Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 May 2023 | 15(5): 23139–23146
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.8480.15.5.23139-23146
#8480 | Received 13 April 2023 | Final
received 06 May 2023 | Finally accepted 20 May 2023
Nesting habitat and nest directionality of the Indian Giant Squirrel Ratufa indica maxima (Schreber, 1784) (Mammalia: Rodentia: Sciuridae)
in the Nelliyampathy Reserve Forest, Western Ghats,
Kerala, India
K. Mohan 1,
Joseph J. Erinjery 2 & Mewa Singh 3
1,3 Biopsychology
Laboratory, Institution of Excellence, University of Mysore, Mysuru, Karnataka
570006, India.
2 Department of
Zoology, Kannur University, Mananthavady campus,
Kannur, Kerala 670645, India.
1 kmohan1992@yahoo.in, 2
joerin@gmail.com, 3 mewasinghltm@gmail.com (corresponding
author)
Editor: Giovanni Amori,
CNR - Institute of Research on Terrestrial Ecosystems, Rome, Italy. Date of publication: 26 May 2023
(online & print)
Citation: Mohan, K., J J. Erinjery & M. Singh (2023). Nesting habitat and nest
directionality of the Indian Giant Squirrel Ratufa
indica maxima (Schreber,
1784) (Mammalia: Rodentia: Sciuridae) in the Nelliyampathy Reserve Forest, Western Ghats, Kerala, India. Journal of Threatened Taxa 15(5): 23139–23146. https://doi.org/10.11609/jott.8480.15.5.23139-23146
Copyright: © Mohan et al. 2023. 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: CSIR-HRDG - SRF grant
(09/119(0206)/2018-EMR-I); SERB-Distinguished Fellowship (SB/S9/YSCP/SERB-DF/2018(1)); SERB Grant (SRG/2021/001098).
Competing interests: The authors declare no competing interests.
Author details: K. Mohan is pursuing PhD at the University of Mysore, Mysuru, Karnataka, India. He is working on the behavioral ecology of the Indian Giant Squirrels in the Western Ghats. His research interest is in ecology, animal behavior, and conservation. He is a recipient of The CSIR-HRDG SRF Grant. Dr. Joseph J. Erinjery is an assistant professor at the Department of Zoology, Kannur University, Kerala, India. He is a spatial ecologist and primatologist interested in species distribution, remote sensing, ecological modeling, animal behavior, and disease dynamics. He has several years of field experience, working on the ecology, behavior, and adaptability of primates and other mammals in the Western Ghats. He has developed species distribution models for several species in the Western Ghats. Prof. Mewa Singh is a
distinguished professor (for life) at the University of Mysore, Mysuru, Karnataka, India. He is a recipient of the SERB-Distinguished Fellowship. He is a fellow of the
Indian National Science Academy, The National Academy of Sciences, and The Indian Academy of Sciences. He is the cheif editor of the Journal Dialogue: Science, Scientists & Society. His research interest is in ecology, evolution, animal behavior, and conservation.
Author contributions: KM - conceived the idea and collected the data; KM, JJE, MS - analyzed the data; KM, JJE, MS - wrote the article.
Acknowledgements: We thank: the Kerala Forest and Wildlife
Department, Thiruvananthapuram for permission (WL10-11666/2016 dated: 18 April
2016) to conduct field work in the forests of Kerala; the Kerala Forest
Development Corporation (KFDC) for all the logistics; and Dr.
Santhosh S. for his critical thoughts and support in developing the concept. KM
thanks CSIR-HRDG for the SRF Grant (09/119(0206)/2018-EMR-I); MS thanks SERB-
Distinguished Fellowship (SB/S9/YSCP/SERB-DF/2018(1)); and JJE thanks SERB
Grant (SRG/2021/001098) for the financial assistance.
Abstract: The information on
selection of nesting habitat and nest directionality for arboreal species is
crucial in developing conservation and management plan for the species. We
studied the factors which affect the nesting habitat selection and the nest
orientation by using the quadrat sampling method in Nelliyampathy
Reserve Forest, Kerala. A total of 119 nest sites were observed on 26 different
tree species in four different habitat types. Around 56.30% and 36.13% of the
nests were sighted in contiguous forests and plantation with native tree shade,
respectively. Of the 119 nests, 112 were in trees of height up to 30 m. Cullenia exarillata, Mesua ferrea, Actinodaphne malabarica,
and Schleichera oleosa
accounted for 45.4% of the nest with 15.9%, 11.8%, 9.2% and 8.4% nests,
respectively. About 24.4% of the nests were directed towards the north-east
direction (n = 29) whereas least preferred direction was the south (n = 05).
This shows that the nests are oriented towards sun rise and to avoid wind and
rainfall of monsoon which is foreseen from the south-west direction.
Keywords: Arboreal, behavior,
conservation, ecology, forest fragmentation, native tree plantation, nest
orientation, nest tree selection, rain avoidance, sunlight preference.
INTRODUCTION
An understanding of
the costs and benefits of choosing a certain nest site and placing a nest
entrance in a specific direction in arboreal mammals is still in its inception.
Nest sites chosen based on specific criteria, most often improve concealment,
which may increase nesting success by lowering the risk of predation ( Pradhan
et al. 2017). Young individuals are highly sensitive to environmental factors,
and by reducing environmental extremes in the nest, adults are likely to
improve the survival rate of their infants, and their own fitness (Murphy 1983;
Webb & King 1983; Bekoff et al. 1987; Webb 1987;
Martin 1988; Martin & Roper 1988). It is predicted that most nests are
orientated in response to environmental conditions such as wind, precipitation,
and in particular sun radiation (Haggerty 1995; Burton 2007).
Avian nesting sites
have shown that there is a wide range of nest orientation and pattern among
species and/or nesting guilds (e.g., open cup, domed nest, primary/secondary
cavity nest). Although some studies have found little to no pattern in nest
orientation (Albano 1992; Rendell & Robertson 1994; Tarvin
& Smith 1995; Mennill & Ratcliffe 2004), many
groups representing a variety of nest architectures do show considerable
preferences in the orientation of their nests (Austin 1974; Walsberg
1981; Martin & Roper 1988; Bergin 1991; Hooge et
al. 1999; Mezquida 2004). Most often, researchers
have discovered that nest orientation is related to either prevailing winds (Norment 1993; Mezquida 2004) or
sun exposure (Viñuela & Sunyer
1992; With & Webb 1993; Yanes et al. 1996; Rauter et al. 2002; Hartman & Oring
2003; Burton 2006), both of which may have an immediate impact on the
microclimate of the nest (Hartman & Oring 2003; Ardia et al. 2006). In hot habitats near the equator,
animals would be predicted to orient their nests to optimize shade during the
day, when the sun and thus temperatures are at their maximum (Maclean 1984).
Mid-latitude nests commonly face eastward since it may be less important to
avoid the mid-day or afternoon sun there. Nests that face east rather than west
may warm up more quickly in the morning (Nelson & Martin 1999), lessening
the potential effects of low overnight temperatures on embryos and young. Nests
may be pointed towards the equator at the highest latitudes to benefit from the
greater insolation and warmth coming from that direction and lessen the
consequences of a cold climate (Ojedat et al. 2021).
But due to variations in nest-site parameters at any given latitude, there may be
a significant difference in the preferred nest orientations within species.
Hence, nest site and orientation are crucial elements of bird reproduction that
may have an impact on embryonic development, hatching success, nestling
growth (Austin 1974; Viñuela & Sunyer 1992; Lloyd & Martin 2004; Burton 2006) and
overall nesting success (Martin & Roper 1988; Filliater
et al. 1994; Rauter et al. 2002). With the context of
avian nesting sites, we would like to investigate the nesting orientation of
the Indian Giant Squirrel as well.
The Indian Giant
Squirrel (IGS; Ratufa indica
maxima) is a diurnal and arboreal species which is found only in peninsular
India (Agrawal & Chakraborty 1979; Corbet & Hill 1992). Despite being
widely distributed within its range, it is found in severely fragmented
populations (Molur et al. 2005). The ecology of
squirrels in Asian countries has received less focus, and available research is
scarce (Pradhan et al. 2012; Borges 2015). It is a solitary species that only
appears in pairs during the breeding season. During a single breeding season,
it usually builds more than one nest, or drey. Recent
studies on nesting tree selection in the IGS have shown that the most common
and abundantly available tree species in the forest were preferred for nesting
over random tree species (Rathod et al. 2022). The nests made of leaves and
twigs are large, globular in shape with a lateral opening which are built on
tall, profusely branched trees in the higher canopy (Ramachandran 1988; Borges
1989; Datta & Goyal 1996; Kumara
& Singh 2006; Pradhan et al. 2017). The nesting trees were comparably
taller species with interlinking crowns which allowed easy access and movement
in the canopy, probably to avoid predators (Ramachandran 1988; Datta & Goyal 1996). At the landscape level, the nest
trees were found predominantly in the contiguous forests of evergreen,
moist-deciduous and deciduous forests with abundant availability of food
resources, and away from the agricultural fields.
Factors that
influence nest-site selection, nest design, nest orientation, and the inter-
and intra-specific variation of these behaviours in
IGS are scarce. As a result, we investigated nest-site selection and nest
orientation patterns in the IGS. Our goals were to find out (1) the nesting
preferences of IGS, and (2) whether there is any directionality to its nest
entrance.
MATERIALS AND METHODS
Study area
We carried out this
study in Nelliyampathy Range of Nelliyampathy
Reserve Forest (10.41–10.30 N & 76.58–76.75 E), Nemmara
Forest Division in the Western Ghats in Palakkad District of Kerala (Figure 1).
It covers an area of about 157 km2 (Erinjery
et al. 2018) with a vegetation of evergreen, semi-evergreen and moist deciduous
forests with interspersed tea, coffee and cardamom plantations (Ramachandran
& Suganthasakthivel 2010). The altitude ranges
500–1,633 m. The average rainfall is about 3,378 mm over a period of 10 years.
The forest mainly consists of Cullenia, Mesua and Palaquims
species (Pascal 1988; Ramachandran & Suganthasaktivel
2010; Erinjery et al. 2015). Some of the arboreal
species which belong to family Sciuridae found here
are the Western Ghats Striped Squirrel Funambulus
tristriatus Waterhouse, 1837, the Dusky Striped
Squirrel Funambulus sublineatus
Waterhouse, 1838., the Indian Giant Flying Squirrel Petaurista
philippensis Elliot, 1839, Travancore Flying
Squirrel Petinomys fuscocapillus,
and the Indian Giant Squirrel (Ramachandran
& Suganthasakthivel 2010; Kumara
& Suganthasakthivel 2011; Babu
et al. 2015).
Data collection
The habitat of the
IGS nests was broadly divided into four different types namely contiguous
forest (45 km2, >60% canopy cover, dominated by evergreen and dry
deciduous forest trees), fragmented forest (8 km2, evergreen/dry
deciduous forest patches are divided between open plantation), plantation with
native tree shade (25 km2, 30–60% canopy cover, mainly included
coffee, cardamom plantations with native trees) and plantation with monoculture
tree shade (17 km2, >30% canopy cover, dominated by monoculture
Teak and Silver Oak). The categorization of the habitat was derived from the
high-resolution vegetation type and land-use map with accuracy >85%
developed from Sentinel2 MSI 10 m spectral bands and Sentinel1 SAR bands, NDVI
and Textural layers (Erinjery et al. 2018). We
obtained the data on IGS nesting by Quadrat sampling method (Heltshe & Forrester 1983). The study area was divided
into quadrats of 0.5 x 0.5 km2. Based on the average home range of
the IGS, the observer walked randomly in each quadrat looking for the nests of
the IGS, and made sure that 75% of the pre-defined habitat types in the quadrat
was sampled without overlapping. We did not conduct surveys in habitats such as
open plantations, rocky mountains and grasslands as these habitat structures
did not consist of any tree species. Thus, a total of 95 km2 was
only sampled and considered for the analysis. Only active IGS nest was
considered for the analysis. It is difficult to differentiate between active
and non-active nests of the IGS unless an individual is sighted using the nest.
Active nests are the ones which are freshly built nests of lush green in colour and are highly dense and compact in structure, which
makes it difficult to sight as they camouflage with the tree canopy. Over a period
as twigs of the nest dry, it becomes easier to identify due to variation in the
nest and canopy colour. Non-active nest consists only
of dry leaves and twigs and the walls of the nest are very loosely arranged and
mostly worn out. Nest location was recorded by using handheld GPS (Montana
650). Data on nesting tree species (the trees in which nests are constructed),
height of the tree and height of the nest from the ground was recorded by using
the laser range finder (HAWKE LRF 900). We collected data on nesting direction
for which nest orientation readings was recorded by holding a compass directly
below the nest and orienting it with the nest entrance. A statistical test was
performed to know the independent variable which is contributing to the habitat
selection between the above mentioned habitat types by chi square test followed
by Marascuilo’s post hoc test. The alpha level for
all statistical tests was kept at 0.05. The average was represented as mean+standard deviation
(SD) to understand the true variation of the data using SPSS 20.
RESULTS
Nest tree selection
A total of 119 nests
(Table 1) of IGS were located on 26 tree species (Table 2). There were more
than one or two nests in a single tree. The tree species with multiple nests
were Culenia exarillata,
Artocarpus heterophyllus
and Mesua ferrea. The
number of nests in contiguous forests, fragmented forests, plantation with
native tree shade and plantation with monoculture tree shade was 67, 4, 43, and
5, respectively, and the density of 1.48 nests/km2, 0.5 nests/km2,
1.72 nests/km2, and 0.29 nests/km2, respectively in these
habitat types varied significantly (χ2 = 45.9 df
= 3 p <.01). Plantation with native tree shade and contiguous forest
accounted for 81% of the nests. IGS were observed to nest on 26 tree species
(Table 2) and the number of nests per tree species varied significantly (χ2
= 95.07 df = 25 p <.01). However, only four tree
species including Culenia exarillata, Mesua ferrea, Actinodaphne
malabarica, and Schleichera
oleosa accounted for 45.36% of the nests with
15.96%, 11.76%, 9.24%, and 8.40% nests, respectively (Table 2). Table 2 shows
the frequency in various class height on which nests were observed. The
squirrels made nests in trees of height classes of <10 m, 10–20 m, 21–30 m,
31–40 m, and >40 m with a frequency of 0, 61, 51, 5, and 2, respectively,
which differed significantly (χ2 = 117.89 df
= 3 p <.01). Likewise, the number of nests in nest height categories of
<10 m, 11–20 m, 21–30 m, 31–40 m, and >40 m was 1, 70, 41, 6, and 1,
respectively, which significantly varied (χ2 = 121.71 df = 4 p <.01). The tree height and the nest height
correlated significantly (Pearson r = 0.96 N = 5 p <.01) (Figure 2) showing
that most nests were in trees of height up to 30 m with similar nest height
numbers indicating that the nests were towards the tree canopies.
.
Nest characteristics
The IGS builds
globular nests out of green leaves, twigs, and branches. The nests were either
round or oval in shape with a lateral opening. The nests were usually
constructed away from the tree trunk where the canopies were interlocked with
the neighboring tree canopies. The entry of the nest was placed horizontal to
the ground. Most of the nests were constructed by using the tender leaves of
the nesting trees itself. However, squirrels also used the leaves of other
plant species such as Mallotus tetracoccus and Pouteria
campechiana in the construction of the nests. The
number of nests in different directions (Figure 3) varied significantly (χ2
= 27.06 df = 7 p <.01). Most of the nests sighted
in the study area were observed facing towards the north-east direction (n =
29) followed by east (n = 23), and south-west (n = 17) whereas least preferred
direction was towards the south (n = 05) (Figure 3). Nests were very often
found at the highest point on the tree (Table 2).
DISCUSSION
Preference for
nesting habitat could depend on factors such as access to nesting material,
nest safety, branching pattern of the tree species, and availability of food. A
total of 26 tree species were recognized as nesting trees of the IGS in Nelliyampathy Reserve Forest of which Culenia
exarillata, Mesua ferrea, Actinodaphne
malabarica, and Schleichera
oleosa were highly preferred for nesting. The
high preference of these species maybe because of the dense canopy and higher
canopy height which facilitates the IGS for easy movement from the nest in all
directions (Ramachandran & Suganthasakthivel
2010). There is also the major advantage to escape from predators and to move
to other parts of the home range for foraging and other activities through the
canopy (Datta & Goyal 1996; Arockianathan
2020). In addition to that, most of the species of trees which is preferred for
nesting accounts for the major diet of the IGS as well. Given the fact that the
IGS prefers to feed as soon as they get out of the nests in the dawn and to
feed before entering the nests in the dusk (Ramachandran 1988), they prefer
mostly to build nests in the same trees which they feed on frequently. This
helps them to reduce the time and energy spent on locomotion and foraging
activities and in turn helps in conserving energy for other activities.
Most of the nests
were sighted in the plantation with native tree shade and contiguous forest,
maybe because of the availability and more abundance of the preferred nesting
trees with highest parts of canopies and canopy contiguity, and the presence of
food throughout the year in these habitat structures. Fragmented forests and
plantation with monoculture tree shade were less preferred because they are
more open and exposed habitat types where the probability of encountering
predators is more, and as they consist less diverse tree species composition,
the food choices are also limited. We observed that the nests were built
towards the tree tops, but these were not built on the extreme top of the
canopy, as the squirrels sought cover above the nest. Such cover might help to
avoid direct heat from the sun and serve as hiding from birds of prey (Datta 1998; Pradhan et al. 2012). We observed that majority
of the nests were built by using the same foliage of the tree in which the
nests were built but, in some instances, they were using different foliage than
the nesting tree species. We could not comprehend the reason behind this kind
of behavior and hence, more specific study is required for knowing as to why
some trees are used for nesting but its leaves are not used for nest building.
We found evidence to
support the hypothesis that the nest orientation is mostly towards the north-east
and east directions. This shows that the species has the cognitive ability to
identify different directions and they preferred to orient most of the nests
towards the sun rise. As the temperature in these forests becomes low in the
nights, the animal receives the early morning sun light from the easterly
direction for the warmth. Further, this region gets its rains primarily from
the south-west monsoons in which the heavy winds and the rains are received
from the west. The easterly direction of the nests therefore helps avoid direct
exposures to winds and heavy monsoon rains.
Table 1. Preferred
habitat types of the Indian Giant Squirrel for nesting.
Habitat type |
Number of nests |
Density of the nest
per sq. km. |
Percentage (%) |
Contiguous forest |
67 |
1.48 |
56.30 |
Fragmented forest |
04 |
0.5 |
3.36 |
Plantation with
native tree shade |
43 |
1.72 |
36.13 |
Plantation with
monoculture tree shade |
05 |
0.29 |
4.20 |
Table 2. Tree species
and nesting height preference by Indian Giant Squirrel.
|
Tree species |
Tree height (m ± SD) |
Nest height (m ± SD) |
Percentage (%) |
1 |
Actinodaphne malabarica |
25 ± 9.98 |
24 ± 9.45 |
9.24 |
2 |
Aglaia bourdillonii |
40 ± 0 |
38 ± 0 |
0.84 |
3 |
Aglaia malabarica |
24.66 ± 1.52 |
22.33 ± 1.52 |
2.52 |
4 |
Artocarpus heterophyllus |
18 ± 3.03 |
17.5 ± 2.42 |
5.04 |
5 |
Cedrela toona |
25.5 ± 4.24 |
22.12 ± 2.69 |
6.72 |
6 |
Cinnamomum malabatrum |
18 ± 0 |
17 ± 0 |
0.84 |
7 |
Cordia gharaf |
22 ± 0 |
21 ± 0 |
0.84 |
8 |
Cullenia exarillata |
23.12 ± 4.96 |
21.73 ± 5.69 |
15.96 |
9 |
Drypetes malabarica |
16 ± 3.46 |
13.66 ± 4.93 |
2.52 |
10 |
Dysoxylum malabaricum |
15 ± 4.24 |
13.5 ± 4.94 |
1.68 |
11 |
Ficus beddomei |
16 ± 0 |
13.5 ± 0.70 |
1.68 |
12 |
Ficus racemosa |
20.66 ± 4.61 |
19.66 ± 3.78 |
2.52 |
13 |
Ficus talbotii |
25.33 ± 4.61 |
23.66 ± 4.16 |
2.52 |
14 |
Garcinia
gummi-gutta |
18 ± 0 |
18 ± 0 |
0.84 |
15 |
Holoptelea integrifolia |
17 ± 4.24 |
15.5 ± 3.53 |
1.68 |
16 |
Macaranga peltata |
16 ± 3.46 |
14.33 ± 3.05 |
2.52 |
17 |
Mangifera indica |
18 ± 0 |
16 ± 1.41 |
1.68 |
18 |
Mesua ferrea |
22.57 ± 7.77 |
21 ± 7.22 |
11.76 |
19 |
Myristica dactyloides |
20.5 ± 5 |
19 ± 4.83 |
3.36 |
20 |
Neolitsea scrobiculata |
23 ± 0 |
21 ± 0 |
0.84 |
21 |
Palaquium ellipticum |
23 ± 10.39 |
20 ± 8.12 |
3.36 |
22 |
Persea macrantha |
22.6 ± 3.43 |
21.6 ± 2.88 |
4.20 |
23 |
Pleurostylia opposita |
20.8 ± 4.60 |
19.6 ± 4.03 |
4.20 |
24 |
Polyalthia longifolia |
23.75 ± 6.13 |
21.25 ± 6.13 |
3.36 |
25 |
Schleichera oleosa |
21.3 ± 5.33 |
19.2 ± 5.63 |
8.40 |
26 |
Vernonia monosis |
21 ± 7.81 |
18.33 ± 7.23 |
2.52 |
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