
Rowley 1996).
2. Sample and methods
2.1. Samples
Fossil resins, lignite and carbonaceous shale samples were
collected from the Vastan lignite mine (21 2547 N: 73 0730 E)
Cambay Basin, western India (Fig. 1). The rock strata are referred to as
the Cambay Formation. The studied section is 30 m thick and is
composed of shale, clay, calcareous clay and lignite beds. The
sediments of the lower half of the Vastan lignite mine section were
deposited in estuarine to lagoonal environments whereas the upper
half was deposited under shallow marine conditions (Sahni et al.
2006). Fossil pollen having afnities with those of the modern
Dipterocarpaceae family have been recorded from the lower part of

64

S. Dutta et al. / Review of Palaeobotany and Palynology 166 (2011) 6368

73
0

32

400 km.

Sabarmati R

CAM BAY

2m

Lignite

INDIA
16
BAY
OF
BENGAL

22

OF CA
MBA

ARABIAN
SEA

8
72

80

88

Narmada R
ANKLESHAR

GULF

Nummulites
burdigalensis (~53 Ma)

24

VAS TAN

Clay stone
Deccan Traps

Calcarerous clay
Clay stone with
shell layer
Shale
Fossil resin
rich layer

Lignite Deposits

S U RAT
0 Km 20

Pollen of
dipterocarps

ti

Paleocene
Eocene

21

73

Fig. 1. Location and lithological column of the Vastan lignite mine section showing the position of fossil resin-bearing layers and pollen of dipterocarps.

the sequence. Fossil resins were collected from both the bottom and
top lignite seams. The age diagnostic benthic foraminifera Nummulites
burdigalensis burdigalensis which occurs in the upper half of the
Vastan sequence (Fig. 1) indicates an early Cuisian (ca. 53 Ma) age of
the fossil resin-bearing horizon (Punekar and Saraswati 2010).
Dinoagellate cysts suggest an early Eocene (ca. 5455 Ma) age for
the resin-bearing lignites (Garg et al. 2008). Recently, Clementz et al.
(2010) reported the second Eocene Thermal Maximum (ETM2; ca.
53.7 Ma.) from the middle part of the mine section based on organic
13
carbon C data which corroborates the biostratigraphic age of the
resin-bearing lignite section. We obtained the extant dammar resin
from a dipterocarp tree (genus Shorea robusta) from a sal forest of
West Bengal, eastern India.

2.2. Palynological preparation

Rock samples were thoroughly cleaned with water. Crushed shale
samples were kept in 40% HF for 34 days. Carbonaceous shale
samples were then kept in HNO3 for 24 h. The macerated samples
were washed with water 45 times by siphoning and were then
sieved with 500 mesh (29 m). Samples having a calcareous
mineralogy were rst kept in HCl for 12 h and were then treated
with hydrouoric acid. Lignite samples were kept in HNO3 for 24
36 h. The samples, when pulverized, were washed 23 times with
water and were sieved. After removal of minerals, samples were
treated with 515% solution of potassium hydroxide for 25 min and
were then washed. Very ne mineral particles remaining in the
macerated residue were removed with the help of heavy liquid (zinc

chloride). Water-free macerated residue was mixed with a few drops

of polyvinyl alcohol and spread uniformly over the cover glass. The
cover glass was oven dried for ca. 30 min and then mounted in Canada
balsam.
2.3. PyrolysisGCMS
Flash pyrolysis was conducted at 600 C/20 s using a Chemical
Data Systems (CDS) analytical Pyroprobe 5150. The pyrolysis
chamber was held at 300 C. GCMS analysis of the ash pyrolysates
was performed using an Agilent 6890 N GC coupled to a Micromass
AutoSpec Ultima magnetic sector mass spectrometer. A
60 m 0.25 mm i.d. 0.25 m DB1-MS capillary column was used
with a split of 15 mL min 1 using helium carrier gas. The GC oven
was an initial 40 C for 2 min, increased at 4 C min 1 to 310 C and
held isothermal for 20 min. Full scan acquisitions were performed
over the range m/z 50600 at ca. 1 scan s 1. Mass spectral
conditions included 70 eV electron energy, 250 C source temperature and a 300 C transfer line temperature. Product identications
were based on mass spectral interpretation and correlation to
previously published data.
3. Results
We recovered well-preserved macroscopic pieces of fossil resin
from 53 Ma old sediments in the Vastan lignite mine (Sahni et al.
2006; Punekar and Saraswati 2010). We previously reported cadinane
and bicadinane distributions in a preliminary study of the Vastan resin

S. Dutta et al. / Review of Palaeobotany and Palynology 166 (2011) 6368

Fossil resin from Vastan

C15 Sequiterpenoids
f
e

C30-31 Sequiterpenoids

g h

M 424-426

65

C2

10

15

20

25

M 408-410

30

35

40

45

50

55

60

65

70

75

80

85

90

Relative Abundance

Extant dammar resin

g k

B
C2

c
d

M 424-426
M 408-410

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

Time (min.)
Fig. 2. Total ion chromatograms resulting from PyrolysisGCMS of (A) early Eocene resin from Vastan lignite mine, Cambay Basin, western India; and (B) extant dammar resin of
Shorea robusta. A list of identied peaks is given in Table 1.

(Mallick et al. 2009). Here we examine this chemistry including a

comparison with extant dammar resin. Both fossil and extant resins
were analyzed by pyrolysis-gas chromatography-mass spectrometry
(PyGCMS) and the major products identied as C15 bicyclic
sesquiterpenoids and their dimers (Fig. 2). These included calamanene, cadalene, 4-isoproply-1,6-dimethyl-1,2,3,4,4a,5,6,8a-octahydro-

Table 1
Major compounds identied from the pyrolysates of early Eocene and extant resins
from India.
Peak Compound name

Molecular
weight

a
b
c

160
156
204

d
e
f
g
h
i
j
k
l

1,5-Dimethyl-1,2,3,4-tetrahydro-naphthalene
Dimethylnaphthalene
1-(1,5-Dimethyl-hexyl)-4-methyl-benzene or
Dihydrocurcumene
Isomer of peak e
4-Isoproply-1,6-dimethyl-1,2,3,4,4a,5,6,8aoctahydro-naphthalene
Isomer of peak e
Isomer of peak e
4-Isopropyl-1,6-dimethyl-1,2,3,4-tetrahydronaphthalene or Calamanene
Calamanene isomer
1-Isopropyl-4,7-dimethyl-1,2-dihydronaphthalene or -Calacorene
Calamenene isomer
Cadalene

206
206
206
206
202
202
200
202
198

naphthalene and their isomers. Several polymethylnaphthalenes were

also abundantly found in the Vastan fossil resin pyrolysates (Fig. 2A;
Table 1). A separate distribution of peaks at higher retention times
mostly comprised C30H48 (MW 408410) and C31H50 (MW 424426)
dimers attributed to C30 bicadinenes and C31 bicadinenes, respectively.
These are diagnostic biomarkers of class II resin which derived from
angiosperm family Dipterocarpaceae (Anderson et al. 1992; Stout 1995;
Murray et al. 1998; Dutta et al. 2009). Pyrolysates of resin samples from
both the bottom and top seams of the Vastan mine section show
identical hydrocarbon distributions. The corresponding data from the
resin of modern dipterocarps (genus Shorea robusta) also comprised a
dominant distribution of C15 bicyclic sesquiterpenoids and lesser
amounts of C30 bicadinenes along with several C31 bicadinanes and
bicadinenes (Fig. 2B). The extant resin also showed some of the same
terpenoid speciation as the fossil resin. For example, calamanene, 4isopropyl-1,6-dimethyl-1,2,3,4,4a,5,6,8a-octahydro-naphthalene, cadalene and their stereoisomers were detected in high abundance from the
extant dammar resin.
We also investigated the palynological assemblages of the early
Eocene samples to verify the occurrence of dipterocarps. Many pollen
grains recovered from the lower part of the Vastan mine section have
strong morphological afnities with extant Dipterocarpaceae pollen
(Plate I). Pollen grains of modern Dipterocarpaceae are 2585 m in
size, subspherical in shape, tricolpate and possess nely reticulate to
grano-rugulate sexine (Tissot et al. 1994). Following the Articial
System of Classication, fossil pollen grains have been described
under Dipterocarpuspollenites retipilatus, Albertipollenites kutchensis
and Foveotricolpites alveolatus.

66

S. Dutta et al. / Review of Palaeobotany and Palynology 166 (2011) 6368

Plate I. Fossil and modern pollen of Dipterocarpaceae (Scale bar = 10 m). (AC) Dipterocarpuspollenites retipilatus Kar, 1992; Slide no. BSIP 14080 and 14082; (D, E) Albertipollenites
kutchensis Mandal and Rao, 2001; Slide no. BSIP 3585; (F, G) Foveotricolpites alveolatus Mandal and Rao, 2001; Slide no. BSIP 3639; (H, I) Extant pollen of Dipterocarpus indicus.

3.1. Systematic descriptions

Genus Dipterocarpuspollenites Kar, 1992
Dipterocarpuspollenites retipilatus Kar, 1992
(Plate I AC)

Description: Pollen grains subcircular in polar view, size range

4356 m, tricolpate. Colpi long, wide, extending almost up to the
poles. Exine about 1 m thick, tectate, tectum perforate, sexine
giving an appearance of nely reticulate pattern. (Recorded from
Sample no. V-04-01).

S. Dutta et al. / Review of Palaeobotany and Palynology 166 (2011) 6368

Afnity Dipterocarpus indicus

Genus Albertipollenites Srivastava, 1969
Albertipollenites kutchensis Mandal and Rao, 2001
(Plate I DE)
Description: Pollen grains prolate in shape, size range 62 40
67 44 m, tricolpate. Colpi long, wide, extending up to the poles.
Exine about 4 m thick, tectate, columellae closely placed forming
reticulate pattern, muri simplicolumellate, 23 m thick, lumina
elongated, 23 m across. (Recorded from Sample no. V-04-03).
Afnity Dipterocarpus indicus
Genus Foveotricolpites Pierce, 1961
Foveotricolpites alveolatus Mandal and Rao, 2001
(Plate I FG)
Description: Pollen grains prolate in shape, size range 47 28
51 x 31 m, tricolpate. Colpi long, extending up to the poles. Exine
2.5 m thick, sexine pilate, pila 2 m long, capita about 2 m broad,
pila heads generally fused forming perforate tectum. Surface
foveolate, giving an appearance of tiliate pattern, lumina circular,
about 1 m across. (Recorded from Sample no. V-04-1A).
Afnity Dipterocarpus indicus
3.2. Palynooral assemblage of the Vastan mine section
A palynooral assemblage comprised of pteridophytic spores,
angiospermous pollen, dinoagellate cysts and fungal remains were
recorded from the Vastan sequence. The angiospermous pollen shares
about 48% of the total assemblage whereas, pteridophytic spores,
comprising about 25% of the assemblage, are sub-dominant. Dinoagellate cysts and fungal remains are more frequent in the upper part of
the sequence. Amongst the pteridophytes, the spores of the families
Osmundaceae (Osmundacidites spp., Todisporites spp.) and Schizaeaceae (Lygodiumsporites spp.) are most common. Amongst the
angiosperms, pollen grains having afnity with the family Bombacaceae are most frequent and are present throughout the section. These
pollen grains have been described as Dermatobrevicolporites dermatus,
Tricolporopollis matanomadhensis and Lakiapollis ovatus. Of these,
Lakiapollis ovatus is most abundant. Pollen grains showing afnity
with the family Arecaceae are represented by Spinizonocolpites spp.,
Spinomonosulcites spp. and Acanthotricolpites spp. In the lower part of
the section Acanthotricolpites spp. are abundant. Pollen grains of the
families Liliaceae (Retimonosulcites ovatus and Liliacidites magnus) and
Annonaceae (Matanomadhiasulcites kutchensis) are moderately frequent throughout the sequence.
4. Discussion and conclusions
In India, the most prolic fossil records of Dipterocarpaceae are
found from the Miocene onwards (Prasad 1993; Guleria 1996; Khan
and Bera 2010). Some researchers hypothesised that the Dipterocarps
migrated into India from SE Asia once the land connection between
Indian and Asian plate was established (Lakhanpal 1970; Sasaki
2006). This proposition is supported by the remarkable species
richness of the family in SE Asia. In contrast, the phylogenetic studies
comparing Dipterocarpaceae and the Sarcolaenaceae, a tree family
endemic to Madagascar, have shown that the Sarcolaenaceae share a
common ancestor with Asian dipterocarps (Ducousso et al. 2004).
This study postulated that Asian dipterocarps drifted away from
Madagascar with the India-Seychelles landmass and then dispersed
through Asia. The India-Seychelles-Madagascar plate separated from
Gondwana in the early Cretaceous, approximately 130 Ma (Scotese
et al. 1988). Following separation, the Indian plate moved north-east
and most recent studies suggest that the collision with the Asian plate
began in the early Eocene at ca. 50 Ma (Scotese et al. 1988; Rowley
1996; Copley et al. 2010). However estimates of its inception have
varied widely from ~ 54 Ma (Patriat and Achache 1984) to ~ 45 Ma
(Dewey et al. 1989).

67

The earliest conrmed fossil dipterocarp so far reported in SE Asia

is from Oligocene (3423 Ma) sediments of Borneo (Muller 1981).
Interestingly, Borneo is the center of extant diversity of dipterocarps
with more than 280 species. Bicadinanes, which are catagentic
products of dammar resins, have been detected in many late Cenozoic
uvio-deltaic oils from across SE Asia (Stout 1995; Murray et al.
1998). Prasad et al. (2009), while attempting to trace the afnity of
some Palaeogene pollen from western and eastern parts of India with
those of the endemic plants of Indian western Ghats; recounted some
palynofossils (Albertipollenites spp., Dipterocarpuspollenites retipilatus,
Foveotricolpites alveolatus, Intrareticulites brevis, Tricolpites reticulatus)
which bear morphological resemblance with pollen of the family
Dipterocarpaceae. Recently, Rust et al. (2010) reported extraordinarily well-preserved biota within the fossilized resins from Vastan. They
also assumed that the resins were produced by dipterocarps based on
the presence of resin canals within the wood. However, this
observation is somewhat ambiguous since resin canals are present
in many resin-producing trees including some gymnosperms.
Based on the class II resin chemistry evident in our pyrolysis
experiments suggests that the appearance of the family Dipterocarpaceae took place during the Early Eocene on the Indian continent.
One potential source of uncertainty is the report of van Aarssen et al.
(1994) who demonstrated that the Mastixia genus (family Cornaceae)
of angiosperm can also produce the polycadinene resins which yield
bicadinanes on maturation. However, the occurrence of Mastixioid
angiosperms in the early Cenozoic sediments of India can be
discounted as the tree family was mainly restricted to Europe and
North America during the Tertiary period (Tiffney and Haggard 1996;
Manchester 1999). It is also noteworthy that Curiale et al. (1994)
reported the occurrence of polycadinene resin from late Eocene (41
34 Ma) sediments of central Myanmar which was part of the Asian
plate prior to collision. Therefore, it is very likely that the dipterocarps
dispersed from India into Asia during the middle Eocene (4941 Ma),
at which time collision between the Indian and Asian plate must have
completed.
Most of the extant dipterocarps are conned to wet climates with a
dry season not exceeding four months (Ashton 1988). The palynological assemblages, dominated by tropical angiosperm pollen,
suggest prevalence of warm humid tropical climate during the
deposition of these lignites. Therefore, it is very likely that the early
Eocene oras of Vastan are recording the earliest tropical rain forests
of Asia.
Our results also have implications for understanding biogeographical patterns in other ora and fauna. The IndiaAsia collision not only
played a signicant role in the dispersal of faunas from India (Bossuyt
and Milinokovitch 2001; Cooper et al. 2001; Clementz et al. 2010 and
references therein) but inuenced the oral distribution during the
Cenozoic Era in Asia. It is believed that many angiosperms did not
originate in the SE Asian region, but dispersed into the area from
western Gondwanaland (Morley 2000; Conti et al. 2002). Our ndings
strongly support this view.
Acknowledgements
DST (SR/FTP/ES-19/2008) is acknowledged for providing nancial
support to S. Dutta. Authors are thankful to Peter Ashton for valuable
discussions. Work at MIT was supported by the NASA Exobiology
Program (NNX09AM88G). C. Colonero is acknowledged for providing
technical support during resin analyses. S. Bera is acknowledged for
providing the modern pollen of Dipterocarpus indicus.
References
van Aarssen, B.G.K., de Leeuw, J.W., Collinson, M., Boon, J.J., Goth, K., 1994. Occurrence of
polycadinene in fossil and recent resin. Geochimica et Cosmochimica Acta 58,
223229.

68

S. Dutta et al. / Review of Palaeobotany and Palynology 166 (2011) 6368

Aiba, S.-I., Kitayama, K., 1999. Structure, composition and species diversity in an
altitude-substrate matrix of rain forest tree communities on Mount Kinabalu,
Borneo. Plant Ecology 140, 139157.
Anderson, K.B., Winans, R.E., Botto, R.E., 1992. The nature and fate of natural resins in
the geosphereII. Identication, classication and nomenclature of resinites.
Organic Geochemistry 18, 829841.
Ashton, P.S., 1982. Dipterocarpaceae, in: van Steenis, C.G.G.J. (Ed.), Flora Malesiana,
Series 1: Spermatophyta 9, pp. 237552.
Ashton, P.S., 1988. Dipterocarp biology as a window to the understanding of tropical
forest structure. Annual Review of Ecology and Systematics 19, 347370.
Ashton, P.S., Gunatilleke, C.V.S., 1987. New light on the plant geography of Ceylon.
Historical plant geography. Journal of Biogeogragraphy 14, 249285.
Bossuyt, F., Milinokovitch, M.C., 2001. Amphibians as indicators of early Tertiary out of
India dispersal of vertebrates. Science 292, 9395.
Clementz, M., Bajpai, S., Ravikant, V., Thewissen, J.G.M., Saravanan, N., Singh, I.B., Prasad,
V., 2010. Early Eocene warming events and the timing of terrestrial faunal exchange
between India and Asia. Geology. doi:10.1130/G31585.1.
Conti, E., Eriksson, T., Schnenberger, J., Systma, K., Baum, D.A., 2002. Early Tertiary outof-India dispersal of Crypteroniaceae: evidence from phylogeny and molecular
dating. Evolution 56, 19311942.
Cooper, A., Lalueza-Fox, C., Anderson, S., Rambaut, A., Austin, J., Ward, R., 2001.
Complete mitochondrial genome sequences of two extinct moas clarify ratite
evolution. Nature 409, 704707.
Corlett, R.T., 2007. What's so special about Asian tropical forests? Current Science 93,
15511557.
Copley, A., Avouae, J.-P., Royer, J.-Y., 2010. India-Asia collision and the Cenozoic slow
down of the Indian plate: implications for the forces driving plate motions. Journal
of Geophysical Research 115, B0310. doi:10.1029/2009JB006634.
Curiale, J.A., Kyi, P., Collins, I.D., Din, A., Nyein, K., Nyunt, M., Stuart, C.J., 1994. The
central Myanmar (Burma) oil familycomposition and implications for source.
Organic Geochemistry 22, 237255.
Dayanandan, S., Aston, P.S., Williams, S.M., Primack, R.B., 1999. Phylogeny of the
tropical tree family Dipterocarpaceae based on Nucleotide sequence of the
chloroplast rbcL gene. American Journal of Botany 86, 11821190.
Dewey, J.F., Cande, S., Pitman, W.C., 1989. Tectonic evolution of the India/Eurasia
collision zone. Eclogae Geologicae Helvetiae 82, 717734.
Ducousso, M., Bena, G., Bourgeois, C., Buyck, B., Eyssartier, G., Vincelette, M.,
Rabevohitra, R., Randrihasipara, L., Dreyfus, B., Prin, Y., 2004. The last common
ancestor of Sarcolaenaceae and Asian dipterocarp trees was ectomycorrhizal before
the India-Madagascar separation, about 88 million years ago. Molecular Ecology 13,
231236.
Dutta, S., Mallick, M., Bertram, N., Greenwood, P.F., Mathews, R.P., 2009. Terpenoid
composition and class of Tertiary resins from India. International Journal of Coal
Geology 80, 4450.
Garg, R., Ateequzzaman, K., Singh, V., Tripathi, S.K.M., Singh, I.B., Jauhri, A., Bajpai, S.,
2008. Age-diagnostic dinoagellate cysts from lignite-bearing sediments of the
Vastan lignite mine, Surat District, Gujarat, western India. Journal of Palaeontological Society of India 53, 99105.
Guleria, J.S., 1996. Occurrence of Dipterocarpus in the Mar Formation of Bikaner,
Rajasthan, western India. Palaeobotanist 43, 4953.
Kar, R.K., 1992. Occurrence of Dipterocarpus type of pollen from the Miocene sediments
of Kerala, South India. Journal of Palynology 28, 7985.
Khan, M.L., Bera, S., 2010. Record of fossil fruit wing of Shorea Roxb. from the Neogene of
Arunachal Pradesh. Current Science 98, 15731575.

Lakhanpal, R.N., 1970. Tertiary oras of India and their bearing on the historical geology
of the region. Taxon 19, 675694.
Mallick, M., Dutta, S., Greenwood, P.F., Bertram, N., 2009. Pyrolytic and spectroscopic
studies of Eocene resin from Vastan Lignite Mine, Cambay Basin, western India.
Geological Society of India 74, 1622.
Manchester, S.R., 1999. Biogeographical relationships of North American Tertiary oras.
Annals of the Missouri Botanical Garden 86, 472522.
Mandal, J., Rao, M.R., 2001. Taxonomic revision of tricolpate pollen from Indian Tertiary.
The Palaeobotanist 50, 341368.
Morley, R.J., 2000. Origin and Evolution of Tropical Rain Forests. John Wiley and sons,
Chichester, USA. 362p.
Muller, J., 1981. Fossil pollen records of extant angiosperms. Botanical Review 47,
1142.
Murray, A.P., Edwards, D., Hope, J.M., Boreham, C.J., Alexander, R., Summons, R.E., 1998.
Carbon isotope biogeochemistry of plant resins and derived hydrocarbons. Organic
Geochemistry 29, 11991214.
Patriat, P., Achache, J., 1984. IndiaAsia collision chronology has implications for crustal
shortening and driving mechanism of plates. Nature 311, 615621.
Prasad, M., 1993. Siwalik (Middle Miocene) woods from the Kalagarh area in the
Himalayan foot hills and their bearing on palaeoclimate and phytogeography.
Review of Palaeobotany and Palynology 76, 4972.
Prasad, V., Farooqui, A., Tripathi, S.K.M., Garg, R., Thakur, B., 2009. Evidence of Late
PalaeoceneEarly Eocene equitorial rain forest refugia in southern Western Ghats,
India. Journal of Biosciences 34, 771797.
Punekar, J., Saraswati, P.K., 2010. Age of the Vastan lignite in context of some oldest
Cenozoic fossil mammals from India. Journal Geological Society India 76, 6368.
Rowley, D.B., 1996. Age of initiation of collision between India and Asia; a review of
stratigraphic data. Earth and Planetary Science Letters 145, 113.
Rust, J., Singh, H., Rana, R.S., McCann, T., Singh, L., Anderson, K., Sarkar, N., Nascimbene,
P.C., Stebner, F., Thomas, J.C., Solrzano-Kraemera, M., Williams, C.J., Engel, M.S.,
Sahni, A., Grimaldi, D., 2010. Biogeographic and evolutionary implications of a
diverse paleobiota in amber from the early Eocene of India. Proceedings of the
National Academy of Sciences USA 107, 1836018365.
Sahni, A., Saraswati, P.K., Rana, R.S., Kumar, K., Singh, H., Alimohammadian, H., Sahni, N.,
Rose, K.D., Singh, L., Smith, T., 2006. Temporal constraints and depositional
palaeoenvironments of the Vastan lignite sequence, Gujarat: analogy for the
Cambay Shale hydrocarbon source rock. Indian Journal of Petroleum Geology 15,
120.
Sasaki, S., 2006. Ecology and physiology of Dipterocarpaceae, in: Suzuki, K., Ishii, K.,
Sakurai, S., and Sasaki, S. (Eds.), Plantation Technology in Tropical Forest Science:
Springer, pp 322.
Scotese, C.R., Gahagan, L.M., Larson, R.L., 1988. Plate tectonic reconstructions of the
Cretaceous and Cenozoic ocean basins. Tectonophysics 155, 2748.
Stout, S.A., 1995. Resin-derived hydrocarbons in fresh and fossil dammar resins and
miocene rocks and oils in the Mahakam Delta, Indonesia. In: Anderson, K.B.,
Crelling, J.C. (Eds.), Amber, Resinite, and Fossil Resins: ACS Symposium Series 617,
pp. 4375.
Tiffney, B.H., Haggard, K.K., 1996. Fruits of Mastixioidae (Cornaceae) from the
Palaeogene of western North America. Review of Palaeobotany and Palynology
92, 2954.
Tissot, C., Chikhi, H., Nayar, T.S., 1994. Pollen of wet evergreen forests of Western Ghats,
India. Institut Franais de Pondichry 35, 1133.