Development of Hyla Annectans Jerdon, 1870 from Nagaland, India

10 Pages·2001·8.5 MB·

by  J Meren Ao

) )
DEVELOPMENT OF HYLA ANNECTANSJERDON, 1870
FROMNAGALAND, INDIA
1
J. Meren Ao2 and SabitryBordoloi3
(Withfourplates
Key words: Hyla annectans, ontogenic stages, normal development, Nagaland,
laboratory condition, metamorphosis
Hyla annectans breeds during May to July in temporary rain pools, terraced fields and
other waterlogged areas (depth 5-7 cm). Normal development has been studied from the
egg through metamorphosis for four breeding seasons (1996-2000) under laboratory
conditions. Thetimerequired forcompletion ofalifecyclevariesunderdifferentweather
conditions in thefield. In the laboratory (16 22°C)the frog completed its lifecycle in 64
days, 14 hours.
Introduction middle ofApril and laststill July end. The frogs
breed in water logged places like temporary
Studiesonthesuccessiveontogenicstages, ponds, rainpools, puddles and terraced paddy
to record the normal developmental table, are fields. Embryonicdevelopmentwasobservedfor
important in understanding the ecology of a a period offour breeding seasons (1996-2000).
speciesandforplanningconservation measures. Laboratory rearing was carried out in the
Various authors (Dutta and Mohanty- DepartmentofZoology,KohimaScienceCollege,
Hejmadi 1976, Mohanty-Hejmadi and Dutta Kohima, Nagaland.
1977, Agarwal and Niazi 1977, Roy and Khare
1978, Mohanty-Hejmadi et al. 1979, 1980, Methodology
KiyasetuoandKhare 1986,Duttaetal. (1990-91
have contributed to the study of normal Amplexing pairs were collected from the
developmental tables ofAnuran amphibians in field and transferred to aquaria or glass
India. containers with water, allowing only halfofthe
InNagaland (25° .15'-27°4’ N, 93° 20'-95° body to be submerged. Amplexing lasts for 3-5
15' E), Hyla annectans is distributed along the hours. Eggs are laid between 0100-0400 hrs in
m
Borail range at various altitudes 1,400-2,440 the aquaria as well as in the field. Development
above msl. Average atmospheric temperature stages were fixed in 5% formaldehyde solution;
during breeding season ranges from 16.5- measurements were taken from preserved
26.6 °C. Relative humidity ranges from 74.33- specimens.PhotographsforPlates 1-3 weretaken
81.63%. Precipitation ranges from 128.23- frompreservedspecimens, whileforPlate4 live
428.33 mm. Water temperature varies from individuals were used.
14-28.5 °C. Breeding activity starts from the
Observations
'AcceptedDecember,2000
departmentofZoology,KohimaScienceCollege, Breeding activity starts from the middle
Kohima797001,Nagaland,India ofApril(12.3-26.6°C)withthemales’ breeding
•EcologyLaboratory,DepartmentofZoology,
CottonCollege,Guwahati781 001 Assam,India call. Females appear only after one or two
,
JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.2001 169
DEVELOPMENTOFUYLA ANNECTANS
showers. In early May, amplexing pairs were Stage 5: Eight cell stage (Age 2.45 hrs;
collectedandkeptinglasscontainersoraquaria. length 1.52 mm). The third cleavage is
Eggs were laid in batches in the field. In the latitudinal, slightly above the equator, which
aquarium, due to the absence of vegetation or forms eight blastomeres. The four smaller
other substratum, thejelly capsules ofthe eggs micromeres of the animal pole are pigmented
adhered together to form a single mass. The darkbrown,whereasthefourbiggermacromeres
number ofeggs laidby one female ranged from ofthe vegetalpole are unpigmented.
590-650. Culture was maintained in clean Stage 6: Sixteen cellstage (Age 3.20 hrs;
enamelled trays and 100 fertilized eggs were length 1.52 mm) (Plate 1, Fig. 4). The cleavage
stockedineachtraytoavoidover-crowding.They furrow is vertical. First, the pigmented
wererearedinthelaboratoryat 16-22°C. Larvae micromeresaredividedintoeightcells,resulting
were fed with Spirogyra, which is common in in twelve cells (i.e. 8 micromeres and 4
the breeding habitats. Tadpoles were staged macromeres).Thisisfollowedbythedivisionof
according to Gosner (1960). the four unpigmented macromeres as the
The no. ofindividuals per tray (Stocking cleavage furrow reaches the vegetal pole,
number = 100) was reduced with progressive resulting in 16 cells altogether.
developmental stages. Stage7: Thirty-twocellstage(Age3.52hrs;
length 1.56mm) (Plate 1,Fig. 5).Thelatitudinal
Embryonic stage cleavage furrows of the micromeres and
Fortysamplesweremeasuredtorecordthe macromeres bring about the formation of 16
mean size ofeach stage (1-46). micromeres and 16 macromeres.
Stage8:Midbiastula(Age7-54hrs; length
Fertilization stages 1.56 mm) (Plate 1, Fig. 6). The number ofcells
Stage 1: Fertilizedegg (Age 0 hrs; length increasedtomorethan64cells. Theblastomeres
1.52 mm) (Plate 1, Fig. 1). The egg is spherical. aremorenumerousandsmallerthanbefore. The
Theanimalpoleispigmenteddarkbrown,paling surface ofthe animalpoleresembles a clusterof
to white at the vegetal pole. beads.
Stage 2: One cell stage (Age 0.55 hrs; Stage 9: Late blastula (Age 12.10 hrs;
length 1.52 mm). A lightly pigmented area, the length 1.56mm). The surface ofthe animalpole
grey crescent appears between the animal and has a granular appearance, which gradually
vegetalpoletowardsthepigmentedhemisphere. becomessmooth. Thepigmentedregionextends
overthevegetalpole,whichmarksthebeginning
Cleavage stages oftheepibolicmovementofthemicromeresonto
Stage 3: Two cell stage (Age 1.50 hrs; the macromeres.
length 1.52mm)(Plate 1,Fig.2).Themeridional
cleavage furrow originating at the animal pole Gastrulation stages
proceeds to the vegetalpole and divides the egg Stage 10: Crescent dorsal lip (Age 15.05
completely into two equalblastomeres. hrs; length 1.56mm)(Plate 1,Fig. 7).Thedorsal
Stage 4: Four cell stage (Age 2.20 hrs; lip of blastopore has formed and is
length 1.52 mm) (Plate 1, Fig. 3). The second crescent-shaped. The unpigmented zone of the
meridional furrow, which starts at the animal vegetal hemisphere is reduced due to continued
pole, extends to the vegetal pole at a right angle migrationofthepigmentedmicromerestowards
to the first. the vegetal pole.
170 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 2001
DEVELOPMENTOFHYLA ANNECTANS
Ao, J. Meren et al.:Hyla annectans Jerdon Plate 1
Fig.1 Fertilizedegg, Fig. 2Twocell stage, Fig. 3 Fourcell stage, Fig. 4 Sixteencell stage,
Fig. 5 Thirty-twocell stage, Fig. 6 Midblastula. Fig. 7 Crescentdorsal lip,
Fig. 8 Horse-shoe shapeddorsal lip, Fig 9. Neuralplate.
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DEVELOPMENTOFHYLA ANNECTANS
Fig. 10Closure ofneural fold, Fig. 11 Neuraltube. Fig. 12, 13 Tailbud stage, Fig. 14 Muscularresponse
stage, Fig. 15 Opercularstage, Fig. 16 Opercularfoldofembryo closed onright side,
Fig. 17 Operculum ofembryo closed on leftside
172 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 98(2), AUG. 2001
DEVELOPMENTOFHYLA ANNECTANS
Stage 11: Horse-shoe shaped dorsal lip ruptures at the anterior end.
(Age 16.50 hrs; length 1.56 mm) (Plate 1, Fig. Stage 17: Tail bud stage (Age 42.50 hrs;
8). The epibolic migration ofmicromeres over Trunk 2.44 mm; Tail 0.47 mm) (Plate 2, Figs.
the vegetal pole reduces the exposed area of 12, 13). Tailbud appears at the posterior end of
unpigmented macromeres, which is surrounded theembryo. Itiswiderthanlong, directeddorso-
by the lateral lips ofthe semicircular or horse- posteriorly and marked offfrom the body by a
shoe shaped blastopore. ventral notch. Stomodeal groove depression is
Stage 12: Smallyolkplug{Age 26.10 hrs; slightly marked by a darker area. Bulges ofthe
length 1.56 mm). The ventral lip ofblastopore gill plates are distinct.
shifts to the posterior end. The uninvaginated Stage 18: Muscular response stage (Age
macromeres, surroundedbytheblastoporal lips, 60.00 hrs; Trunk 2.71 mm; Tail 0.75mm) (Plate
protrude a little and constitute the yolk plug. 2, Fig. 14). Head region is well defined, with
optic bulges and bulges of the gill plates. Oral
Neurulation stages suckers are indicatedby two heavily pigmented
Stage 13: Neural plate (Age 34.30 hrs; elongated areas joined medially by a narrow
length 1.6 mm) (Plate 1, Fig. 9). The embryo is lightly pigmented band below the stomodeum.
slightlyelongated.Thedorsalsurfaceisflattened Stomodeal depression is seen between the oral
to form the neural plate, which is differentiated suckers.Duetogradualelongationoftheembryo,
with the concentration of pigments along its the tail starts curving laterally to right or left,
borders. within the contour ofthe vitelline membrane.
Stage 14: Neuralfold (Age 36.20 hrs; Stage 19: Heartbeatstage(Age70.07hrs;
length 1.65 mm). The neural fold becomes Trunk3.10mm;Tail 1.4mm). Pulsationofheart
distinct, with broad cerebral and nairow spinal is seen below and behind gill bud. Small
cordregionsoftheneuralplate. Theneuralfolds pigmented depression at anterior end marks
gradually approach each other from blastopore olfactory pit. Stomodeal depression becomes
to anterior region. somewhat triangular. External gill buds
Stage 15: Closure ofneuralfold (Age prominent. Embryo coils to mechanical stimuli.
38.20 hrs; length 1.72 mm) (Plate 2, Fig. 10). Stage20: Gillcirculationstage(Age72.29
Theposteriorendoftheembryobecomesbroader. hrs;Trunk3.28mm;Tail 1.5mm). Gillsdistinct,
The neural folds come closer and touch each rudimentarybranchingatdistalend. Oralsuckers
other, both in the cerebral and spinal cord nipple-shaped. Stomodeal pit still a shallow
regions, formingashallowneuralgroove,which triangular depression. Vitelline membrane
is broader in the cerebral region. becomesthinandweak. Anteriorendofthehead
Stage 16: Neural tube (Age 40.50 hrs; pressed against the vitelline membrane.
mm
length2.0 (Plate2,Fig. 11)Theneuralfolds
have fused completely to form the neural tube, Post-embryonic development
which is raised at the mid-dorsal ridge and Stage 21: Lama hatched(Age 102.29 hrs;
demarcated by a darkly pigmented strand. The HeadandTrunk3.2mm;Tail3.2mm).Thehead
head and trunk are well marked. Gill plates causes a bulge in the vitelline membrane at the
appear as faint bulges. The increase in size of anteriorend. At this point the membrane breaks
the embryo along with the associated vitelline and the larva emerges from the mass ofjelly.
capsule, isnotaccompaniedbyasimilarincrease Tailstraight,tailfindusky.Olfactorypitdeepens,
in the length of the first envelope, and thus it corneabeginstobetransparent. Stomodeumnow
JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.2001 173
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DEVELOPMENTOFHYLA ANNECTANS
adeeptriangularpit, whose opening isthe larval Tail 16.0 mm) (Plate 3, Fig. 19). Distal end of
mouth. limb bud slightly conical.
Stage 22: Tailfin circulation stage (Age Stage 29: Length oflimb bud is equal to
1 17.29 hrs; Head and Trunk 3.4 mm; Tail 3.6 one and halftimes its diameter (Age 600 hrs;
mm). Tailfincirculationstartsatbaseofanterior Head and Trunk 9.0 mm, Tail 18.0 mm). Distal
part of dorsal fin, just above the trunk. Mouth end conical.
slightly wider. Upper and lower labial fringes Stage 30: Length oflimb bud is equal to
develop, but without papillation. twiceitsdiameter(Age684hrs;HeadandTrunk
Stage 23: Opercular fold stage (Age 11.0 mm, Tail 19.1 mm). Distal halfofconical
mm
143.59 hrs; Head and Trunk 3.6 Tail 4.5 limbbudslightlybentventrally.Nopigmentation
mm) (Plate 2, Fig. 15). Operculumcoversbases on limb bud.
ofexternalgills.Jawsnotkeratinised.Upperand Stage 31: Footpaddlestage(Age 748 hrs;
lower labial fringes develop papillae and faint Head andTrunk 11.5 mm; Tail 20.1 mm) (Plate
labialridges. Pigmentationontailbegins, cloaca 3,Fig. 20).Thedistalendoflimbbudisflattened
not opened. mediolaterallyto forma footpaddle. Kneebend
State 24: Opercularfoldofembryo closed is prominent. Light pigmentation starts at outer
on rightside (Age 172.29 hrs; Head and Trunk base oflimb bud.
3.7 mm; Tail 5.0 mm) (Plate 2, Fig. 16). Stage 32: First indentation (Age 813 hrs;
Operculumclosed on the right side. Labialjaws Head and Trunk 12.0 mm; Tail 23.0 mm). The
formsupra- and infra-rostrodont. A faintrow of margin of the foot paddle becomes slightly
upper supra-angularkeratodont develops. Other indented on the dorsal side, which marks the
characters as in Stage 23. prominences ofthe future 4th and 5th toes.
Stage 25: Operculum ofembryo closedon Stage 33: Second indentation (Age 853
left side (Age 185.29 hrs; Head and Trunk 4.2 hrs; Head and Trunk 12.8 mm; Tail 24.5 mm)
mm; Tail 6.0 mm) (Plate 2, Fig. 17). Operculum (Plate 3, Fig. 21). The margin ofthe footpaddle
closed and gills disappear; oral suckers becomes indentedontheventralside,behindthe
diminishing. Spiracle formed. Tail lightly prominence of4thtoe, whichmarksthe 3rd, 4th
pigmented.Analtubeopens,tadpolestartsfeeding. and 5th toes.
Stage 26: Hind limb bud stage (Age 356 Stage34: Thirdindentation(Age890hrs.;
hrs; Head and Trunk 7.0 mm; Tail 10.0 mm) Head andTrunk 13.2 mm; Tail 26.0mm) (Plate
(Plate 3, Fig. 18). Appearance ofhind limb bud 3, Fig. 22). The margin offoot paddle becomes
at a groove between the belly wall and base of indented, on the ventral side, behind the
tail oneitherside ofcloacaltailpiece. Lengthof prominence of 3rd toe, which marks the
limb bud less than half its diameter. Dental prominence of2nd, 3rd, 4th and 5th toes.
formulabecomes 1:1 + 1/3.Pigmentationspreads Stage35:Fourth indentation(Age936hrs;
to dorsal and anal fins. HeadandTrunk 13.4mm; Tail 26.4mm.)(Plate
Stage 27: Length oflimb budequalto half 3, Fig. 23). The margin of the foot paddle is
its diameter (Age 420 hrs; Head and Trunk 7.8 indented behind the 2nd toe demarcating the
mm; Tail length 13.0 mm). Length oflimb bud prominence ofthe 1st toe. All the five toes are
equal to half its diameter. The patches of separated from each other.
pigmentation in the tail fin spread considerably. Stage 36: Margin of5th toe web directed
Stage 28: Length oflimb bud equal to its towards the tip of2nd toe (Age 1019 hrs; Head
diameter(Age514hrs;HeadandTrunk8.4mm; and Trunk 13.6 mm; Tail 27.2 mm) (Plate 3,
174 JOURNAL BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 200
DEVELOPMENTOFHYLA ANNECTANS
Ao, J. Meren et al.’.Hyla annectans Jerdon Plate 3
Fig. 18 Hindlimbbud stage, Fig. 19Lengthoflimbbudequal toits diameter, Fig. 20Footpaddle stage,
Fig. 21 Secondindentation, Fig. 22 Thirdindentation, Fig. 23 Fourth indentation,
Fig. 24Margin of5th toe webdirectedtowards thetipof2ndtoe,
Fig. 25 Marginof5th toewebdirectedtowards prehallux.
JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 98(2), AUG. 2001 175
DEVELOPMENTOFHYLA ANNECTANS
Ao, J. Meren et al.:Hyla annectans Jerdon Plate 4
Fig. 26, 27 Cloacaltailpiecereduced, Fig. 28 Both forelimbs emerged,
Fig. 29 Angleofmouthreachedthe middle ofeye,
Fig. 30 Angle ofmouthreachedposteriormargin oftheeye, Fig. 31 Metamorphosedfroglet.
176 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 98(2), AUG. 2001
DEVELOPMENTOFHYLA ANNECTANS
Fig.24).Themarginofthe5thtoewebisdirected labial fringe begins, however angular papillae
towards the tip ofthe 2nd toe. stillremainas a small tuftonbothcomers ofthe
Stage 37: Margin of5th toe web directed mouth, which starts widening. The homy beak
towards the tip of1st toe (Age 1084 hrs; Head is shed. Tail starts darkening. The angle ofthe
andTrunk 13.7mm; Tail28.0mm). Themargin mouth is level with the nostril. The cloacal tail
ofthe 5th toe web is directed towards the tip of piecedisappearsatthisstage, leavingthecloacal
the 1st toe. Pigmentation appears in the 4th and aperturefreebelow.
5th toes along the foot. Toes are longer. A Stage 43: Angle ofmouth between the eye
rudiment of prehallux is indicated by a light and nostril (Age 1434 hrs; Head and Trunk 14
protuberancebehind the 1sttoe. Ventral surface mm; Tail 16.0 mm). The widening angle of
ofthe foot is closer to the ventral fin. mouth has reached a point midway between
Stage 38: Margin of5th toe web directed nostril and the anterior margin ofeye. The tail
towards prehallux (Age 1136 hrs; Head and becomes still darker, the dorsal and ventral fins
Trunk 14.0 mm; Tail 29.00 mm). (Plate 3, Fig. shrink and the length ofthe tail is reduced, but
25). The margin ofthe 5th toe web is directed still longer than the extended hind limb.
towards prehallux. Inner metatarsal turbercle Stage 44: Angle ofmouth reached the
appear as a small outgrowth. Pigmentation middle ofeye Age 1462 hrs; Head and Trunk
(
appears in 3rd, 4th and 5th toe along the foot. 14.00 mm; Tail 5.0 mm (Plate 4, Fig. 29). The
Stage 39: Appearance ofsubarticular widening angle of the mouth has reached the
tubercles in the toes (Age 1210 hrs; Head and levelofthemiddleoftheeye. Dorsalandventral
Trunk 14.3 mm; Tail 29.2 mm). Subarticular finshavedisappeared.Tailresorbedconsiderably
tubercles appear onthe inner surface ofthe toes and is as long as the femur.
as light patches. The inner metatarsal tubercle Stage 45: Angle of the mouth reached
becomes a small oval outgrowth. posteriormargin oftheeye(Age 1498 hrs; Head
Stage 40: Toe pads complete (Age 1274 andTmnk 13.5 mm; Tail 1.0mm) (Plate4, Fig.
hrs; Head and Trunk 14.4 mm; Tail 29.2 mm). 30). The widening angle of the mouth has
Distal ends of the toes thickened. Subarticular reachedtheposteriormarginofthe eye. The tail
tubercles are clearly elevated. The cloacal tail is resorbed to a small triangular stub.
piece is notreduced. Stage 46: Metamorphosedfroglet (Age
Stage 41: Cloacal tailpiece reduced (Age 1550 hrs; Head and Tmnk 13.5 mm; Tail 0.0
1392 hrs; Head and Trunk 14.8 mm; Tail 30.0 mm.) (Plate 4, Fig. 31). The triangular tail stub
mm. (Plate4, Fig. 26, 27). The cloacal tail piece with dark tissue disappears completely.
getsreducedandonlyanarrowstripremainsover Total time taken for completion of
and in between bases of the thigh still attached metamorphosis was 64 days 14 hours.
withventralfindistally.Theforelimbsarevisible
throughtheskin. Greenpigmentationofthedorsal Discussion
surface begins. Tail not dark. Keratodonts start
shedding. Oral papillae remain intact. Hamburger (1947), Gosner (1960), Rugh
Stage 42: Bothfore limbs emerge (Age (1962), Nieuwkoop and Faber (1967) were
1422 hrs; Head and Trunk 14.5 mm; Tail 25.1 referred for preparing the normal table. Gosner
mm) (Plate4, Fig. 28). Bothfore limbs emerge, (1960) proposed 46 stages, with simplified
usuallytherightforelimbemergesfirst,followed criteria for staging developmental events in
after a few hours by the left. Resorbtion ofthe Pelobatids, Bufonids, Ranids and Hylids. In the
JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.200! 177
DEVELOPMENTOFHYLA ANNECTANS
presentstudy,thedevelopmentofHylaannectans embryos occur in stage 21, whenthe corneajust
was divided into these 46 stages. Certain begins to be transparent. However, the cornea
variations have been noted in the development becomesfullyclearonlytowardstheendofstage
ofHyla annectans. 22 and beginning ofstage 23. At stage 22, the
Developmental, stages have been divided circulation in the tail fin begins, but the tail fin
intofifteenmajorsubheadings(1)Fertilizationtwo isnottransparentasinGosnerseriesandremains
stages(2)Cleavagesevenstages(3)Gastrulathree dusky. Narrow cloacal tail piece persists, in
stages (4) Neurula four stages (5) Tail bud one Gosner series - 41 which, however, disappears
stage (6) Muscular response one stage (7) Heart only in stage 42 when both forelimbs have
beat one stage (8) Gill circulation one stage emerged. Rostrodont and keratodonts are shed
(9) Cornea becoming transparent one stage completely at stage 42.
(10)Tailfincirculationonestage(11)Operculum
formation three stages (12) Hind limb bud five Acknowledgement
stages(13)Identationanddevelopmentoftoesten
stages (14) Cloacal tail piece reduced one stage This paper is dedicated to the memory of
(15) Metamorphosis climax five stages. Late Prof. MahendraKumarKhare, who initiated
Characteristic features of development research in Developmental Biology in the North
includethetailbud,initiallyindicatedbyastrong Eastern Hill University, Shillong, Meghalaya,
upward arching ofthe dorsum. Hatching ofthe India.
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