Herpetology Notes, volume 5: 203-209 (2012) (published online on 11 June 2012)
Feeding ecology of the Travancore tortoise
(Indotestudo travancorica) in the Anamalais, Western Ghats, India
Deepak Veerappan and Karthikeyan Vasudevan*
Abstract. The feeding ecology of the Travancore tortoise (Indotestudo travancorica) was studied for four years (2006-2009) in the
Anamalais, Western Ghats, south-western India. The diet of I. travancorica was examined and quantified based on faecal matter
and direct feeding observations. Indotestudo travancorica is a primarily herbivorous species which feeds on herbs, grasses and
fruits. The recovered remains of several animal species including invertebrates (insects, millipedes, molluscs, scorpions and crabs)
and remainders of mammalian hair suggest that a considerable number of live invertebrates and carcasses of larger vertebrates
are also consumed, albeit in lesser quantity, which was confirmed by in-situ observations. The array of diet items consumed by I.
travancorica suggests that its diet is complex and it needs to be taken into consideration in conservation breeding programmes.
Keywords. Diet, feeding, grass, season, tortoise.
Introduction
The diet of a species is key to its survival. An
understanding of the diet of a species also provides
information on the role it plays in the ecosystem.
Numerous studies have attempted to identify factors
that are important in the choice of diet (for review see,
Pyke, 1984). In reptiles, most species are carnivorous
although numerous turtles and lizards are at least partly
herbivorous, of which several are known or suspected
to play a role in seed dispersal (Braun and Brooks,
1987; Macdonald and Mushinsky, 1988; Moskovits and
Bjorndal, 1990; Moll and Jansen, 1995 and references
therein; Mason et al., 1999; Strong and Fragoso, 2006).
However, few in-situ studies have been carried out
with a focus on the diet of reptiles in the oriental region
(Corlett, 1998; Platt et al., 2001; Platt et al., 2007; Ihlow
et al., 2012).
Indotestudo travancorica is a tropical forest-dwelling
tortoise, endemic to the Western Ghats Mountains of
southern India (Iverson et al., 2001; Le et al., 2006). This
species is found in a variety of habitat types composed
of evergreen, moist deciduous, and bamboo forest,
semi evergreen forests and rubber and teak plantations.
Within these habitats I. travancorica can often be found
near marshlands, grass clearings in the forest, and rocky
outcrops close to streams (Vijaya 1983; Bhupathy
Wildlife Institute of India, Post Box 18, Chandrabani, Dehradun, Uttarakhand -248001, India.
*Corresponding author; email: [email protected]
and Choudhury, 1995). Indotestudo travancorica is
considered “Vulnerable” in the recent Red List of
threatened species (Asian Turtle Trade Working Group,
2000). The diet of I. travancorica has been reported to
constitute mainly of grass, bamboo shoots, herbs, fruits,
insects, millipedes and frogs. However, this statement
was solely based on anecdotal observations presented
as brief notes, based on observations of free-ranging
individuals (Vijaya, 1983; Ramesh and Parthasarathy,
2006) and observations in captivity (Das, 1991). In this
study, the diet of I. travancorica was quantified and
described based on direct observations and analyses of
faecal remains.
Materials and Methods
Field studies on the feeding behaviour of I. travancorica were
carried out in the Anamalai and Parambikulam Tiger Reserves,
which are part of the Anamalais mountain range. These two
Protected Areas (PA’s) are located between 10°13- 10°33’ North
and 76°37’-77°21’ East. The study area includes a total of 1244
km2, representing 959 km2 in the Anamalai Tiger Reserve (ATR)
and 285 km2 in Parambikulam Tiger Reserve (PTR). However,
this study was carried out only in middle elevations of the PA’s,
covering an area of 396 km2. The study area is characterized by
a mosaic of vegetation types; southern tropical wet evergreen
forest consist of Dipterocarpus bourdilloni – Strombosia
ceylanica associations (Pascal, Ramesh and De Franceschi,
2004); bamboo forest with mixed deciduous trees dominated
is by Grewia tiliaefolia, Terminalia tomentosa, Lagerstroemia
lanceolata and Cassia fistula; moist deciduous forest is composed
of Terminalia tomentosa, T. bellarica, T. paniculata, Dalbergia
sissoo and Dillenia pentagyna; and reed brakes are composed
of Ochlandra spp (Champion and Seth, 1968). Riparian forests
have been characterised as distinct vegetation types in this region
204
(Sekar and Ganesan, 2003). Thirty three percent of the protected
area is covered by monoculture teak plantations (Wilson, 1973).
Indotestudo travancorica was found in all vegetation types.
Faecal remains have been widely used to examine dietary
components of tortoises (MacDonald and Mushinsky, 1988;
Moskovits and Bjorndal, 1990; Mason et al., 1999; Hailey,
Coulson and Mwabvu, 2001). All captured individuals were kept
individually inside a large airy cotton cloth bag for about 12 h,
during which they usually defecated. The faeces of individual I.
travancorica were collected and dried under a 40 W incandescent
lamp. The dried material was subsequently examined using a 10X
hand-held lens and separated into the following diet components:
grass, scorpions, molluscs, insects, millipedes, seeds, other
plant materials (other than seeds) and sand. Animal hairs were
identified using a Leica EZ4™ microscope at 35 x magnification.
A mammal hair identification key (Bahuguna et al., 2010) was
used for identification. Diet components in individual faecal
samples were scored as; 1 – low, 2 – medium and 3 – high, based
on the relative quantity of dried material found in each faecal
sample.
Whenever possible, opportunistic observations of feeding
behaviour were also made. Ad-libitum observations were made
by a single observer located 8-10 m away from the animal to
avoid disturbance. The plant or animal parts which a tortoise fed
upon were collected and identified.
For subsequent analyses, individuals below 160 mm straight
carapace length were considered as juveniles, while males were
differentiated from females based on the concavity in the plastron
of the adult males. The collected data was separated into diet
components of plant and animal origin. A chi-square test for
proportions (Gibbons, 1971) was used to test for differences in
the proportion of different diet categories between (i) Monsoon
(June to November), Post monsoon (December to February) and
Summer (March to May) and (ii) male, female and juvenile.
Results
Thirty two faecal samples were collected from 8 males,
15 females and 9 juveniles during the study. All faecal
samples contained the remainder of at least one dietary
item. Overall, plant matter had the highest contribution
in the obtained faecal samples, followed by other
classes: 90.6% contained grass and bamboo remains,
93.8% contained other plant materials (leaf remains,
fiber and twigs); 37.5% contained seeds; 75% contained
insect remains; 62.5% contained sand; 21.9% contained
vertebrate remains (hair, bones and scales); 18.8%
contained mollusc remains; 9.1% contained crab remains
(unidentified species) and scorpion (Heterometrus sp.)
remains and 3.1% contained millipede remains. Several
encountered remains were too small to identify and
assign to any of the diet categories. These items were
classified as “unidentified”; 71.9% contained such
remains. Millipede remains were found only in juvenile
faecal samples, while the remaining dietary items were
Deepak Veerappan & Karthikeyan Vasudevan
represented in males, females and juveniles (Fig. 1A).
The proportion of dietary components between sexes did
not significantly differ (Table 1). Plant remains (grass,
seeds & other plant matter) were found across all three
seasons. All diet components were present in samples
obtained during the monsoon. The proportion of the class
other plant material in the diet was significantly different
between seasons (Table 1). However, the proportion
within the grass and seed classes did not significantly
differ across seasons. Scorpion, mollusc and millipede
remains were not found the samples collected during
summer; crabs and millipede remains were not found
in samples collected during post monsoon (Fig. 1B).
There was no significant difference in the animal matter
represented in the obtained samples between seasons.
Seeds obtained from faecal samples were identified to
belong to Grewia tiliaefolia, Lantana camera, Dillenia
pentagyna and Gomphandra sp. These seeds were
present in the samples collected from all the sex classes.
The proportion of seeds found among different sexes
was not significantly different (Table 1). Seeds were
represented across all three seasons and their proportion
of occurrence in the samples was not significantly
different across seasons (Table 1). Ten out of 32 samples
(37.5 %) contained seeds. Overall, 28% of the faecal
samples examined contained animal matter. Twenty six
out of the thirty two faecal samples examined (16.5%)
contained invertebrate remains including millipedes,
insects, mollusc, scorpion and crab. Seven out of the
thirty two faecal samples contained vertebrate remains
(4.4%). These included, scales of skinks, hairs of a stripe
necked mongoose (Herpestes vitticollis) and sambar
deer (Cervus unicolor), and vertebrae of an unidentified
species of rodent. Plant remains such as bamboo shoots
were identified as Bambusa arundinacea.
Twenty-five opportunistic observations of feeding
tortoises were recorded. Details on the date and time of
feeding observations made during the study are given
in Table 2. On one occasion, an adult male and two
juveniles were observed feeding on Basidiomycetes
fungi on the forest floor (Table 2; Fig. 2). On three
different occasions, tortoises scavenged on carcasses
(Fig. 3); a female tortoise was found scavenging
on a sambar carcass and one juvenile was observed
scavenging on a dead rodent. On two occasions, a male
and a female were found feeding on Mimosa pudica and
Synedrella nodiflora which are both exotic shrubs, found
in the study area. On seven occasions I. travancorica
was found feeding on grass (Paspalam sp.).
205
Feeding ecology of the Travancore tortoise
Figure 1. Mean score of different diet item in the 32 faecal samples of I. travancorica: (A) for different sex (B) and in three
seasons in ATR and PTR from 2006-2009.
Table 1. Percentage composition of plant and animal components in the diet of I. travancorica of different sex classes and across
seasons. (* P < 0.05). Abbreviations are: GR (grass), SE (seeds), PM (Other plant matter), IN (insects), SC (Scorpions), CR (Crab),
VE (Vertebrate remains), MI (millipedes) and MO (mollusc).
Categories
Plant components
Sex
GR SE
Male % (n=8)
24.1 33.3 23.3
25.0 33.3 0.0
Female % (n=15)
48.3 50.0 50.0
54.2 33.3 66.7 57.1 0.0
Juvenile % (n=9)
27.6 16.7 26.7
20.8 33.3 33.3 14.3 100.0 50.0
0.3
2.9
2
Ȥ , df = 2
1.5
PM
1.9
Animal components
IN
SC
0.3
CR
1.1
VE
MI
28.6 0.0
0.9
3.1
MO
0.0
50.0
3.1
Season
Monsoon% (n=19)
58.6 66.7 63.3
62.5 66.7 66.7 57.1 100.0 100.0
Post-monsoon % (n = 7) 24.1 16.7 23.3
20.8 33.3 0.0
Summer% (n = 6)
17.2 16.7 13.3
16.7 0.0
33.3 0.0
0.0
0.0
Ȥ2, df = 2
1.1
0.4
1.6
0.7
5.1
0.45 9.2*
0.8
42.9 0.0
3.5
0.0
206
Deepak Veerappan &Karthikeyan Vasudevan
Discussion
Plant matter constituted 45% of the diet of I.
travancorica (Fig. 1A). Previous studies have already
reported that I. travancorica feeds on herbaceous plants
such as Synedrella nudiflora, Desmodium repandum,
Senecio scandens, and Veronica buabaumii (Ramesh
and Parthasarathy, 2006). The proportion of plant
matter within faecal remains of males, females and
juveniles did not significantly differ. However, the
proportion was significantly different across seasons
while the proportion of other plant matter remains
obtained during summer was considerably lower (Table
1). This might be explained by a lesser availability of
other plant matter during the summer compared to other
seasons, due to lesser intake, or both. However, these
hypotheses need to be confirmed through studies that
involve examination of samples collected across season
and sexes. Indotestudo travancorica has additionally
been reported to feed on fallen fruits of Artocarpus spp.,
Dillenia pentagyna and Ficus virens (Vijaya, 1983;
Ramesh and Parthasarathy, 2006). Fallen fruits appear to
be a major food item of tortoises in the Oriental region
including Indotestudo elongata and species belonging
to the genera Manouria and Geochelone (Ernst and
Barbour, 1989). Indotestudo elongata, a species closely
related to I. travancorica, has been reported to feed on
fallen fruits of Dillenia spp, Ficus racemosa, Cyanotis
cristata and Olax scandens (van Dijk, 1998; Platt et al.,
2001). Intact seeds of different plant species in the diet
of I. travancorica emphasize their potential role as seed
dispersers.
Animal matter provides an additional source of high
quality protein, calcium and high sodium potassium in
tortoise diets (Hailey, Chidavaenzi and Loveridge, 1998;
Figure 2. Juvenile # 12 found feeding on fungi in
Orukomban range of Parambikulam Tiger Reserve.
Figure 3. An adult female Travancore tortoise (♀5)
scavenging on a sambar (Cervus unicolor) carcass in Kothala
range of Parambikulam Tiger Reserve.
Manson et al., 1999; Hailey, Coulson and Mwabvu,
2001). Animal matter has been previously reported to
occur in the diet of some herbivorous and frugivorous
tortoises (Van Zyl, 1966; Cloudsey-Thompson, 1970;
Moskovits, 1985; MacDonald and Mushinsky, 1988;
Moskovits and Bjorndal, 1990; Milton, 1992; van Dijk,
1998; Platt et al., 2001). In captivity I. travancorica has
been known to feed on animals such as frogs, insects
and millipedes (Das, 1991, 1995). Similar prey items
have been reported for I. elongata (van Dijk, 1998;
Ihlow et al., 2012). Three independent observations
in the current study found I. travancorica scavenging
on carcasses, which confirm consumption of animal
matter by in-situ I. travancorica (Fig. 1). However, the
contribution of animal matter in general was observed to
be lower than that of plant matter while the proportion
was not significantly different between age-sex classes
or seasons (Figure 1A, Table 1).
Our observations on I. travancorica additionally
revealed fungi (Basidiomycetes) as dietary components
(Table 2; Fig. 2). The Basidiomycetes, which were part
of the diet, grow on the forest floor and are ephemeral
207
Feeding ecology of the Travancore tortoise
Table 2. List of the direct observations on feeding tortoises. Abbreviations are: J = Juvenile and H = hatchling from 2006 to 2009
in ATR & PTR.
S.no
Date
Time
Location
Sex & ID
Feeding
1
08.03.06
18.45
Padaravarai
ƃ1
Pasapalam sp
2
18.04.06
17.00
Varagaliar
ƃ4
Pasapalam sp
3
30.04.06
18.10
Sichali
Ƃ1
Pasapalam sp
4
21.05.06
17.45
Orukomban
ƃ5
Dillenia pentagyna fruits
5
02.06.06
18.00
Sichali
ƃ6
Pasapalam sp
6
02.06.06
18.00
Sichali
Ƃ2
Herbaceous climber (unidentified)
7
30.06.06
17.10
Sichali
ƃ7
grass (unidentified)
8
03.08.06
17.25
Kothala
Ƃ7
Herbaceous climber (unidentified)
9
18.08.06
17.00
Topslip
ƃ 10
Pasapalam sp
10
24.11.06
17.45
Karian shola
ƃ 12
Mimosa pudica
11
09.12.06
18.15
Varagaliar
J2
Pasapalam sp
12
13.12.06
17.52
Orukomban
Ƃ 11
Pasapalam sp
13
23.12.06
18.30
Kothala
Ƃ5
Cervus unicolor carcass
14
29.01.07
18.00
Mandhrimed
Ƃ 12
Synedrella nudiflora
15
30.01.07
17.45
Manamboli
J4
rodent carcass (unidentified)
16
13.06.07
18.10
Karian shola
ƃ8
grass (unidentified)
17
07.05.08
17.16
Karian shola
ƃ12
Strobilanthus sp
18
14.12.08
17.30
Orukomban
ƃ16
grass (unidentified)
19
13.06.09
18.00
Karian shola
J9
grass (unidentified)
20
07.07.09
17.00
Topslip
ƃ 17
grass (unidentified)
21
09.07.09
17.15
Erumapara
ƃ 18
grass (unidentified)
22
10.07.09
17.30
Anaikundhi
H1
grass (unidentified)
23
12.10.09
17.10
Orukomban
ƃ 22
Basidiomycetes (unidentified)
24
12.10.09
17.11
Orukomban
J 11
Basidiomycetes (unidentified)
25
12.10.09
17.50
Orukomban
J 12
Basidiomycetes (unidentified)
and patchily distributed in the study area. Indotestudo
elongata has also been reported to feed on fungi within
(Arakan) state in Myanmar (Platt et al.,
central Rakhine
2007). van Dijk (1998) additionally reported I. elongata
to feed on two species of mushrooms; Termitomyces sp.
and Russula sp.
Sand and small stones are reported from the diet of
many species of tortoises (Sokol, 1971, Moskovits and
Bjorndal, 1990; van Dijk, 1998). The presence of sand
in faecal samples may be due to accidental ingestion
or consumption for the purpose of digestion. However,
sand may also act as an important abrasive agent which
enhances the digestion of fibrous plant material (Sokol,
1971; Luckenbach, 1982; Marlow and Tollestrup, 1982).
1
Such plant material constitutes an important part of the
diet of I. travancorica.
Both I. travancorica and I. elongata conclusively
include a wide range of items in their diet. A similar
omnivorous diet has also been found for Indotestudo
forstenii, a species with similar natural history (World
Chelonian Trust, 2005). Omnivory and opportunistic
feeding therefore seem to be characteristic for the
members of the genus Indotestudo. Indotestudo
travancorica has been identified as a target species
as part of a nation-wide coordinated conservation
breeding programme (Vijayaraghavan, Choudhury and
208
Vasudevan, 2008). The opportunistic feeding behaviour
and the array of diet items consumed by I. travancorica,
as documented in this study, implies that its diet is
complex. This needs to be taken into consideration when
conservation breeding programme is implemented.
Acknowledgements. The Tamil Nadu and Kerala Forest
Departments are thanked for supporting the study. We
acknowledge the support extended from Wildlife Institute of India
Grant-in-Aid to carry out this study. We thank Bivash Pandav for
his inputs in the study. We thank Shruti Sengupta, Utpal Smart,
Silamban, Rajamani, and Karuppasamy for their assistance in
the field. Vinod Thakur from the Vertebrate Museum and Babu
from the herbarium section at Wildlife Institute of India, helped in
identification of mammal hair and plants and seeds.
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