This document presents the Normal Table of Bufo melanostictus Schneider, which details the external morphology and measurements of the various developmental stages. It begins with an introduction providing context about B. melanostictus as a widely distributed toad species in South Asia suitable for classroom and experimental studies. The study aimed to document the normal development of B. melanostictus in detail through observations of externally visible characteristics over 43 numbered stages from the fertilized egg through metamorphosis. Materials and methods describe the collection of eggs and embryos, measurements, fixation techniques, and "staging" system used to define each developmental phase. The Normal Table then describes the key features of each numbered stage.

1. +:Plfl."if,June,1965
A Normal Tableof Bufo melauostictusSchteider
By
MOHAMMAD SHARIF KHAN, M.Sc.
De|arttnentofZoologr. Gove mentCotlege,Lahore..
INTRODUCTION
BufamelatlostirtusSchr,eiderjsrhecommonesrtoad of theIndo_pakistan
Sub-conlinentand occursjn almostall parts of Pakisran(Muray, 1894).
Being so widely distributednnd casily obtainable,Bufo nletanostichts
could well serveasa typefor classroomstudy,andcould alsoprovidematerjal
for experimentalwork on amphibjanErnbryology. In bolh caseshowever,it
would be necessary10 havea detailedaccounlof its normal development.
Deta;led descriptionof rhe varjous sragesof developmenrof over a
dozenamphibians,often arrangedin the form of illusrraledtables,havebeen
publishedbut almostno literatureon ihe developmentof our commontoad,
Bufomelatlostiet8,isavailable. In rhisLaboraroryHashmimadea prejiminary
study ofthe earlydevelopmenralstagesof this toad, but his studieswere
aimed at gellinga generalpiclure of earlyontogenyrather rhad at compiling
a Normal Table of development. Moreover, his findings have remained
unpublishedso1ar.
The presentstudy has beenurdcrrakento preparea Normal Table of
Bufo melanostictusas a prelimina.y 10 frrther embryologicalwork on lhjs
This work was don€ in the laboratoriesof the covernment College,
Lahore,in parlial fulfilment of the requirementsfor the M.Sc. degreeof the
University of the Panjab. I wlsh to acknowledgemy indebrednessto Dr.
AhsanulIslam who suggestedthe problemand underwhose supervjsionrhe
wofk wascompleied.
MATERIAL AND METHOD
(r) Collectiooof Mat€rial
The nalerial for tbe presentstudy wascollectedfrom lhe Govcrnmcnt
Coll€gelawns,lheColBagl, aDdorherIocatirjesjn ihe vicjnjtyoftbe laboralory.
ln order to make surethat the eggsand the othcr developmentalstages,on
rhichthisstudyisbased,reallybetongedto ,rdo melarrrlirlur, it wasnccessary
Dep3rtmentol Zooloey,Talim.rl.hlemCotlege,Rabwah,DistlictJbang,Weslpakistad.

2. BIOfOGTA
to catch the animalsin the act of Iaying eggs. This end was achievedby
tollowing Hashmi's (unpubijshed)technique,wlereby coputaringpairs of
toadsarecollect€dand kept ir a largebasinjn the laboratory. In mostcases
tbe male and femaleremajn in amplexusevenin the collecringb.g; the eggs
are laid whentlre animalsa.e rcleasedjnto thc basinconlainingfresh,water.
In olhercaseswherethemalcandfenalego oLnof amplexuswh€ndislurbed,
they frequentlyresumethe processof copulationwhenput backin water. Ir
eilher caseslringsof eggscan be collectedas soonaslaid.
Itr Lahore copulatingtoads arc comc acrossduring the rainy season
(JLrlyto early September)soon after a downpour,eslecialiyin the morning.
Thepossibilityofkeepinga collec{ionoflive toadsin the laboratory,ard rhus
avoiditrgthe lrouble of searchingfor toads early in the moning, was also
explored. Unfortunatelythe animals k€pt in the frogeary did not go inro
amplexusand,lius, did nol proviCeanymaterialfcr lh€ pres€ntstr:dy.
In the pairswhich werebroughtto the laboratory.llc enlireprocessof
egglayingwasobserved.To beginwilh themalejumpson 10thebackofrhe
femaleandcalcheshold ofher with thc helpof his fore-arms. The1woremain
in this position for a long tine-lhe exaclduralionbeingdiferenr in ditrerenr
cases. The femalemayjump aboutfron placeto place,carryingthe maleon
her back. Wh€n the femaleis about to lay eggs,rhythmic movementswhich
arefrom infrontbackwardscanbeseenexrernallyonherbelly. Thenshesettles
down at the bottom of thetank, andthis is an indica(ionfor hermareio bring
his hindlimbs togeller to make a "pouch". tn that "pouch" the eggsare
Saiheredand the spermaticfluid is ejac lated overthem by the male. When
no more eggsare coming out of the cloacalopening,the f€maleccmes xp
again10lhe surfaceof water. The matewilhdrawshis hind legsard rh€ egg
stringsarelet loose. The eggsarelaid in this wayjn manyinstalmenls.
As mentionedabovcthe femaleconlinuesto changeher positionsothat
tbe eggslringsg€t wound round the waier planlsand ihe stonesl]jng in the
pond. The egg strirgs are very slicky when laid, and readily slick 10 every
object they comein conract!v;th. Sand parljclesadhereto the egg slrinss
which,Iherefore.becomecomouflagedand areprobablysavedfrom ihe preda-
lors, like crowselc.
As soonas laid, the eggslringswerecollectedin a petri dish and were
transfer.edto a large carlhentrough containingtap-water. Someplanls of
Hy.ltilla wercalsoaddedro lhc tfough 1ocnsurea suppiyof Oxygen. In the
earlhetrvesselthe lempcralurcof wateris usuallyabout 2'C belowthe room
temperalure. This is a distinctadvantag€duringthe hot sumrneraf(ernoon.
CeDerallythe eggsof Anrphibiaare sludiedin a salinesolulion,called
M. S. KHAN

3. V o l . 1 1 , 1 9 6 5 NoRMAL TABLE oF ,,/d ,e/andr,i./,r
Hollfreter'ssolulion, whicfiisalsoknown as lhe .standard culrxrc mediurn.,.
Previouswork in l]ris laboratory by Hashmi and cilani (unpubiisjr€d)has,
howevcr,eslabljshedrlral developmentof road (Bufo tnelanoslios) ^^d ftog
(Randtigtina 1lpi.z') clrn 1afteplace normally in tap-water. The lap-warer
supplicdto the ZoologicalLaboratoryis pumpedou1by a rubc-weltnearthe
college,and is sloredin a reservoir. Its pH wasfoundro bc7.0.
Unforlunately,no devicefb. maintaininga conslanlremperaturewas
available. Thc delelopmert was,therefore!allowcdlo pr.oceedat the room
renrpemtxrc,whichwasrecordedat shorl intervals. It wasnotedthat flucrua-
tions wcrerelatiyelystight.and the entire developmenrcould be regardedas
havingt6ken placeat a lcrnperaturcberween29-32.C.
(r) fikation
At first Bouids fluid. recrifiedspirn cnd 5% fo. ali vere usedas
fixatives,brt it wasseenthat early embryos,particutarlyfrom gasrrutaSrage
to tail bud Stage,wcrc nol preservedadequa!€tyin ihesc fixativesand ihe
materialwasdamagedin shapeand form. Boui s fluiddiluledwith anequat
volumeof lvaLerwasseenio bc a good fixllive. 11wasalso seenthat ji the
eggsaredippedin boiling waterand ltrenpreservedin 3% formalin,they keep
thcir shapeand fornl. So both thesemethodswercemploycdfor frxation.
Il was also noliced that coats of jelly could be rcmoled easilyfrom
thoseeggswhich had beenfixed by immersioninto hot water. Afrer a treaF
ment with boiling waler rhoseeggswerekept in 3% tbrnalin overnight,and
thentnc jelly wasremovcdwjth the help of needlesor finelorccps.
For storage,ai firsta mixturcot cqualquanLiriesof 70% Alcoholard
Glycerinewas used,bul jl was seenthat ernbryosfrom gastrulato iail bud
Stagetendedto be deformcd. So, Llesesragesweresroredin 3% formatin.
The rest of the stageswere,howeler, sloredin 70% Alcohol and clyccrin€.
(c) StudJofrnate al
In the coxrseof the presentstudy only the cxtcrnalnrorphologyof the
vafioussLagcsof dcvelo!menlhasbecntakeninio consjderalion.
Thc obscrlalions havc beenr€cordcdin drawing. All the drawings
havebeenmadewith the help ol canreralucida (Abbe model).
For neasuringthe eggsand early slages,a disceyc-pjccemicrometcr
was employed,while the later embryosweremeasurcdwilh tle lteip of a
mechanicalstage.
The dianeter of the eggsand early cteavagestageswasrccordcdalong
the equalorialplane. Theseslagesare completelyspherical. Eachstagewas
measuredal lcasttwice1()ens!rcagains(anypossibililyofefl.or.

4. M. S. KHAN EIOLOGIA
For advancedstagesthe following lengthsweremeasured:-
(l) Thewholelengthofthe animalfrom anteriortip of thebodyio the
tip of thetail.
(tt) The length fiom anteriortip of body to anus.
(iit The length ffom anusto the tip of the tajl.
As mentiotredaboveno thermostaticcon1rolof the water in whichthe
emb(yosweredevelopingcould be ensured.Consequenlly,the age of various
stagesin hourscould not be regardedas a valid criterioDfor recognizingthe
stagesof a Normal Table. Some observationsregardingthe age of early
developmentalslageswere,however,madeandarerecordedin Table I.
TADLEI
Stage Time Temperature
Oviposition
2-C€ll
Blastula
Formationof Blastopore
Consarictionof Blastopore
Closureof Blastopore
Neural plate
Neural folds
8-8.30am.
9-9.15a.m.
10.00a.m.
11.45-12.00a.m.
2.00-4.00p.m.
6.00-6.10p.m.
6.00p.m.
6.30p.m.
29.5'C
29.8'C
30.8'C
31.2'C
31.85'C
32.0'C
3l.5'C
30.3'C
(d) r'staging"
A refereDceto th€ previousliteratureshowsthat thereis no standardized
method of dividing the €arly oniogenyinto stages. Difierent authors have
adoptedtheir ownschemesor systemswith thercsultthat thereis no guarantee
of the comparabilityof sin larly nurnber€dstagesof differentspecies.
In the courseof the presentstudy an attempt has beenmadeto follow
the paitem ofthe NormaJ'fable of Xenoputlaetir (Daudin)editedby Nieuw-
koop & Faber(1956),itr so far asthe externalcharacterscatrbe usedfor the
purpose.
Otrly the distinct, unmistakableand easily recognizablestages of
developmenthave been trumbered,and full care has beentaketr to avoid
multiplication of stagesdue to variatiotrsin the structures. In all 43 stages
have beenr€cognizedin the developmentol Bufo nelattostictus.lt has not
beenlound feasibleto recognizea pre-metamoDhosisperiod and a period
ofmetamorphosis,becauseall chang€s,howeversuddentheymayappearto be,
are gradualin their final analysis. Il may howeverbe mentionedthat stage
25 is relativelystableand if suitableconditjoDsfor metamorphosisar€ trot
available,the animalmay conlinuein ilis stage.

5. Vol.ll, i96t NORMALTABLEOp Bulohetaaosti.tu, j
THE NORMAL TABLE OF BUFO MELANOST]CT{/S SCHNEIDER
Sttg€ l. Unfertilizedegg. (plate t).
Averagediameter: I.3 mm.
Animal (A.P.) and vesetatpoles(vEC. p.) are djsrinct. The anjmal
hemisphereis deepiypigmented,whitethe vegetalpoteis wjthout pigmenrand
app€aNto be almostwhi1e. The pjgmentexlendsup to midwaybelwe€nlle
€quatorof the eggand the vegeralpole_
Stag€2. Fertilizedegg. (plale t).
Averagediamer€r:1.3mm.
This stagedifrers from stageI in havrng,belw€enthe
vegetalhemrspheres,a crescenlicarcawhich js nejtherbtacknor
intennediate(grey) in colour (CRES.).
animal and
white,but is
Thepigmenrisjusr touchingthe€quatoroflh€ egg,but doesnot extend
beyondit towardsthe vegelalpole.
The vilelline membrane(VlT.) whjch adheredvery ctoselyro rhe esg
at Stagel, becomesrather looseafter ferriljzation and can be distjnguished
readily.
Stage3. 2-Cell stage(Pla1et).
Diameter:1.25mm.to L3 mm.
The fi.st cleavageis vertical and divjdesthe fe.titized egg into two
almostequal blastomeres(BL.). The cleavagefurrow is equallydistinct and
deep throughout its length. The animal pole (A.p.) is deepty ptgnenled
whereasthevegctalpole(VEc. p.) iscrean-coloured. Theviteltinem€mbrane
is clearlyindicated. The endsof rhe blastomeresare rounded. The grey
crescentis visible. Ir is broaderon rhefururedorsalsiderhanon the opposite
side. It is like a pigmenredoblique areaexrendingon rhefuture rtorsalside
betweenthe anirnatpole and rhe middle ofthe egg. The yotk appearsro be
Stagc4, 4 Cell.rage(PlareI).
Diamererr1.25mm.to 1.3mrb.
The secondcleavagejs also vertjcaland lies aimostat right angleslo
tfie firct one. The cleavagefurrowssrarrfrom the animatpolebut fail t; rneet
at the vegetalpole. The furrowspassingrhroughthe two blastomeresdo nor
make an angleof 180' with each orher at ihe animat pole. Th;s cleavase
therefore,resultsin the formation of 4 blastoneresrwo of which are smaltcr
lhan lh€ others. The smallerblaslomeresare cut iolvardsthe future dorsal

6. f-
M. S, KI]AN BIOLOGIA
side. Thjsorientationcanbededucedwith referenceto thegrey crescentarea:
the smallerblastomereshavea smallerpigmentedarea(and, hence,a larger
shareof greycrescentarea)than do tbe largerblastomeres.
Stage5. 8-Cell stas€(Platel)
Diameter:1.25mm io 1.3mm.
The third cleavageis horizontal and is almost equjdistantfrorn the
animalpoleand the equatorof the egg lt resultsin the formation of 4 cap-
like micromeres(MI.) placedon top of 4 nacromeres(ME.)
The two largerblastomeresseenon the (future)ventralsideat Stage4,
cut off micromereswhich are larger and more pigmentedthan thoseof tbe
smdler blastomeres.Thevitellinemembraneis discernible,ihough with some
difrculty.
Stlge 6. 16-Cellstage(Platel).
Diameter:1.25mm 101.3mm.
The 4 micromeresand the 4 megamereshavedividedonceto giveris€
to 8 micromeresand 8 megameres.Cleavagefurrowsmakethei. appearance
simultaneouslyjn micro-andmegameres.Thesefurrows do not passexaclly
through the animalpole but alongbolh of its sidesthus cuttingthe respective
megameresin sucha waythat the porlion whichis cut outsideis smallerthan
that which is cut on ;nner side.
The animal pole is darkerthan th€ vegetalpole which is almostwhile.
The vitellinemembraneis discernibleto someextent The grcycr€scentarea
has becomedividedup into cells,and is not easilyrecognjzable
Stage7. Earlycleavage(Platel).
Diameter:I.168mm.to 1.204mrn.
The distinclionbelweenthe micromeresand megamcresis quite sharp
asthe former are much darkerthan the latter. All the micromeresare not
alike: lhe micromeresnearthe animal pole are smallerthan thoselying close
lo th€ middleofthe egg. Similarlythe megamercsnearthe vegetalpole are
largerthan thoselying nearthe marginalzone. The endsof the blastomeres
aretound. The cleavagefurrows areno1pigmented.The vitellinem€mbraDe
is discernible.
The micro' and hegameresarc not delimitedvery clearlyso that ihere
is a gradualincreasein lhe sizeoflhe blastomeresfrom lhe animalto thevegetal
pole,

7. v o l . l l , 1 9 6 5 NORMAL TABLE OF Btlo n.tarostictut
Stage8. Mid-cleavage(Plate1).
Diametcr:1.25'lrm.to 1.3mm.
This slagecan be djstjnguish€dfrom the previous one by its much
6mallermicromereswhosearea is clearlydelimited,Th3pigmentis confined
only to the micromereswhich occupya region in lh€ animal hemisphere,
atovetheequator. The yolk cellsalsoarc muchsmallerand morcnum€rous
tbdr al Srage7.
The megameresare more compaclly arrangedtLan at the previous
stagewhere th€ furrows werc quite promitretrt. The vitelline membraneis
discernible.
Stdge9. Beginningof gastrulation(Plate l).
Diamet€r:1.25mm.to 1.3mm.
The micromeres(MI ) arevery smalland havespreadover the equalor
ofthe eggtowardsthe vegetalpole. Tfie areasocoveledby the mrcromeresrs
darter. 1le descend;ngborder of the micromeresis lessdark like a band
aroundth€€ggjuslbelowtheequelroiallineofih€ egg
Megameres(ME.) white and large ar€ now lisjble only at the vegelal
poleand in 'ome atedarourdir' Thc wholeareaoccupiedbv megamere"i"
called vegelativefield.
The vegetativefield exte ds up to i1h of the distancebetweenanjmal
pol€ and the middle of the egg.
Strge 10. Drrsal lip of blastopore(Plate l)
Diameter:1.25mm.io 1.4mm.
The micromerescan be seento have spreadstill further towardsthe
vegeialpole. In the vegetalhemisphere-ashort distancebelowthe equator'
there app€arsan agg.egaiionof pigmenrwhich ullimarelymarks the dorsal
lip of ihe blastopore(D.L.B.). The blastoporeis usually situaled midway
betweenanimal pole and equator of th€ egg
Strge 11, Crescentshapcdblastopore(Plate 1)
Diameter:1.288mm.to 1.304mln
At this stagelhe laterallips of the blastopore(L L.B.) havealsocome
into being besidesthe dorsal lip (D.L,B.). Thus the blasloporehas become
horse-shoeshaped;andthe micromefesreachup to it.
The dorsal and lateral sidesof the blastoporeare surroundedby the
micromeresand th€ venlral sideby the megameres.

8. E M. S. KHAN
Stag€t2, Yotk plug slage(Plates 1 & 2).
Diameier:1.288mm-to 1 304mm.
BIOLOGI/
At this stagethe micromereshaveapproachedlhe blasioporefrom ihe
ventral side also, so that a ventral lip of lhe blasloporehas also become
recognizable.Throughthe blastopore(B ) while megamerescan be seenlik€
a white circuiar area calledyolk plug (YK.PG )
On the dorsal side,i.?., anlerior to the dorsallip of blastopore,ther€
exlendsa darkish area,in rh€ niddle of which ihere is much darker streak
running to some distance. This js the rcgior of presumptiveneural plate
(Plare2, N.PL.). Thereis a slightflatteningof the treuralplatercgion'
This stagecould, perhaps,be split into two stagesdependingon the
sizeof the blastoporeand the shapeof the presrmpliven€uralplate This
slage,however,variesconsidcrablyand it wasnot deemedfeasibleto basea
stagemerely on the size of blastopore.
Stage13. Neural plate & Closureof blastopore(Plaie2).
Diameter:1.34Jmm.to 1.486mm.
The lateral lips of the blastoporehave met atrd ihe once rounded
blastoporehasconstrictcdto form a shortdark slit (CL B ), which lies dorsally
ina dark groove (N.G.), yolk plug disappears
The medullaryfolds (N F ) (borderingthe neuralplate)are qurtepro-
minentin the rostral regionbut lessso candallv Thereis furlh€r flattenjng
of the neuralregion on eithersideof lhe grooYe
Stage14. Early n€uralfold or earlvneurulastage(Plate2)'
Diameler:L427mm lo 1496mm
The cerebralpan of the neuralplate bendsdown The neuralfolds
(N.F.) ar€ now distinct all along their length enclosinga groove,calledthe
neuralgroove(N.C.). Th€ neuralfoldsin the cerebralor cranialpart havea
blackisharea at thei inner margins.
The dorsalpartoflhe neurula,te., rhereg;onof ih€ neuralplate'shows
elotrgationin the antero-posteriordjreclion, while the ventral part remalns
of the samedimentions.
On the anlero-venlralsidcthereis a c(escentricblack thin llne formed
by the agg.egationof pigrnentlying transversely,it marksthe posi(ionof the
cementelatrd(CE.CL.). The posilion of blastopor€is stjll recognizablelike
a dark streak.
tt

9. vor.11,1965 NORMAL TABLE OF Bpfonelatusti.tus
Stige 15. Mid neurula stage (Plate 2).
D'rmerer:l.q6Inm.Io 2.1mm
Inthe cerebralregionrh€neuralfoldsarepromineniandshowatendency
towardsmeetitrgin ihe mid-dorsalljne. The anteriorpart of ihe neuralplate
isroundish.
ln ceudalregion the neural folds (N.F.) are quite prominent though
norlo lhe 'ameexlerl a' io ll,ecereb'alregon.
Thedarkishline observedon the antero-ventralside,jn Stage14,which
heraldedthe form?tion of the c€mentglandhasbecomemoreprominenrand
ihick (CE.CL.).
Thereis stiLlfnrther elongationin the antero-posteriordireclionon the
dorsalsideof ihe embryo.
Stag€16, .Latc neurula or neural tube (Plaie 2).
D'ameter:2.0mm.to 2.3 mm.
At this stagethe neural groove has clos€dto form the neural rtrbe
(CL N.T.). The cerebralpart of neuraltub€ is much more dilatcd than the
Ther€is an overallincreascin tbe tengthof tll€ embryoitr the antero-
posteriordir€ction. So the embryois more or lessrod like with borh ends
biunt ard rounded.
The cr€scenticband observedin srage14 & 15 is now ,,V,, shaped.
Theanteriorendsof its limbs arebecomjngihick. This is the cemenr gland
(cE.cL.).
The enrbryoshowsmovementsin the jelly and usesc€mentglandfor
arlachmentwith the egg membrane.
On eirhersideofthe headin medianlateratplane,two slight protuber-
ancesmarkingthe position of the externaigills are to be seen.
Lying in the mid-ventml Iine arterior ro the cementgland, the.e is a
siight depressioncaliedthe slomodaealpir (ST.PT.)which narks the posjtion
of lhe luture moulh.
Thc pronephric resion (PRO.) appears
regionon Ihe dorio-larer/lsrde.
Strge 17. Tail bud stas€ (Plate 3).
Total lengrh
betweenthe belly and head
2.2mm.ro 3.00mm.
2.9mm.
0.1 Dtm.
Leng.hfrominrefrorendlo rle endof be !.
LengthofTail brd (from theeDdof bellylo
lhel.pof lail)

10. M. S. KTIAN BIOLOGIA
l0
On the posleriorilorsalend of the embryoa snall bud likc outgrowth'
the tail bud (T.B.D), makesits appearance Dorsalfin (D F') is onlyslighlly
delimited.
The headregionis mrrked off from the trunk region The headreglon
is more massivethan the generalbody
'lhe v-shapedcerem gland{cE CL r "how
$bich ;' darkl) pigmenLed.dirrdingrhearmt inlo a (omodaealaooacauorr
ridge. The stomoalaealpit (ST PT ) is darklv pigmentedand promrne
The myotomesare clearlvvisible Tlre embryoshowsmuscularmov€-
ments;1hat whentheembryoisshakenit hatchesoui ofrhe eggmembranes'
ii" o"J,i"" "r,r'. "v" (E.) can be narked just belowthe middlepart of th€
0.",i"" " t."* ** ." eacLlaleralsideof theh€ad Theseareasareprgment-
Jiiit" tl" g"n".ur bodv surface The gill areas (G AN) and pronephric
reionsaremoreprom'nenlthanal Sragel6
Th€reis an aggregationof pigmenton the posteriorborderof the belly
markingthe posiliotrof flrtu(eembryo'
Strge 18. Fin formation (Plate 3)
Total length : 3-1 mm to 3 8 mm'
Lengthfrom anieriorrip to the endof bellv: 2'9 mm'
Lengthfrom the endof belLyto thc tip ol
r a i l : 0 2 8 m m .
Dorsal and ventralfins arecl€arlvse€n The do^al fin (D F ) extends
anteriorlyalmostio the level of rhe pronephricreqion
losteri:dv li,le:::
i.lJ-ii'" tip .r,r''" ,"il (T.) towardsthe ventralsideand exrendsup ro the
p""".t"t 0".U". "t,n" Oellvthusformingthe venlralfin (V F ) The fins are
The stomodaealpit (ST PT ) is more promjnent'a grooveextendsfrom
it to rfremiaateoftte V-shapedcementgland(CE GL ) A pair ofdepressions
iu,ut"r,* posirionol oa'lopeningtappedronr}le.anrer:or
to'' l1lilll
sidesoirhe hcad(OLF. Pf ) The'eisa cooriderableirc'easeInthe numDer
"i,"n"--t"i"rn*' ftllO.). The tail bud is dislinct from the bellv The
ugg."gutiooof pigm""t at the sile of the future anusis prominent'
Hatching takes placespontaneouslyat this stage The embryowrig'
sles oui of the egg-membranesbv making powerful movemenlswhich
.are
i".""i"a. tn" .G* Oetweenthehead and bellv This resionis bentalter-
r.i"it'i" in".ttni*o teftsid€ssotheembrvocanswimto some€xtentand
""" "lt*g" iis position whilc lying otr the bottom of the conta'ner
or slicling to its sideswith the help of the cementgland'

11. The areaof preslrmptivegills (G AN ) shor'-san anteior biggerand a
.*,*i", ,n)^f'.t ourgrosth'on rhe mcdianlareral'idesof lhe head rlu'
ffi;; ;r.",.'",itill bLd' lheevepo'iriontr"een in "ragerTismole
-"rt"a. tfr" t.ri" *gion showsprosencephatic'mesencephaiicandmyelence-
phalicregions.
Thestomoilaealcleft isnow lying in a squarishareawhich rs more prc-
.ir*,irt". ;. ift" p."ious stage Theanaiopeningisnotrecognizableclearly'
"i'i",t, ln. *"' ''0" cr $hich-rlte rnr"s will appcarcan be 'een The
r,.,a resronr. 'harpr) nrarkcdoR lrom lh€ rrunkregion Pronephricrcgron
il.""",t,"";;.;';."i rhc rail regionbecome'furrherelonsared rhere is
;,; ;i;";"." in rhr t-unk resion Dorsallln (D F t exrendsLpto Iherevel
;;t;l;; of headand b;rlv Dorsalfin is moreprodnent thanvenlml
fin (v.F.).
Vol. 11,1965 NORMAL TABLE OF "/o attatott'ls
Slage19. Gill bud slage (Plale 3)
Total lenglh
Lenglhfrom anlerior1ipto the tip of
anallube
Lengthfrom the1ipof analiube to the tip
oflail
Stage20. Early gill stage(plate 3)'
Total length
Lengthfrom anteriortip to anus
Lcngthfrom anusto theliP of tail
The first or the anterior most gili (G'
the postcrior one, and has its free margins
po'terioror "econdg'I alsohou paprllae
dcclopedr'ranrhetrpillac of rhe6FI gill
: 3.1 mm.to 35mm.
: 2.98mm.
: 0.51mm.
i 4.0 mm io 4.2mm
: 3 2 0 m m .
: 0.88mm.
BD.) is better developedthan
produced itrto Papilla€. Thc
on ils margin which are less
The areaswhereeyeswould appeararequiteprominentandbulg€oul'
fn" "ronloau"urregion has assumedtdangular form' One angle of th's
triangle continuesas a grooveinto the 1wo arms of ihe cementgland The
two ;rrns of the V-shapedcementgland(CE GL ) becorneseparatefiom each
other becauselhe middle portion' whichioins the two armshasbecomeve'y
inconspicuous.The belly bpcomessharter than in 'he previousstage' blrt
tt'r" tu;l "tongot"r. The posteriorpart of belly is narrow€rlhan the anterior
part. Thefururepositionoflhe anusismarkedbypigmentedgranules AnaL
ol, f,qN.pf.) is also discerniblc. The dorsal fin (DF) is better develoFed
andextendsup to the posteriorborderof lhe head'

12. 12 M. S. KII{N
Stage 21. Gill fiLanlent stage (Plate 4).
Total length
Lengthfrom anterjortip to anus
Lengthfrom anusto tiP oftril
At this slagcthe first or anteriorgill (Gl.) is attachedto the sideoflhe
headat a slighilylowerlevelthanthe secondgill (G2) Posteriorto thesecond
gill thereis a slight protuberancenark;ng the posiiion of the third giu The
first and secondgills havetheir marginsproducedinto fingerlike processes-
theseare the gill filaments. ThefiIStgill haslonggill filaments(Plate8), The
nasalpits(OLF.PT.) cometo possessdisijnct boundarieswhicharepigmenled
The eyeregionbecomesvr'elldemarcat€dand definite. Tbe slomodaealclctt
(ST.CL.)is boundedby four ridges,two alotrgthe antero'late|alsideand two
along the postero-lateralside Theseridgeslie obliquelv to thc transverse
axisof the body. Tire stomodaealdeprcssionis deep€rthatrthai of prevjous
stages.
The two armsofthe cenent gland(CE GL.) havebecomeseparat€and
their endstoward the mouth are swollen The anus(AN.PT.) is indicated
at the .iunctionsof the tail and thc belly. The pronephricregion(PRO) is
prominent. The dorsalfin hasexrendedup to thelevelof the first gill Due
to the lossof pigmentin the superficialregions.the outer wail of lhe head
becomesalmostaransparent.
Stage22, 3-gill stageor tail piec€stage(Platc4).
Totatlength
Lengthfrom anteriortip to anus
Lengthfrom anusto tip oftail
BIOLOGIA
: 4.6mm.io 5.0mm
: 3-5mm.
: l.l mm.
: 5.3mm.to 5.5mm.
: 2.8mm.
: 2.6mm.
The third gill bud which wasobservedin Stage21 has developedinto
a gill, the free marginsof which are producedinto papillaelvhichare smaller
than thoseof thc first and secondgills (Plate8) The eyecups(Plate 4 E C )
are'C'-shaped astheventraisidesarenol v€t beenformed.
The cementgland (CE.GL.) had dividedinto lwo at Stage2l Each
moi€tyof lhe cementglandnow is beadshapedand is placedobliquelybelow
Lhevenro-laleralmargtn'of rbemourh. The moulh{MTH } Ir' assumeda
tmnsverseslitlike shapeat this stage. The formalion of intestinebegins at
this stage. At theanterioror rosttaletrdofth€ massof yolk cellsthercappears
a (ran:!ersegrooe$hichiscalledlhe roslrallurrottI ROSFI I Therersal'o
a caudalfurrow (CA.FU ) on the righl sidein the volk mass Posteriorlvthe
belly tapersinto an anal tube (AN-T.) whichopensat atrus(AN ) Theanus
issituat€dat thejunctionofthe ventralfin withthe be1lv. Therail haselongated
grearl).thebell)andlheLailxrealmostolequallenglhlbelarl6nsdreJlso

13. V o l . 1 1 ,1 9 6 5 NOP,M{L TAALE OF Ria helanosti.trs l 3
: 5.5mm.to 6.00Inln.
: 2.8mrn.
: 3.2nm.
: 6.0mm.to 6.8mm.
: 3.0mm.
: 3,8mm.
greallydev€loped.The porrion of rhe skin exrendingfrom the posteriorparl
of lhe belly up 10the anusi5 ca ed rait piece(T.pE.). ln rhjs stagelhe tail
pieceis formed. The pronephricregion(pRO.) is stiil prominent. Thehead
has becomemore transparentttran jn the previous sla8e_ A Ialeral line
(LAT.L|.) can be recognizedas a dark tine running on either sjdefron the
pronephricregionio the posreior endof the raij.
Stage23. Opercularfold stagc(plate4).
Tolal length
Lengtnfrom antefiorendio anus
Lengllrliom anusto rip oftail
At this sragefrom the regionof the headjust in fronLof ltre first gjll
on erthersideariserwo foldlike exrensionsof the body wall caled oFercuiar
1blds. (OP.F.). They progressivetyeiongatetorvardsrhe posterior region
coveringthe gills. The eyo cup (E) ls conplctcly circutar,ltrough stjl the
ventralportionofit isrhin. Theorbirandnasatopenings(NA.Op.)areclearly
r€cogriz:blc. The mourh is surroundcdaboveand belowby flesty areasor
lips ,hich unile togctherto forn1rhe oral cup. The mouth is boundedpre_
orally by a cfescedic lrorny dark brown ridse. On rhe posr_oralside;lso
the.eis a horny ridge. Both theseridgcsform rhebeak(BE.)_ The margins
of theseridgesarenot providedwirh rheteerhlike projectionsasyer. Dental
formula at this stageis l/0. The pre-oralfleshytip hasone compiererow of
denticlesandthepost-orallip hasnot develoleddenriclesyet.
Thc cementgland (CE.CL.) has becomcgrearlyrcducedard thc rcm_
nantsofits two bead-shapcdmoietiesare losing their dark colour. Fourlh
gill bud appcarsjust behindthe third gilt. Pronephricregionwhichwasquile
proninent in previousshgeshasbecomeindisringuishablehere(ptare8). The
dem:rcltion b3tweenthe headand belly has becomefurlher reducedthan aL
Stage22. The regionof headin front of gills showsconsiderabtelarcml ex-
pansion,so that the head appean to be darso-ventrallycompressed.The
regionof belly howeveris round ventrally. The hecdand beltyhavebecome
almosl transparentso that many jnlernal structlrrescan be sccn. Metano-
phoresarc presenlon ihe dorsalsid€of the head,beUl and tail. The lateral
l;ne(LAT.Ll.) is clcarlyindicatedalongtle lileral sidesof the rrrnk andtajl of
the tadpole.
Stege2,r. Lrle opercularstage(Pia1e5).
Total length
Lengthfrom anter;orlip to anus
Lengthfrom anus!o 1ipof rail

14. M. S. KIIAN
At this stagethe operculum(OP F') ahnosttouchesthe bellv on the
right sideand fusewith ii Onlv the gill filaments(G F ) of the right gillsare
to-be seen. (For gills seePlale 8) The evesare b€tterformed than inlhc
previousstagebut still the ventralsideof the evesis thin'
The pre-oral and post-orulridges of beak (BE) are provided with
sharplyserratedma.gins. The pr€-oralileshvlip bearsa completeand ur-
jnterruptearow of dc icles. The post-oralfleslrylip hastwo completerows
"i a*ii"r"'. Thes€denticlestake the form of sharp,spinelike, dark brown
processes,usedin scrappingtheplantor fesh on whichthe tadpolesfeed The
dentalformulais 1, l*1/2. The wholestructureconstitut€sa sucker(S)'
ii" "".".t gr"a (dE.cI-.) is progressiveivin theprocessof reduction lt is
now rcprcsentedby merely small crescenlicdark spois'
Thereis further elongalionin the lengthof lhe body Tail exceedsthe
lengrhof the trunk. The distinciion belweenbelly and headis growrngless
and less. Thc intestinatcoil (INT.CO) can be seenthrough the transparetrl
wall of belly. The anal tube and the cloacallail piecealsobecomefurth€r
elongated.
slag€(Plaiess &8)
BIOLOCIA
Stsge25. Spiraclestageor Hind limb bud
Total length
Lenglhfrom anteriorljP to anus
Lenglhfrom anusto theliP of tail
: 6.8mm.to 7.0mm.
: 3.2mm.
: 3.6mm.
The opercularfold has fused completelvwilh the bodv wall on the
right siile,buionthe left sidethe fusion is inconpleteso that a spi'acle(SP')
is'formed rhrough which the gill filaments(G'F ) prolrlrdeout The sprracle
,s situatedon ventrolalerai sideofthe bodyin thc regionwhereih€ iirst outer
intestinalcoil tums inwardsto the mid-venlralline'
The pre-oral fleshvlip has an outer compl€terow of denliclesand a
secondintetruptedrow. Thepost-oraltip hasgot 3 completerowsofdenticles'
the dentalformulabeing1,1+ l/3. Thedenliclesar€long,spinelike anddark-
brown. on the lateral sid€sof th€ lips there appearpapillaeon eachside'
No$,theoral disciscall€da sucker(S). Mouth (MTH ) occupiessub-lermrnal
posirion. The beak(BE.) is well-formed(Plate8) The two moictiesof the
;ement gland(CE.GL ) havc becomereducedlnerely10 aggregationsof dark
granules. Thesespotsli€ nearthe two endsof mouth on the anterrorventro'
laieral sidesof the bodY.
Smrli hind linb-b ds(H. L. BD.) maketheir appeatanceat thej ncnon
of rhe belly and ihe lail on the dorso-lateralside of the anal tube The
inrc,rinalspiraltlNT.CO ) i' fullv formedal rhissraPe Faecalmatrercan
beeen,o comeourlrom lheandltubellroughanuc(AN l in man)pecimeo

15. V o l . l l , 1 9 6 5 NORMAL TABLE OF rrfo melaaostictts l 5
at this stage. Melanophofcshavedevelopedin Iargenumberson llre dorsal
and lateralaspedsol thc body. ln the belly region,howevcr,melanophercs
arepresenton ihc vcnllo,lateralparr a1so. Ventruilythe belly regionis flat.
Stage26. Lenglirofhind limb bud lesslhanits breadlt. (plales5 & 6).
Tbeoperculunlis complele.The spimcularopen;ng(Sp.)hasno gjlt
iilamcnlscomingout of n. T[e hindlimb buds(H.L.BD.)arcstillveryrmall,
and areshorterin lengththan in breadlh. Tlre dorsalsurfaceofthe tadpolcs
is almostbiackdue to the largen!nber of melanophorcs.Thc melrnophores
ar€seento spreadalongtheouierinlestinalcoiJ. Thc aggrcgationofpisncnt,
markingthe position of the moictiesof cementglitnd arc fainlly represenlcd.
Thereis further increasein the lenglh of Lheradpole. Other chamctersare
the sameas at Stagc25.
Strge27. Length of hind limb bud greaterthan breadth(Platc6).
Total lenglh
Lengthfrom antcriorlip to anus
Lenglhliom anus1orip oflail
Total length
Lengthfrornanleriorlip to anus
Lenglhlrom anusto tip oftail
Stag€28. Lenglhof hind lilnb bud lj limes
Lenglhfromanterioriip to anus
Lenglhfromanusio tip oftail
: 7.1mn.
: 3.1mm.
: 4.0mm.
: 6.95mm.107.2nln.
: 3.7lnm.
: 3.68mm.
The h'nd limb blldsat thissragearelongcrthanbraad. Tlerc is stiil
furtheraggregalionof the melanothoreson lh€ dorsil sideaswellas along
theoutcfborde.oflheoulercoiloflhcintenine- Olhercharaclersar€tlc sirme
as ai SlaSe25.
its breadth(Pla1e6).
: 3.6rnn.
: 3.7mm.
Tolallength : 7.0nnn.to 7.4.mm.
The lenglh of the hind limb bud lA times irs breadrh. The free end
oI lhc bud is roundingo$.
Srage29, The length of hind limb bud l: rimesi1sbreadth.(Plare6).
Lengthfrom anreriortip 10anus : 3.8mm.
Lcngthfrom anusto 1ipoftail : 4.0mrn.
Totallenglh : 7.1mm.to 7.5.mn,
Thelenglhof rhehindlinb budis lg tjmesirsbreadth.11sljp js broad
and is flalteningup.

16. Iii'
1 6 M, S, I(HAN BIOLOGIA
Strse 30. Leng'h of hind limb 2' times its b:eadth(Plate6).
Lenglhfrom anteriortip io anus
Lengihfiom anusto lip of tail
Total lenglh
The hind limb bud js 2+iimeslongerthan it is broad lt js round near
i1sbase(stump),but its greaterpart is flatlenedfrom sideto side(paddle)'
The freeend is no longerround but somewhatnarrow and conicalandfore_
shadowsthe 4th toe. The constriclionbetwcenpaddleand sturnpmarksrhe
posilionof the ankle.
Stage31. 51h1oestage(Plate 6).
Lenglhfrom anterior1jPto anus
Lenglhfrom anus10tiP oflail
Totallenglh
The stumpof thc hind iimb becomessomewhatelongated.Al tbe same
time its breadthbecomesasgr€atasthat ofthe paddleof the hind lirnb. The
nriddler€gion(Ankle constrictjon)is howerer narrower. On the dorsal(or
posFaxial)aspecrof the fool (paddle)there appeaf an indenlationmarkjng
the 5th 1oe. Melanophoresh,tYeappearedon the dorsal side of the limb
stump. They extcnd up to its middle.
Stag€32, 3rd toe stage(PI4te 6).
L€ngthfrom anteriortip to anus
Lenglhfrom anusto tip of tail
Total length
Tnthe hind limb the paddie-likefooi (FT.) is clearlymarkedoff from
the slump. The Iatter is slightly longertban the paddle. Ofl ihe lorxeror
pr€-axialsideof the 4th toe anotherindentalionappearsin lhe treemargin of
rhe paddleand marksthe posilionof lhe 3.d loe Somemelanophoresapp€ar
on the 4th 1oe.
: 3.0mm.
: 4.1 mfi.
: ?.3 mm.10 7.8 mm.
: 3.2mm.
: 4.0mm.
: ?.6mm.to 7.8mm.
: 3.2mm.
: 4.8mm.
: 8.3mm.to 8.5mm.
: 3.4mm.
: 5.6mm.
: 8.9mm.to 9.3 mm.
Stage33, 2nd toe staee(Plate 6).
Lengthfrom anterior1ipto anus
Lenglhfrom anusio lip of tail
Total length
Belowthe indentntionof the 3rd tce (on pfe-axialside)on the paddle-
like foot ih€reappearsanolber;ndenlationmarkinglh€ positionofthe 2ndtoe.
The distanccbetwe€nthe 51h,4th and 3rd loesincreases.The areain between
becomesmuchthinner. Thebasalportion ofthe stumpis broaderthantlrerest
of the limb. Thereare aggregalicnsof melanophores,one on the dorsalsid€

17. vor.11,I96t NOI.MAL -TABLE
OFBuJohelano tctu,
ol rheba,alparrof thesrump.fore,hadowitrerherntgh:anorheraggregarionbelweenthe thigh and tbe paddlemarkingrhe p."iti.n of the sta.k-:";?;;;
:1,::^""--:l*:,t.'
onthepaddternarkinftreo*i,i", .r,r,. r".i
,rra"L,""-
pnoresalso appearbetween4th and 3rd tocs.
Stag€34. lsr roe srage(plare 6).
Lenglhfrom anterjorrip to anus
Lengrhfrom anusfo tjp oftait
Total lenglh
Stage36. Full anatrubesrage(ptares6 & 7),
Lenglh from anleriortjp ro anus
Lenglhfrom anusro iip oftait
Total tengih
On the pre-axial or ventralsjde of the paddlelike foot apDearsanotherindentationmarkinsrle posjrjonofrhe fi*t to", Ua"* rr,. ina"rlir-i.rt
for the secondroe. The various dieits (2,5) ar€ b"t,". d"_"..;;";--;;;
separatedfrom eachother than at Slage33_
..
Tl:_l_hleeparls of rhe Ieg,ririghor stylopodiurn(TH.) strankor zeuso-podium_(sH.)andthefoot or autopodiumirr.i ari "/",r,
'j".",-#1.
:i: il!.f'
*lt
..'-_lwardsarrheknee;ornt. .rhereis anincreasein reptgmentationof the thigh, shankand dorsatsideof foor. Tlr" ,r"" b;;;;the secondand third toeslas alsobeenjnvadedby melanophores.
Sfage35. Fuil rail pieceslage(ptates6 & 7).
Lengthfrom antedorrip to anus : 6.5mm.
Lengthfrom anusro tip oftail : 8.5mm.
Totaltenglh
: 1.2Cm,to 1.5Cm.
_
The.hindtimbs becomefurlher elongated. Ttr€und€rsid€sof the feerare appliedto rhe ventral fn on either side. ff,. n"., ar. l*, ""r*".i".Th€tail piece(T.pE.) extendsup ro rhe ankle. All rhetoesareseparal fr-on
.one
another,the fouflh toe beingthe longest. O, rh" d".r"t,id;;;;;;;;
lh€re-arelhree aggregalionsof oelanophoresshowire rh" p*ili;;;;;;";:
: 7.0mn.
: 7.5mm.
i 1.4cm.to 1.6cm.
lr*#l:ft1it;$r-fu::.l:**]lf#'rJ:**i
;l'r"'.'##*:ilffii::,;:r*ffil{itr{^
: 4.8mm.
: 5.9mm.
: 9.4mm. to 10.3mm,

18. lE
M s KHAN BIoLoGIA
s:deof lhe lirnb. Mel.nothorescan bc eenagarcgalcdin a circleround
i; ;;'.;;.' orrneine+na|oop],::i',';Jli;:,i,T';l;:T'i,':;
r)reselton the dorsalfin also The lengl
i;;;;;;,i';;;i;t",.'nk (tom anlerior end to the anuo are armostequar'
Stage3?. Mid anai tlrbe stage(Plates6 & 7)
il:ili::il::i"Ji:'i'Jiitri",il#,",,".Total lenglh
rheanartubc""*d':' i.'lT'"::i:i"[
-ft
:dL::tH]
angleof 60" with thc longitldinal ax's oI
is in the Processot reducrion'
Thetoesarelurnedslishrrvourwards'anclonlvthet"'lt"ltll
1"'9:l
."."*,'uooi'"0roihesirreorrh€.veni:l,i:
"J*":i*t :i.H:"il'il
:::I.fi:[:"#"il";::'1'J:lT"'5Ji'[],',l.,'#;" 1""0""''"'"""0n*"'
ares€enon the fins The mclanopborcson thc dorsalsideof nasalopenrngs
are spreadingdownwaros
Stnge38. Disappearanceof anal tube slage(Plales6 & 7)
:::flli::l:li"Jl'JiIii,TJ',i111,",,".Total Iength
Tfreanattute hasdisappeared The tail piece(T PE ) is very-narrortr-
ft:l'ji;lffi"H*'l.'i',ffj'i|r"1.}*",;i..ffi$qt+:J:+i**rimb atium. rhe cveshowsa whitish-i:;;;;;-;";"
of ?5. with the
1heposition ol the lens The hind l€gs
*J,''ffiHt/ffi*'*J:;;:"::;"":n::'";';:'l:''li:i'1l:
Ii"ii*''?o'*i'"".*,1"::"'::":"1"i;:il"l"i"ii.?lll
it hasbeencoleredoterby thc abdomrn
entralparrof lhe rrunl becomc:lighllvopaque
Stage3g Left fore-limb stage(Plales6 & ?)
Lcnglhfrom anteriortip to anus : 6 0 mm'
i"n!rt, f,olrr oou' to tip or tuit : 7 0 mm.
Totallenglh
: 135 cm to 145 cm
Th€ left forelimb (LFT H ) has comeout ihrough the spiracle(SP)'
It is smallaod bent downwardso thal whenlhe tadpoleis on its ventralside

19. Vol-I l, 1965 NoRMAL TLBLE OF Btio nela"ostktus
ils palmarsideis towardsthe ground. The fingersand armsarccoveredovcr
by melanophores.Thc tail picce (T.PE.) has becone gr€a1lyreduc€d. The
abdominalmusclescoverthe inleslire complereiy. The skin of ihe body is
translu€entoropaque.Onsomeindividuaisguanophoresmakethejrappearance.
Malatarsalportionof the hindlirnb is still zrpplied1oth€ venlralfn.
Thethighs(femuro do no1corneout at right angleto the iongitudinalaxisof
thebodybt( makeanangleof75'with it asin Slage38.
Stage40. Conipleleforelimb stage(Plales6 & 7).
) 9
The right forclimb hasbrok€ntlrroughthc limb atrium. Thigls make
a rigbt anglewith thelongiiudinalaxisof the body. Thereis completelossof
denljclesand the lorny beak. Thc mo lh (MTH.) is slit like. lt exlends
backwardsup 10a poinr lying midwaybelweenthe nasalopeningand lhe eye
on eilher sidc. Thc eyesprotrude to solneextent. The lail piecc has dis.
appeared. On the dorsalsidean areapossessinglhe characlcrisiicaduh skin
can be recogDizcd.
Stage41, Bcginningof regrcssionof tail (Plates6 & 7).
Lengthfrom antedortip 1oanus : 5.0mm.
Lengthfrom anusto iip of tiil : 2.5mm.
Totallengrh i 7.0mm to 8.00mm.
The lail has consjderablybeenreduced. Tbe dorsal and venlral fins
a.c rudimentary- The co ersof the moulh approachthe anteriorcnd of lhc
cycs. The head region of the body is more massjrelhan the belly part-
Nasaiopcningscometo lie at the lip of the snout. The body is rendered
c.mpl€reryopaque.
Eyesprotrude to a grealercxlent tharl at Slage40. Thc adult skin
areahasspreadconsjderablyon thc dorsalsideof headandthetrunk.
Staee42. St mpy tail srage(Pla1e6).
Lengthfrom anterjortip to anus
Lengthfromanusto tip oftail
Tolallengrh
Len$hlrom anleriortip to inus
Lengrhffom anusto tip of rail
Tolallcngth
: 5 . l m m .
: 7.0mn.
: 1.2cm.101.35cm.
: 4.5mm.
: 0.7mm,
: 5.2mm.to 6.0mm.
Thegapeof moulh wjdeDsup, sothai thc corncrsoflhe morth lie a1 lhe
levelof middleof eye. Thetail jsljleastump. Thedorsalandvcntralfins
carnot be madeout. The tip oflhe tail is crumpledand black. The adult
skin is recognizablcon the whole of lhe body.

20. M. S. KTIAN BIOLOGIA
20
Stage43. Tail lessstage(Plate6).
Lengthfrom ant€riortip to anus : 6 5 rnm'
I-engthofiail: tail is completelyabsorbed'
Tolallenglh :65mm'
The tail hascompletelvbeenabsorbed The gapeofrhe mouthwidens
up so that it ext€nilsbackwar'lsup lo the posteriorend of the eye' or even
beyondit.
The forelimbs and hind-limbs becomefu her elongated The eyes
protrudein the characteristicfashion. The skin showswarts'
COMPARISON WITH NORMAL TABLES OF OIHER AMPHIBIA'
An atlempthasbeenmadcin this part to comparetheNormal Tableof
Bufu melanosticius,as describedabove,with the Normal Tablesof someother
Ampbibians.A similarattempthasbeenmadebv Nieltwkoop& Faber(1956)
wlro lave compareOthe Normal Table of Xenopuslaelriswilh that of olher
l-pfr;U"n.; and by Rossi(1958),who has comparedihe Normal Tabl€ ol
nolo tu|o *it rn"t of nanapipins. brllre prcsentcompadsonhelp has b€en
tuteo ir""ly from the comparativechart prepatedby Nieuwkoop & Faber'
In the above-describedNormal Tabte of Bufo nelanostictusthe rr''
ferlilizedegghasbeentaken asSlage1. SimilarlvDel Conte & sirlh (1952)
i" nrlo *i,o*^; weisz (1945)it Xenopustuevis;Shumwav(1940)in 'Rafla
pipicnti Pallirter & MooreI laJ7,in Ranasflwtha: ald Milller(lalg) in Razl
pipiensrccolni,7theurferrilrTedeggaslhefirstSIUge HoweterRo'ri (1q59)
'ti
nt" Un: Adler (1901) ttt Bufo rulsaist Eakin (1946) in Hlla rcci a;
Nieuwkoop & Faber (1956);n Xrnottus1d?rir;Callien & Houillotr (1951)in
Discoglassuspictus;Kopsch(1952)n Ranofutco; Cambarand Marrol(1954)
in RaiadalmatinaandMoser(1950)in .RandlPmpotaia; do r,ot recogntzetbe
unfertilized€ggas a Stage. Th€reis somcthing10be saidfor this view also'
becaus€,afte;all, the unferlilizedeggc,nnot possiblvberegardedasa develop-
mental stage. Ifl the presentsludy the first view has beenadopledt€cause
the femaleextrudesunferlilized eggswhich djtrer markedlv from fertiliz€d
eggs. In order to facil;tate d€scrjptionand idenlificalionof stages'th€ un-
fe ilized eggmust be taken inio accoun'
The fertiljzedegg(Slage2) hasnot beenrecognizedas a dislinct slage
by workers on sufo iitg.ris, HYta regila, Ranalus&' Ranadalnatina and
Ranatempotafia. As regardstheNormal Tablesof BuIo |ulgaris,Hlla rcgilla'
and'Ranitempotaria,workhasbeendoneonly on advancedslagesofdevelop_

21. Yol,11,1965 NORMAL TABLE oF Bufo ,neta@stictus
In somecases,€.8.in the NorrnalTableof X"rrlrr Zdertr (NieuvkooD
& Faber.l950rrome 'half-srages"
haveatsobeenrecognised.ThusSraeei
of Ihepre5enl,ludyir reprerenredi0 rheirTjrbteby Srageso! rnd ?. Thishas
beendone by relying on very fine characte.ssomeof which are not readitv
visible.Similarl)for Srage|| {Crescenr-shapedbtasropore)of rhe presenr
study, Nieuwkoop & Faber recognjzetwo Stagesand name rhem as
Cresc€ntshapedblastopore(Stage 10+) and horse-shoeshapedblasiopore
tSlrgeI lr. for nid-neu'ulaSlage( I)t of Buh mctanosti,us rhe,eaurhors
recognize3 Stages,yir., No. l5-I8 jn Xmoput laefis dependingupon minute
changesin the neuralfolds. Suchdisrinctionsare not feaslbtein Bufo netano_
rliclrr. A Normal Table doeslosesomeof iis uselulnessif it invotvesthe
recognjlionof obscur€cha.acte$.
It is not €asyto comparetheNormal Tablesofdifferent speciesbecause
any pa icular organ may appearat one stagein one speciesand at quire a
differentstagein anotier species.For exampieopercularfolds appear at
Stage23 in BuIo helon rrrclrr but ar Srage42 in X€rolrr /deljr (Nieuwkoop
& Faber, 1956). Similarly hind'limb buds appear when opercutarfold is
coveringthe gills at Stage46 in Xenopuslaevis,butiL Bto metanostictusrhey
appearat Stage25 whenthe gill filamenlsare10be seencoming our of ihe
spiracl€and the operculumis compicre. The hind limb bfis in BuIo bufo
appearat Stage23(Rossi1959). Thiss.ageof_AuJobufotscomparabjeirralmoit
all chamctersto Slage23 of Bufo nelanastictusexcept that the latter has
not yet acquiredthe hind ljmb buds. Taylor & Kollros (t946) recordrhe
appearanc€ofthe hindlimb b]uds)n Rant pipienrat a sragewhentheoperculum
is compleleand thef€ is no traceof gill filamentscomingout of the spiracle.
This Stagecorrespondsro StaAe26 of Btfo nelanosti.tu|.
Thedevelopm€niofhind limb lakesplacegraduallyinBulanetatlostictus-
The appearanceof toes is also systemalic. Firstly fhe foor stump becom€s
paddtelike (Stage30). The lip of ihis paddleforms the 4th toe. Then on
its post-axialside appearsthe 51htoe (Stage31). Then on pre-axialside
appearthe 3rd,2nd and lst toesrcspectjvely,characlerizingthe Slages32,33
& 34. The samesequencehas beeD noted by Taylor & Kollros (1946)in
Ranapipiens,and by Rossi(1959)in Btfo bufo,markinsrhe StagesVI, VtI,
VllI, lX, X and SlagesIII, lV. V, VI, VIT r€slecriv€ly. Howeverin X?raprr
Ia?].rrNieuwkoop & Faber (1956)recognizeStage53markingth€appearance
ofthe4th and 5th1oe;andStage54 whenall the5loes arerepresented.11
meansthat a crilerionwhichhelpedin recognizingthe difierentstagesof Bir.a
melanostictuswas uselessin Xenoptlslaeris.Most probablyjD the latter the
toesdo not appearone after anotherolherwiseNi€uwkoop& Faber(1956)
would surelyhavemadeuseof this criterion.
2 l

22. 22 M. S.KHAN BIOLOGIA
ln RanapipiensTaylor & Kollros (1946)based7 slages(XI-XVII) on
suchapparenllyinsignificanicharactersasthe apFearanceof melanophoreson
tle toesor the lenglhoflhe toesandth€ distancebetweenthem. Rossi(1959)
however,dcesnot recogniz€suchsiagesin 3rlo ,r/a, and hasconpressedthe
above-mentioned7 stag€sof Ranapipiens;^to only2 (vIl &VlIl);n BuIobtlfo.
ln Bufo nelanostictuswheretoes afe short, the 7 stagesrecognjzedin -Rdta
pi,i?rr correspondIo one 'rdge(Slagel5).
The nexr3 stages(36,37 and 38 in Bufo melarrrl/crr.rarebasedupon
th€lengthofthe analtube. Similary slages(IX, X &Xl) havebe€nrecognized
by Rossj (1959)in nufo bufo. Taylor & Koll.os (1946) recognizedtwo
stagesXVII-XVIII, on the basjsof the tail piece(anal tube)in -Ra'ap?terr.
ln X"nop"r /a€r,sNieuwkoop & Fab€r(1956)do not baseany stageupon
anal iube, but basethe corresJrondingstagesupon the fore limb. In 3r/o
melanostictusI^dpoteswhich arehcavilypjgmenledand in whichthe develop-
menlof tbeforeJinb lakesplacein lh€ limb atfinm,it is not possibleto use
the forelimb for recognizingstages.
In Bufo melanostictus,^nd in B!/a b,/o (Rossi1959)the left arm comes
out of the spiraclemarkingStage39in tbe formef and StageXII in the latter.
Taylor & Koltros (1946)speak of the app€aranceof on€ or both of the
forelegs wbenthey recordthis eventin StageXX in Ranapipiens. ln Bufo
melanostictusand Bufa bufu bolh the forelimbs ar€out at Stage40andXIII
Thenext3slages(Stages4l , 42& 43in Bufo nelarorlicl,r arerecognjzed
on the basisof differenl slagesof lhe regrcssionol the t^il. ln Bu:t'obuJo,
Rossi.(1959)r€cordsonly 1$'o slages(XM XV) on this basjs. In B"/o
melanostiduslhe pracessesof increasein the gale oflhe mouthandregression
of lail go on sideby sidedurins Stases40-43. Nieuwkoop& Faber(19s6)
recordsthcgapingof mouth acconpaniedby redrclion in lhe lengthof rail in
Stages62-64. However,a1 Stage65 th€ mouth has beenformed bu1tajt js
stjll lher€. So the next rwo stages(6{) & 66) are basedon lhe lail. In natr
1?ikrdthe regressionofthe tail (StagesXXIII-XXV) takesplaceafrerlhemouth
hasbeenformedfully (StagesXXI & XXII). Henceihe toadlingcomesinro
beingin Bufo nelanonidus at Slage43, in BuJobufo at SrageXV, and in
Xenopuslaeris at Stage66. In Rana pipiensthe frogling is formedar Srage
XXV.
From the above accountit js clear that the developmentof Br/o
,?p/auarllclrrresemblesverycloselywilh that ofolher anuranAmphibia. The
differencesin the number of stagesand method of sragingjn different
specjesjs due primarily to the facrthat the variousorgansor partsoforgans

23. Yol.10,1964 NORMAL TABLE oF Alh heltuotti.tus 23
do not arisein preciselythe samesequenc€.Suchdiffererces,however,are
very minor, and it can safety be asse(€d that the epigeneticsof Brr.o
melanostictusis ess€ntiallythe sameas in other ranids and bufonids.
SUMMARY
l. Normal developmentof thc common toad Buh mela ostictlts
Schneiderhasb€enstud;ed,from theeggto thestagewhendefinitivebodyform
has been acquired.
2. Th€ ontogenyhas beendivided up inlo stages. For this purpose
only the exterml morphologicalc.iterja bave beenused.
3. The stagesin the Normal Table ol Bufo melanostictus ha.r'ebee
compared1{iththoseof a Iargenumberof toadsand frogs.
4. The Normal Table ol Bulo melanoslictusis seento r€sembleth€
NormaLTablesofother Anura in all essentialrespecls.

24. M- S- KHAN24 BIOLOGIA
'a
E S*^ .== a
; F*Q F.3 *3
i t : : s l F - lE , j ; : _ :
sr SE;Ft iE
i ;
s ! ?
Nameof rie Stages
Unterrilizedegg
Fertilizedegg
2-celistage
4-celJstage
8-cellstage
I6-cellstage
Early cleavag€slage
Mid cleavag€slage
Beginningof gastrularionslage
Dorsallip ofblasroporestage
Crescent-shapedbtastopore
Yolk plug stage
Neuralplatestage
Early reural fold siage
Mid neurulastage
Late nerrula srage
Tail bud slage
Fin formalion slage
Cill bod stage
Eariygill stage
Cill61a11161151u*"
3-gillstage
I , .
2 1
4 3
6 5
7 6
9 8
r 0 9
l l 1 0
1 2 l l
13 12
1 4 1 3
15 14
1 6 1 5
1 7 t 6
l8 17,18
1 9 1 9
20 20
I
2
3
4
5
7
8
9
t 0
] I
l 2
1 3
I4
1 5
t 6
t 7
1 8
l 9
20
2 l
22
I
2
3 )
4 l
5
6
7
8,9
1 5
1 6
1 7
l 8
l 9
20
2 t. 2 3
l

25. Vol. 11, 1965 NORMAL TABLE OF ,r/o fr€lar,rri.rb
F:;;:irEgSE€Eg
3 : 3
. r c d . " ^ E ; ; a a E . . .
Si"E*t 5:?[iE ]i E"$
sE3ssE€iSa;sf5!€!
t
2
3
5
6+,'l
8
9
10,10+
101,l l
12,12,
l 3
14
l5:18
19,20
25-33
34
37-40
4l
I
4 3
s 4
7 6
8 7
9 . .
1 0 8
9
" t , ^ , " 1
l3l I
14 t3
.. 14,15
1 5 . .
1 6 1 6
l7 17-19
18 20
19 21
20 22-25
2t 26-2a
4
5 7-10
6 1l_14
' 1 5
'7
t6
8 1 7
9 1 8
10 t9
lt 20-22
1 2 . .
t3 23
t4 24J6
t5 27-29
I
2
3
4
5
6
,7
8
9
10
u
t2
1 3
t4
15
t 6
1'l
1 8
1 9
20
2 1
22
2-4
I
2
3
4
5
6
7
8
9
l0
u
12
t4
l 5
r 6
t7
l8
l 9
20
I
2
3
4
5
6
I
2
3
4
5
6
7
8
9
IO
l t
, 2
l3
t4
l 5
l6
1 7
1 8
19
20
2l
22

26. 26 M. S K}IAN BIOLOGIA
= d
.:*
F!l
'6
i S*^ .r.
.; isa &EF i 6 - ! "
s " . i : i
FES8,F3
: E
i i
s^i
i "
Name of the Stage
Opercularlold stage
Late opercularstage
Spiraclestage(hindlimb bud stage)
Lengthofhindlimb bud lessthan
its breadth
Lengthof hindlimb bud greater
than breadth
Lengthof hindlimb 1+tjmesi1s
breadth
Lengthof hindlimb 1: timesits
breadth
Lenglhof hiod-lirDb2] timesiLs
oreaoth
5thtoe stage
3rd toe stag€
2nd toe siage
lst toe stage
Full tail piecestage
Full analtube stag€
Mid analtube stag€
Disappearanceof analtubestage.
Left forelimb stage
Completefore-limbstage
BeginDingof regressionof tail
Stumpytail
Disappearanceof tail
23
25
23 l0
l 1
12
13
24
25
23
24
25
t
II
III
IV
VI
YII, VIII
XI
xtI
XIII
xtv
2'1
28
29
30
3 1
33
34
35
3'1
38
39
40
4l
4 2 - .
43 XV

27. Vol. 11, 1955 NoRMAL TABLE OF Balo ,ketanosti.trt 27
= - E ; * a i F a ! ^
sEt:; 1;E c:S"3Si S:rgaS* tF! *i; €,t l ii
$€xffselacss:isgs'i€s;
! €
s?F
$:-s
*ad
43-45
4648
1 1
1 2
l9
20 42
25(5);I 18
(r&K)
II
29
30
30'31
i u I
32,33J I
3440 7-10
2l
22
49
50
31,32
4l
III
IV51,52
45,55
56
51
VI
vIt
VIII
IX
2I
22 43
44
45
x . . 4 6
xl-xylt 24,25 41-50
58
62
64
XVIII
XIX
XXI
27
51
xxtl 28 52
xxrll- 29,30 53,54
XXV
65,66

28. 21NORMAL TABLE OF ?,{o netanosidusVol,11,1965
: "
: d d ^ = r i * : q f . i a t a . .
j?3 -rr;:; {ii ;1!:i $q !:B la3:_: 3r i!: i:;- Si {:6 s; }eu' t;
iz; St:s;S!f8S,?e:!S€cEjc=
2 l
22
23
2330'31
i ^ |
:2,::I i
34 40 1-10
2342 .. 29
4345 . . 30
46-48 22,23 31,32
16
t'7
l 8
l t
12
l 3
t 9
20 42
III
IV
25(5);I
( T & K )
] I4t)
50
43
45
53
45,5s
56
51
VI
VII
VI]I
IX
2 I
22
23
x . . 4 6
xt-xvrl 24,2s 4"r-50
J8
62
63
XVI]I
X]X
XX 27
xxr .. 5r
XXII 28 52
xxlrl- 29,30 53,54
XXV
65,66

29. 2a M. S. KHAN BIOLOGIA
AN.
ANK.
AN.PT.
AN.T.
A.P.
B-
BE.
BL.
CA.FU.
CE.GL.
c.G.
CL.B.
c.L.N.T.
CRBS.
c.T.P.
DEN.
D.F.
D.L.
D.L.B.
D.V./DOR.
E.
E.C.
FI.
F.L.
FT,
G1--G4.
G.AN.
Dorsallip of blastopore
Dorsalview
Eve
Eyecup
FiDger
Fore-limb
Foot
First gill-Fourth gill
Gill
Gill analag€
LIST OF ABBREYIATIONS
Anus'
Ankle.
Anal pit.
Anal tube.
Animal Pole
Antedor view.
Blastopore.
Beak.
Blastomerc,
Caudalfunow,
Cementgland.
Cleavagegroove
ClosedblastoPorc
Closedneulal tube
GreYcrcscent
Cloacaltail Pi€ce
Dentical
Dorsalfin
DorsalliP

30. 1965 NORMAL TABLE
G.BD,
G.F.
H.L,BD.
INT.CO.
K.
LAT.LI.
LFT,H.
L.LP.
L.L.B.
L.V.
ME.
MEL.
MI.
MTH.
MYO.
NA,OP,
N.F.
N.G,
N.PL.
N.T.
OLF,PT.
OP,F.
oR.suc.
POST,DEN.
PRE. DEN.
PRO.
S.
SH.
SP,
ST.CL.
T.
T.BD.
TH.
OF Rafo,nelanosti.tut
Cill bud
clll frlametrt
Hind limb bud
lnt€stionalcoil
Knee
Latral lin€
Left forclimb
Irwer IiP
Latral lip of blastopore
Latral view
Megarneres
Melanophores
Micromeres
Mouth
Myotomes
Nasalopening
Neural folds
N€ural grcove
Neuralplate
Neunl tube
Olfactory pjt
opercular fold
Oral sucker
Posforal denticles
Pre-oral denticles
Pronephricregion
Sucker
Shank
Spiracle
Stomodaealcleft
Tail
Tail-bud
Thigh
b
iI

31. t0 M. S.KTIAN
Tajl piece
Tubcrclc
Upircr Iip
Vegetablcpole
Vcnlr,tlFir
Vitellinerncn]brane
Ventraltip
Yolk plug.
DIOLOG]A
T.PE.
TUB.
U.LP.
VEC.P.
VIT.
v.L.
v.v./vEN,
YK,PC,
REFERENCES
Arllcr,W. Dic Enhvicklungdcr ausscreuKorperformund dcs McsoCcrns
be Bulo yulea'is. InteFnatl. Monats&et Attat. phtsiot., t8 (19ot).
Boulcneer,C. A. Faunaof Btitish I dia, Reptitesan(tBattdchid,Londan.
Taybr and Francs.(1890).
Cambar,R. & Marror, B. Tabic clronologiquedu deveiolmentde la
grenovillc agile (R.t"a lolnlatitn lton.) Brll. Biol. FtancaBctg.,8a,
I68-r77(r954).
Del Conlc, E. & Sirlin,J. L. Pattefnsericsof thc firsr embryonarysragesin
Bufo xt't1atun.,1kat. Rec., 125J3s (1952).
Eakin,R. M. Delerminaljonandregulalionofpolarilyin the retinaof/1rld
t?gi|ld. Uniy. CdliforniaPubl. Zoal., 5r,245 (t946).
Eakin,R. M. & Harris,FL lnconpatabilitybetweenan]fhibianhcsrs and
xenoplaslicgrafrsasrclated1olost age.J. trl. Zaot.,99,35-64(1945).
Callicn.L. & Houillon, G. T.rbJechronologiquedu delebFemcntchez
Discostosstslic ts.Bull. Biot. Ftatm Bets. 95, 373-37s(1951).
Cilani, S. H. GnpLrblished).Tireea.ly devetop eDtof RanatigrahaDadin.
M.Sc. Research.IuDjab University,Depa(mentof Zoology,t957.
Harrison,R. G. Experimenlsonthcdevetophentofforelinrbof lrrrl_lr1or?a.
Joxtn. E.?et. Zool. 25, 413 (1918.
Hashmi,T. H. (unpublished)Tle Anphibjaof Lahorcuirh descriprionof the
early devclopmcntaf Bufo melanontictusSchneider. M.Sc. Research.
Puniab Universiry l)eparhenr of Zootogy, (1955).
Kopsch. F. Die Entwicklxngdes larauncnCrasfroschcsRan| fusca R.e.el
(dargcsrelltin def Art der Normentafetnzuf EnrwicktunSsgeschichte
der Wirbclrhicrc)Stutgarl,(1952).
Miller, D. C. A rnblc for ihe normai delctopmenlof R.tnapipicns. ptoc.
Indiano Acdd. S(i. 49, 209214 (t939).

32. vol. 11, i965 NORMI'L T/ILE OF Btlb hetaroyictus 3 l
Moser,H. Ejn Beitragzur Anaiyseder Thyroxineinwirkungjrr Kau]quap_
penversuchund zur Fragenacl demZustandekonnender Fruhbelcit_
schafr des M€ranorphoseRcaklionssyslcms.Thesis, Basel; Rer.
SuisseZoal,57 (Suppl.,2), t_144(19s0).
ll.urral
r At 'The
knhruk zaoros, of Sint,London: Richardson(1884).
Nicuwkoop,P. D. & Faber,J. NanLtt Tabte "t ,.*r" r*,,, fi)rirji.
- ,,
Norrl-H.,"-ndp.rbl.hi"f( . nrprny.AmJLrodm(r956r.
Polltrler.A. W. & Vuofe.L A. Txbtestor rIe rornat deejopmen,.
^ .
Ranastlfttt,o. .1,,//.n,, . 68. 489 (loJ I.
Ko(s.A. Tr!o'e rrorotoprchederto{ijdppo enbrio,rrteeld,rted. ..A,r'u
^ .
brla (Lt: Mahit"t, 1:uo!//r/.66. tJJ t45itqSUr.
5cflelbel.C. L",lcfin.ziorredeptr,radide .r l4.rrfrorttrosidctR..fr. R,t.
^.
R. ^t!,1 ^a,. Ltn.i 25, jr2.j48 rtoJT).
tnr.YI * S:cpe'_,nrl-enom..Jcerrror.r.,nr at Ra,r?ipi, tl:.t, | tetr"lrorms.Akat. Rec. 7t, t39-j1j ,1940).
Taylor,A, C.
.&
Kollros, J. J. Stages;n rtre normal dcvclopnentof /?dru
^ .
pipienstanl:]ae.Anat. ReL., 94, j 23 (1946).
Tsclernofl,N. D. ZLrrEmbryonalenlwjcktungder trinteren Exlremjlatten
-_.
desFrosches.Akol. Anaz.,30,593-6j2(i902).
Weisz,P. B. TIe nofnral sragesin rhe devctopmenrof lbe So.rthAfrican
ctawcdload,Xero?uslaetis..tnar. Rec.93,161_169(1945).
Wjnterberi,M. p. Z r I'ordre d,appariliol desoriejlset et prcmierdevetopfe_
mentdes memberschez les Anaures. Conet. Refitl., Sor.a.niot.Sg,
576-578(r905).

33. BiOLOCIAM. S. KHAN
e--ry4
2 Ly.
A;v/
40v
@,;Ll
6
@,lF/-ur'
9
@
r VEG.P
ffiY'
/troLF
@*
4 W
@-5DV
@E
@.PLATE I
I
I
l-
ffi5LV
@
,

34. NaRMLL
'TABLE
OF Rrfo nelanontctB 33
,:ff- @@:l3w
,r@$@ffi
r5AV
ffi-'
t6Dr'.
PLATE2

35. BIOLOCIAM. S. KTIAN
c^N cE6L TSO
"'"€
1.-r'
PLATE3
: ,
ft
ii

36. Vol. ll, 1965 NORMAL TABLE OF Bufo nelatusIi.t6
2zLV
PLATE4

37. 36 M, 5. KHAN BIOLOGIA
25LV

38. 37NORMAL TABLE OF ,a/o nela,orric.rrVol, 11,1965
r.---.. ^)"-/ V
2't 26
H> DJ P29 2a
3 4
tu$9-
43
,,.. A .-
P q Y r " V
i / il :i [ *
j ) l
35 36 37 eB 39
PLATE6

39. II. S, KHAN BIOLOCIA
I

40. Vol. 11, 1965 NoRMAL TABLE oF &{o nelanoei'tt6 39
*@e
S U C K E R
PLATE8