243
ACTA ZOOLOGICA BULGARICA
Applied Zoology
Resea ch A icle
Ac a Zool. Bulg., June 2025, 77 (2): 243-248
Published online 20 May 2025
DOI: h ps://doi.o g/10.71424/azb77.2.002788
*Co esponding au ho : [email p o ec ed]
h ps://www.ac a-zoologica-bulga ica.eu/2025/002788
E ec o Habi a D ying on Age and Size a Me amo phosis
in Indian Pain ed F og, Upe odon ap obanicus
(Anu a: Mic ohylidae)
San osh M. Mogali*, Bhagyash i A. Shanbhag & S ini as K. Saidapu
Depa men o Zoology, Ka na ak Uni e si y, Dha wad 580 003, Ka na aka S a e, India; [email p o ec ed]
Academic Edi o : Simeon Lukano
Abs ac : Anu ans ha e complex li e cycles. The key me amo phic ai s in anu ans a e la al du a ion and size a
me amo phosis. In mos o he anu ans, hese wo me amo phic ai s a e la gely in luenced by se e al
bio ic and abio ic ac o s bu mainly by habi a d ying. We s udied he in luence o habi a d ying on
la al du a ion and size a eme gence in Upe odon ap obanicus unde labo a o y condi ions. F eshly
ha ched adpoles (Gosne s age 19) we e exposed o dec easing wa e le els (g adually o apidly) up o
he beginning o me amo phic climax s age (MC, Gosne s age 42; eme gence o o elimbs). The con ol
g oup was ea ed in cons an wa e le els. Tadpoles ea ed in dec easing wa e le els (g adually o ap-
idly) eached MC and comple ed la al du a ion ea lie and me amo phosed a a smalle size han hose
ea ed in cons an wa e le els. Fu he , adpoles expe iencing apid deple ion o wa e eached MC and
comple ed la al du a ion ea lie and also me amo phosed a smalle size han hose expe iencing g adual
deple ion in wa e le els. Thus, adpoles o U. ap obanicus showed adap i e plas ici y in me amo phosis
o habi a d ying. The s udy shows ha U. ap obanicus adpoles a e capable o de elopmen al plas ici y
and exposed o dec ease in wa e le els, he ade-o be ween g ow h and de elopmen is in a ou o
de elopmen , esul ing in ea ly me amo phosis a a smalle size.
Key wo ds: Anu ans, desicca ion h ea , de elopmen , g ow h, me amo phic ai s, adpole
In oduc ion
A p ime goal o li e-his o y heo y is o expec pa -
e ns in he age and size o o ganisms a on ogene ic
swi ch poin s, o example ha ching, me amo pho-
sis and ep oduc i e ma u i y (Wilbu & Collins
1973, S ea ns & Koella 1986, Rowe & Ludwig
1991). In anu ans, pheno ypic plas ici y is a ubiqui-
ous cha ac e is ic a ec ing many li e-his o y ai s,
especially hose ela ed o ep oduc ion and la al
de elopmen (Mine e al. 2005, Wells 2010, Rich-
e -Boix e al. 2011, U ban e al. 2014). Op imal
ime o me amo phosis is de e mined by a ade-o
be ween oppo uni ies o g ow h and isks o mo -
ali y in aqua ic and e es ial habi a s (Wilbu &
Collins 1973, We ne 1986, Rudol & Rödel 2007,
Mogali e al. 2011a). In mos anu ans, he ime and
size a me amo phosis a e ex emely plas ic and
244
Mogali S. M., Shanbhag B. A. & Saidapu S. K.
bo h depend on a se e al ac o s such as wa e le -
el (Loman 1999, Mogali e al. 2017, Székely e al.
2017), la al densi y (Newman 1987, Mogali e al.
2016), p eda o s (Relyea 2007, Mogali e al. 2011b,
2016, Mogali 2018), empe a u e (Gómez-Mes e &
Buchholz 2006, Maciel & Juncá 2009), ood a ail-
abili y (Newman 1998, Mo ey & Reznick 2000,
En iquez-U zelai e al. 2013) and he complex in e -
ac ion be ween hem (G özinge e al. 2014, Mogali
e al. 2016).
In epheme al ponds, desicca ion is a key h ea
and comple ion o me amo phosis be o e he ponds
d y is obliga o y (Newman 1992). Slow g ow h a es
and p olonged la al du a ions in unp edic able hy-
d ope iods o he wa e bodies a e su e o dec ease
he chances o adpoles comple ing me amo phosis
be o e he wa e bodies d y (Newman 1992, Al wegg
& Reye 2003, Johansson e al. 2005). On he con-
a y, a has ened la al de elopmen can lowe la al
mo ali y, bu i is in a iably a he cos o g ow h
esul ing in a smalle body size a me amo phosis
and ha may a ec hei la e su i al as well as
hei ep oduc i e success (Reques & Tejedo 1997,
Mo ey & Reznick 2000, Al wegg & Reye 2003).
Howe e , when la al mo ali y isk inc eases due
o pond desicca ion, an ea ly me amo phosis may be
a ou ed despi e he cos s associa ed wi h a smalle
size (Newman 1992). Hence, pheno ypic plas ici y
in ol ing he ade-o be ween ce ain li e his o y
ai s (e.g., la al g ow h, age and size a ans o -
ma ion) is a use ul s a egy. The o iginal Wilbu and
Collins’ (1973) model o amphibian me amo phosis
p edic s ha in aqua ic sys ems when condi ions a e
a ou able o la al g ow h, adpoles should pos -
pone me amo phosis and ans o m a a la ge body
size. Bu when condi ions o he empo a y ponds
become uns able, a s a egy o adjus he de elop-
men al p ocesses so as o me amo phose ea ly and
eme ge on land is use ul. A de elopmen al s a egy
o pheno ypic plas ici y can dec ease he exposi ion
o isky condi ions and, he eby, enhance he su -
i al a e.
In Sou he n India, many anu an species, in-
cluding he Indian pain ed og Upe odon ap oban-
icus (Pa ke 1934; amily: Mic ohylidae), ep oduce
in epheme al ponds o med du ing he sou h-wes
monsoon. Such ponds a e usually small and o en do
no las e en o a o nigh in he absence o in e -
mi en showe s (Mogali e al. 2011a, 2017, 2023a).
Thus, adpoles o U. ap obanicus p o ide an ex-
cellen model o s udy adap i e plas ici y in la al
de elopmen when hey a e exposed o a ying de-
g ees o d opping wa e le els (desicca ion h ea ).
The p esen s udy explo ed he in luence o g adual
o apid wa e deple ion on he wo majo me amo -
phic ai s, la al pe iod and size a eme gence, in a
labo a o y se up. We hypo hesised ha hose ad-
poles acing ei he low o high desicca ion h ea
would me amo phose ea lie and a a smalle size
han hose de eloping in cons an wa e le els (no
desicca ion h ea ). Second, we hypo hesised ha
hose adpoles acing he apid deple ion o wa e
(high desicca ion h ea ) would me amo phose ea -
lies and a smalle size as compa ed o hose de-
eloping in condi ions o a g adual decline in wa e
le els (low desicca ion h ea ). Impo an ly, he ex-
pe imen al design pe mi ed us o exclude he in lu-
ence o con ounding ac o s such as ood sca ci y,
c owding and p eda o p essu e which gene ally in-
e e e wi h he g ow h and de elopmen o adpoles
in na u e.
Ma e ials and Me hods
F esh eggs o Upe odon ap obanicus belonging
o h ee di e en pa en al lines (< Gosne s age 10,
Gosne 1960) we e collec ed in he ea ly monsoon
season om epheme al ponds in he Ka na ak Uni-
e si y Campus (15.440407º N, 74.985246º E, al i-
ude 750 m a.s.l.) Dha wad, Ka na aka S a e, India.
Soon a e collec ion hey we e b ough o he labo-
a o y and we e placed in a plas ic ub (42 cm diam-
e e and 16 cm deep) wi h 5 dm-3 o aged ap wa e .
All eggs ha ched (Gosne s age 19, Gosne 1960) al-
mos synch onously on he nex day. Tadpoles (Gos-
ne s age 19) we e picked andomly and we e ea ed
in he plas ic ubs (32 cm diame e and 14 cm deep)
wi h 0.5 dm-3 – 3 dm-3 o aged ap wa e un il he on-
se o me amo phic climax s age (Gosne s age 42;
Gosne 1960). Fi een such ubs wi h 10 adpoles in
each we e main ained (in o al 150 adpoles, i.e., 50
adpoles in each g oup). The expe imen al g oups
we e as ollows (see also me hodology o Székely
e al. 2010):
G oup I. Cons an wa e : Tadpoles we e ea ed
in cons an wa e le els (3 dm-3).
G oup II. G adual desicca ion: Tadpoles we e
ea ed in 3 dm-3 o wa e o he i s 4 days and hen
subjec ed o a 0.5 dm-3 dec ease in wa e a 4-day
in e als.
G oup III. Rapid desicca ion: Tadpoles we e
ea ed in 3 dm-3 o wa e o a day and om he sec-
ond day onwa ds 0.25 dm-3 o wa e was emo ed
each day.
In desicca ing wa e g oups when wa e
eached 0.5 dm-3 (a e day 20 in g oup II and a e
day 10 in g oup III) no u he educ ion was made.
The g oups II and III hus simula ed low and high
E ec o Habi a D ying on Age and Size a Me amo phosis in Indian Pain ed F og, Upe odon ap obanicus...
245
desicca ion h ea . Tadpoles o all g oups (i.e., I,
II and III) we e ed on boiled spinach ad libi um.
Wa e was changed on al e na e days and esh ood
was p o ided. The ea ing ubs we e placed on a la
su ace in a oom wi h a na u al pho ope iod (12 h
ligh : 12 h da k) and empe a u e. The posi ions o
ubs we e andomised on an al e na e day o a oid
possible e ec s o posi ion i any. The wa e empe -
a u e (ºC) in ubs was eco ded wice, daily a 10:00
h and 15:00 h. Following he onse o me amo phic
climax (MC, eme gence o o elimbs, Gosne s age
42), he subjec s we e ans e ed o small plas ic
ubs (19 cm diame e and 7 cm deep) co e ed wi h
ine nylon mesh wi h a li le wa e and placed in-
clined hus mimicking he semi- e es ial en i on-
men o acili a e eme gence. The days o each MC
s age and la al du a ion we e no ed o each indi-
idual. A e comple ion o me amo phosis (Gosne
s age 46), snou - en leng h (SVL in mm; measu ed
using digi al callipe , accu acy 0.01 mm) and body
mass (in mg; measu ed using an elec onic balance,
accu acy 0.001 g; KAB7L, Wensa Company, India)
we e eco ded. No a single adpole died du ing he
cou se o he expe imen . A e comple ion o ex-
pe imen s, he ogle s we e eleased nea na u al
wa e bodies. Da a on days o each MC s age, la al
du a ion, SVL, body mass o ogle s and wa e em-
pe a u e we e analysed using a one-way ANOVA
ollowed by he Tukey’s pos -hoc es . All s a is ical
es s we e pe o med using SPSS e . 16.0.
Resul s
The days equi ed o each me amo phic climax
s age (F 2,147 = 137.852, P < 0.05), comple e la al
du a ion (F 2,147 = 160.722, P < 0.05), snou - en
leng h (F 2,147 = 505.364, P < 0.05) and body mass
(F 2,147 = 394.791, P < 0.05) di e ed signi ican ly
among di e en ea men g oups (Table 1). Tad-
poles which we e ea ed in ea men s wi h declining
wa e le els (i.e., g oup II and III) eached MC and
comple ed la al du a ion ea lie (P < 0.05. Table 1)
and also me amo phosed a a smalle size (as quan i-
ied h ough SVL and body mass) (P < 0.05, Table
1) han hose adpoles which a e ea ed in condi ions
wi h cons an wa e le els (i.e., g oup I, Table 1).
Fu he , adpoles expe iencing apid deple ion o
wa e (g oup III) eached MC and comple ed la al
du a ion ea lie (P < 0.05) and also me amo phosed
a smalle size (P < 0.05) han hose expe iencing
g adual deple ion in wa e le els (g oup II, Table 1).
The daily wa e empe a u e o a ious ea ing
ubs luc ua ed be ween 22.5-23.0 ºC and as such did
no di e signi ican ly h oughou he cou se o he
expe imen s (mo ning: F2, 147= 1.951, P = 0.146; and
a e noon hou s: F2, 147 = 0.662, P = 0.517).
The equency dis ibu ion da a showed ha
all indi iduals (100%) om he apid desicca ion
g oup (g oup III) me amo phosed a < 8.00 mm
SVL, bu none o he indi iduals subjec ed o g ad-
ual desicca ion (g oup II) o cons an wa e le el
(g oup I) me amo phosed a compa able SVL (Fig-
u e 1). Fu he , all indi iduals (100%) om he ap-
id desicca ion g oup (g oup III) and 38% indi idu-
als om he g adual desicca ion g oup (g oup II)
me amo phosed a a smalle body mass (< 75 mg)
while all o he indi iduals subjec ed o cons an
wa e le els (g oup I) me amo phosed a di e en
body masses (Figu e 2). The da a on days needed o
each MC showed ha all indi iduals (100%) ea ed
in unde condi ions o apid desicca ion (g oup III),
also 72% indi iduals om he g adual desicca ion
Table 1. Snou - en leng h (SVL), body mass o me amo phs, days equi ed o each me amo phic climax s age (Gos-
ne s age 42) and leng h o la al du a ion (days, Gosne s age 46) in Upe odon ap obanicus ea ed in wa e s wi h
di e en le els o desicca ion. Da a ep esen mean ± SD (minimum-maximum) and analysed by one-way ANOVA
ollowed by Tukey’s’ HSD pos -hoc es . N = 50 adpoles o each g oup (150 adpoles in o al); dissimila supe sc ip s
(a, b, c) indica e signi ican di e ence be ween he g oups in he same column; signi icance le el was se o 0.05.
Rea ing g oups SVL
(mm)
Body mass
(mg)
Days equi ed o
each MC
La al du a ion
(days)
I. Cons an wa e 10.01 ± 0.49 a
(9.15-10.90)
121.54 ± 16.20a
(100-149)
26.96 ± 1.29a
(25-31)
29.68 ± 1.36a
(27-33)
II. G adual desicca ion 8.86 ± 0.35 b
(8.45-9.98)
80.78 ± 10.76b
(67-113)
24.93 ± 0.94b
(23-27)
27.56 ± 1.12b
(25-30)
III. Rapid desicca ion 7.60 ± 0.26 c
(7.09-7.99)
57.32 ± 4.74c
(50-65)
23.38 ± 0.96c
(20-25)
25.48 ± 1.00c
(22-27)
F and P alues F 2,147 = 505.364,
P < 0.05
F 2,147 = 394.791,
P < 0.05
F 2,147 = 137.852,
P < 0.05
F 2,147 = 160.722,
P < 0.05
246
Mogali S. M., Shanbhag B. A. & Saidapu S. K.
g oup (g oup II) bu only 10% o he indi iduals
om he cons an wa e -le el g oup (g oup I) ook
< 25 days o each MC (Figu e 3). The da a on
la al du a ion (days) showed ha all indi iduals
(100%) ea ed unde apid desicca ion condi ions
(g oup III), also 48% o he indi iduals om he
g adual desicca ion g oup (g oup II) bu only 6% o
he indi iduals om he cons an wa e -le el g oup
(g oup I) ook < 27 days o comple e la al du a ion
(Figu e 4).
Discussion
Ou esul s con i med ou wo hypo heses. The
labo a o y designed s udy con i ms ha adpoles o
U. ap obanicus show pheno ypic plas ici y by be-
ing able o ime hei me amo phosis in esponse o
a ailabili y o wa e . Tadpoles aised in cons an wa-
e le els me amo phosed la e and a a la ge size
han adpoles ha we e aised unde condi ions o
dec easing wa e le els (g adual o apid). Tadpoles
ha we e aised in g adual deple ion wa e le els
me amo phosed la e and a a la ge size han hose
adpoles om apid deple ion wa e le els. Hence,
adpoles o U. ap obanicus clea ly showed adap-
i e plas ici y in me amo phosis o pond d ying. Ou
esul s a e in acco dance wi h Wilbu and Collins’
model (1973) o amphibian me amo phosis, which
model assumes ha adpoles encoun e ing a ou -
able condi ions (e.g. cons an wa e le el o his
s udy) pos pone me amo phosis, capi alising on he
oppo uni y o addi ional g ow h, while adpoles
exposed o hos ile condi ions ( apid desicca ion
h ea o his s udy) de elop quicke and lea e he
wa e body ea lie , also suppo ing ea lie s udies
(C ump 1989, Newman 1989, Den e e al. 1998,
Loman 1999, Mogali e al. 2011a, 2017, 2023b
Székely e al. 2017).
Fig. 3. Pe cen adpoles o Upe odon ap obanicus pe
class (days) o each me amo phic climax (MC, Gosne
s age 42) in di e en ea ing g oups
Fig. 4. Pe cen me amo phs o Upe odon ap obanicus
pe class (days) o comple e i s la al du a ion (Gosne
s age 46) in di e en ea ing g oups
Fig. 2. Pe cen me amo phs o Upe odon ap obanicus
pe body mass class (mg) in di e en ea ing g oups
0
20
40
60
80
100
1 2 3 4
Cons an wa e
G adual desicca ion
Rapid desicca ion
50-74
Pe cen me amo phs
75-99 100-124 125-149
Body mass class (mg)
Rea ing g oups
Fig. 1. Pe cen me amo phs o Upe odon ap obanicus pe
snou - en leng h class (mm) in di e en ea ing g oups
E ec o Habi a D ying on Age and Size a Me amo phosis in Indian Pain ed F og, Upe odon ap obanicus...
247
The mechanisms p oposed o explain he ac-
cele a ion o me amo phosis in anu an adpoles
acing he isk o desicca ion di e . Ele a ed em-
pe a u e (Newman 1992, Tejedo & Reques 1994)
o a dec ease in ood (Al o d & Ha is 1988, New-
man 1994), mo ali y o conspeci ic membe s has
been a ibu ed o lowe ed g ow h a e wi h ac-
cele a ed de elopmen al a e. Ye , o he s udies
indica ed ha he empe a u e has no in luence on
de elopmen al a e (Loman 1999, Lau ila & Ku-
jasalo 1999, Má quez-Ga cía e al. 2009, Mogali
e al. 2017). In he p esen s udy, he e was no di -
e ence in wa e empe a u e (bo h in he mo n-
ing and a e noon) o he ea ing con aine s in all
ea men g oups and ood was p o ided in excess
quan i y. Also, he e was no mo ali y and, he e-
o e, he dea h o g oup membe s pe se could also
be uled ou as a ac o in speeding up me amo -
phosis. Hence, he key ac o in luencing accele -
a ed me amo phosis o adpoles o U. ap obanicus
is he desicca ion h ea . In conclusion, adpoles o
U. ap obanicus clea ly showed adap i e plas ici y
in me amo phosis o habi a d ying.
Acknowledgmen s: SMM is hank ul o UGC’s DSKPDF, New
Delhi. BAS and SKS a e hank ul o INSA, New Delhi o he
inancial suppo . The s udy was conduc ed acco ding o e hical
guidelines o he Commi ee o he Pu pose o Con ol and Su-
pe ision o Expe imen s on Animals (CPCSEA), New Delhi,
unde Regis a ion No. 639/02/a/CPCSEA.
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Recei ed: 06.11.2024
Accep ed: 03.04.2025
