!.

tt

I .......-.

THE FIRST JURASSIC TURTLE FROM INDIA

Dy p.l,t. o,qrr,q., P. MANNA. s.c. GHosH anr/ o. p. nes

ABSTRACT. A primitive cryptodirao turtle. /IzlorfieA.t s2dtrrla/rr gcn. et sp- nov.. i\ dcsclihcd tl-olr the Early Jurassic

similar ro ihc oklest knowll Etrly Jurirssic cryplodirc, Kd\'!:nt chel\s, horD thc KayenliL l'_ornlation ol-Arizona.

The new fhmily lndochelyidae is proposed, which pr-obably, has the same phyletic strtus as thxt ol K^ycntachelyidae,

\\'i!h bolh evolving simultancously in different regions.

THt onjI]n oi lurtles is srriJ tiispr.rtccl. bLrt [r-ccnt lnirl]::c's ilrt'e sugflestcd eil\el att ttrigin ironl lhe Pcrmian

parciasuirrs (;c. 1995, l',D: ;.. f;<tt; tttc di:lp:,iu i.y,,rts li{icppel rtrii DuLragr i996; D';id-:- nnli

ilicppel 1997: Hedges anC poling 1999). presunrirbly also durir'rg the Pernrian. 'l'he earlicsl known

testudine is Prsgutto<-helys cluenstctlti lrolll the Latc Triassic o[ Gernran]'. ancl is t'cgatcied as the sister

group of all other rurtles (Gaffney and Meeker I 9tl3). A second species of Prr,,ga nochclys, P. ruclne . is the

oldeit turtle from Asia, found in the Norian Huai Hin Lat Formation ol Thailancl in associatior withostracodes, dipnoans, actinopterygians, capilosaurs, plagiosaurs, phytosaurs ancl nunretous estheriids and

bilalves (De Broin et rr1. 1982: De Broin 1984). An '.tnnamed proganochelyid sku)l is also recotded from

the Earl), Jurassic Elliot Formation of South Aflica. and is the earliest knorvn Aliican turtle (Caffney

1986\. Prolcto(het'si.t rol)usta, thc oldest kt':o$'n lllcurociirc. is recorclctl llolr the samc holizon ilGerrnany as PRt,?ut1o(hcL,..s tlucttstctlti (Fraas 1913r. thr!. demonstrating the piesenc.r olthc Pleutorlilain the Llpper t'rilssic. 'l'he earliest cryptodire an.i $i. oldest kno*'n tutle ll'om North Ainerica is

Kaye ochell,s nprlr, a primiiive cryptodire irom the Early Jurassic Kayenta Fonnation ol Arizona(Gaffney er al. 1981). There are numerous Late Jurassic turtle records from llorth America (Gaffney

1979), Europe (Romer 1957) and China (Yeh 1983).From the Cretaceous onwards, the record of fossil turtles becomes more prolific throughout the world.

From the Indian subcontinent, pelomedusid turtles are known from the Infra- and Intertrappean

Beds (Carter 1852; Hislop and Hunter 1855; Lydekker 1890; Sukeshwala 1941l' lain 197?) of central

India. These beds are considered to be Upper Cretaceous (Pascoe 1964; Robinson 1967). Pelomedusids are

also known from the Upper Eocene and Oligocene ofthe Fayum, Egypt, and fiom the Miocene (Burdigalian)

ofSaudi Arabia. These Asian and African petomedusids appear to represent a branch off the common African-South American pelomedusid stem, with Cdrtercn^s pisdurctisis Jain, 1977, as their earliest representative

(De Broin 1987). Finally, the Early-Middle Eocene carettochelyid cryptodire. Chorhkkicheh's shqhi, is

known from carapacial and plastral scutes flom the Kuldana Formation of Pakistan (De Broin 1987).

The purpose of this paper is to describe the shell morphology of the oldest non-proganochelyidchelonian discoyered in the Early Jurassic Kota Fomralioll of the Upper Condwana Group in the Pranllita

Godavari Valley, Deccan, India.

G]]OI-OGICAI- B AC KCROUND

The Kota Formation, a member of the Upper Gondwana Group of rocks. crop-( out on the east and west

banks of the Pranhita and Godavari rivers. and is characterized by the presence of a highly tbssiliferousiiirioslollc o€li a5 \^ cii a" tire tegutar. lurrsrstcnl a\s< ir,ordgcs uf ltul tuttre aLnq iacLlstrlne sancistone anq clays(King l88l ; Kutty 1969; Rudra 1972). A generalized stlatigraphy of this outcrop js shown in Text-flgure l,and the lithological succession as exposed in the 'Kota' type area is shown in Text-figure 2.

lPaheonlology, VoI.,13, Pltrl l, 2000, pp.99-1091 O Thc Ihlreontological Asloci.tlior

\

PAI-AEONTOI-OGY. VOLUME .I3

TE1T,F1G. L General geology of rhe arca showing the fossil locality in the Kota Formation (inset: location ofKota village).

The Kota Formation is well known for its rich fossil content. Fossil fishes include the semionotid genera

Lepidotes, Paradapedium, Tetragonolepis, the halecomorph geruts Pholidophorus (Jain 1973; Yadagiri

and Prasad 1977), and the coela canth htdocoelacanthris (Jain 1974). On the basis of these fossil fishes, the

age of the formation is considered to be Early Jurassic (Jain 1983). Seven estheriid-bearing units in the

Kota argillaceous limestone at the type locality indicate a Jurassic age for the formation (Tasch €l r1.

1973). Govindan (1975) found the freshrvater ostracode Dan irelki at Kota, and suggested that the age ofthe fomation may extend into the Middle Jurassic. The presence of the primitive sauropod dinosaur

Barapasaurus tagorei (JaiI. et al, l9'15), and the kuehnotheriid mammals Kotatlxeri|lt lruldatrei lDatta.1981,), Trishulotheriurn kotqensis and Indotheriut pranhitai (Y adagiri, 1984) also suggest an Early

Jurassic age for the formation. As well as the above-mentioned vertebrates and ilvertebmtes, the formation

has also yielded numerous charophytic gylogonites. wood and fragmentary insects. The fauna that itcontains suggests a dominantly fleshwater depositional enYironment.

MATERIAL AND METHODS

The specimen studied was lbund lrr .sirri, embedded in the hard limestone of the Kota Fornation at I l 5 I11

abovJ the bed of the Pranhita River. Both the carapace and plastron of the specimen were found to bc wellpreserved apafi from the tips ofthe posterior region, the mid-posterior marginal fiinge on the left side and rhe

marginal part on the right side. The central part of the plastron is also missing. No skull or postcraniai ntatcrials

ID A.T'l l\ ET A L.: JURASSIC TURTLE

TEx't.Ftc-4.h1tlochel:-tsPdatlaldgen.et.sp.nov.,hololype'GSl203S0,interpletivedrawinSA'dorsalview;ts'rentralview. Solid lines (except outlineirepresent the scutc bordcrst Sen'ated lines lepresent sutures; broken lines rcpresent

inconspicuor.rs scute borders; hatched areas represent btoken parts of_the. specimen and dotted ar-eas represent space

filled with matrix: for abbreviatioos. see Table l ' Scale bar represents 20 mrn'

incuryed atrterior margin; no supramarginal scutes (shared wlth Kayent\chebts); eleven paired. squarish.

;arginal scutes; four'paired, pieural icutes longer than broad (shared w\th Proganochells); plastron

relaively small and narrow with truncated anterior margin not plotruding beyond the cervical scute; a very

small ga'p between carapace and plastron all along the anterior margin; plastron having eleven elements

including a pair ofmesoplastra; e;toplas[on broxd, spatulate, extending to the anterior margin of the shell

and sepi'rating the epipiastra (sharcd wirh Kqentachelysl'. paired gular scutes extend to entoplastrcn

almost a thirJ of its length; one pair of mesoplastra. rarrowing towards the mid-line; plasto-carapace

bridge one-third of shell length.

Descnplion. This monotypic genus is represenred onl) by the holotype (TexFfig. 3). The shell is low,fleaftshaped,

227.5 mm long and 2l3.dmm-wide. The maximum shell height is 48.1 mm atrained at the sixth neural. Both carapace

,na pfuaoo, a"re smooth and without any omamentation except for- two small prominences on the third and eighth

neurals. The dimensions of individual elements are given in Table 2'

CaraPace(Text-figs44'5A).Theanteliormargjnofthecanpaccisincurvedinthenuchalregion;otherwise,the,nu.gin i. n"u.ty ,iooth, eniire and sharp. The p;sterior region is mostly serrated, but is slightly notched in the pygal

regi|n. The sh;ll is slightly depressed Gngitudinally all along the neural region. The anterior half of the carapace

.i|fe. ao*n "onria",abiy

from ihe first neural, thus forming a small hump over it. This probably indicates the point of

att;chment of the last ceivical with the first fixed thoracic vertebra. However, the area enclosed by this anterior part oI

the carapace and the plastron below provides little space for relraclion of the head and neck. The posterior half of the

carapac; rs slrghrly s;alloped at the poste or end. Ihe nuchai regron consrsls oI a iarge nuchal bone which ls almosl

"ou"."d by th-e fiist vertibral scute. The neural series consists of nine broad, drurn shaped to hexagonal neurals

(n1-n9)l tiie second neural is alnost spherical in outline and the eighth is comparatively small'

Eighi paired costals (cl -c8) are almost parallel to each other and are separated by the neurals medially. Each costaL

exten?s to the peripherals and none of them forms costoperipheral fontanelles. Each costal is narrower towards ihe

Scutes

PALAEONTOLOCY.

Bones

VOL UM E 43

Scutes Bones

TE*T-FIC.5. Restored shell morphology of-hrlachely spatulutu gen. et sp. nov. A, do6al view: B. ventral view:conventions as for Text-figure 4. Scale bar tepresent-< 20 mm.

neurals and inlxdcf at the peripherals except ibr the iirt costJ jr utich thc sjruati,lil isjusi rhe reverse. The llrsl costalarricuratci only with rhe second and third peripherats rather than with rhe n..tit.orgi, ttrird peripherais.- - " -- '"^

Eleven paired peripherals (pl-pl l) are ;ll squarish or rectangular in ir,up" u.o "u"i

l, ,uppo.tJa "itr,".

ry u ,ingr"costal or by two adjacenr ones; p2 is supported by borh cl ana rn"e nucrniwrie.eas pt is ruseainly to th;r.ili i#;;are two suprapygcls The anterior suorapygal (spl) is triangular and attaches to th; last costar t&l and tne lasine,iaifn9). The posrerior suprapygat fspzr is cieic.oti'" "ra

i, for'r"rio ;;i.d";l;;. pygrl. rhe panty paired pl0 ind thepaired pl l The pygal was probably broad.The epidermal scutes are thin and preserved in only a few patches. The interior relief of the scure marsin is clearlvimpressed as grooves on the exrerioi relief of rhe dirmat bones. Th"r.;;"-;;; ;;:';;;,il'.J]fjffiT;i:{(v1-v5) The third (v3) is widest and covers aboul53 per cent. ofthe width oithe shell. The second, third and fourthvertebBl scutes are hexagonal in outline. They have almost straigtrt marglns whereas the first one has a mushroom-shaped outline with irs Dosterior marpin undulated. ffre nftfr vJreUiaiis-Uo*l_rt rp"a

""d i,; ;";;;;;;;j;j;enclosed by rhe lasr pairid marginals fnd rhe paired supracaudals.

TABLE l. Abbrcviations.

Dorsal elentettsScutes

Ventral elenpntsScutes

Pn, cervicaiV, vertebr'alL, pleuralM marginalSc, supracaudal

nu, nuchal bonen, neuralc. costalp. periphr:ralsp, suprapySalpy, pygal

Gu, gularHu, humeralPec. pectoral

Fem, femoralAn, analAx, axillaryIn, inguinal

ep, epiplastronen, entoplastronhyo. hyoplasrron5tp. hJpophsirolmes, mesoplastronxip, xiphiplasrronb, br-rttress

/DLTTA ET AL., J URAS S lC TURTLE

TABLE 2. Measurements

Maximum shell lengthMaximum shell width

Bones

Nuchal (nu)Neural (n)Suprapygal (spl )Suprapygal (sp2)

Pygal (py)Costal (c)Peripheral (p)Epiplastron (ep)Entoplastron (en)

Hyoplastron (hyo)Mesoplastron (mes)

Hypoplastron (h)"p)

Xiphiplaston (xip)Bridge (b)

Scutes

Cervical (Pn)Vertebral (V)Supracaudal (Sc)

Pleuml (L)Supramarginal: noi - iearMarginal (M)

First neural (nl)Second neural (n2)Third neural (n3)Fourth neural (n4)Fifth neural (n5)

Sixth neural (n6)

Seventh neural (n7)Eighth neural (n8)

Ninth neural (n9)

Vertebrals

First vertebral (Vl)Second verteb.al (V2)Third venebral (V3)Fourth vertebral (V4)Fifth venebr.ri (\5,

Number Paired./Unpaired Size in mm,length x reidth

24.2x82.1variable18.6 x 55.216.5x'71.'723.0 x 35.0variablevariable19.0x 14.035.Ox22.550 1x50.0

9.5 x 1!.146.7 >< 46.1

31.3x31062.5x34.2

Size in mm,

lenBthxtridth

13.9x 14 0variablevariable

variable

Meximufi median,width inmm

15.512.813.012.613.815.818.218.22t.o

Maximum median,Length in mnt

82.6102.5114.598.5i40

22'7.5 ntu Maximum shell height 432'0 mm

213.0mm Maximum median length of plastron 169 5 mm

I9I1

l622

2I222222

Number Paired./Unpaircd

unpaired

8 pairedl1 paired1 pairedunpaired1 paired

unpaired

I paired4 paired

l1 paired

I52ti

22

Maximum median,length in mm

25.211.122.616.215.0t7.t13.08.9

14.1

Marimum median,length in mm

18.145.052.64t.1ri i

106 PALAEONTOLOGY. VOLUME 13

The well preselved cervical scute (Pn) is triangu]ar with an incurved anterior margin. and extends tuedially into thc

first vertebral scule. Four asymmet cal pleural scutes (Ll-L4) are present on either side ol lhe vertebral scutcs. Each

pleural scute originates from the pointed marginal ends ofthe two adjacent vertcbral scutes and is irregularly elongated.

Eleven paired, squarish marginal scutes (Ml-M t l) Iie along the periphery of the carapace. The second, third, fourthand part of the fifth border the first pleural scute; the fifth and sixth border the second pleural; the seventh, eighth and

ninth border the third pleural; and pan of the ninth and tenth border the fourth pleural scute. The first marginal scule(Ml) and the last (Ml1) are attached to the first and fifth vertebrals respectively.

Plasrlo,i (Text-figs 48, 5B). The plastron is comparatively small and narrow; its maximum length is 169 5 mm. There is

a wide gap between the plastron and the posterior margin of the shell. This probably provided free space for the

moverncnt of the hind limbs ard the tail. The most proninent clcnrcnt in thc plnstron is the entoplastron (cn). The

cntoplastral suture is hi-qhly sefated and its lateral margin oo cach side ol lhe midlinc of thc shell rLlns staaiSht t() Incel

the extrcmc alnlcrior rnlr gin of thc shell and separatcs tllc paircd epiplastr-a (ep) on each side of it. Thc niddlc ol lhe

cnloplaslrou is $,idest. and gradually tapers posteriorly lo l]rcct thc nlid line, impiir{ing a spatulate shitpc to dte

enloplastron. Two cpiplastra, one on each side ol tlie entoplastron. are distinct in the specinrcn- The epiplastral suture

runs liorn tlrc phstr:ll nlargin as the hyo-epiplastral suture straight for sonle distance towards the mid line of the

plastron and suddenly tums anteriorly to merge with the enloplastral suture. Thus the paired epiplastra rcnlain

separated by thc rostral paocess of the entoplastron. Thin, indistinct. gular scutes cover almost a third of the

enlnplastron. However. the intcfgulrr':.Lu.s arc n61 1'sry rllcarh seen in lhi!'oecimcn.The hyoplaslron (hyo) is broader ther, rhe hypoplasrron (h_y"p)- Paired razor-shaped mesoplastra (mes) are pres,)nt ;n

the fiid-region of ihe plastron. Pans ol the hyoplastron. hypoplaslron and rnesoplaslron are not discemibiu )rr the

rniddle part of the platform owing to the effect of ercsion. Their probable outlines are depicted in Text-Iigurc 5B. 'fhe

xiphiplastron (xip) is nluch wider afld longer than the epiplaslron, and the irnpression of the pehic ankylation is notIbrmed. The humer.al scutes (Hu) are much wider than the fernorals (Fcm) rvhereas the pectoral (Pec) and abdonlinal(Abd) scures are almosl equal in size.

Thc plastro carapacc bridge is 625mm long and is about 54 per cent. of the total shell length. Therc is an

impression of the axillary scutes (Ax) and inguinal scutes (ln) which are presutued to be the traces of inframarginalscutes (Text-fig.48). Horvever, no other inframarginal scutes aae visible in lhe present spccimen.

Thc possibility of delelopnenl of a burrress (b) in the specjmen is conjectlred because of the tcndcncy of jnward

hor1, gr,-*.lth ol the antero lateral plaslral !.he to\tiiids ihe a\in-.!r:!ilxrixl !',rgior of lhe \hell

DISCUSSION

It is evidelt from the above description that IndocheLys spatulata prcserves many of the dermal and

epidermal characte$ of carapace and plastron. In the following discussion we attempt to determine the

relationship of this gerus, and to place it systematically based on comparisons ofthe shell morphology. Inthe absence of a skull, we cannot take cranial morphology into considemtion.

One attempt has been made to classify the Testudines using shell morphotogy. Zangerl (1969) proposed

a classification of the order Testudines (as Chelonia) on the basis of progressive adaptive characters of the

dermal and epidermal shield cover. Four major levels of shell organization were recognized, namelyAmphichelydia, Mesochelydia, Metachelydia and Neochelydia, and each given subordinal status.

Comparison of Indocheb,s with these four sub-ordinal levels demonstrates that it possesses most of the

characters associated with the Mesochelydia in that it has nine neurals, five moderately large vertebral

scutes, eight paired. costal bones, eleven paired peripherals, four paired, moderately large, pleurals, one

pair of mesoplastra, and gular scutes covedng about one third of the entoplastron. On the other hand, the

absence of supramarginal and inframarginal scutes, and the reduction of the marginal scutes to 22 suggest

an approach to the more advanced metachelydian gradc. However, because the information contcnt ofturtle skulls is so much greater, Zangerl's classification has not been widely used subsequently and '. . can

be viewed as an abenation in the development of natural groups of turtle taxa and has been .justifiablyignored by other workers. .' (Gaffney 198a, p.292).

Recent phylogenetic analyses of the turtles are based almost entirely on cranial moryhology. ln theabsence of a skull. we are uxahle t. elace Indochehs preci'.elv in the cladogram of higher ca'egorie" nfturtles and their near relatives as proposed by Gaffney and Meylan (1988). However, an attempt is made toplace it among the higher categories and determine its nearest relatives by utilizing some features of the

shell morphology.

DATTA E7 Aa.: JI,'RASSIC TURTLE 107

It is now recognised that the head retmction mechanisms of pleurodires and cryptodires arose witllinthese groups, anA that primitive members of both groups lacked this ability. ln Indochelys, the anterior

shell opening all along the antedor margin barely provides enough space to accommodate a retracting head

and neik, whether the head is withdrawn by a sideways bending of the neck, as in pleurodires, or stmight

back into the shell, as in cryptodires (Roner 1957). It is probable that the head of /ndoc,/zelys still could not

be rvithdrawn, as in primitive turtles like Proganochelys and the most primitive pleurodires and

cryptodires.The she1l morphology of Progonochclls cliffers substantiully fiom that ol Indochel\"s. Although lhe)'

share gcneral or primitive testudinatc cltalacters. srrch as lirur pairs of pleurlll sculcs lnd one pail ol'

nresopiastra (Galfney 1985), in otl'rcr lespccts Ptogunoclttlrl diffcrs from lntlochclys in all aspecls ofcarapacial and plastral bones and scutes.

The shell morpholo gy of Indoclrclls spalll1ata more closely resembles that of the primitive cryptodiran

Kalentachelys aprix (Gaffney et ol. 1981) fron't lhe Early Jurassic of Arizona. The most stliking shared

cllaracters of these two genera arc found in the anterior region of the plastron. In both genera, the

enrr'11rstron is the sanrc shape a-lcl size e{lending up to the tip of the :nteri()r plastral nrargin and thus

separrtins the paired epiplastra for nearl-,- their ontilc length. In both g(:nera. the r'ntoplaslrorl is a brord,invened, leaf like structure. and modified into a spirtuiate shape. Ktlye toclvh's has a pair ol guiar and

intergular scutes. The gular scutes are conrpletely separatcd by the intergular scutes, rvhiclr meet fbr most

of rheir length along the mid line and cxtend from the epiplastron onto the entoplastron. In 1,ldocre1-),t, tilegulars are distinct. The iDtergulam are less distinct but probably confonn to the same shape and sizc as

ihose of Kayentachelys. In both htdochelys and Kalentachelys supramarginal scutes are absent.

In other respects ,?dor:,/re1ys may be more derived than Kay,enlaciel-vs. lt has eight pailed costals (nine in

Kq.ent(rchelys), the first costal sutures with the second and third peripherals (first three in ,&'al e,,1.Ic,l?ell .t).

There are two wide suprapygals (one small suprapygal in Ka\entochelrs) and apparently no infranrarginalscutes (foul pairs il Kclcrlacliells)

Fr,ril '.he cornparati\ e stu'il,' oI lnorlhoiogicrl aharactcrs. it is r.':-r- .lear lhat the Eelll' Jurassic

ltuiocirclls shares sevelal sheli charactels lvith the No h American Late Jurassic genus Diitoclie1t's(Gaffney 1979). Alrhough the similarity o f lndochel) s to Dinocfte/,ys is greater than to any other MesozoicEuropean and North American genera (GlyptoPs. Trinitichelys, Pleurostenton, Plesiochelys), it differsfrom Dinochelys nt that its third vertebral scute is widest, covering about 53 per cent. of the shell-width. InDfuochelys the third vertebral scute is the widest, covering about 62 per cent. of the shell-width' The

plastrorlin Indochelys also has a smoothly rounded anterior edge whereas in Dinocielys it is produced intolobes owing to indentation of the gular and intergular s\lci. Dinochelys is usually a thick-shelled turtle(Gaffney 1979) whereas Indochelys is thin-shelled.

Thus the shell morphology of different Mesozoic genera belonging to different families ranging fromthe Late Triassic to Late Jurassic demonstrates lhal lndochel)-s has certain characters that are notcomparable with any other Mesozoic forms. Consequently, the new family lndochelyidae is suggested

for this genus. However, there are certain characters in the new family that are shared with other families.This is because the shell has a structural complexity with limited potential differentiation. Thus.

evolutionary modifications in the turtles have tended to lead to the same features in virtually all the

phyletic lines. This has resulted in a great deal of parallelism among them.The inverted Tlike entoplasfton is slightly modified into a 'spatulate' shape rn lndoclrclys spahtlata.

This feature and others, namely the ischia articulating to the xiphiplastron, the presence of a cervical scute,

the absence of mid-plastral fontanelles, the immovable anterior plastral lobe, the strongly ossified shell,

and a possible buttress in the specimen, are all cryptodiran in nature. The retention of mesoplastraprecbdes Indochelys from membership of the Eucryptodira (Eucryptodire character 2: Gaffney and

ileyl"r, 1988, p. i,'1i ai,d pialcs it \uiihin th(r basal glade of lryptudires *ith thc Laycntautrciyids,pleurostemids and baenids.

The vertebral scutes are much broader than long (the pleural scutes being longer), the nine neurals

sutured to vertebrae, the epiplastra separated ventrally by the rostral process of the entoplastron, and onlyone pair of mesoplastra meeting along the midline. are all rather archaic in character. These prove

reasonably beyond doubt that we are dealing here with a relatively archaic cryptodiran turtle. It is.

I03 PALAE()NTOI.OCY. VOLUME,13

theref'ore, postulated that lndochel)'s antl Ka\entachel\'s belonged to two distinct lineaces ancl evolvcd inparallel, perhaps cnretging simultaneouslv within the szLme incestral clyprodiran siock ol the sinclcmonophyletic heterogenous Casichelydia in two geogl.aphically isolated land masses.

. The spherical low-domed carapace, plano-convex flat ihell, moderately sltort Ien-rth ofthe bridge, sn.rall

plasron size, and broad and upcurved posrerolateral peripherals indiiate that itlc,clrctys ,pititnro l,undoubtcdly an amphibious or aquaric tu(le, likc Kayintaihelys (Gaffney c, a/- 19g7).

Ackn.o.wledgentents. We are Srateful to the Director Ceneral, Geological Survey ol lndia for giving permission topublish this paper, and to Mr T. C. Lahiri. Direcror, palaeontology and StratiSriph), bivision, Geololical--survey of India.fbr his cr-itical examination of the rrlrnusc.ipt. '1hc ldvice ancl cirnstant tcchn-iclil guirlance of pr.ol- i. K Roychoq.cl6urv.Indian,Statistjcal Institutc, Geololi.ll Studies Unit, Calcula, is gratcfull), a:knL.tvlcd!ed. I hrnl\ rrc rlx, .lue r,,Mr P. K. Basu. Geologist (Senior) t,i rhe,Cgrlogical Survcl, o ndia, for his heI and li.uitiitl discussions. ].ol obrait)in-qreluted repdnls we lharrk Dr S. L. Jain. Indian Stllistical Institute, Geological Studies Unir, C.rlcutra. Dr \1. J. Bentoirof lJristol-uni\ersiry' Dr tsugene Gaft'ney ol rhe American Museum olf Naturar Historv-, ancr Dr A. R. Mirner orBirkbeck College London are thanked for their clitic.rl conlments on our ear'lier manuscript and constant encour.agc-m(nt und .uege"tions lor intlrovinF thc rn.lrlts. r.fl.

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a it aj \I-IAS. C GHOSH

D, P, DAS

Geological Survey of India,Palaeontology Division,Calcutta-70o 016, India

Typescript received .]0 Januarv l99gRevised lypescript received Ii Augusr I999

P, MANNA

Department of Zoology,Rishi Bankim Chardra Collese.

Naihari, West Bengal. Indii

(1

DATTA ET A L.: JURAssIC TURTLE r09

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Typescripr received 30 Januarv lgg8Revised rypescript received Ii August I g99

i:. l.i lr.\iTA

s. c. cHosH

D. P, DAS

Geological Survey of India,Palaeontology Division,Calcutta-7oo 016, India

P. MANNA

Department of Zoology,Rishi Bankim Chardra College,

Naihati. West Bengal. lndia