Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 February 2021 | 13(2): 17630–17638
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.6628.13.2.17630-17638
#6628 | Received 27 August 2020 | Final
received 06 February 2021 | Finally accepted 19 February 2021
Faunal diversity of an insular
crepuscular cave of Goa, India
Pratiksha Sail 1, Manoj Ramakant Borkar 2, Ismat Shaikh 3 & Archana Pal 4
1–4 Biodiversity Research Cell,
Department of Zoology, Carmel College of Arts, Science & Commerce for
Women, Nuvem, Goa 403713, India.
1 pratikshasail2210@gmail.com, 2
borkar.manoj@rediffmail.com (corresponding author), 3 ismatshaikh27@gmail.com,
4 pal.archana9272@gmail.com
Editor: Jayant
Biswas, National Cave Research and Protection Organization, Raipur, India. Date
of publication: 26 February 2021 (online & print)
Citation:
Sail, P., M.R. Borkar, I. Shaikh & A. Pal (2021). Faunal
diversity of an insular crepuscular cave of Goa, India. Journal of
Threatened Taxa 13(2): 17630–17638. https://doi.org/10.11609/jott.6628.13.2.17630-17638
Copyright: © Sail et al. 2021. 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: Self-funded.
Competing interests: The authors declare no competing interests.
Author details: Ms. Pratiksha
Sail has a master’s degree in Zoology
from Goa University. She has been conducting research in the area of ecology
and conservation science working on impact of linear infrastructure projects
and EIA. Dr.
Manoj Ramakant Borkar, senior academic and zoologist with 33 years
of teaching and research experience, has served on Goa’s State Biodiversity
Board, State Wildlife Board and State Experts Appraisal Committee for EIA.
Presently serving as Associate Professor & Head, Department of Zoology at
Carmel College for Women, Nuvem Goa. Ms. Ismat Shaikh
has a master’s degree in Zoology from Goa University. Ms.
Archana Pal has a master’s degree in Biotechnology from Birla Institute
of Technology (BITS), Mesra. She has been pursuing
research in the area of microbial degradation.
Author contribution: PS—Field work, laboratory analysis, computation of
data and writing first draft; MRB—Conceptualization and supervision of field
& laboratory work, data collation and interpretation, writing final
research manuscript and editing; IS—Field work and laboratory analysis;
AP—Field work and laboratory analysis.
Acknowledgements: Senior corresponding author Dr.
Manoj R. Borkar expresses gratitude to Dr. Pradeep Sarmokadam, Member
Secretary of Goa State Biodiversity Board for encouragement towards
biodiversity listing of hitherto unexplored ecosystems in Goa. Ms. Andrea Sequeira
is acknowledged for assistance in field
photography. Thanks are also due to
Superior General of Apostolic Carmel Congregation Sr. Maria Nirmalini
AC for upgrading facilities at the Biodiversity Research Cell of Carmel College
of Arts, Science & Commerce for Women, Nuvem,
Goa.
Abstract: This study is the first attempt to document
troglofaunal diversity of crepuscular cave ecosystem from the state of
Goa. Twelve faunal species (seven
invertebrates and five vertebrates) have been documented from an insular
crepuscular cave which measures 18.62m in floor length and shows a transition
of light and hygrothermal profile between its
entrance and dead end. Absence of
primary producers, thermal constancy, high humidity, poor ventilation, and
competitive exclusion due to limited food resources restricts the faunal
diversity of this cave; though trophic linkages are interesting yet
speculative, as is typical of subterranean ecosystem. Among the macro-invertebrates, cavernicolous Whip Spider is a significant species here;
whereas the important vertebrates encountered are the Fungoid frog and the
Indian Cricket frog, besides roosts of the Rufous Horseshoe bat. Eco-energetic subsidy, possibly offered by
crickets and bats that regularly feed outside this oligotrophic cave ecosystem
is discussed. The need to document the
unique and vulnerable troglofauna of this sensitive
ecosystem from the conservation perspective is highlighted.
Keywords: Crepuscular, eco-energetic subsidy, insular cave, troglofauna.
INTRODUCTION
Caves are natural or man-made subterranean cavities
that may also be formed in the face of a cliff or a hillside. Their speleogenesis is attributed to various geologic processes;
inclusive of a combination of chemical processes, erosion by water, tectonic
forces, microorganisms, pressure and atmospheric influences (Ford &
Williams 2007).
Cave as an ecosystem has its unique features characterized
by gradients of darkness, narrow range of temperature differences and high
humidity with limited air currents as some of its abiotic factors. These features make the cave ecosystem unique
and the study of its biota fascinating (Biswas 2009). Bio-speleology is a relatively nascent area of
ecology in India; the dearth of bio-speleological data being on account of the
relatively hostile and secluded nature of these ecosystems.
The cave environment creates an ecosystem, which the
fauna living in epigean conditions find difficult to adapt to (Biswas
2010). Nonetheless, caves support and
sustain a good faunal diversity within the limited geophysical parameters. Organisms attracted to live in caves show
high degree of morphological, behavioral, and
physiological adaptations necessary to endure and thrive in such an ecosystem (Vandel 1965; Barr 1968; Biswas 1992; Gunn 2004). Species colonise caves for various reasons
for at least part of their life cycles for temporary shelter or due to low
predation pressure and easy availability of prey (Biswas 2009).
This study is the first attempt at charting out the
faunal diversity of crepuscular cave ecosystem of Goa’s insular landscape. The term insular is used here to indicate the
cave’s presence on an island. Divar is the third largest of the seven islands of Goa;
roughly triangular in shape and measures an average 5.80km long and about
3.00km wide, with an area of around 17.56km2. A cursory mention of this cave ecosystem of Divar Island appears in the book “Island Biodiversity Goa”
(Sawant & Jadhav 2014).
Present study is particularly intended to obtain data
on the troglofaunal diversity inhabiting the cave. In the present case the crepuscular cave
investigated is a relatively small subterranean ecosystem with a floor length
of 18m and average roof height of 3.4m; as such the environmental variables
typical of a classical cave resulting in distinct zonation as proposed by
Vermeulen & Whitten (1999) cannot be applied here. Nonetheless, from the cave entrance towards
the dead end it is possible to demarcate three zones using variation in hygrothermal profile and diminishing light intensity as
zone boundaries. The organic carbon and
phosphate content of the soil through the floor length of the cave have been
analysed. The data gathered addresses an
important gap in our state specific biodiversity of sensitive ecosystems.
MATERIALS AND METHODS
The cave investigated in this study is located at
15.5350N & 73.9240E on Divar
Island towards the north of Old Goa, 9.50km from Panaji and surrounded by the Mandovi estuary and its network of tributaries (Image 1a,b
). The island is dominated by agricultural
landscape and a few aquaculture units (Image 1c).
The cave was physically measured for various speleometric dimensions and a schematic diagram was hand
drawn by mapping the layout of the cave and later digitized to add details like
measurements and sampling points.
Physical parameters such as light (Luxmeter
HTC Model LX-103), temperature and humidity (Digital Hygrometer-Thermometer HTC
Model 103-CTH) were recorded throughout the cave.
Soil samples from various points within the cave were
collected in pre-sterilized zipped plastic pouches using a spatula and analysed
for organic carbon (Walkley & Black 1934) and phosphate content (Adelowo et al. 2016).
Monthly visits were made for seven months from August 2018 to
February 2019 for documentation of troglofauna in
relation to three zones of the cave.
Only direct evidences were considered to compile an inventory of troglofauna. Most
observations were made between 09.00h and 17.00h, but occasional observations
were made at night to account for any night cave visitors. Species were photo-documented and collection
was avoided to uphold conservation ethics, unless absolutely necessary for
taxonomy.
The various troglofaunal species observed and
photo-documented in this cave were identified based on taxonomic guides, as
also consultations with taxon experts (Chopard 1970; Bastawade 1995; Whitaker 2006; Srinivasulu
et al. 2010; Gururaja 2012; Keswani
et al. 2012; Baidya & Chindarkar
2015). Notwithstanding that much larger
sample sizes instill confidence for detecting small
changes between sampling times or sites, in this exercise semi-quantitative
ACFOR scale was used for the rapid assessment of abundance of the cave
fauna. The utility of ACFOR scale in
coarse assessment of abundance, both accurately and quickly, has been proven
(Hawkins & Jones 1992).
RESULTS & OBSERVATIONS
The investigated cave on Divar
Island in Goa is a natural subterranean vault at the base of a cliff (Image
1). This crepuscular cave is lateritic
and damp with progressively diminishing light from the entrance to the end
chamber, totaling a floor length of 18.62m (Image 2
& Fig. 2). Schematic diagrams (Fig.
1 & 2) depict the measurements of the cave.
The cave entrance is 3.52m wide and opens up horizontally into a chamber
typed as the zone A (proximate to the entrance), since it had less light and
vegetation than outside and the temperature and humidity were found to be
30.5°C and 80%, respectively (Fig. 2 & 3, sampling point C). The next cave segment with light intensity
decreasing further was considered to be the zone B (middle zone) with
temperature of 30.4°C and humidity at 97% (Fig. 2 & 3, sampling point
D). The zone C (end zone) is characterized by complete darkness with
99% humidity and temperature ranging between 29.1–30.1°C (Fig. 2 & 3,
sampling points E & F). The
intensity of light decreased with increasing distance from the cave entrance to
the end zone, average light of 1782.13 lux and 0 lux being recorded
respectively at the two extreme points.
The organic carbon and phosphate content of the soil from sampling sites
along the three cave zones is shown in Fig. 5 & 6 and correlates well with
roost positions of the bats (Fig. 4), values being higher in the zone B and C
as compared to zone A.
In the present investigation, 12 faunal species were
recorded from various zones of the cave; (See Table 1 and Image 3). Two species of frogs have been encountered in
the cave, namely, Fungoid Frog Hydrophylax malabaricus and Indian Cricket Frog Fejervarya
limnocharis; the sightings of the former being
more frequent after the rains extending in distribution from the zone A to Zone
C, but the latter more confined to zone A.
Often called the Paddy Frog, Fejervarya
limnocharis is a common species of croplands
here. Brooke’s Gecko Hemidactylus brookii occurs
from the entrance through the zone A of the cave co-inhabited by Scutigeromorph Centipede Scutigera
coleoptrata, Woodlouse unidentified species,
Daddy Longlegs Puria dorsalis, Humped
Spider Zosis geniculata
and Whip Spider Phrynichus phipsoni. The Cricket Itaropsis
parviceps and Long-necked Sugar Ant Camponotus angusticollis were
associated with zone B of the cave. The
Whip Spider Phrynichus phipsoni
population of this cave exhibits site fidelity, with individuals dispersing at
night on the cave walls from their
hideouts in crevices.
The Rufous Horseshoe Bat Rhinolophus rouxii is an important mammalian constituent of troglofauna in this cave, its population and distribution
varying seasonally. In the wet months
dense bat roosts occupied the zone B and C.
In the zone A, five individuals of Rufous Horseshoe Bat were seen
clustered on extreme right wall, where light intensity was perceptibly
less. Bats preferred to roost on the
walls rather than the roof of the cave.
In the month of November 2018, the bat density decreased and the roosts
were restricted to zone B and C. By
February 2019, the numbers dropped further; barring a small cluster of bats in
the distal end of the zone B and a roost of predominantly sub-adults was
confined to the end part of the cave.
Such seasonal shifts in the distribution and number of the bat roosts
within this cave (see Fig. 4) accounts for absence of a thick bulky guano
deposit in this cave and the guano is loosely scattered on the cave floor.
A single individual of Common Krait Bungarus caeruleus
documented from this cave during the night survey is most likely to have been a
straggler or an opportunistic predator that had been attracted to the cave,
after seeing concentrated food supply in form of roosting bats. Such opportunistic predations on cave
roosting bats have also been recorded by Tanalgo et
al. (2020).
All the species listed in this study are secondary
consumers and saprotrophs. The smaller
invertebrates help in remineralization of dead organic matter, as also
constitute the prey base for lower rungs of secondary consumers such as daddy
longlegs, Scutigeromorph Centipede, humped spider,
and whip spider. The herpetofauna and
bats operate at the next trophic level, whereas the Common Krait could either
be a straggler or an opportunistic predator of frogs, lizards, and bats. From the troglofaunal inventory compiled in
this study, it is evident that this cave has a truncated biodiversity typified
by absence of phototrophic primary producers and presence of fewer apex
predators in the food chains operating here.
DISCUSSION
Perusal of literature confirmed a glaring gap in
documentation of troglofauna in the state of
Goa. Deciphering the composition and
dynamics of cave communities has been a key challenge for speleobiologists. The limited organization of a cave ecosystem
has been attributed to permanent darkness and competitive exclusion due to
resource scarcity (Fernandes et al. 2016).
Despite a relatively hostile cavernous environment characterized by
diffuse or complete absence of light, constant temperature and high humidity,
poor air circulation and severely constrained food supplies; caves do support
and sustain a unique assemblage of biota, whose density however is lower than
epigean habitats (Mitchell et al. 1977; Parzefall
1983). The relatively poor faunal
diversity of this cave ecosystem could also be attributed to the absence of
phototrophs; resulting in scarcity of food, reducing the number of predators as
well as the overall biodiversity of the cave ecosystem (Gibert
& Deharveng 2002). Species might colonise cave environments for
reasons such as the need for temporary shelter or in order to escape from
persistent adverse environmental conditions on the surface, whereas others may be
temporary visitors with limited reliance on the cave environment.
Several researchers have conclusively shown that phenology
of species and in particular that of ectotherms, is influenced by air
temperature and water availability (Kearny et al. 2013; Amarasekare
& Coutinho 2014; Sheldon & Tewskbury
2014) and variations in these features in epigean environments often force
animals to search for ‘shelter microhabitats’ of stable environments such as
caves that offer the most suitable and stable conditions (Seebacher
& Alfrod 2002; Papaioannou
et al. 2015).
Expectedly, the variations in microclimatic attributes
across the various cave zones create an environmental gradient, which
influences distribution of various faunal elements. Such patterns of macro-invertebrate
distribution in a cave ecosystem have also been confirmed by Mazebedi & Hesselberg
(2020). The basic tenet of
eco-energetics requires linkages between biodiversity and the abiotic
components; and the species richness is limited by productivity (Ricklefs & Schluter 1993). As is also the case with the cave
investigated here, macro-invertebrates usually play a critical role in cave
ecosystem functions because of their relatively high diversity compared to the
vertebrate biota. Such an opinion has
also been corroborated by Lavoie et al. (2007) and Moseley (2009). Crickets and bats subsidise the consumer
community dynamics of this oligotrophic cave ecosystem. The crickets are known to take regular
nocturnal feeding sorties on vegetation outside the cave (Benoit et al. 2004)
and the bats offer guano subsidy by feeding in epigean habitats (Iskali & Zhang 2015).
In the present case too, both these species offer a definite trophic
connect to the cavernicoles of this cave, with the
epigean resources.
Epigean populations of Phrynichus
phipsoni have been reported from Goa earlier by Borkar et al. (2006).
As documented from this cave; whip spiders also inhabit the crevices and
cracks in the subterranean ecosystems that commensurate well with their dorso-ventrally flattened body contour (Chapin 2015). Their site fidelity and homing behavior as has been observed in this investigation, has
been well documented (Hebets 2002).
Herepetofaunal constituents of this cave are two species of frogs,
one species of gekkonid lizard belonging to the Hemidactylus
clade complex of brookii group (Bauer et
al. 2010a,b) and the Common Krait. The
association of the Fungoid Frog with subterranean caves of Western Ghats is
well established (Chari 1962) and has also been observed in the caves of Kanger Valley National Park, Chattisgharh
(Biswas 2010; Biswas & Shotriya 2011). Caves are known to accumulate heat and create
a microclimate favouring macrofaunal poikilotherms such as the amphibians and
reptiles, and the abundant invertebrate species here remove limitations of food
for these opportunistic predators (Turbanov et al.
2019). A single observation of Common
Krait in this cave presumably for opportunistic feeding is corroborated by
Sinha (1999), who has also reported Banded Kraits preying on Bats from Siju caves in Meghalaya.
The presence and roosting habits of Rufous Horseshoe
Bat in caves of the Western Ghats have been previously reported by Korad et al. (2007).
The variations in numbers and size of the bat roosts observed in this
study is speculated to be driven by the species-specific social structure and
foraging behaviour (Kunz & Lumsden 2005). The relatively sparse and scattered guano in
this cave correlates well with shifting positions and density of the bat
roosts. Similar observations have also
been reported by Biswas & Shrotriya (2011). Bat guano may support guanophile
communities that in turn could attract predators of these guanophages
to the cave (Encinares 2019). Perhaps in a small subterranean cave like the
one investigated here the species diversity may seem small, because all micro
crustaceans and cavernicolous guanophiles
have not been included.
CONCLUSION
On Divar Island where this
cave is located, locals have consciously resisted urbanization thus far; but
the place is a popular location for Bollywood film shoots. Tourists have been steadily pouring in to
relish the rustic countryside. Influx of
tourists will open up the hitherto inviolate areas for exploration and
exploitation. Ecological studies of cave
ecosystems and charting out their troglofauna are a
prerequisite for management and conservation of sensitive and fragile
subterranean ecosystems (Schneider & Culver 2004). Conservation of cave ecosystems is vital not
only because they shelter unique and vulnerable biodiversity (Mammola 2019), but more so because their stable
environments provide natural laboratories for testing doctrinal evolutionary
concepts of adaptation and speciation (Culver & Pipan
2019). Also, collection and collation of
a standardised data for a long term referral purpose is crucial for species
conservation assessment (Lunghi et al. 2020).
Table 1. Zone of occurrence and abundance of faunal
diversity of the crepuscular cave in Divar, Goa.
|
Common
name |
Species |
Zone
of occurrence |
Abundance
ratings |
1 |
Scutigeromorph
Centipede |
Scutigera coleoptrata Linnaeus,
1758 |
A |
F |
2 |
Woodlouse |
Unidentified
Isopod |
A |
F |
3 |
Cricket |
Itaropsis parviceps Walker,
1869 |
B |
C |
4 |
Long-necked
Sugar Ant |
Camponotus angusticollis Jerdon,
1851 |
B |
C |
5 |
Daddy
longlegs |
Puria
dorsalis Roewer,
1914 |
A |
C |
6 |
Humped
spider |
Zosis geniculata Olivier,
1789 |
A |
O |
7 |
Whip spider |
Phrynicus phipsoni Pocock,
1900 |
A, B &
C |
F |
8 |
Fungoid Frog |
Hydrophylax malabaricus Tschudi,
1838 |
A, B &
C |
F |
9 |
Indian Cricket Frog |
Fejervarya limnocharis Gravenhorst,
1829 |
A |
R |
10 |
Brooke’s
Gecko |
Hemidactylus brookii Gray,
1845 |
A |
O |
11 |
Common
Krait |
Bungarus caeruleus Schneider,
1801 |
A |
R |
12 |
Rufous
Horseshoe Bat |
Rhinolophus rouxii
Temminck,
1835 |
B & C |
A |
Abundance rating are based on the semi-quantitative
visual estimates: A—Abundant | C—Common | F—Frequent | O—Occasional | R—Rare.
For
figures & images - - click here
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