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Diversity and
distribution of anurans in Phansad Wildlife Sanctuary
(PWS), northern Western Ghats of India
Unmesh Katwate1, Deepak Apte 2 & Rupesh Raut 3
1,2 Bombay Natural History
Society (BNHS), Hornbill House, Opposite Lion Gate, Shaheed Bhagat Singh Road, Mumbai, Maharashtra 400001, India
3 Department of Zoology, Elphinstone College, Mumbai, Maharashtra 400032, India
1 theunmesh@gmail.com
(corresponding author), 2 spiderconch@gmail.com, 3 rupesh.raut@gmail.com
doi: http://dx.doi.org/10.11609/JoTT.o3038.3589-602 | ZooBank: urn:lsid:zoobank.org:pub:08F331D2-4CCB-4476-93FD-6CC7C934C75D
Editor: Sanjay Molur,
ZOO/WILD, Coimbatore, India Dateof publication: 26 February 2013 (online & print)
Manuscript details: Ms # o3038
| Received 16 December 2011 | Final received 08 December 2012 | Finally
accepted 31 December 2012
Citation: Katwate,
U., D. Apte & R. Raut(2013). Diversity and distribution of anurans in Phansad Wildlife Sanctuary (PWS), northern Western Ghats of
India. Journal of Threatened Taxa 5(2): 3589–3602; doi:10.11609/JoTT.o3038.3589-602
Copyright: © Katwateet al. 2013. Creative Commons Attribution 3.0 Unported License. JoTTallows unrestricted use of this article in any medium, reproduction and
distribution by providing adequate credit to the authors and the source of
publication.
Funding: BNHS Conservation Department
fund.
Competing Interest: None.
Author Contribution: UK and
RR have contributed equally in extensive field studies, analysis of data and preparation
of manuscript. DA gave important suggestions during field studies and
preparation of manuscript.
Acknowledgements: We
thank our co-researchers and field assistants from Bombay Natural History
Society (BNHS) for their contribution in field surveys. We are grateful to our
enthusiast wildlife photographer friends – Ajinkya Sawant, Saurabh Rane, Kunal Salunkhe,Aditya Sawant, Abhijit Gharat and Pritesh Nandvikar from Wild
Explorers (WE) for their contribution in photo documentation of different amphibian
species from study area. We are thankful to RFO and DFO, Maharashtra Forest
Department for permit (Letter No. D-4/Land/1290, Dt. 15 October 2010) and field
support; the museum and library authorities and the Director, Bombay Natural
History Society for providing reference facilities and the constant
encouragement, respectively; the Principal, ElphinstoneCollege for necessary support; Neelesh Dahanukar for providing valuable guidelines; and the
anonymous reviewers for constructive comments on the manuscript.
Author Details: Unmesh Katwate is a
zoology postgraduate currently working as Scientist-A in BNHS. He is studying
ecology and conservation aspects of freshwater fishes and amphibians in
northern Western Ghats and currently working on diversity and ecology of costal
plateaus of Konkan in Western Ghats.
Deepak Apte a marine biologist and conservationist,
currently a Chief Operating Officer (COO) of BNHS.
Rupesh Raut is a head of Department of Zoology, Elphinstone College. He works on ecology, systematic
biology and molecular phylogeny of freshwater fishes and amphibians of Western
Ghats.
Abstract: In global consequences of rapidly changing climate and increased
amphibian population decline, mapping amphibian diversity in biodiversity hotspots
is essential. In this study we have systematically studied anurans of Phansad Wildlife Sanctuary in terms of species diversity,
population structure, threat status and distribution. We recorded a total of 22
anuran species, of which 11 species are endemic to Western Ghats biodiversity
hotspot. Family Dicroglossidaewas found to be more species-rich. Spatial and temporal variation in anuran diversity was observed by using
Shannon diversity and evenness indices. Most of the endemic and threatened anuran
species are found to be associated with evergreen undisturbed forest
patches. Habitat parameters like
humidity, forest type, canopy coverage, riparian canopy coverage, stream
persistence and litter depth are found to be major variables governing species
diversity and distribution. Major
anthropogenic threats to amphibians of PhansadWildlife Sanctuary are discussed along with future conservation
objectives. With range extension of
species like Fejervarya caperataand Minervarya sahyadrisfurther north in the Western Ghats, taxonomic ambiguities recorded during study
are discussed briefly.
Keywords: Amphibians, diversity, endemism, species
composition, threat status.
The publication of this article is supported
by the Critical Ecosystem Partnership Fund (CEPF), a joint initiative of l’Agence Française de Développement, Conservation International, the
European Commission, the Global Environment Facility, the Government of
Japan, the MacArthur Foundation and the World Bank.
For
figures, images, tables -- click here
Introduction
The Western Ghats of India,
ranked among 34 biodiversity hotspots (Mittermeier et
al. 2005), are well known for rich amphibian
diversity. Around 6,771 amphibian
species are listed throughout the world, of which 333 species are recorded from
India (Frost 2011). With the
addition of one Raorchestes (Seshadri et al. 2012) the total number of amphibian species
listed in the Western Ghats are 183, among them 162 (88.5%) are endemic (Gururaja 2012). Deforestation, changes in land use, urbanization and industrialization
are major threats for the amphibians of Western Ghats (Daniels 1991; Ghate & Padhye 1996). Recent record of pathogenic fungus is
also alarming (Nair et al. 2011). Out of total 160,000km2 area of the Western Ghats only 9%
area is protected under national parks and wildlife sanctuaries (Gunawardene et al. 2007) therefore management of biological
resources is critical in the Western Ghats. To implement strong policies for amphibian conservation in the Western
Ghats, systematic study, spatial mapping and long term monitoring of amphibian
species are needed (Padhye & Ghate2002; Dahanukar & Padhye2005).
Amphibians of northern
Western Ghats are relatively less studied as compared to the central and the
southern Western Ghats. Species
discoveries like Gegeneophis nadkarnii (Bhatta & Prashanth, 2004), Indotyphlus maharashtraensis (Giriet al., 2004), Gegeneophis goaensis (Bhatta et al.,
2007), Pseudophilautus amboli(Biju & Bossuyt, 2009)
and Nyctibatrachus danieli(Biju et al., 2011) are significant findings from
this region. Taxonomic as well as
certain ecological studies by Daniel (1974), Yazdani& Mahabal (1976), Ravichandran& Pillai (1990), Daniels (1992), Ghate & Padhye (1996), Sekar (1999), Padhye et al.
(2002), Dahanukar & Padhye(2005), Biju & Bossuyt (2009), Kumbar & Patil(2010) and Biju et al. (2011) in Maharashtra region
of northern Western Ghats are mostly confined to Bhimashankar,Mulshi region of Pune District, Satara,Sangli District and Amboliregion of Sindhudurg District. Knowledge of amphibian diversity and its
species assemblage structure is still unknown from other parts of northern
Western Ghats, which is especially true for the Konkanregion.
Gokhale & Velankar(1996) demarcated Phansad Wildlife Sanctuary (PWS) as
a highly diverse region in Western Ghats because of its unique evergreen type
of vegetation. Presence
of regionally endangered and endemic semi-evergreen to evergreen plant
communities rank this area to be one among the highest conservation
priority zones of Maharashtra (Rodgers & Panwar1998). Although forests of PWS are
not directly connected to the main Western Ghats range but the similarity in
the occurrence of flora and fauna is striking (Rodgers & Panwar 1988; Pande & Pathak 2005). Diversity and distribution of amphibians in PWS is not fully understood
yet. In this study we have
systematically documented diversity and distribution pattern of anurans in
PWS. To our knowledge this is the
first systematic effort ever taken to document amphibian fauna in PWS.
Materials
and methods
Study area
PWS is situated on the lower
hills of the Konkan coast, west of Sahyadri and it covers about 52.66km2 area in
the Raigad District, Maharashtra. It lies between 18.33–18.50N
& 72.90–73.06250E with altitude ranging from 20m to
300m. Annual rainfall ranges
between 2162mm and 3469mm. Parts of
the study area have perennial sources of water bearing old evergreen type of
forest. Small hilly regions and
slopes show Memecylon umbellatumtree dominating the stunted evergreen type of forest. Plateau regions shows grassland type of
habitat whereas boundary line areas of PWS are dominated by degraded moist
deciduous type of forest. We have selected a total of eight different
localities in the sanctuary area representing different habitats (Table 1 &
Fig. 1). We surveyed this area from
June to September 2011. Even though our study period was limited to four
months, significant efforts were taken to give a broad exposure on status of
amphibian fauna of PWS.
Data collection and analysis
Anuran
species diversity in PWS was studied using two different methods. Ad-hoc search method was used to prepare
a checklist of species present in the study area and belt transect method was
used to quantify anuran diversity in different locations in PWS. Surveys were conducted mostly during
late evening and early night; some areas were also searched during day time. At all
sampling locations different habitat features like floor litter, underside
rocks and boulders, tree barks and leaves, water pools, rocky crevices near
flowing streams were documented and scanned for anurans. Sampling was performed by hand picking
the specimen. Efforts were
maintained to identify specimens up to species level in the field,body measurements of species taken with a digital caliper and individuals were
released back at original location. In case of unidentified species
representative voucher specimens were collected, fixed in 4% formalin and
stored in 70% alcohol. IUCN
guidelines were followed for scientific collection of some threatened anuran species
(IUCN 2011). Total of 3260
individuals were recorded of which only 17 were preserved for further
investigation and others were released in the same location. Voucher specimens were deposited at Konkan field station museum of Bombay Natural History
Society, Ratnagiri (accession numbers BNHS A1 to BNHS
A17). Identification of specimens
was done using standard taxonomic literatures like Boulenger(1890), Annandale (1919), Daniel (1963 a,b;
1975), Daniel & Sekar (1989), Bossuyt& Dubois (2001), Biju & Bossuyt(2009), and Biju et al. (2011). The threat status of the anuran species
is adopted from IUCN Red List of Threatened Species-Version 2011.1 (IUCN 2011).Site specific anthropogenic threats observed in the
study sites are listed in Table 1.
Biological
parameters like percentage canopy cover, forest type, forest floor litter
depth, riparian canopy cover (10m on both sides of the main stream) and
physical parameters like altitude, atmospheric temperature and relative
percentage humidity were recorded for each species during the study. Temperature and relative percentage
humidity were recorded using digital hygro-thermometer. Canopy cover was estimated using
spherical densiometer. All habitat variables were graded from
lowest to highest values. Details
of habitat guild classification are depicted in Table 2. Canonical Correspondence Analysis (CCA)
has been proven to be one of best multivariate statistical tool to extract
correlation between biological assemblage of species and their environment (Gururaja & Ramachandra2012). We analyzed the relationship
between biological and physical habitat parameters, study sites and species
abundance data through CCA by using PAST ver1.98 (Hammer et al. 2001). Quantification of anuran diversity was
performed by using the belt transect method. During the study period, two 100x5
m belt transects were laid at each study location every month. Transects were
laid in such a way that they would overlap all possible breeding habitats. Active search for
amphibians on transect were conducted by four people from 1700–2300 hr. Number of species and abundance of each
species were recorded for each transect by visual encounter. Actual number of individuals encountered
during transect sampling were used to calculate percent species abundance at
each study location in every month of the study period. Quantitative data
generated from transect studies were used to calculate diversity and evenness
indices (Magurran 1988). The Shannon index of
diversity was calculated using the equation:
H’
= –∑pi (lnpi),
where pi = ni/N and ni is the number of individuals of ith species and N = ∑ni.
Evenness index was calculated by the equation:
E
= H’/ln S.
Cluster
plot indicating species composition similarity between different study
locations was generated using Bray-Curtis similarity index (Magurran1988). Observed species abundance
may not always represent the true abundance of the communities therefore
adequacy of sampling efforts was assessed by using species accumulation curves.
Here, we have used EstimateS (version 8.0.0) software
developed by Colwell (2006) for generating species accumulation curves, which
uses Monte Carlo simulations of random samples drawn from the total set of
samples for estimating the average species richness. We ran 200 randomizations for a given
number of samples to estimate the species richness values and their means which were used for plotting species accumulation
curves with respect to number of samples as well as against number of
individuals recorded during study period. Species accumulation curves were found to be best
fitted by using Michaelis-Menton equation
Sobs= Smax n/ (B + n),
where Smax is the maximum number
of species, n is the number of individuals in the set of samples so far, and B
is the number of individuals needed to get half the maximum number of species, ie. When n = B, Sobs = Smax/2. Geographical map of study site (Fig. 1)
was prepared by using DIVA-GIS (Hijmans et al. 2002).
Results and Discussion
In
the four-month study period we documented a total of 22 anuran species belonging
to seven families and 15 genera (Table 3, Images 1–12). Family Dicroglossidaewas found to be more species-rich covering about 41% (nine species) of the
total anuran species recorded. Family Rhacophoridae was the second most
species-rich family in our study area covering 23% of total anuran species
(five species of arboreal frog). Family Microhylidae and Ranixalidaecontributed 13.63% and 9.09%, respectively to the total anuran species found in
the study area. Bufonidae,Nyctibatrachidae and Ranidaecontributed 4.5% individually to total anuran species. We have reported total 11 anuran species
(50% of total) from PWS showing exclusive endemism to Western Ghats. In the transect study we documented
total 22 anuran species with an abundance of 3260 individuals in 64
samples. Our study period was
restricted to four months only therefore there are possibilities of missing
some rare and cryptic species during sampling efforts. Figs. 2a & 2b shows species
accumulation curves plotted against number of samples (a) and number of
individuals (b). The curves were found to best fit (R2 =0.89) with Smax value at 22.
Study
site A and F reported 15 and 14 species respectively. Minimum species were
observed at sites G and H with 10 species each. Shannon H’ diversity index
values also predict the same pattern. Site A has highest H’ value (1.01), while
site G and H have lowest index values with 0.86 and 0.87. Despite of low species richness sites C,
D and E have more evenly distributed species abundance with an E index value
(S=11, E=0.94), (S=11, E=0.93) and (S=11, E=0.89) (Table 4). Anurans at site D
and E show maximum species endemism with seven species followed by site F and A
having six endemics of Western Ghats. Table 5 reveals that species richness and H’ index value are found to be
almost equal throughout June, July and August with (S=21, H’=1.14), (S=20,
H’=1.15) and (S=21, H’=1.11), respectively. A drastic fall in species richness
and H’ index value was observed in the month of September (S=15, H’=0.92).
Similar pattern was found to be followed by evenness in anuran abundance with
E= 0.78.
Our
study period covers wet period of northern Western Ghats as it starts from June
as south-west monsoon and ends in September. It has been already known that frogs in
northern Western Ghats reproduce mainly in June and July because of longer dry
period (Dahanukar & Padhye2005). Maximum
abundance was found in the month of June, about 1167
individuals were recorded. It is possibly because June marks the onset
of south-west monsoon and it’s a peak time for
amphibian reproduction. Furthermore
no drastic fluctuations in abundance were observed from July to September but
species diversity in the month of September declines drastically, only 15
species were observed in this month (Table 5). Fall in species richness, Shannon
diversity and evenness index values in September (Table 5) indicates that the
frogs in this region prefer earlier three months of wet period to reproduce and
at the end of September their breeding activity drastically declines. We have never observed any calling
activity of direct developing frogs like Pseudophilautuscf. amboli, Raorchestes bombayensis and Raorchestes cf. bombayensis as well as Wrinkled Frog, Nyctibatrachus humayuniin September. At broad level this
might explain that these species prefer highly humid early and mid wet seasons
for reproductive activity.
Habitat features plays an important role in governing species diversity and distribution
of amphibians (Becker et al. 2007; Santos-Barrera & Urbina-Cardona
2011). Alteration and loss of
habitat structure by multiple anthropogenic activities and changes in land use
have been identified as the most critical factors affecting amphibian survival
(Cushman 2006; Gardner et al. 2007; Urbina-Cardona
2008). In our study period we have
recorded different habitat features at each study site. Table 1 shows that study areas in PWS
were divided into three major types of habitats. Supegaon, Barshiv and Par Gaan bears
degraded moist deciduous forest, Chakacha Maal and Ghunyacha Maal plateau showing typical grassland habitat while core
areas of sanctuary like Chikhal Gaan,Phansad Gaan and Savarat Gaan showing Memecylon umbellatum dominated
evergreen type of forest. Table 1
reveals that areas like Supegaon, Barshivand Par Gaan are facing anthropogenic threats like
human encroachment in sanctuary areas, extensive grazing, monotypic teak wood
plantation, deforestation for firewood, slash and burn for shift cultivation
(agriculture purpose) etc. In comparison
of some physical and biological habitat parameters (Table 1) the Chikhal Gaan, Phansad Gaan and Savarat Gaan stands unique among other study sites in PWS because
of atmospheric temperature of these areas ranges between 26–29 0C
(vs. 28–33 0C in other areas), Humidity varies between
88–97 % (vs. 68–86 %) except Chakacha Maal and Ghunyacha Maal, canopy cover stands between 60–91 % (vs.
20–60 %), riparian canopy cover varies between 75-91% (vs. 0–63 %)
and forest floor litter depth stands between 70–210 mm (vs. 0–70 mm
at other sites). For implementation
of systematic conservation efforts in the Western Ghats, biodiversity hotspot
prioritization of highly diverse areas is essential (Myers 1990; Myers et al.
2000; Dahanukar & Padhye2005; Mittermeier et al. 2005). Analysis of species assemblage
similarity (Fig. 3) between study areas reveals Chikhal Gaan and Phansad Gaan forming a single and distinct cluster while other
study areas remains separated.
Environmental
parameters play an important role in species distribution, therefore
understanding relationships between habitat characteristics and species
distribution pattern is vital for developing effective conservation strategies
for threatened taxa (Boyd et al. 2008). Anthropogenic activities like deforestation, construction of dams, human
encroachment, slash and burn techniques for shifting cultivation, road
construction and mining. had been known to be possible
causes of changes in the habitat structure at landscape level which ultimately
affects species diversity and their assemblage (Gururajaet al. 2008). It is already known
that habitat destruction due to several anthropogenic activities has had a
negative impact on amphibian diversity and distribution in Western Ghats
(Krishnamurthy 1996, 2003; Padhye et al. 2002; Dahanukar & Padhye 2005;
Krishnamurthy & Reddy 2008). Relationship between anuran species, study sites and habitat variables
were predicted by using canonical correspondence analysis (CCA). Axis 1 & 2 represents species points
which are determined by weighted average of axis values of preferred habitat
variables and study sites in which they were occurred. CCA analysis extracted different axis
scores for species point, study site and for habitat variables. Two best axis scores representing
maximum variation in data were selected to draw the ordination plot (Fig.
4). Species points in this triplot formed their unique niche clusters with respect to
different study locations and habitat variables. Nyctibatrachus humayuni, Sphaerotheca dobsonii, Indirana beddomii, I. leithii, Polypedates cf. maculatus,Pseudophilautus cf. amboli,Raorchestes bombayensis andR. cf. bombayensis forms a distinct
species cluster showing two dimensional unique niche because they share similar
type of habitat and prefers more humid evergreen forest patches. Like species points, similar habitat
sharing study sites also form their clusters. CCA ordination triplotindicates variables like canopy cover, riparian canopy cover, forest floor
litter depth, humidity, temperature, streampersistence and forest type shows high negative correlation with axis 1. Species and study sites present in
ordination with axis 1 have high influence of environmental variables. Variable
like altitude found to be positively correlated on axis 2. Species like Duttaphrynus melanostictus, Euphlyctis cyanophlyctis, Fejervarya cf. keralensis, F. syhadrensis,Hoplobatrachus tigerinus, Minervarya sahyadris, Sphaerotheca breviceps, Ramanella mormorata and Uperodon globulosus were
found to be present in human disturbed areas like Supegaon,Barshiv and Par Gaan as
well as it has been found that they are independent from habitat variable
influence.
On
a broad scale our current analysis of anuran distribution in PWS with respect
to different study localities and habitat parameters (Fig. 4) indicates that
anuran species like Nyctibatrachus humayuni, Indirana beddomii, I. leithii, Polypedates cf. maculatus,Pseudophilautus cf. amboli,Raorchestes bombayensis andR. cf. bombayensis were restricted to
evergreen forest areas like Chikhal Gaan, Phansad Gaanand Savarat Gaan because of
their specific habitat requirements like dense canopy coverage, maximum forest
floor litter, low atmospheric temperature and high humidity. Whereas species like Duttaphrynus melanostictus, Euphlyctis cyanophlyctis, Fejervarya cf. keralensis, F. syhadrensis,Hoplobatrachus tigerinus, Minervarya sahyadris and Sphaerotheca brevicepsshowing maximum occurrence in human disturbed areas like Supegaon,Barshiv and Par Gaan having
minimum canopy cover, minimum forest floor litter, low humidity and
comparatively high temperature. CCA
analysis results indicate that maximum conservation priority should be given to
regions having high influence of habitat variables. Priority regions for long term anuran
species and habitat monitoring in PWS should be Chikhal Gaan, Phansad Gaan and Savarat Gaan.
In
this study we have recorded the occurrence of Minervarya sahyadris (Image 13) from PWS. Identification
characteristics like rictal gland present just behind
the mouth commissure, white horizontal band along the upper lip, smaller snout
to vent length, pointed snout, fold from eye to shoulder, tympanum dark brown
in color with inferior border white, digit tips rounded, very less developed
webbing and reddish-brown mid dorsum color were observed among specimens
recorded from PWS. This endemic
species was earlier recorded from Calicut in Kerala and Gundia,
Jog, Dandeli, Castle Rock in Karnataka (Dubois et al.
2001; Gururaja 2012). Occurrence of M. sahyadrisin PWS shows the northward range extension of this species in Western Ghats.
Another endemic dicroglossid Fejervarya caperata (Image 14) was also recorded during
study. Characters like pointed snout, 20–35 mm snout to vent length,
dilations absent on figure or toe tips, presence of fejervaryanline on the sides of abdomen, three distinct cross bars on thigh and four
distinct longitudinal ridges on the back were observed among F. caperata specimens sampled during the study. Earlier records of this species were
found from Karnoor, Bajpe, Shimoga, Jog, and Dandeli in
Karnataka parts of Western Ghats (Kuramoto et al.
2007; Gururaja & Ramachandra2012; Gururaja 2012). Record of this species in PWS also marks
its range extension further north in Western Ghats.
We
have reported some frog species from study area having ambiguous taxonomic
status. Biju & Bossuyt (2009) described Pseudophilautus amboli from Amboli, Sindhudurg District of Maharashtra. Our PWS collections of Pseudophilautus cf. ambolidiffers from Pseudophilautus amboli (Biju & Bossuyt 2009) sensu stricto in the ratio of head width vs. head length as well
in the proportion of third finger disc diameter vs. finger width. In
consideration of such ambiguities further study is required to obtain presence
of this species in PWS. Some
population of Raorchestes bombayensis from our collection differs from original
description of Annandale (1919) and latest reviewed description given by Biju & Bossuyt (2009).Raorchestes bombayensis (Annandale 1919; Biju & Bossuyt 2009) sensu strictodiffers from our PWS collection in having head width greater than head length
(vs. head width almost equal to head length) and smaller eye length. Occurrence of Fejervaryacf. keralensis also needs further
confirmation as it lacks spinular projections on
thigh region. We have also
collected some specimens of tree frogs Polypedatesfrom our study area. Although some
of our specimens matches very closely with description given by Daniel & Sekar (1989) for Polypedates maculatus, but they also differ in having
comparatively smaller snout to vent length, tympanum almost equal to eye
diameter and distinct striped banding pattern on dorsum.
Global
threat status analysis of anurans recorded in this study (Table 3) shows that
about seven species (32%) are threatened (Vulnerable (VU), Endangered (EN) or
Critically Endangered (CR)). Fourteen species come under Least Concern (LC) category while one is
Data Deficient (DD). Occurrence of
CR species like Pseudophilautus cf. amboli as well as some EN and VU species like Minervarya sahyadris,Nyctibatrachus humayuni,Indirana leithiiand Raorchestes bombayensisis important from conservation point of view.
In
conclusion occurrence of around 50% endemic anuran species of which 32% are
globally threatened marks PWS as key site for amphibian diversity in Western
Ghats biodiversity hotspot. Implementation of strong conservation measures and
policies are needed in PWS for amphibian conservation. This study would be beneficial in this
context.
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