Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 August 2026 | 18(8): 29459–29469
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.10360.18.8.29459–29469
#10360 | Received 02 January 2026 | Final received 22 July 2026 | Finally
accepted 04 August 2026
Community structure and
environmental determinants of amphibians in the Western Ghats watersheds of
eastern Goa, India
Mayur M. Gawas 1 , Sharan S. Sawal 2 & Nitin S. Sawant 3
1–3 Department of Zoology, Goa
University, Taleigao, Goa 403206, India.
1 mithilgawas0987@gmail.com, 2
sharanssawal2@gmail.com, 3 nitin.sawant@unigoa.ac.in
(corresponding author)
Editor: Anonymity
requested. Date of publication: 26 August 2026 (online &
print)
Citation: Gawas,
M.M., S.S. Sawal & N.S. Sawant (2026). Community structure and
environmental determinants of amphibians in the Western Ghats watersheds of
eastern Goa, India. Journal of Threatened Taxa 18(8): 29459–29469. https://doi.org/10.11609/jott.10360.18.8.29459-29469
Copyright: © Gawas
et al. 2026. 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: Mr. Mayur
M. Gawas is PhD Scholar at Department
of Zoology; School of Biological
Sciences and Biotechnology,
Goa University, Taleigao Plateau, Taleigao,
Goa 403 206, India. Mr. Sharan S. Sawal was a M.Sc. Student at Department of Zoology; School of Biological
Sciences and Biotechnology,
Goa University, Taleigao Plateau, Taleigao,
Goa 403 206, India. Dr. Nitin S. Sawant is affiliated with the School of Biological Sciences
and Biotechnology, Zoology, Goa University, with
research interests in ecology,
climate change, biodiversity,
Wildlife and conservation biology. He is currently involved in major research projects funded by the Ministry
of Environment, Forest and Climate Change (MoEF&CC), Government of India, and the Goa State Research Foundation (GSRF), Government of Goa. He has published
around 23 research papers in ecology,
biodiversity, and wildlife conservation and is actively engaged in research and conservation initiatives in collaboration with the Goa State Biodiversity Board,
Goa Forest Department, MoEF&CC, GSRF, and the University of Porto–CIBIO, Portugal.
Author contributions:
MG-—field data collection,
data analysis, manuscript preparation and writing. SS—field data collection. NSS—conceptualization
of the research, research methodology
and study design,
manuscript preparation, review, proofreading
and supervision.
Acknowledgements: We sincerely
thank the Goa Forest Department for granting necessary
permissions, Honourable MLA
Smt. Deviya Rane for support in facilitating permissions, and our fellow colleagues for their valuable assistance during fieldwork.
Abstract: We studied the amphibian
diversity and the influence of abiotic factors across 12 watershed zones in the
Western Ghats region of Goa, India. Surveys conducted at 102 sampling sites
recorded 22 amphibian species, including eight Western Ghats endemics. Species
richness and diversity were highest in watersheds with intermediate canopy
cover and deeper leaf litter. Pearson correlation analysis identified leaf
litter depth, humidity, and temperature as key factors influencing amphibian
diversity. The findings highlight the importance of forest floor microhabitats
and microclimatic conditions in shaping amphibian communities and emphasize the
need to conserve structurally diverse forest habitats in the Western Ghats.
Keywords: Abiotic factors, alpha diversity,
Canopy cover, ecology, Leaf litter depth, Microclimate, Species richness.
Introduction
The Western Ghats is recognised as one of the world’s 34 biodiversity hotspots
and supports high amphibian diversity and endemism (Myers et al. 2000). Goa
forms part of the Goa Gap, a bioclimatic and vegetational transition zone
within the Western Ghats (Pascal 1988; Qureshi 1965; Barboni
et al. 2003; Reddy et al. 2016; Ramachandran et al. 2017; Biswas & Karanth 2021; Chaitanya & Meiri
2022). Despite the rich amphibian diversity of the Western Ghats, Goa remained
comparatively understudied for many decades because the state was under
Portuguese rule during the period when extensive faunal surveys were being
conducted in British India (Kulkarni et al. 2013).
The earliest published accounts
of amphibians from Goa were provided by Abdulali
& Sekar (1988), followed by studies by Sekar (1991, 1992), Das & Whitaker (1997, 1998), Bhat
& Desai (1998), and Sarkar & Ray (2004). Kamble
(2008) carried out one of the last detailed taxonomic studies on Goan amphibians. Later, Kulkarni et al. (2013) reported 36
species from Goa, while Dinesh et al. (2015) updated the checklist and reported
34 species from the state. Although these studies contributed significantly to
the knowledge of amphibians in Goa, information regarding amphibian community
structure, spatial distribution, and the influence of abiotic factors on
amphibian assemblages remains limited.
Understanding amphibian
diversity, habitat associations, and community structure is therefore essential
for developing effective conservation strategies (Sawant & Jadhav 2013; Bowalkar & Sawant, in press). Therefore, the present
study was conducted to document amphibian species from the forests in the study
area, examine their community structure, and assess the influence of selected
abiotic factors on their communities.
Methods
The study was carried out in the
Western Ghat region of Goa covering Mhadei Wildlife Sanctuary, Bhagwan
Mahavir Wildlife Sanctuary, Mollem National Park,
Netravali Wildlife Sanctuary, Cotigao Wildlife
Sanctuary, and parts outside these sanctuaries amounting to a total
geographical area of 1335.95 km2 (Image 1). The surveys were
conducted across watersheds associated with Valvanti
river (W1), Kotrachi nadi
(W2), Madhei river – Satrem
tributary (W3), Madhei river (Main) (W4), Ragada river (W5), Dudhsagar
river (W6), Uguem river (W7), Netravali river (W8), Kushawati river (W9), Agonda
river (W10), Talpona river (W11), and Kali river
tributary (W12) (Table 1). The study was carried out during the monsoon season
from June–September in dense forested areas, defined as regions having tree
canopy cover greater than 40% (Forest Survey of India 2023). The entire study
area is divided into 12 watershed zones (W1–W12) using QGIS software and SRTM
data from U.S. Department of the Interior (Image 2). Each watershed zone was
further subdivided into three elevational zone low (0–200 m), mid (200–400 m),
and high (above 400 m) and sampling was conducted uniformly in each of the
elevational zone to minimize sampling bias. In total, 102 sampling sites were
established (three sites per elevation in each watershed) (Image 1, table 1).
At each site, three replicate line transects (30 × 20 m; 10 m on either side)
were laid. Amphibians were surveyed using standardized visual and acoustic
encounter surveys for 30 minutes per observer, covering all available
microhabitats (Crump 1994; Rödel & Ernst 2004).
Surveys were conducted at night between 2000 h and 0300 h, coinciding with peak
amphibian activity.
Along with species data,
environmental, climatic, and habitat-related variables were collected to
understand the relationship between amphibian communities and environmental
factors. Seven key abiotic variables were recorded: tree cover (%) using a
densitometer, leaf litter temperature (0C) using an infrared
thermometer, leaf litter depth (cm) using a scale, leaf litter ratio by
counting number of leaves, twigs and fruits in 1 m2, understory
vegetation (count in 5 m2), ambient temperature (0C)
using a thermometer, and relative humidity (%) using a hygrometer.
Results
A total of 22 amphibian species
were recorded from the study area. Of these, two species are classified as
‘Endangered’, two as ‘Vulnerable’, one as ‘Near Threatened’, one as ‘Data
Deficient’, and 16 species as ‘Least Concern’ according to the IUCN Red List of
Threatened Species. Eight of the recorded species are endemic to the Western
Ghats of India, highlighting the region’s unique and irreplaceable biodiversity
(Image 2; Table 2).
Alpha diversity
Species richness ranged from 9–19
species, with least species in W12 (9 species) and highest species in W3 (19
species) (Image 3A; Table 3). The Shannon diversity index (H′) was highest at
W1 1 (2.35) and lowest at W12 (1.82), with elevated H′ values at Sites 1 and 11
indicating a diverse and well-distributed amphibian community in these
watersheds (Image 3B; Table 3). Simpson Index (1−D) ranged from 0.79 in W12 to
0.89 in W11, watersheds such as W5 and W11 which exhibited high Simpson values,
indicates low dominance and higher probability of encountering different
species (Image 3C; Table 3). Evenness (J’) was highest at W5 (0.92), showing a
balanced species distribution, while W3 (0.74) had lower evenness despite high
richness possible due to dominance by few species (Image 3D; Table 3).
Environmental Variation
Tree cover: Tree canopy cover was
generally high across all watersheds, ranging from 71.11% SE = 6.33 at W10 to
100% at W5 and W12. Watersheds in the northern and central regions (W1–W7)
consistently exhibited dense canopy cover (>90%), whereas southern
watersheds showed variable tree canopy cover between 71% in W10 to 100% in W12,
indicating canopy openness or disturbance in south (Image 4A; Table 4).
Leaf Litter Temperature: This was
relatively stable across the sites, ranging from 22.70 0C SE = 0.65
in W8 to 24.76 0C SE = 0.19 (W11) (Image 4B; Table 4).
Leaf litter depth: It varied
across the gradients, with W3 recording the highest mean depth (3.50 cm SE =
0.54), followed by W2 (3.22 cm SE = 0.67), and W9 (3.11 cm SE = 0.46) (Image
4C; Table 4).
Leaf Litter Ratio: ratio remained
consistent across watersheds, generally ranging between 0.73 and 0.80, except
in W10 (0.67 SE = 0.05), which had the lowest value. Most northern and central
watersheds had a ratio of 0.76–0.80, indicating high-quality litter habitats
(Image 4D; Table 4).
Understory Vegetation: This
showed high spatial variability. The densest vegetation was observed in W6
(173.33 SE = 12.69), and W3 (160 SE = 17.16), while W10 had the sparsest
understory (82.78 SE = 14.89). Northern watersheds such as W1–W6 also had
moderate to dense understory with an average of 140.04, contributing to
microhabitat complexity (Image 4E; Table 4).
Ambient Temperature: This varied
significantly along the gradient. The coolest site was W8 (23.53 0C,
SE = 0.49), while the warmest was W7 (28.90 0C, SE = 1.08), and W10
(28.14 0C, SE = 0.30). Southern watersheds, particularly W7–W12,
exhibited higher temperatures compared to northern ones (Image 4F; Table 4).
Humidity: This was generally high
across the watersheds. W4 and W9 recorded the highest mean humidity (92.33% SE
= 1.48 and 91.11% SE = 1.44) while W7 recorded the lowest (74.56% SE = 5.40)
(Image 4G; Table 4).
Overall, southern watersheds were
characterized by higher ambient temperatures, variable humidity, and relatively
sparse vegetation, whereas northern and central watersheds exhibited denser
canopy cover, cooler microclimates, and deeper litter layers. These
environmental gradients likely influence the observed patterns in amphibian
community composition and beta diversity.
Pearson correlation
Pearson correlation analysis
revealed leaf litter depth as the strongest driver of amphibian alpha diversity
across 12 Western Ghats watersheds (Image 5). Species richness was strongly
positively correlated with leaf litter depth (r = 0.65) and humidity (r =
0.37), but negatively with temperature (r = -0.35) and tree cover (r = -0.27).
Shannon and Simpson indices were highly intercorrelated (r = 0.97) and
moderately tied to richness (r = 0.67, 0.57), with weak negative links to
temperature and understory vegetation. Pielou’s
evenness showed a strong negative correlation with leaf litter depth (r =
-0.47), indicating dominance in deeper litter, and weak positive ties to tree
cover (r = 0.21) and leaf litter ratio (r = 0.21). Environmental variables were
tightly linked: tree cover & leaf litter ratio (r = 0.82), understory &
leaf litter ratio (r = 0.63), and temperature & humidity (r = -0.54).
Overall, leaf litter depth, humidity, and temperature were the primary
predictors, with depth exerting opposing effects on richness (+ve) and pielou’s evenness (-ve).
Intermediate canopy cover (80–95
%) and deeper leaf litter emerge as key drivers of amphibian alpha diversity
across Western Ghats watersheds, with peak richness occurring where
microclimatic conditions optimize the litter–forest floor interface. While full
canopy closure maintains evenness through generalist dominance, it reduces
habitat suitability via litter compression and thermal stress. These findings
highlight the value of mid-successional forest patches for conserving anuran
biodiversity amidst ongoing habitat alteration in this global biodiversity
hotspot. To safeguard anuran biodiversity amid escalating climate change and
habitat fragmentation, conservation priorities must emphasize preserving undisturbed
leaf litter layers and intermediate vegetation structure in tropical montane
streams, ensuring both species coexistence and long-term community stability.
Discussion
Amphibian alpha diversity across
the 12 Western Ghats watersheds exhibited moderate variation, with species
richness ranging from 9 (W12) to 19 (W3) and Shannon diversity from 1.82 (W12)
to 2.35 (W1). Highest richness and Shannon values occurred in watersheds with
intermediate canopy cover (80–95 %) and deeper leaf litter (W1, W9, W3, W8,
W11), consistent with studies showing that canopy closure and litter depth are
key predictors of anuran diversity in Western Ghats forests (Katwate et al. 2013) and that amphibian richness tends to
decline under very dense canopy in forested wetlands (Skelly et al. 2005).
In contrast, full‑canopy, shallow‑litter sites (W5, W12) supported
lower richness despite occasionally high Pielou’s
evenness (W5 = 0.92), a pattern similar to sites with low richness but evenly
distributed abundances reported from Phansad Wildlife
Sanctuary (Katwate et al. 2013). Simpson diversity
remained consistently high across watersheds (0.79–0.89), indicating low
dominance and relatively even assemblages; this is in line with work showing
that elevated local alpha diversity and evenness can persist across habitat
conditions while masking underlying shifts in species composition and regional
diversity (Dehling & Dehling
2023). Together, these results suggest that fine‑scale variation in canopy structure and forest‑floor litter depth modulates local richness within broadly forested
catchments, whereas community evenness remains high, as expected for
assemblages dominated by a mix of habitat generalists and a smaller set of
specialists.
Pearson correlation analysis
confirmed leaf litter depth as the strongest predictor, positively driving
richness (r = 0.65) but reducing evenness (r = −0.47), indicating that greater
forest‑floor microhabitat complexity
promotes species accumulation while fostering dominance by leaf‑litter specialists (Scott 1976; Kouamé et al.
2018). Humidity positively influenced richness and Shannon indices (r = 0.37,
0.32), and temperature showed negative associations (r = −0.35, −0.26),
consistent with amphibians’ reliance on cool, moist microclimates, and narrow
physiological tolerances to desiccation and heat (Wells 2007). The strong
inverse relationship between temperature and humidity (r = −0.54) therefore
highlights a key microclimatic trade‑off, where drier, warmer
conditions are likely to restrict activity and occupancy of many forest‑floor anurans despite otherwise suitable structural habitat (Wells
2007).
Environmental variables show some
distinct patterns. Tree cover ranged from 71.11% (W10) to 100% (W5, W12), with
leaf litter depth peaking (3.0–3.5 cm) under moderate canopy and declining
under dense closure (~2.2 cm) due to compression and reduced litterfall (Wells
2007; Cardelús 2010). Understory vegetation was
densest in closed‑canopy sites (W6, W5, W12),
potentially impeding amphibian movement or prey access as reported from Western
Ghats forests. Ambient temperature was elevated (>27 0C) in high‑canopy watersheds, likely from reduced ventilation and heat trapping (Spranger et al. 2024), while humidity dipped lowest in
W7 and W5, coinciding with moderate canopy and high thermal stress (Wells 2007;
Seshadri 2014).
Despite structural roles in
habitat provision, tree cover, and understory vegetation showed weak or
negative correlations with amphibian diversity metrics, as dense canopies
suppressed leaf litter depth (r = -0.40) and elevated temperature, indirectly
constraining occupancy. This pattern supports tropical studies documenting peak
anuran diversity in mid-successional habitats rather than closed-canopy
forests, where intermediate structural complexity optimizes microclimate,
litter accumulation, and foraging opportunities (Skelly et al. 2005;
Wells, 2007). The near-perfect correlation between Shannon and Simpson indices
(r = 0.97) across watersheds confirms their redundancy for characterizing these
assemblages, with Pielou’s evenness contributing only
modestly to overall diversity variation (Magurran
2003). These integrated findings demonstrate that intermediate canopy cover
optimizes the litter–microclimate interface, maximizing species richness and functional
stability (Skelly et al. 2005). Full canopy closure, while enhancing
evenness in some cases, reduces habitat suitability through litter compression
and thermal elevation (Wells 2007; Rödel & Ernst
2004).
Table 1. Details of sampling
sites, elevation categories, associated watersheds, and hill ranges surveyed in
the Western Ghats region of Goa.
|
Watershed code |
Site names |
Watershed/river basin |
Hill range |
||
|
Low elevation (0–200 m) |
Mid elevation (200–400 m) |
High elevation (above 400 m) |
|||
|
W1 |
1_A1, 1_A2, 1_A3 |
1_B1, 1_B2, 1_B3 |
1_C1, 1_C2, 1_C3 |
Valvanti river basin |
Morlegad and vagheri hill range |
|
W2 |
2_A1, 2_A2, 2_A3 |
2_B1, 2_B2, 2_B3 |
2_C1, 2_C2, 2_C3 |
Kotrachi Nadi river basin |
Chorla ghat range |
|
W3 |
3_A1, 3_A2, 3_A3 |
3_B1, 3_B2, 3_B3 |
3_C1, 3_C2, 3_C3 |
Madhei river – Satrem tributary basin |
Surla ghat range |
|
W4 |
4_A1, 4_A2, 4_A3 |
4_B1, 4_B2, 4_B3 |
4_C1, 4_C2, 4_C3 |
Madhei river basin |
Sosogad hill range |
|
W5 |
5_A1, 5_A2, 5_A3 |
5_B1, 5_B2, 5_B3 |
5_C1, 5_C2, 5_C3 |
Ragada river basin |
Anmod ghat range |
|
W6 |
6_A1, 6_A2, 6_A3 |
6_B1, 6_B2, 6_B3 |
6_C1, 6_C2, 6_C3 |
Dudhsagar river basin |
Dudhsagar-Kuveshi hill range |
|
W7 |
7_A1, 7_A2, 7_A3 |
7_B1, 7_B2, 7_B3 |
7_C1, 7_C2, 7_C3 |
Uguem river basin |
Dhargini-Patiem hill range |
|
W8 |
8_A1, 8_A2, 8_A3 |
8_B1, 8_B2, 8_B3 |
8_C1, 8_C2, 8_C3 |
Netravali river basin |
Netravali-Ravon
dongor Hill range |
|
W9 |
9_A1, 9_A2, 9_A3 |
9_B1, 9_B2, 9_B3 |
9_C1, 9_C2, 9_C3 |
Kushawati river basin |
Corla hill range |
|
W10 |
10_A1, 10_A2, 10_A3 |
10_B1, 10_B2, 10_B3 |
10_C1, 10_C2, 10_C3 |
Agonda river basin |
Karmal ghat range |
|
W11 |
11_A1, 11_A2, 11_A3 |
11_B1, 11_B2, 11_B3 |
11_C1, 11_C2, 11_C3 |
Talpona river basin |
Cotigao-Ravon dongor hill range |
|
W12 |
Nil |
Nil |
12_C1, 12_C2, 12_C3 |
Kali river tributary basin |
Netravali-Ravon dongor Hill range |
Table 2. Table showing presence
(1) absence (0) of species across watersheds, * Endemic to Western Ghats of
India.
|
|
Species |
W1 |
W2 |
W3 |
W4 |
W5 |
W6 |
W7 |
W8 |
W9 |
W10 |
W11 |
W12 |
|
1 |
Duttaphrynus melanostictus |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
|
2 |
Pedostibes tuberculosus* |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
|
3 |
Euphlyctis cyanophlyctis |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
4 |
Minervarya gomantaki |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
|
5 |
Minervarya goemchi |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
|
6 |
Minervarya cepfi |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
|
7 |
Hoplobatrachus crassus |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
8 |
Sphaerotheca maskeyi |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
9 |
Microhyla ornata |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
|
10 |
Uperodon globulosus |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
11 |
Uperodon mormorata |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
|
12 |
Nyctibatrachus petraeus* |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
|
13 |
Clinotarsus curtipes* |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
14 |
Hydrophylax bahuvistara |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
|
15 |
Indosylvirana caesari* |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
|
16 |
Indirana salelkari* |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|
17 |
Polypedates maculatus |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
|
18 |
Pseudophilautus amboli* |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
19 |
Raorchestes bombayensis* |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
20 |
Rhacophorus malabaricus*
|
1 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
|
21 |
Ichthyophis davidi |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
|
22 |
Ichthyophis bombayensis |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
Table 3. Table showing alpha
diversity indices across watersheds.
|
Site |
Richness |
Shannon |
Simpson |
Evenness |
|
W1 |
15 |
2.35 |
0.88 |
0.87 |
|
W2 |
13 |
2.10 |
0.85 |
0.82 |
|
W3 |
19 |
2.19 |
0.85 |
0.74 |
|
W4 |
11 |
1.92 |
0.81 |
0.80 |
|
W5 |
12 |
2.28 |
0.88 |
0.92 |
|
W6 |
12 |
1.98 |
0.81 |
0.80 |
|
W7 |
13 |
2.07 |
0.85 |
0.81 |
|
W8 |
14 |
2.32 |
0.88 |
0.88 |
|
W9 |
14 |
2.17 |
0.86 |
0.82 |
|
W10 |
13 |
2.09 |
0.85 |
0.81 |
|
W11 |
15 |
2.32 |
0.89 |
0.86 |
|
W12 |
9 |
1.82 |
0.79 |
0.83 |
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