See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/362962470

Oral and Buccophryngeal Morphology of the Tadpoles of Polypedates

maculatus with Notes on their Diet pattern

Article in Current Herpetology · August 2022

DOI: 10.5358/hsj.41.180

CITATIONS READS

0 110

3 authors:

Jasmin Rout Susmita Mahapatra

Utkal University Maharaja Shriram Chandra Bhanj Deo University
7 PUBLICATIONS 17 CITATIONS 10 PUBLICATIONS 41 CITATIONS

SEE PROFILE SEE PROFILE

Gunanidhi Sahoo
Utkal University
22 PUBLICATIONS 219 CITATIONS

SEE PROFILE

All content following this page was uploaded by Susmita Mahapatra on 14 May 2023.

The user has requested enhancement of the downloaded file.

Current Herpetology 41(2): 180–195, August 2022 doi 10.5358/hsj.41.180
© 2022 by The Herpetological Society of Japan

Oral and Buccophryngeal Morphology of the Tadpoles of

Polypedates maculatus with Notes on their Diet pattern

Jasmin ROUT1, Susmita MAHAPATRA2, and Gunanidhi SAHOO1,*

1Department of Zoology, Utkal University, Bhubaneswar 751 004, INDIA

2Post graduate Department of Zoology, Maharaja Sriram Chandra Bhanja Deo University,
Baripada 757 003, INDIA

Abstract: Oral morphological features and the diet of the tadpoles of

Polypedates maculatus were described in this study. The tadpoles were collected
from various water bodies of the state of Odisha over three consecutive
breeding seasons (2018–2020). Oral and internal buccal characters were
analysed independently for Gosner developmental stages 25 to 46. The upper
labium has four labial tooth rows with a medial gap in the three rows proximal
to the mouth, whereas the lower labium has three rows with a medial gap in
the row proximal to the mouth. The order of length of denticle rows is
A2>P1>P2>A1>P3>A3>A4 with a dental formula of A4(3)/P3(1). The food
spectrum of the tadpoles included mostly phytoplanktons (represented by six
classes and 51 genera) followed by zooplankton (two classes and four genera).
Three Bacillariophyceae genera (Navicula, Pinnularia, and Synedra)
contributed most of the gut contents. Scenedesmus was the major food items in
Chlorophyceae family and had the highest numeric frequency (7.63%) among
all the food items. Other prevalent food items included Zygnematophyceae,
Cyanophyceae, Euglenophyceae, Rotifera, Ulvophyceae and Copepoda.

Key words: Diet; Larvae; Oral disc; Scanning electron microscopy

Introduction The family Rhacophoridae includes 441 spe‐

Anurans are highly diversified and special‐
ised amphibians that have thrived in the terres‐
trial environment unlike other amphibian such
as, salamanders and caecilians (Mohapatra et
al., 2017). The Indian subcontinent harbours
around 417 species of amphibian (376 anurans,
two salamanders, and 39 caecilians) represent‐
ing about 5% of the total amphibian species
diversity of the world (AmphibiaWeb, 2021).
* Corresponding author.
E-mail address: gunanidhi.nou@gmail.com
cies worldwide of which 112 are known from
India. In the state of Odisha, the family is rep‐
resented by six species in four genera, specifi‐
cally Chirixalus (one species), Philautus (two
species), Raorchestes (one species) and Poly‐
pedates (two species) (Mohapatra et al., 2017).
The genus Polypedates is distributed from
southern China to Sri Lanka, and from south‐
western and northeastern India across Indo-
China and Indo-Malaya (Frost, 2021).
Polypedates contains 26 species of which 10
species occur in India (Dinesh et al., 2017) and
two species occur in Odisha (P. maculatus and
P. teraiensis) (Dutta et al., 2009; Mohapatra et
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES
al., 2017). The former is widely distributed in
the state whereas the latter has a patchy distri‐
bution in Mayurbhanj, Cuttack and Khordha
districts of Odisha (Dutta et al., 2009). Polype‐
dates maculatus is classified as Least Concern
(LC) (Dinesh et al., 2009).
Tadpoles, the aquatic larvae of anurans,
differ from their adult forms morphologically
(Altig and McDiarmid, 1999). The external
structure and anatomy of tadpole mouthparts
often provide useful diagnostic characters for
tadpole identification (Altig and Pace, 1974;
Lee, 1976; Inthara et al., 2005). The shape,
size, and number of keratodont rows in the oral
apparatus of amphibian tadpoles vary greatly
among species. However, tadpole mouthparts
for many species, including the Rhacophori‐
dae, are still poorly documented. The internal
buccal and the pharyngeal anatomy of tadpoles
vary across species (Bowatte and
Meegaskumbura, 2011). Hora (1922) presented
pioneering work in the field of buccophryngeal
research followed by Das (1994), Grosjean
(2003) and Grosjean et al. (2004). The bucco‐
phryngeal anatomy of tadpoles of three Poly‐
pedates species from Borneo (P. macrotis, P.
colletti, and P. otilophus) was examined by
Inger (1985). However, similar information is
not available for any of the Indian Polypedates
species.
Anuran tadpoles are important primary con‐
sumers in many freshwater habitats and play a
crucial role in the food chain. The growth of
tadpoles is directly influenced by the amount
of food consumed (Kiffney and Richardson,
2001) and the quality of that food (Kupferberg
et al., 1994). Differences in the internal oral
features among species may be linked to tad‐
poles’ preferences for food particles of varying
sizes (Wassersug, 1980).
Dietary data for anuran larvae from India
over the last three decades are relatively lim‐
ited (Khare and Sahu, 1984; Ao and Khare,
1986; Sekar, 1990; Saidapur, 2001; Sinha et
al., 2001; Khongwir et al., 2003). The majority
of research has been focused on the Western
Ghats (biodiversity hotspot) and the west coast
of India and other areas remain understudied
181
(Aravind and Gururaja, 2011). The diet of tad‐
poles including that of P. maculatus, particular‐
ly from the eastern part of India, is poorly
documented. The present study describes the
oral features of P. maculatus along with its diet
spectrum in a natural habitat.
Materials and Methods
Rearing and preservation of tadpoles
Foam nests of P. maculatus were collected
from various water bodies (permanent ponds,
temporary ponds, and canals) located in differ‐
ent parts of Odisha, India (21°37'52" N,
80°43'38" E and 155,707 km2) during three
consecutive breeding seasons (July to Septem‐
ber, 2018–2020). Tadpoles of this species are
available in the wild from July till September.
Individual foam nests were kept in separate
plastic pots (30 cm diameter, 9 cm depth) 75%
filled with pond water under room temperature
(25–30°C) in the laboratory. The tadpoles
(hatched in the laboratory and a few collected
from natural habitats) were maintained in glass
aquaria (46×31×31 cm) half-filled with pond
water. They were fed with algae collected from
the sampling locations and boiled spinach.
Water was changed in every three days or as
per requirement (if bubbles formed on the
water). The aquaria were checked daily for any
dead individuals.
Ten tadpoles of each developmental stage
(stages 25 to 46; Gosner, 1960) were preserved
in 10% formalin for oral and buccophryngeal
analyses. Individuals that reached metamor‐
phic stages (42 to 45) were transferred to
netted glass jars with a small amount of water
and few pebbles, maintained till metamorpho‐
sis after which they were released back into
their natural habitat. Staging of the embryos
and larvae were carried out following Gosner
(1960). Labial tooth formulae follow Altig and
McDiarmid (1999).
Light Microscopy
Oral morphology was examined under a
stereomicroscope (Helmut Hund, 008.0403.0)
fitted with a Nikon 6X wide optical zoom
182
camera. Oral disc diameter (ODD) was meas‐
ured using a MitutoyoTM dial digital vernier
calliper to the nearest 0.1 mm. For bucco‐
phryngeal analyses, specimens were dissected
following Wassersug (1976) to separate the
buccal roof and floor and stained with methyl‐
ene blue solution (2%) to enhance contrast.
Scanning Electron Microscopy
The oral apparatus were cleaned with double
distilled water to remove any unwanted materi‐
al prior to scanning electron microscopy. The
samples (n=5, Gosner stage 36) were fixed
with 2.5% glutaraldehyde solution in 0.1 M
phosphate buffer at room temperature for four
hours, and then post-fixed in 1% osmium tetr‐
oxide buffer for 1 hour at 4°C. The fixative and
buffer were kept at a pH of 7.4 throughout the
experiment. The dehydrated samples (through
ascending ethanol grades) were fixed on the
appropriate stubs, and a conductive coating
sputtered with Gold/Palladium for about 20
minutes using an ion sputter coater. The tissues
were studied using a Hitachi (model S3400N)
scanning electron microscope in the secondary
electron emission mode at an accelerating volt‐
age of 20 KV.
Morphological Abbreviations
AL, anterior labium; ANP, anterior narial
wall papilla; BFA, buccal floor arena; BFAP,
buccal floor arena papillae; BP, buccal pockets;
BRAP, buccal roof arena papilla; BRAPU,
buccal roof arena pustules; DV, dorsal velum;
GC, gill clefts; GZ, glandular zone; LJ, lower
jaw sheath; LP, lingual papillae; LRP, lateral
ridge papilla; M, mouth; MP, marginal papil‐
lae; MRP, median ridge papilla; SMP, sub-
marginal papillae; OD, oral disc; ODD, oral
disc diameter; PL, posterior labium; PLA, pre-
lingual arena; PLP, pre-lingual papilla; PNA,
pre-narial arena; PNR, pre-narial ridge; PNP,
posterior narial papilla; PNW, posterior narial
wall; SEM, scanning electron microscopy; TA,
tongue anlage; UJ, upper jaw sheath; VV, ven‐
tral velum.
Current Herpetol. 41(2) 2022
Study of the gut contents
To analyse the dietary and feeding behaviour
of tadpoles, tadpoles (n=10 for each develop‐
mental stage [Gosner stage 25–41]) were pre‐
served in 10% formalin immediately after field
collection in order to prevent the digestion of
ingested food particles. The gut of each tadpole
was carefully removed, and the length of each
intestine was measured with a digital vernier
callipers (MitutoyoTM to nearest 0.1 mm). Diet
analysis focused on the contents of first four
centimetres of the gut. The gut contents were
cleansed with distilled water, transferred to a
Sedgewick rafter chamber, and studied under a
compound microscope (Laboscope, CMS-2)
coupled to a Sony cyber shot camera (5.1
megapixels, DCSW5) to capture pictures of the
gastrointestinal contents.
The food items were recognised and quanti‐
fied up to the genus level using standard proce‐
dures (Edmondson, 1959; Shamsudin and
Shazili, 1991; Battish, 1992; Smith, 1994). The
food items of tadpoles were assessed using
numeric frequency (NF%=total number of
food items of a specific food group consumed/
total number of items of all food groups con‐
sumed×100) and frequency of occurrence
(FO%=number of guts in which the specific
food item was present/total number of guts
with these food items×100). The importance
index was calculated following Colli et al.
(2003). The term detritus was used to classify
unidentified items that formed a mass of
organic material. The gut contents of tadpoles
were found to contain 69.62% of the plankton
genera recorded from representative water
samples of the study area (Table 1). The com‐
parison of species diet diversity was evaluated
using an individual-based rarefaction method,
because genus (plankton) richness is strongly
related to the number of observations (Gotelli
and Colwell, 2001). Individual guts were used
as samples and the diversity of plankton was
used as the food richness. A total of 170 tad‐
poles were included from across the sampling
locations and statistical analyses used the
PAST software 2.14 (Hammer et al., 2017).
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES 183

Table 1. Comparision of planktons of the tadpoles of Polypedates maculatus in wild and gut (n=170)
from Odisha.
Habitat Gut of tadpoles
Class Genus
%NF %FO I %NF %FO I
Bacillariophyceae Achnanthes + 0.86 69.57 35.21 – – – –
Achnanthidium + 2.03 100 51.01 – – – –
Actinella + 0.57 66.87 33.72 – – – –
Amphipleura + 1.96 100 50.98 + 2.60 93.02 47.81
Amphora + 0.41 70.75 35.58 + 0.53 73.56 37.04
Asterionella + 1.05 74.53 37.79 – – – –
Aulacoseira + 0.83 88.42 44.62 – – – –
Bacillaria + 0.15 74.56 37.35 + 0.13 77.23 38.68
Brachysira + 0.71 78.21 39.46 – – – –
Cocconeis + 1.04 80.13 40.58 + 0.73 86.42 43.57
Craticula + 1.12 82.31 41.71 + 2.31 93.16 47.73
Cyclotella + 1.79 86.55 45.06 + 5.08 100 52.54
Cymatopleura + 1.01 82.68 41.84 – – – –
Cymbella + 1.35 86.57 43.96 + 4.32 100 52.16
Diadesmis + 1.6 92.10 46.85 + 2.38 100 51.19
Diatoma + 1.7 95.32 48.51 + 0.64 67.23 33.93
Ditylum + 0.16 74.25 37.20 + 0.12 65.62 32.87
Eunotia + 1.55 92.23 46.89 + 0.31 81.42 40.86
Fragilaria + 1.48 88.17 44.82 – – – –
Fragilariforma + 1.06 82.63 41.84 – – – –
Frustulia + 0.98 70.04 35.51 + 1.14 100 50.57
Geminella + 1.62 89.22 45.42 – – – –
Gomphoneis + 1.77 85.93 43.85 – – – –
Gomphonema + 1.75 92.87 47.31 + 1.41 86.43 43.92
Gyrosigma + 1.1 95. 63 48.36 + 0.13 72.76 36.44
Navicula + 3.8 100 51.9 + 6.37 100 53.18
Nitzschia + 1.8 100 50.9 + 4.37 93.02 48.69
Pinnularia + 1.12 100 50.56 + 6.93 100 53.46
Placoneis + 1.08 96.72 48.9 – – – –
Sellaphora + 0.92 78.83 39.87 – – – –
Stauroneis + 0.52 61.87 31.19 + 3.12 86.02 44.57
Synedra + 1.52 88.20 44.86 + 5.58 97.07 51.32
Tabellaria + 0.12 56.50 28.31 + 0.06 50.05 25.05
Chlorophyceae Ankyra + 1.01 98.26 49.63 – – – –
Anadyomene + 0.98 66.32 33.65 – – – –
Tetraedron + 0.16 52.43 26.29 – – – –
Scotiella + 1.02 86.50 43.76 – – – –
Tetrastrum + 0.88 75.12 38 – – – –
Cladophora + 2.09 100 51.04 – – – –
Klebsormidium + 0.33 73.42 36.87 – – – –
Ankistrodesmus + 3.06 100 51.53 + 1.90 100 50.95
Actinastrum + 0.22 69.32 34.77 + 0.08 23.4 11.74
Chlamydomonas + 0.37 72.67 36.52 + 2.36 100 51.18
Coelastrum + 2.34 100 51.17 + 5.05 98.09 51.57
Kirchneriella + 1.21 95.09 48.15 + 0.69 78.43 39.56
184 Current Herpetol. 41(2) 2022

Table 1. (continued)
Habitat Gut of tadpoles
Class Genus
%NF %FO I %NF %FO I
Microspora + 0.84 57.84 29.34 + 0.61 78.22 39.41
Oedogonium + 1.75 91.59 46.67 + 0.10 19.23 9.66
Oocystis + 1.01 78.66 39.83 + 0.41 90.42 45.41
Pandorina + 0.46 82.50 41.48 + 0.35 84.76 42.55
Pediastrum + 1.41 90.02 45.71 + 0.03 7.8 3.91
Scenedesmus + 1.6 70.62 36.11 + 7.63 60.62 34.12
Spirogyra + 3.63 100 51.81 + 0.67 68.76 34.71
Volvox + 0.07 12.80 6.43 + 0.04 7.9 3.97
Zygnema + 1.43 99.64 50.53 + 4.41 86.82 45.61
Zygnematophyceae Cosmarium + 0.68 85.21 42.94 + 3.19 81.16 42.17
Closterium + 0.9 91.67 46.28 + 5.16 84.22 44.69
Desmidium + 2.02 100 51.01 + 3.16 88.03 45.59
Euastrum + 0.6 65.20 32.9 + 0.74 86.36 43.55
Mougeotia + 1.61 69.03 35.32 + 1.94 66.83 34.38
Pleurotaenium + 1.18 68.10 34.64 + 0.37 47.62 23.99
Staurodesmus + 0.8 67.23 34.01 + 0.31 57.82 29.06
Staurastrum + 0.42 64.87 32.64 + 0.52 69.22 34.87
Xanthidium + 0.12 53.74 26.93 + 0.10 43.63 21.86
Cyanophyceae Anabaena + 0.26 62.53 31.39 – – – –
Gloeothece + 0.08 54.92 27.5 – – – –
Gomphosphaeria + 0.16 73.23 36.69 – – – –
Chroococcus + 0.83 68.12 34.47 + 0.56 62.76 31.66
Microcystis + 1.93 96.65 49.29 + 0.17 42.83 21.50
Merismopedia minima + 2.77 100 51.38 + 2.03 80.22 41.12
Oscillatoria + 3.2 100 51.6 + 0.34 81.16 40.75
Euglenophyceae Euglena caudata + 1.1 85.66 43.38 + 0.36 85.62 42.99
Phacus acutus + 1.98 95.42 48.7 + 2.30 72.83 37.56
Trachelomonas dubia + 1.77 83.20 42.48 + 0.41 99.56 49.98
Ulvophyceae Ulothrix tenuissima + 2.26 100 51.13 + 4.12 99.01 51.56
Rotifera Monostyla + 0.14 82.95 41.54 – – – –
Lecane + 1.27 86.62 43.94 + 0.13 70.6 35.36
Lepadella + 0.21 83.28 41.74 + 0.60 65.56 33.08
Platyias + 0.11 82.49 41.3 + 0.26 80.20 40.23
Copepoda Eucyclops + 7.2 100 53.6 + 0.64 68.03 34.33
Detritus + + + + + – 100 –

+ = Occurrence, – = Non occurrence

Results ners; both labia were of equal size. Jaw sheaths

Oral morphology
The oral disc was anteroventral in position
and emarginated. On both the labia and the lat‐
eral corners, there was a single row of MP; two
to three SMP were present at the lateral cor‐
were well developed, moderately keratinized,
completely serrated with uniform-sized serra‐
tions; the supra-rostrodont was arch-shaped
with thin lateral process; the infra-rostrodont
was U-shaped, convex laterally and concave
medially. The formula for the labial tooth row
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES
Fig. 1. Oral disc of Polypedates maculatus
tadpoles (Gosner stage 36) under (A) Light
Microscope and (B) Scanning Electron Microscope;
(C) Labial teeth showing distinct head (H), neck (N)
and base (B).
was A4(3)/P3(1) (Fig. 1A, B). The order of the
length of denticle rows was A2>P1>P2>A1>
P3>A3>A4. Denticles were elongated, fairly
spaced, and sharply curled towards the mouth
at their apex. The sheath and the body were
thin. Serrations on the jaw sheath were spaced
with a wide base and short triangular pointed
head (Fig. 1C). The jaw sheaths and kerato‐
donts disappeared post the emergence of the
forelimbs at stage 42. From stage 42 onwards,
the mouth gradually opens wider, and by stage
44, the angle of the mouth has reached the
middle of the eye. After stage 42, feeding
activity generally ceases, and the larvae fast
during the period of intestinal remodelling.
Buccopharyngeal morphology
Buccal roof (Fig. 2A): The PNA consisted
of an arched pustulose ridge with pustulations,
with six pustules on either side posterolateral
to the ridge. Internal nares are transversely ori‐
185
entated; both nares were separated posterome‐
dially by a distance of about half the length of
each naris; smooth ANW with a few pustules
and a tall conical papilla originating near the
wall’s lateral corner; the PNW was tall,
smooth, and valvular with a flap-like part at
the medial. The post-narial arena was made up
of two tall conical papillae that were placed
linearly and orientated medially behind the
PNW; the papillae near the roof’s middle plane
were larger, while those near the lateral corners
were smaller; the front margin of the larger
papillae was irregular, and was the largest of
all the papillae on the roof; few pustules (three
to four) were found in front of the MR. The
MR was made up of a triangular flap with a
pustulated edge and a lengthy protrusion medi‐
ally with a pustulated tip. On each side of the
lateral wall, perpendicular to the MR, a long
LRP with a corrugated anterior surface was
found. Two pairs of long conical papillae on
the lateral edge of the roof demarcated the
BRA; approximately 30–40 small pustules
covered the entire BRA. The anterior boundary
of the prominent GZ was defined by small pus‐
tules that run parallel to the posterior boundary
of BRA. Secretory pits were present all across
the GZ. The DV margin was raised, fractured
medially, and slightly pustulose on the surface.
Buccal floor (Fig. 2B): The dorsolateral part
of the PLA had four pairs of pustules, and the
posterolateral corners had an equal number of
palps. A pair of stubby papillae on the PLA are
positioned posteromedially between the two
posterior palps. The posterolateral corner con‐
tained four palps; the posterior most palps
being the longest, with a slightly dilated tip
and a corrugated surface with numerous pustu‐
lations; the palp just front of it was smaller, but
it had all of the morphological characteristics
of the former; the remaining two palps were
stubby and short. The TA was broad and rose;
at the centre of the tongue, there are two
smooth long LP with a broad tip. Ten BFAP
defined the arena on the BF; the papillae of the
BFA began at the lateral corners of the roof,
anterior to the BP, and converged down to the
floor’s posteromedial region; these papillae
186
Fig. 2. Buccophryngeal cavity of tadpoles of
Polypedates maculatus (Gosner stage 36): (A)
buccal roof; (B) buccal floor.
were short, stubby and unequal in size; the tips
of several papillae were dilated. About 20 pus‐
tules were scattered around the BFA, primarily
at the posterior end. There were a few papillae
and pustules beyond the lateral borders of the
BFA. There were 20–25 pustulations on each
side of the space between the TA and the BP.
The pockets in the buccal mucosa were oblique
and broad; no papillae found in the pockets. A
few pustules were gently scattered medially
behind the BFA and along the margin of the
VV. The VV was sinuate and broad with six
projections at its ridge. The three median pro‐
jections were close together and centred; the
distance between the outer three projections
was considerable. There was a conspicuous
median notch; with several secretory pits, the
outer border was granular. The glottis was par‐
tially hidden underneath the VV. Below the
VV; there were four pairs of brachial basket
arches.
Current Herpetol. 41(2) 2022
Analysis of gut contents
Tadpoles of P. maculatus began feeding at
stage 25 (Appendix I) by which they had
developed a prominent mouth and complete
operculum, features essential for feeding. A
majority of the diet of tadpoles was composed
of detritus followed by phytoplankton, green
algae, and diatoms. Zooplankton in the diet
included Rotifers and Copepods. A total of
1268 food items were identified, which were
divided into 55 taxonomic groups: 20 Bacillar‐
iophyte genera, 14 Chlorophyte genera, nine
Zygnematophyte genera, four Cyanophyte
genera, three Euglenophyte genera, one Ulvo‐
phyte genera, three genera of Rotifera and a
single Copepoda genus (Appendix I).
The Copepod Eucyclops were the most
abundant plankton in our study area (Table 1),
but tadpoles least prefered them as a food
source. Plankton such as Achnanthes, Achnan‐
thidium, Actinella, Asterionella, Aulacoseira,
Brachysira, Cymatopleura, Fragilaria, Fragi‐
lariforma, Geminella, Gomphoneis, Placoneis,
Sellaphora, Ankyra, Anadyomene, Tetraedron,
Scotiella, Tetrastrum, Cladophora, Klebsormi‐
dium, Anabaena, Gloeothece, Gomphosphae‐
ria, and Monostyla were present in the study
area but tadpoles did not choose to them as
their diet.
Pinnularia (NF%=6.93) was the major food
item found among Bacillariophyceae in the gut
of tadpoles (Table 1) as its %NF is significant‐
ly higher than the frequency expected from its
availability in wild (NF%=1.12). Scenedesmus
among the Chlorophyceae had the highest
numeric frequency (NF%=7.63). Among the
Zygnematophyceae; Closterium were promi‐
nent (NF%=5.16). Merismopedia in the Cya‐
nophyceae, Phacus in the Euglenophyceae,
and Ulothrix in the Ulvophyceae were other
important items in the gut. Scenedesmus had
the highest numeric frequency (7.63%) among
all the food items, while Pediastrum had the
lowest (0.03%) (Table 1). Detritus was de‐
voured by all the tadpoles (Table 1). Based on
the importance index scores of over 50,
Ankistrodesmus, Chlamydomonas, Coelastrum,
Cyclotella, Cymbella, Diadesmis, Frustulia,
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES
Fig. 3. Individual-based rarefaction curve of food items of Polypedates maculatus tadpoles. Mid line is
the rarefaction curve of plankton communities from the study sites. Upper and lower curves represent the
upper and lower limits of 95% confidence intervals.
Navicula, Pinnularia, Synedra, and Ulothrix
fragments were important food items (Table 1).
Individual-based rarefaction curve gives the
information about how well the species have
been sampled (Hughes et al., 2001). The more
concave-downward the curve, the better sam‐
pled the species (Hughes et al., 2001). We ana‐
lysed the data sets by the number of food items
recorded in the study area to compare the
shape of the individual-based species accumu‐
lation curves. A stabilizing curve (curved
downward) was observed suggesting the sam‐
pling effort to be complete (Fig. 3).
Discussion
The elements of oral apparatus are universal
to all anuran tadpoles. The keratinized labial
tooth of Polypedates maculatus is derived from
cells at the base of the tooth ridge and consists
of three indistinct regions: a distal head, an
intermediate body, and a basal, hollow sheath.
Throughout their larval development, they
have a single tooth row per ridge (=uniserial).
Most tadpoles have a single tooth row per
ridge (=uniserial) but tadpoles with bi- (e.g.,
discoglossids) and even tri- or multi-serial
rows have been observed (Altig and
McDiarmid, 1999). The jaw sheaths have a
broad base and a short serrated edge, like those
187
of a typical tadpole. The lateral margins of jaw
sheaths are formed by the fusion of palisades
of keratinized cells. Jaw sheaths with 30–80
serrations per millimetre are common. The cur‐
rent oral apparatus arrangement in the tadpoles
of P. maculatus (stages 25–41) (keratinized
jaw sheaths and oral disc with keratinized
tooth rows) is most likely a permanent trait
(Altig and Johnston, 1989). The antero-ventral
position of oral disc indicates that they have
generalized mouthparts, primarily detritus
feeders, but may also graze on algal vegetation
and filter feeding on the planktonic bloom of
ponds (Khan and Mufti, 1994). The keratinized
rows of labial teeth and jaw sheaths together
remove materials from the substrate during
active grazing (Johnston, 1982; Carr and Altig,
1991; Taylor et al., 1995; Wassersug and
Yamashita, 2001; Larson and Reilly, 2003). In
addition, the number and density of labial
teeth, as well as the arrangement of tooth rows,
may represent adaptation to different foraging
modes (Altig, 2006). The bases of submarginal
papillae of P. maculatus are circular which
supports the views of Altig and McDiarmid
(1999) who proposed that the bases of submar‐
ginal papillae are usually circular. The margin‐
al papillae are thought to have chemosensory
and tactile sensors (McDiarmid and Altig,
1999), though the significance of the papillary
188
Table 2. Comparison of oral apparatus among of Polypedates maculatus, P. teraiensis, P. macrotis, P.
colletti, and P. otilophus.
Observation P. maculatus P. teraiensis
Chakravarty et al.,
2011
Pre-narial arena 6 3–4
papillae
Postnarial papillae 2 2
Buccal roof arena 4 12
papillae
Buccal roof arena 30 50
pustules
Lingual papillae 1 pair 1 pair
Buccal floor arena 10 12
papillae
NA=Not Available
arrangement variation is unknown (Bonacci et
al., 2008).
One of the most common methods of identi‐
ficating anuran tadpoles is the labial tooth row
formula (LTRF). The oral structure of P. macu‐
latus was found to be A4(3)/P3(1) in the cur‐
rent study. The most common configuration is
a labial tooth row formula (LTRF) of 2/3, but
the number can range from 0/0 to 17/21 in
many different combinations throughout Anura
(Altig, 2007). The variation in oral morpholo‐
gy of anuran tadpoles demonstrates that the
structure of mouth and buccal cavities of
anuran larvae is highly adaptive and correlated
with feeding ecology. The current findings
show that the LTRF is consistent throughout
the developmental stages of P. maculatus. At
stage 23, the oral disc and keratodont rows
begin to differentiate, and by stage 25, the ker‐
atodont rows are fully distinct. Through stages
26 to 41, the upper labium has one uninterrupt‐
ed row and three interrupted rows of kerato‐
donts, whereas the lower labium already has all
three rows, with a large gap in the first row
(A4[3]/P3[1]). Such a gap in the posterior jaw
sheath has also been reported for P. macrotis,
P. colletti, and P. otilophus (Inger, 1985), as
well as P. megacephalus (Tesia et al., 2017).
Buccopharyngeal characters are important in
Current Herpetol. 41(2) 2022
P. macrotis P. colletti P. otilophus
Inger, 1985 Inger, 1985 Inger, 1985
3–4 3–4 3–4
1 1 1
NA NA NA
NA NA NA
1 pair 1 pair 1 pair
5–10 5–10 5–10
comparative studies because adult and tadpole
external morphology have not provided
enough information to diagnose species of
Polypedates in India. Therefore, the current
findings may be used to augment existing
taxonomic features among rhacophorids (Table
2). Although both the rhacophorid species P.
maculatus and P. teraiensis in Odisha shared
some similar features (Appendix II), but the
adults and tadpoles of both these species are
difficult to distinguish, so buccophryngeal
characters can help differentiate P. maculatus
from other species (Table 2).
The morphology of the oral and buccophar‐
yngeal structures of anuran tadpoles reflects
their feeding preferences. The results of the gut
content analyses revealed that, aside from a lot
of debris, the tadpole diet was mostly made up
of microalgae and diatoms, an observation
similar to Lajmanovich (2000) and Rossa-
Feres et al. (2004). Prey items were mostly
from classes Bacillariophyceae, Chlorophy‐
ceae, Zygnematophyceae, Euglenophyceae,
Cyanophyceae, Ulvophyceae, Rotifera, and
Copepoda. Consumption of a large number of
algal species places tadpoles in an important
position within the food web in which they
may have an important influence on energy
flow. Members of the class Bacillariophyceae
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES
were shown to be the most important prey cat‐
egory in the diets of P. maculatus tadpoles, a
finding comparable to Sinha et al. (2001) and
Lalremsanga et al. (2013). Bacillariophyceae
members may be the most abundant food
resource in their habitat or may be higher in
calories due to their fat content, and their
intake is more convenient than filamentous
algae (Kupferberg et al., 1994). Bacillario‐
phytes have also been cited as the most fre‐
quently consumed item in the diets of other
tadpoles of lotic (Santos et al., 2016) and lentic
(Rossa-Feres et al., 2004; Echeverría et al.,
2007; Bionda et al., 2012; Huckembeck et al.,
2016; Mahapatra et al., 2017; Rout et al., 2018)
environments, indicating that high levels of
diatoms in intestinal contents of tadpoles can
be considered a general phenomenon. This
information is relevant to understanding food
preferences of anurans and environmental
monitoring, including studies that evaluate the
presence or absence of certain diatomaceous
groups which can be considered indicators of
environmental quality (Cejudo-Figueiras et al.,
2010; Antón-Garrido et al., 2013). Chlorophyt‐
ic algae are also significant food items due to
their rich carbohydrate content (Bold and
Wynne, 1985; Volkman and Brown, 2005).
Carbohydrates are the important source for the
development of hind limbs and the growth of
tadpoles (Emmett and Allen, 1919). Narzary
and Bordoloi (2012) also reported that among
zooplankton Rotifers and Copepods are impor‐
tant food items for tadpoles. Sinha et al. (2001)
identified tadpoles as phyto- and zooplankton
feeders, which is consistent with the findings
of our study. Knowledge of food and feeding
behaviour of the tadpoles is important because
the early life history of amphibians depends on
the availability of the food items in their habi‐
tat (Sinha et al., 2001). The oral morphological
features and feeding habits may prove useful in
the species identification and the future conser‐
vation efforts, especially in selecting suitable
habitat for this species and captive breeding.
189
Acknowledgments
The authors are thankful to SERB, Govern‐
ment of India for financial assistance to GS
and CSIR, Government of India for Junior
Research Fellowship (File No. 09/173(0134)/
2020-EMR-I) to JR. We acknowledge the help
of the Central Instrumentation Facility, Orissa
University of Agriculture & Technology, Bhu‐
baneswar, Odisha for SEM analyses of the
samples.
Literature Cited
Altig, R. 2006. Discussions of the origin and evo‐
lution of the oral apparatus of Anuran tadpoles.
Acta Herpetologica 1: 95–105.
Altig, R. 2007. A primer for the morphology of
Anuran tadpoles. Herpetological Conservation
and Biology 2: 71–74.
Altig, R. and Johnston, G. F. 1989. Guilds of
anuran larvae: relationships among developmen‐
tal modes, morphologies, and habitats. Herpeto‐
logical Monographs 3: 81–109.
Altig, R. and McDiarmid, R. W. 1999. Body plan.
Development and morphology. p. 24–51. In: R.
W. McDiarmid and R. Altig (eds.), Tadpoles. The
Biology of Anuran Larvae. University of Chicago
Press, Chicago.
Altig, R. and Pace, W. L. 1974. Scanning electron
photomicrographs of tadpole labial teeth. Journal
of Herpetology 8: 247–251.
AmphibiaWeb. 2021. University of California, Ber‐
keley. https://amphibiaweb.org (accessed 7 De‐
cember 2021)
Antón-Garrido, B., Romo, S., and Villena, M. J.
2013. Diatom species composition and indices
for determining the ecological status of coastal
Mediterranean Spanish lakes. Anales del Jardín
Botánico de Madrid 70: 122–135.
Ao, M. and Khare, M. K. 1986. Diagnostic fea‐
tures of Hyla annectans Jerdon tadpoles (Anura:
Hylidae). Asian Journal of Experimental Science
1: 30–36.
Aravind, N. A. and Gururaja, K. V. 2011. Theme
paper on “Amphibians of the Western Ghats”
Report submitted to Western Ghats Ecology
Expert Panel, MoEF, Govt. of India.
190
Battish, S. K. 1992. Freshwater Zooplankton of
India. Oxford and IBH Publishing Co., New
Delhi.
Bionda, C. D. L., Luque, E., Gari, N., Salas, N.
E., Lajmanovich, R. C., and Martino, A. L.
2012. Diet of tadpoles of Physalaemus biligoni‐
gerus (Leiuperidae) from agricultural ponds in
the central region of Argentina. Acta Herpetolog‐
ica 8: 141–146.
Bold, H. C. and Wynne, M. J. 1985. Introduction
to the Algae. Structure and Reproduction. 16th
Edition. Prentice-Hall, Englewood Cliffs.
Bonacci, A., Brunelli, E., Sperone, E., and
Tripepi, S. 2008. The oral apparatus of tadpoles
of Rana dalmatina, Bombina variegata, Bufo
bufo, and Bufo viridis (Anura). Zoologischer An‐
zeiger 247: 47–54.
Bowatte, G. and Meegaskumbura, M. 2011. Mor‐
phology and ecology of tadpoles of Ramanella
obscura (Anura: Microhylidae). Ceylon Journal
of Science 40: 109–120.
Carr, K. M. and Altig, R. 1991. Oral disc muscles
of anuran tadpoles. Journal of Morphology 208:
271–277.
Cejudo-Figueiras, C., Blanco, S., Álavarez-
Blanco, I., Ector, L., and Bécares, E. 2010.
Nutrient monitoring in Spanish wetlands using
epiphytic diatoms. Vie & Milieu—Life and Envi‐
ronment 60: 89–94.
Chakravarty, P., Bordoloi, S., Grosjean, S.,
Ohler, A., and Borkotoki, A. 2011. Tadpole
morphology and table of developmental stages of
Polypedates teraiensis (Dubois, 1987). Alytes 27:
85–115.
Colli, G. R., Mesquita, D. O., Rodrigues, P. V.
V., and Kitayama, K. 2003. Ecology of the
Gecko Gymnodactylus geckoides amarali in a
neotropical savanna. Journal of Herpetology 37:
694–706.
Das, I. 1994. The internal morphology of some
anuran larvae from south India: a scanning mi‐
croscopic study. Amphibia-Reptilia 15: 249–256.
Dinesh, K. P., Radhakrishnan, C., Gururaja, K.
V., and Bhatta, G. K. 2009. An annotated
checklist of amphibia of India with some insights
into the patterns of species discoveries, distribu‐
tion and endemism. Records of Zoological Survey
of India, Kolkata 302: 1–153.
Current Herpetol. 41(2) 2022
Dinesh, K. P., Radhakrishnan, C.,
Channakeshavamurthy, B. H., Deepak, P., and
Kulkarni, N. U. 2017. Checklist of Amphibia of
India. http://mhadeiresearchcenter.org/resources
(updated till April 2017)
Dutta, S. K., Nair, M. V., Mohapatra, P. P., and
Mohapatra, A. K. 2009. Amphibians and Rep‐
tiles of Similipal Biosphere Reserve. Regional
Plant Resource Centre, India.
Echeverría, D. D., Volpedo, A. V., and Mascitti,
V. I. 2007. Diet of tadpoles from a pond in
Iguazu National Park, Argentina. Gayana 71: 8–
14.
Edmondson, W. T. 1959. Freshwater Biology. 2nd
Edition. John Wiley and Sons Inc., New York.
Emmett, A. D. and Allen, F. P. 1919. Nutritional
studies on the growth of frog larvae (Rana
pipiens). I. Journal of Biological Chemistry 38:
325–344.
Frost, D. R. 2021. Amphibian Species of the World:
an Online Reference version 6.1. American
Museum of Natural History. https://amphibian
softheworld.amnh.org/index.php (accessed 31
March 2021)
Gosner, K. L. 1960. A simplified table for staging
anuran embryos and larvae, with notes on identi‐
fication. Herpetologica 16: 183–190.
Gotelli, N. J. and Colwell, R. K. 2001. Quanti‐
fying biodiversity: procedures and pitfalls in the
measurement and comparison of species richness.
Ecology Letters 4: 379–391.
Grosjean, S. 2003. A re-description of the external
and buccopharyngeal morphology of the tadpoles
of Ophryophryne microstoma Boulenger, 1903
(Megophryidae). Alytes 21: 45–58.
Grosjean, S., Vences, M., and Dubois, A. 2004.
Evolutionary significance of oral morphology in
the carnivorous tadpoles of tiger frogs genus
Hoplobatrachus (Ranidae). Biological Journal of
the Linnean Society London 81: 171–181.
Hammer, Ø., Harper, D. A. T., and Ryan, P. D.
2017. PAST: Paleontological statistics software
package for education and data analysis, version
3.16. Available via http://folk.uio.no/ohammer/
past/
Hora, S. L. 1922. Some observations on the oral
apparatus of the tadpoles of Megalophrys parva
Boulenger. Proceedings of the Asiatic Society of
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES
Bengal 17: 9–15.
Huckembeck, S., Alves, L. T., Loebmann, D., and
Garcia, A. M. 2016. What the largest tadpole
feeds on? A detailed analysis of the diet composi‐
tion of Pseudis minuta tadpoles (Hylidae, Den‐
dropsophini). Anais da Academia Brasileira de
Ciências 88: 1397–1400.
Hughes, J. B., Hellmann, J. J., Ricketts, T. H.,
and Bohannan, B. J. M. 2001. Counting the un‐
countable: Statistical approaches to estimating
microbial diversity. Applied and Environmental
Microbiology 67: 4399–4406.
Inger, R. F. 1985. Tadpoles of the forested regions
of Borneo. Fieldiana: Zoology, New Series 26: 1–
89.
Inthara, C., Lauhachinda, V., Nabhitabhata, J.,
Chuaynkern, Y., and Kumtong, P. 2005.
Mouth part structures and distribution of some
tadpoles from Thailand. Journal of Thailand Nat‐
ural History Museum 1: 55–78.
Johnston, G. F. 1982. Functions of the Keratinized
Oral Features in Anuran Tadpoles with an Analy‐
sis of Ecomorphological Tadpole Types. Unpub‐
lished master’s thesis. Mississippi State Univer‐
sity, Mississippi.
Khan, M. S. and Mufti, S. A. 1994. Oral disc mor‐
phology of amphibian tadpole and its functional
correlates. Pakistan Journal of Zoology 26: 25–
30.
Khare, M. K. and Sahu, A. K. 1984. Diagnostic
features of Rana danieli (Anura: Ranidae) tad‐
poles. Amphibia-Reptilia 5: 275–280.
Khongwir, S., Iangrai, A., and Hooroo, R. N. K.
2003. Development of mouth parts and food
choice in the tadpoles of Rhacophorus maximus.
Uttar Pradesh Journal of Zoology 23: 101–104.
Kiffney, P. M. and Richardson, J. S. 2001. Inter‐
actions among nutrients, periphyton, and inverte‐
brate and vertebrate (Ascaphus truei) grazers in
experimental channels. Copeia 2: 422–429.
Kupferberg, S. J., Marks, J. C., and Power, M. E.
1994. Effects of variation in natural algal and de‐
trital diets on larval anuran (Hyla regilla) life-
history traits. Copeia 1994: 446–457.
Lajmanovich, R. C. 2000. Interpretación ecológica
de una comunidad larvaria de anfibios anuros. In‐
terciencia 25: 71–79.
Lalremsanga, H. T., Hooroo, R. N. K., and
191
Lalronunga, S. 2013. Oral morphology and
feeding behaviour of Euphlyctis cyanophlyctis
tadpoles (Schneider, 1799) with notes on feeding
behaviour. Science and Technology Journal 1:
58–69.
Larson, P. M. and Reilly, S. M. 2003. Functional
morphology of feeding and gill irrigation in the
anuran tadpole: electromyography and muscle
function in larval Rana catesbeiana. Journal of
Morphology 255: 202–214.
Lee, J. C. 1976. Rana maculatus Brocchi, an addi‐
tion to the herpetofauna of Belize. Herpetologica
32: 211–214.
Mahapatra, S., Rout, J., Sahoo, G., and Sethy, J.
2017. Dietary preference of Euphlyctis cyano‐
phlyctis tadpoles in different habitats in and
around Simlipal biosphere reserve, Odisha, India.
International Journal of Conservation Science 8:
259–268.
McDiarmid, R. W. and Altig, R. 1999. Tadpoles:
The Biology of Anuran Larvae. The University of
Chicago Press, Chicago.
Mohapatra, P. P., Mahapatra, C., and Dutta, S.
K. 2017. Studies on Amphibians of Odisha: an
overview. p. 11–32. In: A. Das (ed.), Diversity
and Ecology of Amphibians of India. Wildlife In‐
stitute of India, Dehradum.
Narzary, J. and Bordoloi, S. 2012. A study on
certain common pond breeding anurans and their
tadpoles in a pond of western Assam, India. In‐
ternational Journal of Advanced Biological Re‐
search 2: 342–348.
Rossa-Feres, D. C., Jim, J., and Fonseca, M. G.
2004. Diets of tadpoles from a temporary pond in
south-eastern Brazil (Amphibia, Anura). Revista
Brasileira de Zoologia 21: 745–754.
Rout, J., Prusty, S., and Sahoo, G. 2018. Assess‐
ing feeding habits of tadpoles of common Asian
toad Duttaphrynus melanostictus. Pranikee Jour‐
nal of Zoological Society of Orissa 30: 47–57.
Saidapur, S. K. 2001. Behavioural ecology of
anuran tadpoles: The Indian Scenario. Proceed‐
ings of the Indian National Science Academy 6:
311–322.
Santos, F. J. M., Protázio, A. S., Moura, C. W.
N., and Juncá, F. A. 2016. Diet and food re‐
source partition among benthic tadpoles of three
anuran species in Atlantic forest tropical streams.
192
Journal of Freshwater Ecology 31: 53–60.
Sekar, A. G. 1990. Notes on morphometry, ecology,
behaviour and food of tadpoles of Rana curtipes
Jerdon, 1853. Journal of the Bombay Natural
History Society 87: 312–313.
Shamsudin, L. and Shazili, N. A. M. 1991. Micro‐
planktonic bloom in a brackish water lagoon of
Terengganu. Environmental Monitoring and As‐
sessment 19: 287–294.
Sinha, B., Chakravorty, P., Borah, M. M., and
Bordoloi, S. 2001. Qualitative analysis of food
spectrum of five species of anuran tadpoles from
Arunachal Pradesh, India. Zoos’ Print Journal
16: 514–515.
Smith, G. M. 1994. Manual of Phycology: An Intro‐
duction to the Algae and their Biology. Scientific
Publishers, Jodhpur.
Taylor, C. L., Altig, R., and Boyle, C. R. 1995.
Can Anuran tadpoles choose among foods that
vary in quality? Alytes 13: 81–86.
Tesia, C., Ohler, A., and Bordoloi, S. 2017. De‐
velopment of Polypedates megacephalus (Hallo‐
well, 1861) (Anura, Rhacophoridae). Alytes 34:
Current Herpetol. 41(2) 2022
49–60.
Volkman, J. K. and Brown, M. R. 2005. Nutri‐
tional value of microalgae and applications. p.
407–457. In: S. Rao (ed.), Algal Cultures, Ana‐
logues of Blooms and Applications. Science Pub‐
lishers, Enfield.
Wassersug, R. J. 1976. Oral morphology of anuran
larvae: terminology and general description. Oc‐
casional Papers of the Museum of Natural His‐
tory, University of Kansas 48: 1–23.
Wassersug, R. J. 1980. Internal oral features of tad‐
poles from eight anuran families: functional, sys‐
tematic, evolutionary and ecological considera‐
tions. Miscellaneous publication, Museum of
Natural History, University of Kansas 68: 1–146.
Wassersug, R. J. and Yamashita, M. 2001. Plasti‐
city and constraints on feeding kinematics in
Anuran larvae. Comparative Biochemistry and
Physiology A 131: 183–195.
Accepted: 18 May 2022
ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES 193

Appendix I
Diets of Polypedates maculatus tadpoles in different stages of development.
Food items Stage 25 Stage 32 Stage 38 Stage 41
Phytoplanktons
Bacillariophyceae Amphipleura – – – +
Amphora – – + +
Bacillaria – + + +
Cocconeis + + + +
Craticula – + + +
Diadesmis + + + +
Diatoma – + + +
Ditylum – + + +
Eunotia + + + +
Frustulia + + + +
Cyclotella + + + +
Cymbella + + + +
Gomphonema – + + +
Gyrosigma + + + +
Navicula + + + +
Nitzschia + + + +
Pinnularia + + + +
Stauroneis + + + +
Tabellaria – + + +
Synedra – – + +
Chlorophyceae Actinastrum – + + +
Ankistrodesmus – – + +
Chlamydomonas – + + +
Coelastrum – + + +
Kirchneriella – – + +
Microspora – + + +
Oedogonium + + + +
Oocystis – + + +
Pandorina + + + +
Pediastrum – + + +
Scenedesmus + + + +
Spirogyra + + + +
Volvox + + + +
194 Current Herpetol. 41(2) 2022

Appendix I
(continued)
Food items Stage 25 Stage 32 Stage 38 Stage 41
Zygnema + + + +
Zygnematophyceae Cosmarium + + + +
Closterium + + + +
Desmidium + + + +
Euastrum – + + +
Mougeotia – + + +
Pleurotaenium + + + +
Staurodesmus – + + +
Staurastrum – + + +
Xanthidium – + + +
Cyanophyceae Chroococcus – + + +
Merismopedia + + + +
Microcystis – + + +
Oscillatoria + + + +
Euglenophyceae Euglena + + + +
Phacus – + + +
Trachelomonas – + + +
Ulvophyceae Ulothrix + + + +
Zooplankton
Rotifera Lecane + + + +
Lepadella + + + +
Platyias – + + +
Copepoda Eucyclops + + + +
Detritus + + + +

+ = Occurrence, – = Non occurrence

 ROUT ET AL.—ORAL MORPHOLOGY AND DIET OF LARVAL POLYPEDATES 195

Appendix II
List of similarities of oral and buccophryngeal characters between Polypedates maculatus and P. teraiensis
tadpoles.
Characteristic features P. maculatus and P. teraiensis
Oral morphology Location of the oral disc Antero-vental
Oral disc Emarginated
Marginal papillae Narrow dorsal and ventral gaps
Rows of marginal papillae Single row dorsally and double row ventrally
Sub marginal papillae Present on the lateral corners
Jaw sheath Keratinisation Moderate
Jaw sheath serration Serrated
Buccal roof Pre-narial arena ridge Arched medially ridge with many secondary
lateral papillae
Plane of internal nares Transverse
Internal nares Moderately separated (>half the length but less
than the length of an individual nare)
Papillae on the anterior narial wall Present
Post narial papillae One pair
Medial ridge Triangular flap
Lateral ridge papillae Present
Buccal floor Infralabial papillae 1 pair
Infralabial palp 2 pair
Tongue anlage Broad and raised
Pustules in BFA Few (<30)
Buccal pocket Oblique
Buccal pocket papillae Absent
Ventral velum Wide and continuous
Glottis opening Posterior to the ventral velum
Branchial basket arch 4 Pair

View publication stats