Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 July 2025 | 17(7): 27242–27248
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.9370.17.7.27242-27248
#9370 | Received 18 November 2024 | Final received 24 March 2025 |
Finally accepted 24 June 2025
Diet composition of three syntopic, ecologically divergent frogs (Euphlyctis,
Minervarya, Polypedates)
from paddy fields of Kohima, Nagaland, India
Thejavitso Chase 1 &
Santa Kalita 2
1,2 Department of Environmental
Science, Tezpur University, Napaam,
Tezpur, Assam 784028, India.
1 thejachase@gmail.com, 2 santa@tezu.ac.in
(corresponding author)
Editor: S.R. Ganesh, Kalinga
Foundation, Agumbe, India. Date of publication: 26 July 2025
(online & print)
Citation: Chase,
T. & S. Kalita (2025). Diet
composition of three syntopic, ecologically divergent
frogs (Euphlyctis, Minervarya,
Polypedates) from paddy fields of Kohima,
Nagaland, India. Journal of Threatened Taxa 17(7): 27242–27248. https://doi.org/10.11609/jott.9370.17.7.27242-27248
Copyright: © Chase & Kalita 2025. Creative Commons Attribution 4.0 International
License. JoTT allows unrestricted use, reproduction,
and distribution of this article in any medium by providing adequate credit to
the author(s) and the source of publication.
Funding: None.
Competing interests: The authors declare no competing interests.
Author details: Thejavitso Chase is currently pursuing PhD from the Department of Environmental Science at Tezpur University, Assam, India. Dr. Santa Kalita is an assistant professor in the Department of Environmental Science at Tezpur University, Assam, India and the PhD supervisor of Thejavitso Chase.
Author contributions: First author (TC) collected the diet content from the field and identified the diet content for analysis and drafted the manuscript. Corresponding author (SK) supervised the work, checked the manuscript and communicated it to the journal.
Acknowledgements: We would like to thank Nisapi Pucho, Sedevikho Chase, and Methaheto Chase for providing accommodation which facilitated this work. We thank the people of Viswema, Nerhema, Kohima, Jotsoma, Khonoma, and Dzüleke for their support, and cooperation.
Abstract: Monitoring indicator species like
amphibians is crucial to assess habitat health. The diet of 129 anurans
belonging to the three most abundant species found in the paddy fields of
Kohima district in Nagaland, northeastern India—the aquatic Euphlyctis
adolfi, the terrestrial Minervarya
nepalensis and the arboreal Polypedates
himalayensis—was studied. Results revealed
302 intact prey items belonging to 11 prey categories, gleaned through the
stomach-flushing method. While Coleoptera was the
most abundant prey found in all three species; Clitellata
(terrestrial earthworms), Diptera, and Orthoptera
were also important prey items. The high degree of overlap in the dietary niche
of the three species despite their diverged microhabitat associations, could be
the result of abundant prey items and the segregation of microhabitats. Lastly,
as these frogs share a common prey base, they evidently segregate their
foraging microhabitats to avoid competition.
Keywords: Aquatic, arboreal, class, index
of relative importance, northeastern India, order, terrestrial.
INTRODUCTION
Anurans (frogs & toads) are
the most diverse order of amphibians and are ecological indicator species that
require close monitoring (AmphibiaWeb 2025). India is
home to a vast number of little-known, threatened, and endemic amphibians,
despite harbouring a very high human population and
this is particularly true for the northeastern India that is one of the country’s
three biodiversity hotspots (Dinesh et al. 2024). The Kohima District of
Nagaland has a hilly terrain and very less naturally occurring standing water.
Rice terrace cultivation is a widely practiced form of agriculture in this
region. Paddy fields serve as crucial habitats for anurans, providing essential
standing water for breeding and supporting tadpole development, especially in
regions with limited natural aquatic environments (Elphick
2000). Despite the high anuran diversity in this region (Talukdar &
Sengupta 2020), a comprehensive literature review revealed only three published
studies on the diet of adult anurans in northeastern India, indicating a
significant research gap in this area (Chanda 1993; Ao
et al. 2001; Sarkar & Dey 2022). Despite the reduced
habitat heterogeneity in paddy fields, resilient generalist species inhabit
these fields (Piatti et al. 2010). Paddy fields serve
as surrogate habitats for aquatic species (Elphick
2000), including anurans from surrounding areas (Seshadri et al. 2020).
While some taxa demonstrate a
restricted trophic niche, relying on a limited range of prey items, others
exhibit a broader diet, consuming a diverse assemblage of prey organisms.
Primarily, anurans feed on arthropods and they can be important pest control
agents in agro-ecosystems (Khatiwada
et al. 2016). Anurans play a crucial role in the food chain due to the diet
they consume and also because they are prey to animals in the higher trophic
levels. Niche overlap does not equate to an increase in competition among
species when there are enough resources for all species (Pianka
1974). Niche partitioning studies can give insights into a community’s species
diversity, abundance, and distribution (Toft 1985).
Information on diet helps in the understanding of ecology, natural history
(Donnelly 1991), niche partitioning (Toft 1985), and
community structure (Toft 1980). The present study focussed on the following two parameters: (i) to assess the composition of anurans in paddy fields;
(ii) to compare the diet of the three most abundant species observed in the
local paddy fields, with respect to three syntopic,
ecologically-dissimilar frog species.
MATERIALS AND METHODS
Study species
Three co-occurring or syntopic frog species that have divergent habitat utilisation patterns were chosen for the study. They were:
the aquatic skittering frog Euphlyctis adolfi (Günther, 1860), the terrestrial cricket
frog Minervarya nepalensis
(Dubois, 1975) and the arboreal tree frog Polypedates
himalayensis (Annandale, 1912). These species
depend on stagnant water for breeding and other vital life processes including
metamorphosis (Chanda 2002). These species use the water from embankments for
breeding during summer. While E. adolfi primarily
inhabits water, M. nepalensis, and P. himalayensis occur primarily in the periphery of
embankments on land, and on vegetation, respectively. For taxonomic definitions
of the studied frog species see Sanchez et al. (2018), Saikia
et al. (2020), and Dufresnes et al. (2022).
Study sites
Six paddy fields, one each from
five villages and one sub-urban locality in Kohima District, Nagaland, were
surveyed. The six paddy fields were located in Nehrema
Village, Kohima Town, Viswema Village, Jotsoma Village, Khonoma Village,
and Dzüleke Village. The closest paddy fields were
2.46 km apart.
Sampling
Sampling was carried out from
March to June, i.e., pre-monsoon to monsoon during 2021–2022. Stomach-flushing
was done following Solé et al. (2005) immediately
after capture of each individual frog from 1800 h to 2100 h. Following the
stomach-flushing, all individuals were released back into the environment. Each
stomach was flushed thrice. The stomach content was stored in 70% ethanol in
screw cap vials. Diet content of 129 individuals of anurans belonging to three
species- Euphlyctis adolfi
(n = 45), Minervarya nepalensis (n = 51), and Polypedates
himalayensis (n = 33) were examined during the
study. Diet contents were identified up to the order level under a dissecting
microscope. Partially digested food items, stones, and plant materials were
categorized as miscellaneous and were not considered for analysis. A
significant amount of diet contents observed was either partially digested or
partially eaten; hence, intact bodies of prey items were a representation of
the total prey consumed. Identification keys for diet contents were taken from
Gibb & Oseto (2006). Prey items were measured
with Mitutoyo 505–730 dial calipers (0.02 mm accuracy). Data analysis was done
using MS Excel and RStudio.
Data analysis
Vacuity index was measured as the proportion of empty
stomachs to the total number of individuals of each species sampled. The volume
of prey items was calculated using the formula for ellipsoid bodies (Colli & Zamboni 1999):
Where, V is the volume, L is the length, and W is the
width of a prey item.
The importance of diet contents was determined by ranking
them using the index of relative importance (IRI) (Pinkas
1971):
Where IRI = index of relative importance, N = numerical
percentage, V = volumetric percentage, and F = frequency of occurrence
percentage. Trophic niche breadth was calculated using the pliang
non-Wiener index (Shannon & Weaver 1949):
Where H' is the Shannon-Weaver index, pi is
the proportion of individuals found to consume prey i. The H' value was standardized using the evenness index (Shannon
& Weaver 1949):
Where J' is the measure of evenness and n is the number
of species. Species were paired to calculate niche breadth by following Pianka’s niche breadth formula:
Where Ôjk is Pianka’s measure of niche overlap, P̂ij
is the proportion of ith resource
used by jth species and P̂ik is the proportion of ith resource used by kth
species.
RESULTS
Out of the 169 individual anurans
belonging to the three species that were examined, 129 individuals contained
food items in their stomachs. A total of 302 intact prey items were recovered
which belonged to three classes (Insecta, Clitellata and Malacostraca) and 11 categories (Araneae, Coleoptera, Diptera, Orthoptera, Blattodea,
Hemiptera, Lepidoptera (larva), Hymeniptera, Trichoptera, Clitellata, Decapoda), respectively. It was observed that several
individuals had empty stomachs: 21 individuals of Minervarya
nepalensis (vacuity index = 29.58%), 14
individuals of Euphlyctis adolfi (vacuity index = 23.73%), and five individuals
of Polypedates himalayensis
(vacuity index = 13.16%). Partially digested prey was observed in
several individuals of anurans while intact prey was relatively fewer. Results
showed that E. adolfi consumed prey of eight
categories while M. nepalensis and P. himalayensis consumed prey of nine categories,
respectively. Statistical analysis revealed that the difference in the total
number of prey consumed among the species was not
significant (Kruskal-Wallis chi-squared = 2, df = 2,
p = 0.3679).
Euphlyctis adolfi consumed the highest number of prey followed by P. himalayensis
and M. nepalensis. Polypedates
himalayensis on average consumed the highest
number of prey per individual (Table 1). There was a
statistically significant difference between the total number of prey consumed by the individuals of the three species
(Kruskal-Wallis test = 28.232, df = 2, p <0.05). Coleoptera was the most common prey item in all the three
species (relative occurrence: 34.88% relative occurrence in E. adolfi, 32% in M. nepalensis
and 48.98% in P. himalayensis).
Niche breadth and niche overlap
Dietary niche breadth was
broadest in M. nepalensis and narrowest in P.
himalayensis (Table 2). Niche overlap was
highest between M. nepalensis and P. himalayensis and lowest between M. nepalensis and E. adolfi (Table
3). There was a high degree of overlap in the dietary niche of the three
species.
Index of relative importance
Coleoptera (beetles) were the most abundant
prey order found to be consumed by all three species studied. Prey categories Coleoptera, Orthoptera, and Clitellata
were the highest contributors to the IRI value by volume for M. nepalensis (Table 5). In P. himalayensis,
the diet volume was contributed mostly by class Clitellata
(terrestrial earthworms) (Table 6). On the other hand, the largest volume
contributors to the diet of E. adolfi were
the orthopterans (Table 4). For all three species, coleopterans had the
highest score for the Index of Relative Importance (IRI). Other important prey
orders for E. adolfi were Diptera
and Orthoptera. Orthoptera and Araneae were the
highest contributors to IRI values in both M. nepalensis
and P. himalayensis. The total prey volume was
the highest in E. adolfi (568.36 cm3,
n = 45), while M. nepalensis, and P. himalayensis had similar volume (189.95 cm3,
n = 51 and 276.41 cm3, n = 33, respectively).
DISCUSSION
Each of the three studied species
have wide distribution across northeastern India (Chanda 2002; Ao et al. 2003; Dinesh et al. 2024) and was found to be the
most abundant species in paddy field habitats in the studied areas. Due to
their resilience and generalist behaviour, these
species can thrive in this altered habitat. Other co-occurring species, viz.,
Hyla annectans, Duttaphrynus melanostictus, Microhyla sp., Zhangixalus
burmanus, and Zhangixalus
smaragdinus were excluded from this study due to
small sample size present in our observations.
The vacuity index reveals a
relatively high proportion of individuals with empty stomachs. A similar study
found that anurans feed at a lower intensity during drier periods (Das 1996a).
The high degree of dietary niche overlap is attributable to the similarity of
IRI ratings of prey items among the three species. Coleoptera
was the most important prey order according to the IRI values across all
species. Diptera and Orthoptera ranked second and
third in IRI values for E. adolfi respectively;
while Orthoptera and Araneae ranked second and third
in IRI values for M. nepalensis and P. himalayensis, respectively. Clitellata
was absent in the diet of E. adolfi owing to
the anuran’s aquatic habitat. Though P. himalayensis
is a tree frog, it is often observed on the ground in paddy fields during the
breeding period. We have observed that they consume prey of Clitellata
(terrestrial earthworms) during this period.
Das (1996) reported that the
related, peninsular Indian species P. maculatus feeds both on ground and
trees and classified it as a terrestrial feeder. Polypedates
himalayensis have been reported to deposit eggs
on forest floors. Individuals of this species were observed calling from holes
in the ground and paddy fields (Rangad et al. 2012),
indicating that this species spends its breeding period on ground, descending
from the nearby bushes. Therefore, niche overlap values indicate a high degree
of overlap in the diet of these anurans. Diptera and Trichoptera were found only in E. adolfi
while Clitellata, Hemiptera, and Decapoda were found only in M. nepalensis
and P. himalayensis. The decapod prey items
observed were freshwater shrimps.
Although several studies have
reported the presence of stones and plant materials in the diet of anurans, the
cause for ingesting such materials has not been ascertained (Modak et al. 2018; Bahuguna et
al. 2019). The presence of such materials may be attributed to accidental ingestion.
This study also reveals that all the three observed species lack specialization
in the food intake and are hence considered generalists in their feeding habit.
Previous studies on E. adolfi also
reported that coleopterans occupied the highest volume percentage amongst all
arthropod prey items consumed (Das & Coe 1994; Das 1996b).
It was observed that although
there is a high dietary niche overlap among the species, the three species
occupied different microhabitats, thus minimizing the chances of competition
between species. E. adolfi individuals were
primarily observed swimming or floating on water. Polypedates
himalayensis were recorded from microhabitats
with less water, such as wet soil, and moist edges of embankments within paddy
fields. Minervarya nepalensis
individuals were observed to be wide-ranging, their microhabitats overlapping
between E. adolfi, and P. himalayensis. Within the embankments, M. nepalensis was seen at the edges and did not swim /
float unless while escaping from the observer.
CONCLUSION
In this study eight species of
anurans were recorded from paddy fields; out of which three were studied for
their diet preferences. The study site has a hilly terrain with several
torrential streams. The landscape has limited areas of wetland habitats, which
make paddy fields a vital refuge for anurans as they require wetlands for
breeding, larval development, and a source of food for both adults, and
tadpoles. While some species may use the paddy field areas for breeding only,
the studied species have been found outside their breeding period in this
habitat. This indicates that these three species are resilient generalists (Piatti et al. 2010). Among the three species, E. adolfi was the only species that had been studied
previously (Das & Coe 1994). The present study revealed a high degree of
overlap of prey among the three species with a low number of ingested prey.
The niche overlap and coexistence
of the species suggest two hypotheses. Firstly, the interspecific competition
caused by the niche overlap is not enough to drive any species to competitive
exclusion due to the abundance of prey base. Secondly, the existing competition
has not lasted long enough for species to evolve different diets. These have
been supported by Pianka (1974) and Piatti & Souza (2011). Although the dietary niche
overlap is high among the species, the overall niche may be differentiated
according to observations in microhabitat usage. Future studies are recommended
to include prey diversity studies and extend the sampling period through the
monsoon to the post-monsoon seasons. To determine the overall niche
differentiation among these three syntopic frog
species, we suggest the inclusion of other niche dimensions such as aural
niche, in addition to spatial, and trophic niches studied here.
Table 1. Average prey consumed per individual of each species.
|
Frog species |
No. of anurans |
No. of prey (n) |
Mean |
SD |
|
E. adolfi |
45 |
129 |
2.867 |
2.06 |
|
M. nepalensis |
51 |
75 |
1.471 |
1.17 |
|
P. himalayensis |
33 |
98 |
2.97 |
1.49 |
|
|
||||
Table 2. Niche breadth values measured with Shannon-Weaver index and
evenness measure.
|
Frog species |
H' |
J' |
|
M. nepalensis |
1.87 |
0.851 |
|
E. adolfi |
1.67 |
0.805 |
|
P. himalayensis |
1.59 |
0.722 |
|
|
||
Table 3. Niche overlap values measured with Pianka’s
measure.
|
Frog species |
M. nepalensis |
E. adolfi |
P. himalayensis |
|
M. nepalensis |
1 |
0.728 |
0.949 |
|
E. adolfi |
0.728 |
1 |
0.765 |
|
P. himalayensis |
0.949 |
0.765 |
1 |
|
|
|||
Table 4. Index of relative importance and its variables for Euphlyctis adolfi.
|
Prey Order / Class |
Volume (%) |
Frequency (%) |
Number (%) |
IRI |
|
Araneae |
3.19 |
15.56 |
9.30 |
194.38 |
|
Coleoptera |
9.41 |
42.22 |
34.88 |
1870.27 |
|
Diptera |
6.74 |
31.11 |
30.23 |
1150.36 |
|
Orthoptera |
42.20 |
20 |
9.30 |
1030.05 |
|
Blattodea |
28.96 |
11.11 |
4.65 |
373.50 |
|
Hemiptera |
0 |
0 |
0 |
0 |
|
Lepidoptera (larva) |
5.94 |
4.44 |
2.33 |
36.73 |
|
Hymenoptera |
3.05 |
13.33 |
6.98 |
133.68 |
|
Trichoptera |
0.50 |
6.67 |
2.33 |
18.83 |
|
Clitellata |
0 |
0 |
0 |
0 |
|
Decapoda |
0 |
0 |
0 |
0 |
Table 5. Index of relative importance and its variables for Minervarya nepalensis.
|
Prey Order / Class |
Volume (%) |
Frequency (%) |
Number (%) |
IRI |
|
Araneae |
12.41 |
19.61 |
17.33 |
583.29 |
|
Coleoptera |
22.07 |
35.29 |
32.00 |
1908.42 |
|
Diptera |
0 |
0 |
0 |
0 |
|
Orthoptera |
20.62 |
25.49 |
20.00 |
1035.47 |
|
Blattodea |
4.55 |
7.84 |
5.33 |
77.53 |
|
Hemiptera |
8.30 |
5.88 |
4.00 |
72.35 |
|
Lepidoptera (larva) |
6.25 |
7.84 |
9.33 |
122.21 |
|
Hymenoptera |
0.85 |
9.80 |
6.67 |
73.69 |
|
Trichoptera |
0 |
0 |
0 |
0 |
|
Clitellata |
23.14 |
3.92 |
2.67 |
101.20 |
|
Decapoda |
3.54 |
3.92 |
2.67 |
24.34 |
Table 6. Index of relative importance and its variables for Polypedates himalayensis.
|
Prey Order / Class |
Volume (%) |
Frequency (%) |
Number (%) |
IRI |
|
Araneae |
7.53 |
30.30 |
13.27 |
630.21 |
|
Coleoptera |
29.69 |
72.73 |
48.98 |
5721.20 |
|
Diptera |
0 |
0 |
0 |
0 |
|
Orthoptera |
13.72 |
39.39 |
17.35 |
1223.66 |
|
Blattodea |
0.98 |
3.03 |
1.02 |
6.05 |
|
Hemiptera |
3.76 |
6.06 |
4.08 |
47.50 |
|
Lepidoptera (larva) |
4.83 |
9.09 |
6.12 |
99.53 |
|
Hymenoptera |
0.28 |
6.06 |
3.06 |
20.24 |
|
Trichoptera |
0 |
0 |
0 |
0 |
|
Clitellata |
30.18 |
12.12 |
4.08 |
415.30 |
|
Decapoda |
9.05 |
6.06 |
2.04 |
67.22 |
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