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\begin{document}

\title{Feeding ecology of a lesser-known arboreal giant; Grizzled Giant
Squirrel (\emph{Ratufa macroura}), Southern Western Ghats, India.}



\author[1]{Kiran Thomas}%
\author[2]{Marek  Šmejkal}%
\author[3]{Paingamadathil Ommer Nameer}%
\affil[1]{University of South Bohemia in Ceske Budejovice Faculty of Science}%
\affil[2]{Biology Centre Czech Academy of Sciences}%
\affil[3]{Kerala Agricultural University}%


\vspace{-1em}



  
  \date{May 25, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
Animal dietary choices help understand a species' feeding niche and are
particularly relevant in conservation management. In this study, we
aimed to gather knowledge on food niche and the foraging ecology of the
Grizzled Giant Squirrel (\emph{Ratufa macroura}) in Chinnar Wildlife
Sanctuary, Southern Western Ghats, Kerala, India. Specifically, the
objectives were to identify food composition, the seasonal variation in
food choice and feeding technique. Through focal animal sampling, the
Grizzled Giant Squirrel in Chinnar Wildlife Sanctuary was found to feed
on 30 plant species belonging to 18 families. The most utilized plant
family was Fabaceae, with eight species, followed by Moraceae (four
species) and Anacardiaceae (two species). The feeding species included
22 trees, four climbers, one liana, one paraphyte, one shrub and one
succulent species. The maximum duration of feeding was observed on
\emph{Bauhinia racemosa} (19.79\%), followed by \emph{Tamarindus indica}
(14.08\%) and \emph{Nothopegia beddomei} (9.89\%). The squirrel's choice
of diet was primarily affected by the availability of food tree species
and food items rather than the season and non-tree species also have
been reported from the diet of Grizzled Giant Squirrel. Even though the
Grizzled Giant Squirrel shows plasticity in the food niche, the
availability of mature trees and plants appears important for its
conservation in the fragmented riparian forest in the Western Ghats,
southern India.%
\end{abstract}%



\sloppy


Kiran Thomas \textsuperscript{1,2*}, Marek
\selectlanguage{polish}Š\selectlanguage{english}mejkal\textsuperscript{2,}and Nameer Paingamadathil
Ommer\textsuperscript{3}

\textsuperscript{1} Faculty of Science, University of South Bohemia,
\selectlanguage{polish}Č\selectlanguage{english}esk\selectlanguage{ngerman}é Bud\selectlanguage{polish}ě\selectlanguage{english}jovice, Czech Republic

\textsuperscript{2} Institute of Hydrobiology, Biology Centre of the
Czech Academy of Sciences, \selectlanguage{polish}Č\selectlanguage{english}esk\selectlanguage{ngerman}é Bud\selectlanguage{polish}ě\selectlanguage{english}jovice, Czech Republic,
kiranzthomas@gmail.com, marek.smejkal@hbu.cas.cz

\textsuperscript{3} Faculty of Forestry, Kerala Agricultural University,
Thrissur. India, nameer.po@kau.in

*Corresponding author

Received:~\emph{~~~~~~~~~~~~} ; Revised:\emph{~~~~~~~~} (optional);
Accepted:\emph{~~~~~~~~~~~~} .

\textbf{Abstract}

Animal dietary choices help understand a species' feeding niche and are
particularly relevant in conservation management. In this study, we
aimed to gather knowledge on food niche and the foraging ecology of the
Grizzled Giant Squirrel (\emph{Ratufa macroura} ) in Chinnar Wildlife
Sanctuary, Southern Western Ghats, Kerala, India. Specifically, the
objectives were to identify food composition, the seasonal variation in
food choice and feeding technique. Through focal animal sampling, the
Grizzled Giant Squirrel in Chinnar Wildlife Sanctuary was found to feed
on 30 plant species belonging to 18 families. The most utilized plant
family was Fabaceae, with eight species, followed by Moraceae (four
species) and Anacardiaceae (two species). The feeding species included
22 trees, four climbers, one liana, one paraphyte, one shrub and one
succulent species. The maximum duration of feeding was observed
on\emph{Bauhinia racemosa} (19.79\%), followed by \emph{Tamarindus
indica}(14.08\%) and \emph{Nothopegia beddomei} (9.89\%). The squirrel's
choice of diet was primarily affected by the availability of food tree
species and food items rather than the season and non-tree species also
have been reported from the diet of Grizzled Giant Squirrel. Even though
the Grizzled Giant Squirrel shows plasticity in the food niche, the
availability of mature trees and plants appears important for its
conservation in the fragmented riparian forest in the Western Ghats,
southern India.

\textbf{Keywords:} Sciuridae-Rodentia, foraging behaviour, habitat use,
riparian forest, conservation





\section*{1. INTRODUCTION}

{\label{introduction}}

The ecological role of an animal in an ecosystem can be revealed by
understanding its diet and feeding behaviour (Bookhout, 1994;
Dell'Agnello et al., 2019). The feeding behaviour might be specific to
the place where an animal lives. It influences all the decisions of the
animal about food resources available along with predator avoidance
strategies (Pyke, 1984; Sih, 1980). Feeding has a considerable impact on
animal fitness through its effect on animal growth, survival, activity
patterns and reproductive ability (Foley \& Cork, 1992; D. Raubenheimer
\& Simpson, 1997; David Raubenheimer et al., 2015), thus it is an
important aspect of animal conservation. The availability of food in an
area is vital in supporting a minimum viable population for the animal
to thrive (Palmer \& Koprowski, 2014). The variation in food
availability by the seasonal fluctuation can challenge the animal to
meet its requirements. Seasonal shift or facultative adjustment in food
material preference gives an animal large dietary breadth and
flexibility to cope with situations of resource scarcity (Dunn et al.,
2010; Masi et al., 2015).

Herbivorous mammals may impact the plant resources near its foraging
sites either negatively or positively and can affect the composition and
structure of plant communities (Olff \& Ritchie, 1998; Palmer \&
Koprowski, 2014). Small mammalian herbivores, such as rodents, modify
their environment by consuming plant biomass and redistributing
materials across the landscape (Moorhead et al., 2017). In particular,
small mammals can stimulate soil nutrient cycling through the faecal
deposition (Bakker et al., 2004). Tree squirrels (order Rodentia) are
cosmopolitan in distribution on almost every continent except Australia
and Antarctica (Koprowski \& Rajamani, 2008; Thorington Jr et al., 2012)
and play key ecosystem services, including seed dispersal and
pollination (Miyaki, 1987; Steele et al., 2005; Zong et al., 2010). The
giant squirrels are tree squirrels and the largest in the world, and
they belong to the genus \emph{Ratufa} (Thorington \& Cifelli, 1989).

The Grizzled Giant Squirrel (hereafter GGS) belongs to the
genus\emph{Ratufa} , has three distinct subspecies and India harbour
only one,\emph{Ratufa macroura dandolena} and it is one among the other
three species of the giant squirrels in India (Ellerman, 1961; Johnsingh
\& Nameer, 2015; Menon, 2014). This Near Threatened squirrel is
primarily diurnal, but their increased activity has been observed during
the early and late hours of the day (Paulraj, 1991). The GGS shows one
of the most important examples of isolated populations, and the habitat
of the animal is extremely unique and is confined primarily to a narrow
stretch of riparian vegetation along the Pambar and Chinnar rivers and
their major tributaries in the Chinnar Wildlife Sanctuary (Ramachandran,
1993), in the southern Western Ghats. In India, Chinnar Wildlife
Sanctuary in Kerala supports the second-big population of GGS. To
protect and improve the conservation measures directed to an animal,
especially a habitat specialist like the GGS, a proper understanding of
the habitat requirement is vital (Mills, 1992). We focused on the
foraging ecology of the GGS, to understand the seasonal variation in
food preference, food composition and feeding technique. The diet and
feeding behaviour of the GGS is not well known so far thus, it would
help in making specific management prescriptions for protecting the
Near-Threatened GGS in one of its prime refugia in India.



\section*{2. METHODS}

{\label{methods}}

\subsection*{2.1 The study area}

{\label{the-study-area}}

The study area, Chinnar Wildlife Sanctuary, is located between 10\selectlanguage{ngerman}°15'-
10°21'N latitude and 77°16'- 77°5' E longitudes in the Idukki district
of Kerala state (Figure 1) and has a total extent of 90.44
km\textsuperscript{2}. The significant variation in altitude and
rainfall leads to a wide array of habitat types like deciduous forests,
dry thorny scrub, riparian forests, shola forests and grasslands that
are interspersed with plains, hillocks, rocks and cliffs, which provide
microhabitats for varied forms of life (Management Plan of Chinnar
Wildlife Sanctuary., 2012).

\subsection*{2.2 Observation methods}

{\label{observation-methods}}

The information on the composition and seasonal variation in the GGS
diet was collected through foraging observations. Focal Animal Sampling
was used following (Altmann, 1974). Each encountered individual was
followed, and observation on the time spent on feeding, plant species
eaten, plant part eaten, pick up rates of plant parts such as fruit,
leaves and flowers were also recorded. The activities occur in bouts
that are periods of, for example, feeding activity within a food source
or movement between two sources. During the period spent in the food
source, some of the time was allotted to searching for a food item,
selecting by smelling, remaining inactive or other grooming activities.
The length of the bout was judged as the period between entry into and
exit from the source. The incidence of feeding on different food items
across different seasons was categorised. This was done for the three
different seasons summer (December to May), southwest monsoon (June to
September) and northeast monsoon (October to November).

Observations were made for 10 months between April 2013 to May 2014, and
within a month, at least one week was spent in the field. Five transects
were selected after reconnaissance, and observations were made for each
transect both in the forenoon and afternoon for an equal duration.
Squirrels were observed using 8x40 Olympus standard binoculars (Olympus
Global, Tokyo, Japan). Different feeding postures and feeding techniques
used by the GGS were interpreted by direct observation during the
sampling.

\subsection*{2.3 Data analysis}

{\label{data-analysis}}

To understand the food composition of GGS, collected data were analysed
by three complementary approaches. The percentage contribution of
different food items to the squirrel diet was analysed based on the
duration of feeding on a particular item and the number of times of
feeding incidence on a particular item. The non-parametric test, ``Mann
Whitney U'' was performed to determine whether there was any significant
difference in the feeding bout and duration between the forenoon and
afternoon hours. \selectlanguage{english}?\textsuperscript{\emph{2}} test for association was
used to find out the seasonal variation on the preferred tree species
and the plant parts fed by the GGS across different seasons\emph{.} A
constrained analysis, species as response variable against the
environmental variables (Residualized predictors were permuted),
canonical correspondence analysis (CCA) was performed to check the food
choice and season using Canoco 5 (Braak \& Smilauer, 2012; Jiangshan et
al., 2014).







\section*{3. RESULTS}

{\label{results}}

\subsection*{Food composition}

{\label{food-composition}}

The GGS found to be feeding on different plant parts including the
leaves, seeds, flowers and sap of different trees and other vegetation
(Table 1).

A total of 1314 minutes (in 10 months) in 42 encounters of feeding
observation were taken during the study. A total of 30 different species
of plants \emph{per se} , which included 22 tree species, four climbers,
one liana, one paraphyte, one shrub and one cactus species, formed the
diet. The maximum duration of feeding was observed on \emph{Bauhinia
racemosa} (19.79\%), followed by \emph{Tamarindus indica}
(14.08\%),\emph{Nothopegia beddomei} (9.89\%), \emph{Strychnos
potatorum} (7.23\%) and \emph{Terminalia arjuna~} (6.47\%; Table 1).

The feeding varied by the food availability in various months. The GGS
fed on \emph{Terminalia arjuna} (January, April and September)
and\emph{Tamarindus indica} (May, November and December) in three
different months, followed by \emph{Bauhinia racemose} (February and
March)\emph{, Ficus macrocarpa} (February and November) and
\emph{Syzygium cumini}(August, May) in two different months. However,
the other species were found fed only once during the study period. The
climbers that formed part of the diet were \emph{Derris brevipes,
Diplocyclos palmatus} and\emph{Cayratia trifolia} . The GGS also used
shrubs like \emph{Hibiscus rosa-chinensis} , cacti - \emph{Euphorbia
trigona,} and liana -\emph{Entada rheedii} for feeding. These species
come under 18 different families. Among them, the most preferred family
was Fabaceae, with eight species found to be part of the GGS diet. This
was followed by Moraceae, four species, and Anacardiaceae, two species.

\subsection*{3.2 Contribution of different vegetation
parts}

{\label{contribution-of-different-vegetation-parts}}

The total feeding incidence across the whole study period was found to
be 62. Out of it, 30 incidences were on leaf feeding (48.39\%), followed
by seed feeding (27.32\%), flower feeding (16.13 \%), and sap-feeding
(8.06 \%) (Figure 2. A). The duration of feeding on different items is
on seeds (52.12 \%), followed by leaves (32.55 \%), flowers (10.58 \%)
and sap (4.76 \%) (Figure 2. B).

\subsection*{3.3 Diurnal variation in feeding
habit}

{\label{diurnal-variation-in-feeding-habit}}

The feeding recorded for the whole study period was 1314 minutes,
including 915 feeding bouts. 42\% of the total feeding duration was
contributed by forenoon observation and the rest 58\% by afternoon
observation, sampling hours are equal for both times of the day. when
the feeding bout took into consideration forenoon hours, 321 bouts,
while the afternoon had around twice the value, 594 bouts. However, the
GGS was found to be inactive in a food source for some time before
commencing the feeding or after feeding, before going elsewhere in
search of a new source or for other activities. In some cases, the GGS
was found to be inactive or resting at a food source between periods of
intensive feeding. The GGS were not significantly different in the
duration of feeding between the forenoon and afternoon (U = 189.5 p
=0.319) as well as in the feeding bout (U = 205.50 p =0.542).

\subsection*{3.4 Seasonal variation in
diet}

{\label{seasonal-variation-in-diet}}

The feeding incidence on all food items was maximum in the summer,
followed by the North-East and South-West monsoon. Univariate analysis
was done in the absence of a correlation between different variables
under consideration; there was no effect of season and time of the day
on the duration of feeding and the feeding bouts. To check if the animal
prefers food items with respect to different seasons,\textbf{?}
\selectlanguage{english}\textsuperscript{2} test for association was performed. However, no
significant association (\textbf{?} \selectlanguage{english}\textsuperscript{2}= 6.32, p= 0.70),
could be observed.

The canonical correspondence analysis (CCA) (Figure 3) to check the
association of species and season is also done for selected species
based on maximum incidences and duration of feeding and found no
significant association. The food species choice of the GGS was found
not affected by the seasons, but it is based on the availability of
various food resources across different seasons.

\subsection*{Feeding technique}

{\label{feeding-technique}}

The GGS were found to be highly selective in picking the food items. The
selection of food items appears to be done by the smell of the items
(Figure 4A). The GGS was found to be handling its food with both mouth
and forelimbs (Figure 4B). The GGS goes to the tip of the branch or
other areas of the crown of the tree according to the availability of
food sources and cuts the pod, fruit, leaves or flower with its mouth
(Figure 4C). Occasionally the food item was brought to the mouth with
the help of the forelimbs. Holding the fruit in the mouth, the squirrel
then moves to the thick horizontal branch to sit firmly, sometimes the
feeding started at the place of harvest itself. The squirrel holds the
branch with the help of the claws of the hind limbs and the tail hanging
down, which gives further balance to the body (Figure 4D). It then gnaws
the epicarp of the fruit to extract the seed to be consumed. During
feeding, the forelimbs were used effectively to manipulate food items,
whether long pods of \emph{Bauhinia racemosa} or small fruit of

\emph{Grewia tiliifolia.}

\subsection*{3.6 Feeding posture}

{\label{feeding-posture}}

Mainly three postures were observed during the present study. The most
common feeding posture is the squirrel perching itself on a horizontal
branch of the tree and feeding, second posture is that of the animal
hanging upside down from small branches, supporting with the hind limbs
and the tail, and reaching down to the food in the hanging posture. The
third feeding posture had the squirrel on the bole of the tree and
feeding upside down.

During most of the instances of observations on feeding posture, the
squirrels were found feeding by sitting on the branch, holding the
branch with hind limbs and the tail hanging down. The hanging tail helps
the animal in balancing the body. This kind of posture was observed
during the feeding incidence of \emph{Bauhinia racemosa} and\emph{Grewia
tiliifolia} (Figure 5A). The second body posture consisted of GGS
feeding while hanging upside down on the tree branch with the help of
claws of hind limbs, while the tail was kept curled over the branch.
This posture was observed mainly in instances of feeding clustered
fruits and flowers on small branches which cannot support the body
weight of the squirrel. Sometimes this posture was used only for
harvest. This was observed while GGS was feeding on \emph{Strychnos
potatorum} and \emph{Nothopegia beddomei} (Figure 5B).

The third feeding posture observed was a GGS lying on the vertical bole
of the tree, anchoring onto the tree trunk with the claws of the hind
limbs and manipulating the food with mouth and forelimbs (Figure 5C).
The observed instances are while feeding on the climbers found on the
trees or the barks.



\section*{4. DISCUSSION}

{\label{discussion}}

The present study observed that GGS fed on items from 30 different
species of vegetation including 22 tree species and eight species
including climbers, lianas, paraphytes, shrubs and cacti in Chinnar
Wildlife Sanctuary, Western Ghats, south India. It is higher compared to
a previous report of 21 tree species (Kumar et al., 2007). They were
found primarily feeding on the tender leaves, flowers, bark and seed of
these species, including lianas, climbers, shrubs and even one cactus.
While in the earlier studies, there has not been any mention of the
plant or plant parts being fed by the GGS. The squirrels were observed
to feed on tender leaves and flowers of the plant species such
as\emph{Derris brevipes} and \emph{Diplocyclos palmatus}
(climbers),\emph{Entada rheedii} (liana), \emph{Hibiscus rosa-chinensis}
(shrub) and \emph{Euphorbia trigona} (cactus). Joshua (1992)\emph{,}
reported the GGS feeding on the tender leaves and flowers of two
trees,\emph{Tamarindus indica} and \emph{Bauhinia purpurea.}

\subsection*{4.1 Food composition}

{\label{food-composition-1}}

The optimal foraging theory proposed by (Pyke, 1984) suggests that a
forager should eat only the most preferred or highest-ranked item if
there is an adequate amount of that item to fulfil its daily diet
requirements. As the highly preferred food item is depleted, the forager
should include the next ranked item in its diet. The percentage
contribution of each item based on the duration of feeding showed a
trend as maximum by the seeds, followed by leaves, flowers and sap
(Figure 2). This indicates that the GGS choose to have more seeds in
their diet, switching to leafy food items, flowers and sap when seeds
are scarce or unavailable. However, the present observation corroborates
the findings of (Thorington \& Cifelli, 1989) on \emph{R. indica} at
Mudumalai Wildlife Sanctuary and Ramachandran (1992) at Periyar,
Parambikulam and Silent Valley. In the present study, we observed that
the GGS at Chinnar acts as an important seed-dispersing agent for many
of the riverine tree/plant species.

The rodents depend heavily on hoarding for survival and reproduction
(Vander Wall, 1990; Wang et al., 2018). Unlike a report on hoarding
by\emph{R. indica} (Somanathan et al., 2007), there was no instance of
hoarding by GGS during the study period. This may be either a clear
indication of the optimal availability of food in the habitat (Ando et
al., 1985) or that the GGS does not possess this habit. More studies are
required on the energetics of the GGS as this animal is a generalist
plant feeder feeding on seeds, sap, leaves and flowers.

\subsection*{4.2 Diurnal variation in feeding
habit}

{\label{diurnal-variation-in-feeding-habit-1}}

The feeding observations across different hours of the day show that the
animal was highly active in the morning and evening hours of the day,
and the resting occurred in the mid-day time. Thus, the animal's feeding
activity appeared affected by the weather factors in the area. In the
hours of high sunlight and rainfall, animal activity was found to be
very low. Temperature dependant activity patterns to avoid extreme
weather conditions are already observed in diurnal squirrels (Lee et
al., 1990; Skibiel et al., 2002).

\subsection*{4.3 Feeding posture and
technique}

{\label{feeding-posture-and-technique}}

The mouth and forelimbs are the organs that help in manipulating the
food item, while hind limbs and tail play roles as supporting organs for
body balancing during feeding. All the three feeding postures, such as
sitting, hanging and clinging postures, were found to be effectively
used by the animal for feeding. This was found according to the
availability and nature of food items foraged. Similar feeding postures
were reported by (Ramachandran, 1992) on \emph{Ratufa indica} .

The Chinnar wildlife sanctuary, southern Western Ghats, India, has a
very small patch of 1.6 km\textsuperscript{2} of riparian habitat which
holds a very small population of the GGS and is already under serious
threats of various forms (K. Thomas \& Nameer, 2018). Further
fragmentation and deterioration of the habitat will lead to food
scarcity and increased predation risk (Gurjar et al., 2013; Justus
Joshua \& Johnsingh, 1994; K. Thomas et al., 2017). Food scarcity
persuades the animal to increase the duration of feeding, which is
positively correlated with stress (Dunn et al., 2010; Laurance et al.,
2011). Habitat fragmentation accelerates the mortality of large and
mature trees (Laurance et al., 2000), which holds a greater quantity of
food resources (Chapman et al., 1992) and potentially can limit the
availability of food of choice for GGS. The GGS was also found to be
associated with many of the riparian species for their drey construction
(Kiran Thomas \& Nameer, 2021). A detailed investigation of the
energetics of GGS along with habitat restoration, including species
associated with GGS ecology, can ensure long-term survival possibilities
for this mammal in this part of the world.

\section*{5. Tables}

{\label{tables}}

Table 1: The details of each feeding incident of the grizzled giant
squirrel on a particular species.\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
\begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Species\strut
\end{minipage} & \begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Family\strut
\end{minipage} & \begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Month\strut
\end{minipage} & \begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Part eaten\strut
\end{minipage} & \begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Total duration (minutes)\strut
\end{minipage} & \begin{minipage}[b]{0.16\columnwidth}\raggedright\strut
Duration of feeding (\%)\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Mangifera indica}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Anacardiaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
October\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
15\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.14\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Nothopegia beddomei}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Anacardiaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature seeds\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
130\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
9.89\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Commiphora caudata}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Burseraceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Bark and tender leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
10\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.76\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Terminalia arjuna}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Combretaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
April, September, January\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves and fruits\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
85\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
6.47\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Diplocyclos palmatus}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Cucurbitaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
November\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature leaves and seeds\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
45\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.42\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Hopea parviflora}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dipterocarpaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
February\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
8\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.61\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Euphorbia trigona}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Euphorbaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
August\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
27\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.05\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Acacia spp.}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Seeds of immature fruit\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
35\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.66\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Albizzia lebbeck}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
36\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.74\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Bauhinia racemosa}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
February, March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature seeds\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
260\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
19.79\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Derris brevipes}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
17\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.29\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Entada rheedii}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
15\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.14\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Pongamia pinnata}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
April\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
23\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.75\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Tamarindus indica}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
November, December, May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
seeds of immature fruit and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
185\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
14.08\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Cassia fistula}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fabaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Bark\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.23\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Strychnos potatorum}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Loganiaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
January\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature seed\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
95\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
7.23\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Macrosolon capitellatus}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Loranthaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature seeds\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
55\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
4.19\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Hibiscus rosa --chinensis}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Malvaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
15\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.14\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Melia dubia}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Meliaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
September\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
37\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.82\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Ficus albiphyla}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Moraceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
February\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and bark\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
40\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.04\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Ficus microcarpa}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Moraceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
November, February\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves and immature seed\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
29\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.21\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Ficus spp.1}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Moraceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
November\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves and immature seed\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
16\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.22\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Ficus spp.2}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Moraceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
September\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
16\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.22\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Syzygium cumini}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Myrtaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
August, May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
seeds of immature and mature fruits and leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
52\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.96\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Psychotria subintegra}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Rubiaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
September\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves, flowers and sap\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
13\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.99\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Aegle marmelos}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Rutaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
October\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.23\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Santalum album}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Santalaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
November\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Immature leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
9\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.68\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Grewia tiliifolia}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tiliaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
October\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves, flowers and fruits\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
25\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1.90\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Grewia spp.}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tiliaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
May\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leaves\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.38\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\emph{Cayratia trifolia}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vitaceae\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tender leaves and flowers\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
10\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.76\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\section*{6. Figure legends}

{\label{figure-legends}}

\textbf{Figure 1.} The study site and the other locations from where the
Grizzled Giant Squirrels are reported from the Western Ghats,
India.,\textbf{Figure 2.} The contribution of each food item to total
feeding based on (A) the duration of feeding on a particular item, (B)
the number of times feeding on a particular item occurred,
\textbf{Figure 3.} The canonical correspondence analysis shows the
association of season and time of the day to the species fed (Eight
species are given in the figure, selected based on the number of months
and more feeding duration observed). \textbf{Figure 4.} Feeding
Technique, A) Selection by smell B) Making the food reach to mouth with
the help of forelimbs C) Cutting the fruit with the mouth D) Feeding the
seeds by sitting on a firm branch, \textbf{Figure 5.} Feeding Postures,
A) Sitting posture B) Hanging posture C) Clinging posture.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

ACKNOWLEDGEMENTS

The authors would like to thank the Chief Wildlife Warden, Kerala State
Forest Department, for the study permit (No. WL10-947/2013). We also
thank the Kerala Agricultural University for the financial support for
the conduct of the research.

\textbf{Author Contribution Statement}

\textbf{Kiran Thomas} : Investigation, Data curation, Writing- Original
draft, Visualization. \selectlanguage{polish}\textbf{Marek Šmejkal}\selectlanguage{english} : Supervision, Writing-
Reviewing and Editing. \textbf{Nameer Paingamadathil Ommer:}Methodology,
Supervision, Writing- Reviewing and Editing, Funding acquisition.

\textbf{Conflict of interest statement}

On behalf of all authors, the corresponding author states that there is
no conflict of interest.

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{\label{references}}

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Ellerman, J. R. (1961). Review of ``the Fauna of India Including
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Foley, W. J., \& Cork, S. J. (1992). Use of fibrous diets by small
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