Status of gymnosperms in the Indian Tertiary flora

(1)

Status of gymnosperms in the Indian Tertiary flora

R.c.

Mehrotra &N. Awasthi

"vlchmtr:, HC ," Awa.,thi N 199'). Slatu, of gymnosperms in Ihe Indian Teniary nora.PUkIL~lh(J/lIni\'143(]): H2-AA.

Gymno,penn,. the mosl dominant group of planls during Mesozoic. were me<lgrely repre,ented in the Indian Teniary nora.

The Tertiary gymnosperms occur in the form of petrified woods, cone, :lnd leaf-impression.,. be.'ide., pollen. refer:,hle to the f:lmilie' Podocarpaceae. AraUClriaceae and Cycadaceae. Unlil Late Crelaceou,-Early l':lIaeocene the gymnosperms were widely distributed in the Indi:m peninsula. However, during Tenia,y they declined to such an extentth:;ttowards the close of Pliocene Ar:wc,ri:,ce:,e to["lIv disappeared from lhe Indian subcontinent. The Podocarp:Keae[00.i, now on the vergc' of eXlinClion a' il occur., in the restricted :'rea.' in nOl1heast India and Nilgiri Hills in southern India. The GIUSt', of decline of these tropical gymno.sperm., and :,dvenl of the nonhern temper:lle members in the Indi:1n suhcontinent during Tenia,y :Ire discussed.

Key-words - I'al:,eohotany. Gymnosperm" Teni:Hy (lndi:,).

R.C. Mehrotrct&N.Alms/hi, Birhu/ Suhni Institu/e of Pu/aeoho/(/ny, 5.3Uniuersitjl Road. Lucknoll'226007, Illdia

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GYMNOSPERMS were dominant during Palaeozoic and Mesozoic times. The Lower Gondwana period was dominated by Glossopteris and its allies. The Middle Gondw;Jna was dominated by Dicroidium which was associated with some Lower Gondwana elements in the early part of Triassic. During its middle and upper parts it was associated with Ginkgoales, Bennettitales, Cycadales and Con- iferales. The Upper Gondwana Period is said to be the best period for Indian gymnosperms,when the flora was dominated by diverse members of Ginkgoales, Pentoxylales, Cycadales, Bennettitales, Coniferales and Taxales and seemingly had covered extensive areas in the Peninsular India.

During the early part of the Post Gondwana,

probably during Middle to Late Cretaceous, the gymnosperms as a group suffered a drastic decline both numerically and geographically from which they never recovered to regain their former pristine pOSi- tion (R3manujam, 1976). There are only a few reports of gymnospermous remains from the Late Cretaceous of Trichinopoly area of Tamil Nadu (Aiyengar &.

Jacob, 1952; Varma, 1955). The pa per is mainly aimed to bring together all the unequivocal megafossil records of gymnosperms known from various Ter- tiary localities of India (see Map1).

TERTIARY GYMNOSPERMS

Gymnosperms. occupying a subordinate posi-

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MEHROTRA & AW ASTHI - GYMNOSPERMS IN THE INDIAN TERTIARY FLORA 1\3

tion in the IncJian TertialY flora, were represented by Cycada les ancJ Coniferales (see Table 1)

CycadaJes

There is a solitalY record of fossil cycad. Paradkar (976) described a petrified ovule as

Gymnouulites deccanii

from the Deccan Intertrappean becJs (Early TertialY) of Mohgaonkalan showing resernblance with the ovules of

Cycas.

The ovule is characterized by narrow curvecJ neck, a flask-shaped lower part and two layered integuments.

Coniferales

The Coniferales were represented by the members of PocJocarpaceae, Araucariaceae, Pinaceae and Cupressaceae.

Podocarpaceae

The fossils of Podocarpaceae comprise leaf, ovule and woods. There is only one record of fossil

leaf of

Podocarpus -

P. oligocenicusdescribed by Awasthi

et al.

(992) from the Oligocene of Makum Coalfield near LecJo, Assam. Among modem. species of

Podocatpus

the fossil leaf is velY similar to that of

Podcxarpus neri!!olius

D. Don.

The only fossil ovule"

Podocarpoovulites chitale..yi

Sheikh & Kolhe1982of the family PocJocar- paceae was cJescribed from the Deccln Intertra ppea n beds of Nagpur. The woocJs of this family commonly occur in the Indian Tertiary sediments. They have been described under the genus

Podocarpoxylon

or

Mesembrioxylon

from the Deccan Intertrappean beds (Mahabale & Rao, 1973; Lakhanpal

et al., 1975;

Bande & Prakash, 1984; TrivecJi & Srivastava, 1989), Mio-Pliocene of CudcJalore Sandstones (Krausel, 1949; Ramanujam, 1953,1954;Agashe, 1969;Trivedi

& Srivastava, 1989),?Oligo-Miocene ofRajahmuncJry

area (Mahabale & Satyanarayana,1978)and Neogene of Kachchh (Lakhanpal

et aI., 1975)

and Rajasthan (Guleria, 1986) Gothan (905) instituted the genus

Podocarpoxylon

for the fossil woods of

Podocarpus

and

DaClydium,

while Seward (919) instituted another genus

Mesembrioxylon

saying that the use of

Table I-List of b'Ymnosperm megafossils from the Tertiary of India

Tax:1 Mouern comparahle Form"lion, Loc:dit)'& Age

form

Ikference

Cycadalcs

G}'IIl,wl"t!i/es c!eccclllii'''Iraubr C)'cadaceae Deccan Imenrappean heu,s of Mohgaon

'Kabn; Early Teniary

I'ar"dbr.197(, Coniferales

Tox()dituylol1 cuc!c1ct!()rel1se 1(;lmanui~m 'T"xouiaceae

Cupresslls'p.d,C. I()nt!()sa Cupressaceae

Me.,elllhnr IX)'!r In(ilsi(i JrmeS;,lln i l'odc)Glrpace;le .II.dudlll.!urel1seM:dl:lh;deO< Ibo l'oc!oGlrpaceae

At rojlllllh{/lel1se.l:1in l'oc!oc:Jrp,lCeae

,llewlI/iJri().q·lfIIlsp,d ..'11. dlldllkllrel1.wMahahale I'odocarpaceae ,".; Ibo

f.J(I(!rn{/ljJ().q'I()I1I'IA'rcIl1JiiI\anue 0< Prakash I'ouocarpaceae r d['(ulI1el1si,Trivedi 0< Snvas[;\v;1 l'ouoGlrpace:Je

I'.A'llIcbel1sisL'kh:lnp:llelal. l'oc!oG,rp;,ceae

Cuc!ualore Series of Tiruvakkarai; Mio- Pliocene

Lower Karewa heds of Hirpur; Mio-Upper Pliocene

DecGm lnlenr"ppeans of j(ajahmundry;

Early Tertbry

DecunIm~nr;Jppeansof1~;ljahmuIld,)';

Early Teniary

Haiahmunury are:J; 'Oligo-Miocene Rajahmundry area; 'Oligo-Miocene Decc:Jn Intertrappean, of I\ombay; Early Teniary

Decc:Jn Intertrapre"n helLs of Mohgaon K;Ii;ln; Early Ternary

K'lnkaw'lli Series of Dh;Jneti (KUlCh):

Pliocene

I~amanuj~m,1960, 1976

Awa.,thi IX Guleria,19H2 Mahabale& '~a(),1973 Maluhale& I~<IC),1973 Mahahale&

Satyanarayana,197H Mahahale 0<

Satya narayana,1978 I\anc!e 0< Prakash,19H4 Trivedi&Srivasl::lva,

'9H9

Lakhanpalel al.,197') Con/d,

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84 THE PALAEOBOTANIST

Podocarpus Podocarpaceae

Podocarpoxylol1 .'chmidianum (Salmi)Kr~usel Podoc:<rpaceae Syn.M. scbmidianumSalmi

P. tiruuakkarainum (Hamanujam) Trivedi& Podocarpaceae SrivaSlava Syn.M. tiruuakkarainu m Hamanujam

P. sahnii (Hamanujam) Trivedi&Srivastava Syn.111 Podocarpaceae Sahnii Hamanujam

P. specio.wm (Ramanujam) Trivedi&Srivastava Podocarpaceae Syn.M ..,peciosum I\amanujam

P mahahalei (Agashe) Trivedi&Srivastava Syn. Podocarpaceae 111 mahahalei Agashe

Podocarprx)uuliles chilaleyi Sheikh&Kolhe Podocarpaceae

Podocarpuso/igocenicus Awasthi et a/. Podocarpus

Araucarioxylon deccani (Shukla) Trivedi& Araucaria-Agathis Srivastava Syn.Dadoxylon deccani Shukla

A. resinosum (Shukla) Trivedi&Srivastava Syn.D Araucaria-AgatbZs re.,'inosum Shukla

A. eocenum (Chitaley) Trivedi&Srivastava SylT.D. Araucaria-AgCllhis eocenum Chitaley

A. Cbhindwarensi,· (Billimoria) Trivedi&Srivastava Araucaria-Agathis Syn.D. chhindwarensis Billimoria

A. shuklae (Singhai) Trivedi&Srivastava Syn.D. Araucaria-Agathi, shuklae Singhai

A. kerienseTrivedi&Srivastava Araucaria-Agathis

A. mohgaoensis Lakhanpal et al. Araucaria-AgathZs A. hikanerense Harsh&Shanm Araucaria-Agathis

Araucariw.ylon sp. Srivastava&Prakash Araucaria- Aga/his

Araucaria - Agathis Araucaria-Aga/hi,

Dadoxylon sp. cf. D. harakarense Surange& Araucaria-Aga/hZs Saxena

Dadoxylon sp. cf. D. jamudhiense Maheshwari Araucaria-Agathis

Mohgaostrohus sahnii Prakash Ardllcariaceae

Indos/mhus hifidolepZs' Salmi Araucariaceae

Tak/ios/rohus ala/us Sahni Araucariaceae

Pityos/mhus crassites/a Sahni Araucariaceae

Harrisos/mhus in/erlrappea Chitaley&Sheikh Araucariaceae

Pinus wallichiana PinuS

A hies sp. cf. A. pindmw Ahies

Shumar Formation of)aisalmer. Itljasthan;

Miocene

Cuddalore Series of Tiruvakkarai; Mio- Pliocene

Cuddalore Series ofTiruvakkarai; Mio- Pliocene

Cuddalore Serie.' ofTiruvakkarai; Mio- Pliocene

Cuddalore Series of MlIrdttandichavadi:

Mio-Pliocene

Cuddalore Series ofTiruvakkarai: Mio- Pliocene

Deccan Intertrappe,m beds of Nagpur;

Early Teniary

/'vlakum Co,i1field near Ledo Assam;

Oligocene

Deccan Intenrappeans: Early Tertiary

Deccan Intenrappeans: Early Teniary

Deccan Intenrappeans of Mohgaonkalan:

Early Teniary

Deccan Intenrappeans of Mohgaonkalan;

Early Teniary

Deccan Intenrappean beds of Keria; Early Tertiary

Deccan Intenr<lppean heds of Mohg<lonkalan; Early Teniary Mar Formalion of nikaner, Rajasthan;

Pliocene

Shantiniketan near Ilolpur. West Ilengal;

Miocene

Shu mar Form<llion of)aisalmer, ]\ajaslhan;

Pliocene

]\:'jahmllndry area; 'Oligo-Miocene

!\aj:Jhrnundry area:' Oligo-Miocene

Early Teniary

Deccan Intenrappean beds; Emly Teniary Deccan Intenrappean beds ofTakli; Early Teniary

Deccan Intenrdppean beds ofTakli; Early Tertiary

Deccan Intertr2ppeans of Mohgaonkalan;

Early Teniary .

Lower Karewa beds of Hirpur; Mid- Upper Pliocene

Lower Karewa heds of Hirpllr; Mid- Upper Pliocene

Guleria. 19H6

Kr~lIsel,1949

Tri\'edi&Srivastava.

19H9

Trivedi&Srivastava, 19H9

Trivedi&Srivast:",a, 19H9

Sheikh&Kolhe, 19H2

Awaslhiet al.1992

Trivedi&Sriv:'stav<l, 19H9

Lakhanpale/al.,1977

Harsh&Sh:<rma, 19HR

Srivast<lva& Prakash, 19H4

Guleria, 19H6

Mahabale&

Satyanarayana,197R M<lh<lhale&

S<ltyan<lrayana, 1978 Prakash, 1957 1962

Salmi, 1931 Sahni, 1931

Sahni, 1931

Chitaley&Sheikh, 1973

Awasthi&Guleri<l,19R2

Awasthi&Guleria.19R2

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MEHROTRA & A W ASTHI - GYMNOSPERMS IN THE INDIAN TERTIARY fLORA 85

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Map - Showing distribution of gymnosperms in India during Teniary.

Gothan's name implied affinities to recent genera for which there were no adequate reasons. Further, Bose and Maheshwari (974) pointed out that since

Podocarpoxylol1

of Gothan is a validly and legitimately published name, it is inadvisable to replace it by

Mesembrioxylon.

Therefore, recently Trivedi and Srivastava (989) transferred the Indian species of

Mesembrioxylon

to

Podocarpoxylon

ex- cepting a few which probably they forgot to do so.

Podocarpaceous woods are generally characterized by growth rings, wood parenchyma mostly without

resin contents, predominantly uniseriate xylem rays, number of cross-field pits 1-2 with apertures vertical or oblique, radial pits bordered and 1-2 seriate, op- posite and rounded.

Araucariaceae

In the Indian Tertiary rocks the remains of Araucariaceae consist of woods and cones. The woods are described as

Araucarioxylon

Kraus or

Dadoxylon

Endlicher (Table 1)from the Deccan In-

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H6 THE PALAE0130TANIST

tenrappea ns (Lakhanpal

et al.,

1977;Trivedi & Srivas- tava, 1989), Neogene of Rajasthan (Guleria, 1986;

Harsh & Sharma, 1988), West Bengal (Srivastava &

Prakash, 1984) and Rajahmundry area (Mahabale&

Satya narayana,1978)The imponant features of these woods are the presence of growth rings, 1-2seriate xylem rays, absence of resin canals, bar of Sanio and xylem parenchyma, cross-field pits1-10,r8dial pining 1-3 or 4 seriate, bordered, alternate and hexagon8!.

For a long time all the fossil woods having such characters from Palaeozoic were assigned ro

Dado.y.vlon,

while those from the younger formations were pbced under

Araucarioxylon,

because the former were thought to belong ro Cordaitales and the latter ro Araucariaceae. According ro Poronie(902) and Gothan (905) the name

Dadoxylon

should be used for the secondary wood with araucaroid type of tracheidal pitting irrespective of the age of fossil woods. Lepekhin8(972)has given a classification of fossil woods with araucaroid pining. According to her there is no difference in the secondaly wood of

Araucarioxylon

and

Dadoxylon. Dadoxylon

shows endarch primary xylem and rather large non-septate pith without secretory canals, while

Araucahoxylon

is based on secondary wood only. On this basis Trivedi and Srivastava(989)transferred some of the species of

Dadoxylon

to

Araucahoxylon.

The fossil comparable ro that of

Araucaria

of Araucariaceae is represented by

Mohgaostrohus sahnii

Prakash 1957, 1962, described from the Deccan Intenrappean beds of Mohgaonkalan.

Pinaceae and Cupressaceae

There are no fossil record of Pinaceae and Cupressaceae from any Teniary sediments of peninsular India. However, from extra-peninsular India AW8sthi and Guleria(982)have reponed the woods of Pi11US, Cupressus and

Ahies

from the Lower Karewa beds (Upper Pliocene) of Kashmir.

Gymnospermsof'uncertain affinities

Besides, there are cones of uncenain affinities described' from the Deccan Intenrappean beds.

These are

111dostrohus hifidolepis, Takliostrohus alatus, Pizyostrohus crassitesta

(Sahni, 1931) and

Harhsostrohus i11tertrappea

(Chitaley & Sheikh,

1973) Indostrohus

is characterized by distinct bract and ovuliferous scale bearing a pair of inverted ovules at a distance from the cone axis. The locality from where it was collected is not known.

Tak- liostrohusand Pityoslrohushave

been reponed from Takli near Nagpur. In the former, the bract is fused with the expanded ovuliferous scale along the mid- line but laterally free and each ovuliferous scale has two ovules. The laner possesses double cone scales and cylindrical ovules with extremely thick integu- ment, the greater parr of "vhich is sclerotesta.

Har- risostrohus

is another cone described from the Moh- g80nkalan chen of the Deccan Intenrappean beds.

The unique character of this cone is the presence of six ovuliferous scales per bract scale, one anached on bract scale and the rest on cone axis. The cone is compact, not woody and bears protecting scales and two ovules per ovuliferous scale. The affinities of all these cones with those of the extant conifers are not definitely known.

In addition, a wood of Taxodiaceae,

Taxodioxylon cuddalorense,

was described by Ramanujam(960) from Cuddalore S8ndsrones. But the author (976) later on was doubtful about its affinities with Taxodiaceae.

DISTRIBUTION OF MODERN GYMNOSPERMS IN INOlA

Gymnosperms occupy extensive tracts all along the subtropical and temperate areas of Himalaya and hilly areas of Kashmir, Assam and Arunachal Pradesh.

They are represented by the members of Cycadales, Coniferales, Taxales and Gnerales. Ginkgoales do not occur in India. Among nine modern genera of Cycadaceae, only

C:vcas

is found in India. The conifers are found predominantly in the Himalayas and are panicularly rich in nonh-west Himalayas. The imponant genera are

Ahies, Cedrus, Larix, Picea, Pinus, Tsuga

(Pinaceae),

Cephalotaxus

(Cephalotaxace8e),

Cupressus,.Iu niperus

(Cu pres- saceae) and

PodocaJpus

(Podoc8rpaceae).

Taxodiaceae is towlly absent. The only conifer of Southern Hemisphere origin found in India is

Podocarpus.

The order Taxales is represented by

Taxus

and

Amenlolaxus,

while Gnetales includes

(6)

MEHROTRA & A WASTHI - GYMNOSPERMS IN THE INDIAN TERTIARY FLORA 87

Ephedra

and

Cnetum.

Most of these taxa are of Northern Hemisphere. The Southern Hemisphere forms mostly introduced are only

Araucaria

and

Agathis

(Sahni, 1990).

DISCUSSION

Even a cursory comparison of the Tertiary conifers of peninsular India with the modern conifers of the extra-peninsular Himalayan region would not fail to indicate a sharp difference between the two. The former were dominated by Podocarpaceae and Araucariaceae, while the latter by Pinaceae. The cumulative effect of the following factors could be ascertained for it:

(a) Appearance of angiosperms

(b) Northward movement of the Indian Plate and uplift of Himalayas.

The Indian landmass (Peninsular India) was the integral part of the then Southern Hemisphere Su per- continent "Gondwanaland" for the major part of the Mesozoic Era (Smith & Briden, 1979), However, during Late Cretaceous the Indian Plate got separated from the main landmass and started moving northward. This was the time when the angiosperm flora started appearing in India. During Palaeogene angiosperms became quite dominant as a result of their progressive diversification and expansion all over the peninsula thus reducing the gymnosperms to an extent that only Cycadaceae, Podocarpaceae and Araucariaceae could be observed. Towards the end of Palaeogene the Indian Plate collided with the Asian Plate resulting in the formation of mighty Himalayas which had adversely affected the gymnosperm population of peninsular India astheycould not- survive under changing conditions. As a result Araucariaceae suffered most and became totally ex- tinct from India after Neogene (Ramanujam, 1978) while Podocarpaceae too is at the verge of its extinc- tion (Rao, 1963) as it occurs in restricted areas only, especially in northeast India and Nilgiri Hills in southern India. However, establishment of land con- nections between India and neighbouring continents and the orogeny of Himalayas after Palaeogene also facilitated the entlY of northern temperate conifers to the extra-Peninsular region. The palynological data from Siwaliks do suggest that the conifers, such as

Pinus, Abies, Cedrus, Picea

and

Tsuga,

etc. (Baner-

jee, 1968; Lukose, 1969; Nandi, 1972, 1975; Mathur, 1973; Saxena& Bhattacharyya, 1987; Singh & Saxena, 1980,1981; Saxena and Singh, 1980, 1982a~b; Singh

& Sarkar, 1984; Saxena

et

at., 1982) had already reached the higher reaches of the newly emerged Himalayas from central and west Asian mainland by Miocene times (Ramanujam, 1976). There is also a report of pinaceous pollen in the Oligocene sediments of Himachal Pradesh (Mathur, 1984). How- ever, absence of their megafossil record in the Siwalik sediments clearly indicates that they did not descend to the foot-hills zone. Thus it is evident that their pollen might have been transported from higher reaches in the north down into the Siwalik Basin. The occurrence of a northern temperate gymnosperm wood of Pinaceae from the Lower Karewa beds (Late Pliocene) of Kashmir (Awasthi & Guleria, 1982) sug- gests that by this time the Himalayas attained suffi- cient altitude to provide hospitable climate for rapid spread of this family.

REFERENCES

Agashe SN 1969. Studies on the fossil gymnosperms of India-PartI.A new species of Mesemhrloxylcm, M. mahahalel sp. nov.

Palaeoholanisl17(3) : 312-316

Aiyengar NKN & jacoh K 1952. A preliminary notes on Pseudo- cycadeoidea, a new genus of plant fossil> from the Trichinopoly District, Madras.Rec .geo/. Sum. [ndla 82 (2) : 325-34 I.

Awasthi N&GuleriajS19HZ.Megaplant remains from the Lower Karewas of Kashmir. Ahslracl [nl. Workshop on Ihe Lale Cenozoic Palaeoclimalic changes in Kashmi1' emd Central Asia. Ahmedahad.

Awasthi N, Mehrotra HC&Lakhanpall{N 1992. Occurrence ofPodocmpus andMesua in the Oligocene sediments of Makum Coalfield, Assam, India.Geophylo/ogy 22: 193- 19H.

Ilande MIl&Prakash U 19H4. A podocarpaceous fossil wood from the Deccan Intertrappean beds of Malabar Hills, Bomhay. Geophylol- ogy14(2) : 171-17H.

Ilanerjee D 196H. Siwalik microflora from Punjab, India.Rev. Palaeobol.

Palyno/.6: 171-17R.

Ilose MN& Maheshwari HK 1974. Mesozoic conifers. [n Surange KR, Lakhanpal RN&Ilharadwaj DC (Editors) - Aspects&appraisal of [ndicm palaeoholany 212-223. Ilirbal Salmi Institute of Palaeobotany, Lucknow.

Chitaley SD&Sheikh MT 1973.Hanisoslrohus il1lertrappea gen. et sp.

nov., a petrified gymnospermous cone from the Deccan Intenrap- pean beds of India.PalaeonlUgrapbica 144B : 25-30.

Gothan W 1905 Zur Anatomie lebender und fossiler, Gymnospermen- Holzer.Ahh preuss. geol. Landesansl 44 : 1-IOH.

GuleriajS 19H6. Fossil woods from the Tertiary sediments near jaisalmer, Hajasthan and their bearing on {he age of Shu mar Formation.Spec.

[ndtan Geopbytologtcal Conf, (Abstract), Pune.

GuleriajS 1992. Neogene vegetation ofpeninsular India.Palaeoholanisl 40 : 285-311. .

(7)

HH THE PALAEOBOTANIST

Harsh 1< &Sh~lrm~111019HH./1rallcarioxylon hikemerensesr. nov. from theTeni~lryo!'llik;lner, Rljasthan, Inuia.Phylonullphol0!l..Y38 (2-3) : III-lIS

KrCiusel 1< 19-19. Oie fmsilen Koniferen-Hiilzer (unter mr"chluss von ArC/llu/ri{)x)'lr!11Kr'lus) II. Teill Kritische unter.suchungen Zur ui;lg- nosllk lebender und fo.s.,i1er Koniferen- Hiilzer.Pulaeolllop,raphica 89B : K3-20.3

Ltkh~mr~d I<N, GuleriJ .IS c'X Awasthi N 197'5. A rodoclqxlceous wood from the Pliocene of Kutch.Ccophytolo!l..Y 5 :172-177

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