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Original Research Article
Folia Primatol 2017;88:57–74
DOI: 10.1159/000477581
Diet and Activity of Macaca assamensis
in Wild and Semi-Provisioned Groups in
Shivapuri Nagarjun National Park, Nepal
Sabina Koirala a Mukesh K. Chalise b Hem Bahadur Katuwal a
Raju Gaire
a Bishnu Pandey c Hideshi Ogawa d
a Small Mammals Conservation and Research Foundation and
b Central Department of
Zoology, Tribhuvan University, Kathmandu , and
c National Trust for Nature Conservation,
Lalitpur , Nepal; d School of International Liberal Studies, Chukyo University, Nagoya , Japan
Keywords
Activity budget · Diet · Human food · Assamese macaque · Nagarjun forest ·
Frugivorous diet
Abstract
Studying the behavioural flexibility and adaptability of macaques to different hab-
itats is one approach to designing a conservation plan. To determine the activity budget
and feeding behaviour and evaluate the effects of seasonality in wild and human-
altered habitats of Assamese macaques ( Macaca assamensis ), we conducted this study
in the Nagarjun forest of Shivapuri-Nagarjun National Park (SNNP) in central Nepal. We
also updated the list of plant food items of Assamese macaques in the SNNP. Using scan
and all-occurrence sampling, we recorded the diets and activities of Assamese ma-
caques in 2 social groups, a wild-feeding group (WG) and a semi-provisioned group
(SPG), throughout the year from August 2013 to July 2014. Both groups spent most of
their time in feeding activities and were quite arboreal, but there were significant differ-
ences in the activity budgets and diets between the groups. Human food was the main
component of the diet for the SPG, whereas it was fruit for the WG, indicating a normal-
ly frugivorous diet. Furthermore, the activity budget and diet composition varied in re-
sponse to the season. These results indicate that provisioning alters the activity and
feeding behaviour of macaques, and can also increase human-macaque conflict and
disease transmission. © 2017 S. Karger AG, Basel
Received: September 2, 2016
Accepted after revision: May 17, 2017
Published online: July 12, 2017
Sabina Koirala
Small Mammals Conservation and Research Foundation
PO Box 9092
Sundhara, Kathamandu (Nepal)
E-Mail sabina.koirala2 @ gmail.com
© 2017 S. Karger AG, Basel
www.karger.com/fpr
E-Mail karger@karger.com
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58 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
Introduction
Non-human primates are flexible in diets and activity budgets, so they can survive
in variable habitats altered by humans [Alami et al., 2012]. Variability in activity bud-
gets depending on habitat differences provides an opportunity to investigate the adap-
tive strategies of a species [Sarkar et al., 2012]. Knowledge of the diet and activity of a
species is an important requirement in designing a conservation plan [Gupta, 2005].
Assamese macaques ( Macaca assamensis ) spend most of their time in feeding
activity [Chalise, 2003; Schulke et al., 2011; Sarkar et al., 2012]. They have been de-
scribed as omnivorous [Boonratana et al. , 2008] because they feed on leaves, flowers,
seeds, barks, shoots and caterpillars, as well as fruits [Mitra, 2002; Chalise, 2003; Chal-
ise et al., 2013a; Koirala and Chalise, 2014], and they raid crops such as maize, rice,
wheat, millet, potatoes and cultivated fruits [Chalise, 2003, 2010; Regmi et al., 2013].
They sometimes even feed on other animals, including mammals, birds, reptiles, am-
phibians, molluscs and spiders [Schulke et al., 2011]. However, some studies con-
cluded that Assamese macaques were highly folivorous [Chalise, 2003; Zhou et al.,
2011]. In contrast to these previous studies, however, it has been found that the As-
samese macaques in Nagarjan forest prefer fruits to other plant parts [Koirala and
Chalise, 2014].
The dietary composition of non-human primates depends on the availability of
food resources [Chalise, 1999; Riley, 2007; Jaman and Huffman, 2013; Koirala and
Chalise, 2014]. It may also be affected by habitat quality, depending upon foraging
options [Poulsen et al., 2001; Jaman and Huffman, 2013]. Feeding habits are affected
by seasonal changes in food availability in the habitat [Chalise, 1999]. Furthermore,
primates select their foods depending on their digestive system and the food nutrients
they require [Chalise, 1999]. The distribution of food resources directly affects the
activity pattern of a species [O’Brien and Kinnaird, 1997; Sarkar, 2000; Poulsen et al . ,
2001; Mitra, 2002; Riley, 2007; Sarkar et al., 2012; Jaman and Huffman, 2013]. For
example, semi-provisioned Barbary macaques ( M. sylvanus ) adopted a lower-energy
foraging strategy, allowing more resting time and less foraging and moving time com-
pared with wild Barbary macaques [Alami et al., 2012]. The feeding time of rhesus
macaques ( M. mulatta ) in a rural group was longer than that in an urban group in
response to the availability of food resources [Jaman and Huffman, 2013]. Artificial
foods provided by humans changed the behaviour of hamadryas baboons ( Papio
hamadryas ) [Kamal et al., 1997]. Savannah baboons ( P. cynocephalus ) in a semi-pro-
visioned group (SPG) spent less time feeding than wild groups did [Altmann and
Muruthi, 1988]. The foraging time of M. tonkeana was greater in a heavily altered
habitat than in a minimally altered habitat [Riley, 2007].
Food provisioning to macaques has potentially harmful consequences [South-
wick et al., 1976; Sugiyama and Ohsawa, 1982; Zhou and Deng, 1992]. Provisioned
food significantly increased the duration and frequency of aggressive interactions
between humans and Formosan macaques ( M. cyclopis ) in Taiwan [Hsu et al., 2009].
A similar pattern was found between Barbary macaques and humans in Gibraltar
[O’Leary and Fa, 1993]. Fuentes [2006] found significantly more contact interactions
between Barbary macaques and humans due to food provisioning in the same area.
Similarly, Wheatley and Harya Putra [1994] reported that aggressive behaviour of
long-tailed macaques ( M. fascicularis ) toward visitors via food handouts was posi-
tively reinforced in Bali; the frequency and intensity of aggressive behaviour of the
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Feeding and Activity of M. assamensis
59
Folia Primatol 2017;88:57–74
DOI: 10.1159/000477581
macaques was positively correlated with the quantity and quality of available food.
Similarly, long-tailed macaques in Singapore were significantly more likely to inter-
act with humans when food was present [Fuentes et al., 2008]. On Mount Emei,
humans were sometimes injured by Tibetan macaques ( M. thibetana ) because of
close interactions between them; in an 8-year period, 10 tourists died as an indirect
result of human-macaque interactions [Zhao and Deng, 1992]. Furthermore, Mc-
Carthy et al. [2009] reported that the close contact between humans and macaques
can have deleterious consequences on the health of both species. Macaques can
transmit simian foamy virus, herpes B virus, simian T cell lymphotropic virus and
simian retro virus to humans [Engel et al., 2002; Wolfe et al., 2004; Jones-Engel et
al., 2005], while humans can transmit measles, influenza and respiratory pathogens
to macaques [Jones-Engel et al., 2001]. Thus, information on ecology and behaviour
is crucial for the conservation of a species in human-modified environments [Alami
et al., 2012].
The local population of Assamese macaques in Nepal differs morphologically
between the co-existing subspecies, eastern Assamese macaques ( M. assamensis as-
samensis ) and western Assamese macaques ( M. assamensis pelops ) [Molur et al.,
2003]. Because of this taxonomic confusion, they are referred to as the “Nepal popu-
lation.” The Nepal population is endemic to Nepal, inhabiting subtropical hills in Sal
forests, mixed deciduous forests, temperate broadleaved forests with rocky outcrops,
and steeply sloped riverside forests at high altitudes [Molur et al., 2003]. However,
they are endangered because of their restricted distribution, the small number of
individuals in fragmented patches, and the reductions in their habitats and popula-
tion caused by human activities [Molur et al., 2003; Boonratana et al., 2008].
Despite past research on Assamese macaques in Nepal [Chalise, 2003, 2010; Chal-
ise et al., 2013b; Koirala and Chalise, 2014], detailed knowledge about their activity
budgets and feeding behaviour in relation to the availability of human food and in dif-
ferent seasons is still lacking. Thus, we assessed Assamese macaques’ behavioural flex-
ibility in response to the season and the availability of human food in their habitat. We
observed 1 SPG and 1 wild-feeding group (WG) of Assamese macaques in the Nagar-
jun forest of Shivapuri-Nagarjun National Park (SNNP), as a continuation of a previ-
ous study by Koirala and Chalise [2014]. We worked on 2 hypotheses: (1) the diet com-
position and activity budgets of Assamese macaques are affected by the availability of
human food, and (2) the diet composition and activity budgets change according to the
season. We also updated the total food plants of the macaque from the SNNP, Nepal.
Methods
Study Area
We conducted field research in the Nagarjun forest (27°43 ′ to 27°46 ′ N; 85°13 ′ to 85°18 ′ E)
in SNNP, north of the Kathmandu valley, central Nepal ( Fig.1 ). The SNNP has 2 separate forests,
Shivapuri and Nagarjun. The Nagarjun forest occupies an area of 16 km
2 , ranging between 1,350
and 2,100 m above sea level [Pokhrel et al., 2011].
The Nagarjun forest contains 4 different types of vegetation, namely Schima wallichii forest,
mixed broadleaved forest, pine forest, and dry oak forest, which occupy 61.3, 27.9, 9.1, and 1.7%
of the total area, respectively [Nagarkoti, 2006]. The mean monthly rainfall ranges from 3.43 to
444.56 mm (July, August, and September have the most precipitation) [Bhandari and Chalise,
2014]. The mean monthly temperature ranges from 3.9 to 20.4
° C (minimum) and 18.6 to 29.6 ° C
(maximum) [Koirala and Chalise, 2014].
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60 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
Two species of non-human primate, the Assamese macaque ( M. assamensis ) and rhesus
macaque ( M. mulatta ), inhabit the area. Other mammalian fauna includes at least 6 species of
bats [Malla, 2000; Thapa et al., 2010], the orange-bellied Himalayan squirrel ( Dremomys lokriah ),
Irrawaddy squirrel ( Callosciurus pygerythrus ), Chinese pangolin ( Manis pentadactyla ), Eurasian
wild boar ( Sus scrofa ), common leopard ( Panthera pardus ), barking deer ( Muntiacus muntjak ),
and sambar deer ( Cervus unicolor ) [Wada, 2005; Nagarkoti, 2006; Bhandari and Chalise, 2014].
Sampling Design and Data Collection
We observed 2 social groups of Assamese macaques: an SPG at Fulbari (elevation 1,372 m,
N 27.74397°, E 85.29482°) and a WG at Raniban (elevation 1,360 m, N 27.74112°, E 85.28718°),
0 20 40 80 120 160
km
0 0.2 0.4 0.8 1.2 1.6
km
Raniban
Fulbani
Protected areas
Study sites
Nagarjun forest
N
S
WO
Fig. 1. Map of the Nagarjun forest in the SNNP, indicating the distribution of the study groups
of Assamese macaques.
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Feeding and Activity of M. assamensis
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Folia Primatol 2017;88:57–74
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following Koirala and Chalise [2014]. The total numbers of individuals in the WG and SPG were
34 and 47 at the beginning, and 37 and 44 at the end of the study period, respectively ( Table1 ).
The groups occupied similar habitats, and both were somewhat habituated to humans due
to frequent visits by tourists and local people. However, the SPG preferred an army camp area in
the Fulbari forest, which is characterised by the presence of garbage and frequent movement of
security personnel and park staff. The monkeys in SPG consumed waste food at a large kitchen
area of the army camp. They typically consumed garbage from a drum where army staff disposed
of leftovers from their meals, near a tap area where army staff washed dishes, and from a dump-
ing site where army staff threw away excess raw vegetables. Garbage at the army camp was always
available, with no seasonal change in availability. By contrast, the WG was usually found in a for-
est that was minimally disturbed by humans. The levels of “artificial” food availability varied
between the groups [Koirala and Chalise, 2014].
One author (S.K.) collected all the data during 4 seasons from August 2013 to July 2014 (20
days in spring, 11 days in summer, 18 days in autumn, and 18 days in winter). Before data collec-
tion, both groups of macaques were habituated to S.K. Daytime observation generally began at
7:
00 a.m. and ended when the study group was lost and could not be relocated or had settled into
its sleeping site. Because it was more difficult to find and follow the WG than the SPG, we made
more efforts during observation of the WG to collect comparable data. After finding and before
losing the study group, however, we used the same sampling method with both groups. Specifi-
cally, we used a scan sampling method to collect data on diet and activity [Martin and Bateson,
1993] and recorded the activities and feeding items of macaques every 15 min; i.e., watching the
macaques for 5 min, followed by 10 min of non-recording until the next scan. We observed each
individual for 5 s using binoculars, and recorded the predominant behaviour during the 5-s pe-
riod. We did not record the behaviour of infants <1 year old. We inevitably could not find some
individuals in the study groups because of the dense vegetation, so we sampled parts of the group
during most scans.
We categorised the activities of the Assamese macaques as feeding, resting, moving and so-
cial behaviours (social grooming, playing, aggression and sexual behaviour) for each scan
[O’Brien and Kinnaird, 1997]. We also recorded plant parts such as fruit (ripe or unripe), leaves
(young or mature), flowers, shoots and others (young cone, petiole, tendril, bark, tuber, root,
etc.), insects (and their secretions and eggs) and mushrooms when macaques were feeding during
the scan. If an individual was feeding while engaging in another activity, such as moving and rest-
ing, the behaviour was recorded as feeding. S.K. also recorded the locations of macaques and
whether they were in trees or on the ground during the scan. In addition, we used the all-occur-
rences sampling method to record new food items and plant species during the interval between
5-min scans [Martin and Bateson, 1993].
We calculated the fruit availability of plants based on the plots established by Koirala and
Chalise [2014]. In total, there were 44 plots, each measuring 10 × 10 m, at each of the 2 study sites,
Raniban and Fulbari. The researchers identified and measured the diameter at breast height
(DBH) of 231 trees with a diameter ≥ 10 cm in each plot. We observed the phenological state of
Group Class August 2013 July 2014
WG Adult male 8 7
Adult female 11 12
Immature 15 18
Total 34 37
SPG Adult male 9 8
Adult female 10 9
Immature 28 27
Total 47 44
Table 1. Age and sex
composition of the study
groups
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62 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
each tree, including the food trees of the macaques, on predetermined dates during the 4 seasons
via visual examination using binoculars. We recorded the percentages of young and mature
leaves, flowering buds, flowers, and unripe and ripe fruits from the proportion of the total cano-
py covered by the item being measured, applying an abundance score of 0–4, where 0 = none of
the canopy, 1 = 1–25% (average 12.5%), 2 = 26–50% (average 37.5%), 3 = 51–75% (average
62.5%), and 4 = 76–100% (average 87.5%) [Riley, 2007].
Data Arrangement and Data Analysis
First, we updated the food plants of Assamese macaques; S.K., H.O. and B.P. recorded the
food plants and items eaten by Assamese macaques of the WG, SPG and other groups in the en-
tire SNNP via ad libitum sampling from April 2011 to March 2016, and reviewed previously pub-
lished reports.
We prepared herbariums of unidentified plant species and identified them in the National
Herbarium and Plant Laboratories, Godawari, Lalitpur, Nepal. The Department of National
Parks and Wildlife Conservation, Nepal, provided permission for all fieldwork and data collec-
tion within the SNNP. No live animals were handled during the fieldwork.
Second, we calculated the proportion of time spent on each activity by each group in each
season separately. We used repeated-measures multivariate analysis of variance (rMANOVA) to
analyse the differences in the activity budgets of the 2 groups and the influence of the seasons.
We used 1-way ANOVA to analyse the seasonal influence on the activity of each macaque in the
2 groups. Additionally, based on the number of behaviour events in each activity, we used a χ
2
test to examine the significance of differences in dietary composition and the locations of the WG
and SPG to check seasonal differences in activity budgets and diet composition. For all statistical
tests, we considered p values ≤ 0.05 to indicate statistical significance.
Third, we calculated the percentage of feeding performed on each plant and each plant part
consumed by macaques from the number of feeding records for the plant species or the food item,
divided by the total number of feeding records, and multiplied by 100. We calculated the fruit
availability in each season using the following index: total DBH
2 × the proportion of fruits in the
total canopy of each tree (12.5, 37.5, 62.5, or 87.5%). We examined the differences in fruit avail-
ability according to season using the Kruskal-Wallis test, and we checked the relationship of ma-
caque feeding times and the proportion of fruit feeding with fruit availability according to season
using the Spearman rank correlation test.
R e s u l t s
Food Plants of Assamese Macaques in SNNP
Based on this study and published reports, we recorded 123 plant species, repre-
senting 89 genera and 56 families, as food plants of Assamese macaques in SNNP
( Table 2 ). Tree species accounted for the highest percentage (57.1%), followed by
climbers (16.8%), shrubs (18.5%), herbs (5.0%), and a few species of mushrooms,
ferns and parasitic plants.
Observation Time and Number of Records
We recorded a total of 21,836 behaviour events via 2,008 scan samplings during
508 h of observations. We recorded 7,666 and 14,170 behaviour events for the WG
and SPG, respectively. In total, we recorded 4,961 feeding events in the WG and SPG
during the total study period.
Effect of Human Food on the Diet of Macaques
The overall diet composition of the WG and SPG varied significantly (df = 8,
χ
2 = 1992.2, p < 0.05). The SPG animals were heavily dependent on human food
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Folia Primatol 2017;88:57–74
DOI: 10.1159/000477581
No. Family Species Local name Plant type Parts eaten Source
1 Aceraceae Acer oblongum Wall. ex DC. Phirphire Tree Fl This study; Chalise et al.
[2013b]
2 Anacardiaceae Choerospondias axillaris
(Roxb.) B.L. Burtt & A.W.
Hill
Lapsi Tree Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
3 Anacardiaceae Rhus sp. Bhalayo Tree Shoots This study; Koirala and
Chalise [2014]
4 Anacardiaceae Rhus succedanea var.
succedanea L.
Rani bhalayo Tree Fr This study
5 Anacardiaceae Semecarpus anacardium L. f. Kag bhalayo Tree Unknown Chalise et al. [2013b]
6 Araliaceae Unidentified species 1 Climber YL, L This study
7 Asclepiadaceae Periploca calophylla (Wight.)
Falc.
Climber YL This study
8 Asclepiadaceae Unidentified species 2 Climber L This study
9 Asteraceae Crassocephalum crepidioides
(Benth.) S. Moore
Anikale jhar Herb L, YL, bunches
of flowers and
young stems
This study
10 Berberidaceae Berberis aristata DC. Chutro Shrub Fr, YL This study; Chalise et al.
[2013b]
11 Berberidaceae Berberis asiatica Roxb. ex DC. Chutro Shrub Fr, YL, FL This study
12 Berberidaceae Mahonia napaulensis DC. Jamane
mandro
Shrub Fr This study
13 Betulaceae Alnus nepalensis D. Don Uttis Tree Fr, exudates This study
14 Betulaceae Betula alnoides Buch.-Ham.
ex D. Don
Saur/lekh
painu
Tree YL, L, leaves
with aphids,
shoots
This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
15 Bombacaceae Bombax ceiba L. Simal Tree Bud, Fl, Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
16 Boraginaceae Ehretia sp. Tree YL, shoots This study
17 Convallariaceae Polygonatum sp. L This study
18 Cucurbitaceae Solena heterophylla Lour. Ban kankro Herbaceous
climber
YL, L This study
19 Cucurbitaceae Trichosanthes tricuspidata
Lour.
Indreni Woody climber YL, L This study
20 Cucurbitaceae Trichosanthes wallichiana
(Ser.) Wight.
Indreni Herbaceous
climber
YL, L, tendrils This study; Koirala and
Chalise [2014]
21 Dioscoreaceae Dioscorea prazeri Prain &
Burkill
Kukur tarul Climber YL This study
22 Ebenaceae Diospyros kaki Thunb. Haluwabed Tree Fr Koirala and Chalise [2014]
23 Ebenaceae Diospyros malabarica (Desr.)
Kostel.
Kalo tindu Tree Unknown Chalise et al. [2013b]
24 Elaegnaum Unidentified species 3 Climber YL This study
25 Ericaceae Lyonia ovalifolia (Wall.)
Drude
Angeri Tree Unknown Chalise et al. [2013b]
26 Ericaceae Rhododendron sp. Guras Tree Fr, Fl This study; Chalise et al.
[2013b]
27 Euphorbiaceae Bridelia retusa (L.) Spreng. Gayo Tree L This study
28 Euphorbiaceae Euphorbia royleana Boiss. Siundi Tree Unknown Chalise et al. [2013b]
29 Euphorbiaceae Macaranga indica Wight Mallato Tree Unknown Chalise et al. [2013b]
30 Euphorbiaceae Phyllanthus emblica L. Amala Tree Fr This study; Chalise et al.
[2013b]
31 Fabaceae Albizia procera (Roxb.) Benth. Seto siris Tree Unknown Chalise et al. [2013b]
32 Fabaceae Albizia julibrissin Durazz.Siris Tree Unknown Chalise et al. [2013b]
33 Fabaceae Bauhinia malabarica Roxb.Tanki Tree Bark This study; Koirala and
Chalise [2014]
34 Fabaceae Bauhinia variegate L. Koiralo Tree Unknown Chalise et al. [2013b]
35 Fabaceae Entada phaseoloides (L.)
Merr.
Pangra Climber Young seeds This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
36 Fabaceae Indigofera pulchella Roxb. Phusre ghans,
mirmire
Shrub L This study
37 Fabaceae Lathyrus sativus L. Khesari Climber L This study
38 Fabaceae Shuteria involucrata var.
glabrata (Wall.) Wight & Arn.
F This study
Table 2. Food plants of Assamese macaques of SNNP based on this study and the literature reviews
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64 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
No. Family Species Local name Plant type Parts eaten Source
39 Fagaceae Castanopsis indica (Roxb.)
Miq.
Dhalekatus Tree Fr This study; Chalise et al.
[2013b]
40 Fagaceae Castanopsis tribuloides (Sm.)
A. DC.
Musure katus Tree Fr, Fl, young
nuts
This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
41 Fagaceae Lithocarpus elugans (Blume)
Hatus ex Soep.
Arkhaulo Tree Fl, L, petioles This study; Chalise et al.
[2013b]
42 Fagaceae Quercus glauca var. glauca
Thunb.
Phalat Tree L, YL, bunches
of young fruits
This study
43 Fagaceae Quercus lanata Sm. Baanjh Tree Fr This study; Chalise et al.
[2013b]
44 Fagaceae Quercus semecarpifolia Sm. Khasru Tree Fr This study
45 Fagaceae Castanopsis hystrix Miq. Patale katus Tree Fr This study; Chalise et al.
[2013b]
46 Juglandaceae Juglans regia L. Sano okhar Tree Unknown Chalise et al. [2013b]
47 Lamiaceae Colebrookea oppositifolia Sm. Dhursuli Shrub Unknown This study
48 Lamiaceae Anisomeles indica (L.) Kuntze Rato charpate Herb L This study
49 Lardizabalaceae Holboellia latifolia Wall. Malkati Climber Fr This study
50 Lauraceae Cinnamomum tamala (Buch.-
Ham.) Nees & Eberm.
Tejpat Tree Shoots This study; Chalise et al.
[2013b]
51 Lauraceae Dodecadenia grandiflora var.
grandiflora Nees
Panhele Tree Fr, YL This study; Koirala and
Chalise [2014]
52 Lauraceae Lindera nacusua (D. Don)
Merr
Panhelo
khapate
Tree Fr This study; Koirala and
Chalise [2014]
53 Lauraceae Litsea doshia (Buch.-Ham. ex
D. Don) Kosterm.
Paheli Tree L This study; Chalise et al.
[2013b]
54 Lauraceae Litsea monopetala (Roxb.)
Pers.
Kutmiro Tree Unknown Chalise et al. [2013b]
55 Lauraceae Litsea sp. Tree Petioles This study
56 Lauraceae Neolitsea pallens (D. Don)
Momiy. & H. Hara ex H. Hara
Tree Shoots This study
57 Lauraceae Persea duthiei (King ex
Hook.f.) Kosterm.
Jhangrikath/
mahilo kaulo
Tree Fr This study; Koirala and
Chalise [2014]
58 Lauraceae Persea odoratissima (Nees)
Kosterm.
Seto kaulo Tree Fr, YL, young
stems
This study
59 Linaceae Reinwardtia indica Dumort. Pyauli/
bakhre ghas
Shrub Unknown Chalise et al. [2013b]
60 Loranthaceae Scurrula parasitica L. Ainjeru Hemiparasitic,
epiphytic shrub
Fr, L, petioles This study; Koirala and
Chalise [2014]
61 Loranthaceae Scurrula sp.Ainjeru Epiphytic shrub Petioles This study
62 Loranthaceae Helixanthera parasitica Lour. Liso Parasite YL This study
63 Loranthaceae Loranthus sp. Ainjeru Parasite Fr This study; Chalise et al.
[2013b]
64 Lythraceae Woodfordia fruticose (L.)
Kurz
Dhainyaro Shrub Unknown Chalise et al. [2013b]
65 Melastomataceae Oxyspora paniculata (D. Don)
DC.
Shrub YL, L This study
66 Menispermaceae Cissampelos pareira L. Bahule
pati
Climbing shrub YL, L This study
67 Moraceae Ficus lacor Buch.-Ham. Kabhro Tree Fr, buds This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
68 Moraceae Ficus religiosa L. Peepal Tree Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
69 Moraceae
Ficus semicordata Buch.-Ham.
ex Sm.
Khaniya Tree Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
70 Moraceae Ficus sp. 1 Bedu Climber Fr This study
71 Moraceae Ficus sp. 2 Climber Fr, L This study
72 Moraceae Ficus sp. 3 Pagri Climber YL This study
73 Moraceae Ficussp. 4 Banpeepal Tree Fr, YL, shoots This study
74 Moraceae Ficus subincisa Buch.-Ham.ex
Sm.
Bedulo Tree Fr, YL This study
75 Moraceae Maclura cochinchinensis
(Lour.) Corner
Damaru Tree Fr, L, YL,
shoots
This study; Koirala and
Chalise [2014]
Table 2 (continued)
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No. Family Species Local name Plant type Parts eaten Source
76 Myricaceae Myrica esculenta Buch.-Ham.
ex D. Don.
Kafal Tree Fr This study; Chalise et al.
[2013b]
77 Myrsinaceae Ardisia macrocarpa Wall. Damai
phal
Shrub Fr, L, bark This study
78 Myrsinaceae Maesa chisia Buch.-Ham. ex
D. Don
Bilaune Tree Fr, Fl, L This study; Chalise et al.
[2013b]
79 Myrsinaceae Myrsine capitellata Wall. Seti kath Tree L This study; Chalise et al.
[2013b]
80 Myrsinaceae Myrsine semiserrata Wall. Kalikath Tree Fr, YL, L This study; Chalise et al.
[2013b]
81 Myrsinaceae Unidentified species 4 Tree YL, young stem This study
82 Myrtaceae Syzygium cumini (L.) Skeels Jamun Tree Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
83 Nephrolepidaceae Nephrolepis cordifolia (L.) K.
Presl
Paniamala Fern Tuber This study
84 Oleaceae Jasminum sp. Climber YL, L This study
85 Oleaceae Ligustrum confusum Decne. Kanike
phool
Shrub Fr This study
86 Oleaceae Ligustrum indicum (Lour.)
Merr.
Keri Shrub Shoots This study
87 Oleaceae Unidentified species 5 Tree Fl, bud This study
88 Oleaceae Fraxinus floribunda Wall. Lankuree Tree Unknown Chalise et al. [2013b]
89 Orchidaceae Vanda cristata Lindl. Harchur Epiphytic herb This study
90 Orchidaceae Kingidium taenialis (Lindl.)
P.F. Hunt
Epiphytic herb Whole plant This study
91 Pinaceae Pinus roxburghii Sarg. Khote
sallo
Tree Young cone This study; Chalise et al.
[2013b]
92 Poaceae Thamnocalamus aristatus
(Gamble) E.G. Camus
Nigalo Herb Shoot This study
93 Rhamnaceae Zizyphus incurva Roxb. Hadebayar Tree Fr This study; Koirala and
Chalise [2014]
94 Rhamnaceae Zizyphus mauritiana Lam. Bayar Tree Fr This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
95 Rosaceae Eriobotrya elliptica Lindl. Mayo Tree L This study
96 Rosaceae Rubus acuminatus Sm. Saanu ainselu Shrub Fr, L, shoot This study
97 Rosaceae Rubus ellipticus Sm. Ainselu Shrub Fr This study
98 Rosaceae Prunus cerasoides D. Don Payou Tree L, YL, Fl,
leaves with
aphids and
gum
This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
99 Rubiaceae Hedyotis scandens Roxb. Bakhri lahara Herbaceous
climber
L, young stem This study
100 Rutaceae Euodia fraxinifolia (D. Don)
Hook. F.
Kanukpa Tree Fr This study
101 Rutaceae Toddalia asiatica (L.) Lam. Main kanda Climber L This study
102 Sabiaceae Meliosma simplicifolia (Roxb.)
Walp.
Tree L, Fl This study
103 Sambucaceae Viburnum mullaha Buch.-
Ham. ex D. Don
Molo Climber Fr Koirala and Chalise [2014]
104 Sapotaceae Madhucalongifolia (Koenig)
Mac.
Mahuwa Tree Fr, Fl This study
105 Saurauiaceae Saurauia napaulensis DC. Gogan Tree Fr, shoots This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
106 Smilacaceae Smilax aspera L. Kukur daino Climbing shrub YL This study
107 Smilacaceae Smilax lanceifolia Roxb. Kubiraina Climbing shrub YL This study
108 Symplocaceae Symplocos sumuntia Buch.-
Ham. ex D. Don
Hakulal Tree Fr, Fl This study; Chalise et al.
[2013b]
109 Symplocaceae Symplocos sp. Baklepat Tree YL This study
110 Taxaceae Taxus wallichiana Zucc. Lauth salla Tree Fr This study
111 Theaceae Eurya acuminate DC. Saano
jhingane
Tree Fr This study; Chalise et al.
[2013b]
Table 2 (continued)
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66 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
( Table3 ). We recorded more feeding events in the SPG (2,520 events) than in the WG
(2,441 events). Nevertheless, the WG consumed more plant species (59 plant species,
44 genera, 33 families, excluding 2 unidentified species) than the SPG did ( Table3 ).
Although human food accounted for 52.9% of the total diet of the SPG, they also fed
on 50 plant species (41 genera and 35 families, excluding 5 unidentified species) dur-
ing our study ( Table3 ). Assamese macaques, especially in the WG, were highly fru-
givorous; the WG spent 65.6% of total feeding events on fruits, including nuts and
seeds, while in the SPG only 22.5% of feeding events were of fruits. Similarly, we re-
corded both groups feeding on plant parts (flowers, young cones, leaves, shoots, pet-
ioles, barks, roots, buds, cactus spines) and animal content (insects, eggs, caterpillars,
vaginal secretion, semen; Table3 ).
We found that macaques in both groups were quite arboreal, although the SPG
animals spent more time on the ground than the WG did (df = 1, χ 2 = 927.7, p < 0.05).
The WG spent 98.5% of their total recorded events in trees, and only 1.5% on the
ground, whereas the SPG spent 85.7% of their total recorded events in trees and 14.3%
on the ground.
Effect of Human Food on Macaque Activity
The proportion of records involving feeding, 0.55 in the WG and 0.37 in the SPG,
was the highest among the 4 activity categories in both groups, while the proportion
of social behaviours was the lowest in the WG (0.12) and the proportion of moving
behaviours was lowest in the SPG (0.14). rMANOVA showed a significant difference
between WG and SPG in activity budgets (Pillai trace = 0.77, F(4, 21) = 17.41, p <
0.05). The results of separate univariate analyses showed that feeding and moving
were significantly higher, and resting and social behaviour were significantly lower in
the WG compared with the SPG (all p < 0.05; Fig.2 ).
No. Family Species Local name Plant type Parts eaten Source
112 Theaceae Schima wallichii (D.C) Korth. Chilaune Tree Buds, Fr, YL,
Fl, petioles
This study; Chalise et al.
[2013b]; Koirala and
Chalise [2014]
113 Thymelaeaceae Edgeworthia gardneri (Wall.)
Meisn.
Argali Shrub L This study
114 Tiliaceae Grewia subinaequalis DC. Phalsa Tree Fr, L This study; Koirala and
Chalise [2014]
115 Ulmaceae Celtis australis L. Khari Tree Fr, L, YL,
petioles
This study; Koirala and
Chalise [2014]
116 Urticaceae Pilea scripta (Buch.-Ham. ex
D. Don) Wedd.
Chiple Shrub YL This study
117 Verbenaceae Caryopteris sp. Khorsane
ghas
Shrub Fl This study; Koirala and
Chalise [2014]
118 Verbenaceae Unidentified species 6 Buds This study
119 Verbenaceae Lantana camara L. Masino
kanda
Climbing shrub Fr, Fl This study; Koirala and
Chalise [2014]
120 Vitaceae Ampelocissus sikkimensis
(M.A. Lawson) Planch.
Pureni Fr This study
121 Vitaceae Tetrastigma serrulatum
(Roxb.) Planch
Charchare Woody climber Fr, YL, L,
stems, tendrils
This study
122 Vitaceae Unidentified species 7 Climber Tendrils This study
123 Zingiberaceae Cautleya spicata (Sm.) Baker
in Hook
Pani saro Herb L, stems, tubers This study; Koirala and
Chalise [2014]
Fl, flowers; Fr, fruits; L, (mature) leaves; YL, young leaves.
Table 2 (continued)
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Table 3. Food items (plant parts) consumed by Assamese macaques in each season
Group Season Food items, %
Fr Fl YC YL L insect OS HF other UN
WG Spring 82.40 0.40 0.00 4.50 0.10 7.70 0.00 0.00 4.20 0.60
Summer 72.70 0.00 0.00 0.00 0.00 25.00 0.00 0.00 2.30 0.00
Autumn 88.30 0.20 0.00 6.60 0.90 2.50 0.50 0.00 0.80 0.30
Winter 40.80 1.00 25.9 1.10 16.60 5.50 0.10 0.10 6.00 2.80
Overall 65.50 0.60 11.40 3.50 7.60 5.70 0.20 0.04 4.00 1.40
SPG Spring 18.00 0.40 0.00 2.70 1.30 5.50 0.70 70.10 0.70 0.50
Summer 33.20 0.00 0.00 6.20 1.00 6.20 0.30 51.10 2.00 0.10
Autumn 23.00 3.00 0.00 22.0 1.50 0.80 1.30 46.80 0.60 0.20
Winter 13.70 0.00 32.10 1.70 4.30 1.60 0.70 45.50 0.40 0.00
Overall 22.50 0.60 8.90 7.30 2.00 3.73 0.60 52.90 1.00 0.20
Fr, fruits; Fl, flowers; YC, young cones; YL, young leaves; L, (mature) leaves; OS, own
secretions; HF, human food; UN, unknown.
Feeding
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
Moving
Activities
Proportion of activities
Resting Social
ଶ WG
ଶ SPG
Fig. 2. Activities of Assamese macaques in the WG and SPG. The proportion of activities was
calculated as the number of behaviour records for each activity didvided by the number of be-
haviour records for all activities.
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68 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
Effect of Seasonality on the Diet of Macaques
Despite the consumption of human food by the SPG being a large proportion of
their year-round diet, there was variety in the plant species and food items consumed
by both groups with respect to the seasons (df = 15, χ 2 = 832.7 in WG, χ
2 = 831.3 in
SPG; p < 0.05; Tables 3 , 4 ). The percentage of fruit feeding was higher in spring, sum-
mer and autumn in the WG, and in spring and autumn in the SPG. The number of
plant species in each season varied from 5 in the summer to 33 species in the spring,
and from 14 in the winter to 23 species in the summer in the WG and SPG, respec-
tively. The major food items also differed with season; they fed frequently on flowers
Table 4. Main plant diet of Assamese macaques in each season
Group Season Plant PE, %1
WG Spring Ficus religiosa L. 30.54
Persea duthiei (King ex Hook.f.) Kosterm. 27.75
Ficus lacor Buch.-Ham. 13.96
Bombax ceiba L. 9.20
Summer Celtis australis L. 42.42
Choerospondias axillaris (Roxb.) B.L. Burtt & A.W. Hill 24.24
Ficus semicordata Buch.-Ham. ex Sm. 24.24
Unidentified species 6.06
Autumn Castanopsis tribuloides (Sm.) A. DC. 69.83
Zizyphus incurva Roxb. 6.48
Choerospondias axillaris (Roxb.) B.L. Burtt & A.W. Hill 5.69
Betula alnoides Buch.-Ham. ex D. Don 5.37
Winter Schima wallichii (D.C) Korth. 37.03
Pinus roxburghii Sargent 28.18
Prunus cerasoides D. Don 6.00
Bombax ceiba L. 5.39
SPG Spring Ficus semicordata Buch.-Ham. ex Sm. 28.80
Unidentified species 21.60
Persea duthiei (King ex Hook.f.) Kosterm. 19.20
Schima wallichii (D.C) Korth. 8.80
Celtis australis L. 7.20
Summer Celtis australis L. 57.27
Trichosanthes wallichiana (Ser.) Wight. 10.00
Autumn Betula alnoides Buch.-Ham. ex D. Don 26.25
Celtis australis L. 16.67
Prunus cerasoides D. Don 14.58
Choerospondias axillaris (Roxb.) B.L. Burtt & A.W. Hill 12.50
Myrsine semiserrata Wall. 7.08
Maclura conchinchinensis (Lour.) Corner 5.00
Winter Pinus roxburghii Sargent 61.64
Schima wallichii (D.C) Korth. 23.84
1 Percentage of the plant feeding records with respect to total plant feeding records in the
season.
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and young leaves in the spring, mature leaves and insects in the summer, fruits in the
autumn and young cones of a coniferous plant, Pinus roxburghii , in the winter.
Effect of Seasonality on the Activity of Macaques
rMANOVA showed that the season had a significant impact on the activity bud-
gets of the macaques in both the WG (Pillai trace = 1.35, F(12, 33) = 2.25, p < 0.05)
and the SPG (Pillai trace = 1.31, F(12, 33) = 2.15, p < 0.05; Table5 ). For each activity
budget, ANOVA showed that the season affected the feeding (F(3, 12) = 4.63, p <
0.05) and movement (F(3, 12) = 6.44, p < 0.05) activity in the WG, while the effect of
season was not significant for either macaque activity in the SPG.
Relationship between Fruit Availability and Feeding Time
The proportion of time spent feeding was higher in the winter (0.66) and spring
(0.61) in the WG and in the autumn (0.42) in the SPG (df = 9, χ
2 = 256.0 in WG, χ
2 =
207.0 in SPG; p < 0.05; Table2 ). Fruit availability in the food plants of our study site
represented 1,430 cm
2 in spring, 3,003 cm
2 in summer, 7,820 cm
2 in autumn, and
6,993 cm
2 in winter. The fruit availability differed among the seasons (df = 3, H = 19.8,
p < 0.05). However, we found no significant correlation between fruit availability in
each season and the feeding time of the macaques ( ρ = 0.098, n = 8, z = 0.26, ns) or
fruit availability in each season and the proportion of time spent feeding on fruit
( ρ = 0.049, n = 8, z = 0.13, ns).
Discussion
In this study, we compared the activity budgets and diets of 2 social groups of
Assamese macaques in response to (1) provisioned food and (2) seasonality in order
to understand the influence of a human-modified habitat and seasonal changes in
habitat on behaviour. The results showed that the SPG consumed more human food
and that the WG consumed more plant species and fruit. Another result indicated
that the WG spent more time feeding and less time in social behaviour than the SPG
Group Season Proportion of time
feed ing moving resting social
WG Spring 0.61 0.13 0.12 0.14
Summer 0.48 0.21 0.17 0.15
Autumn 0.47 0.27 0.16 0.10
Winter 0.66 0.16 0.10 0.08
Average 0.55 0.19 0.14 0.12
SPG Spring 0.38 0.09 0.38 0.15
Summer 0.35 0.16 0.30 0.20
Autumn 0.42 0.14 0.24 0.20
Winter 0.35 0.17 0.30 0.18
Average 0.37 0.14 0.31 0.18
Table 5. Proportion of
activity times of WG and
SPG Assamese macaques in
each season
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70 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
did. The availability of human food might allow the SPG to spend less time feeding
and more time in social behaviour. Both the dietary compositions and time spent on
different activities varied between the 2 groups, supporting hypothesis 1. Hypothesis
2 was also supported by the finding that the percentage of fruit feeding was higher in
spring, summer, and autumn in the WG, and in spring and autumn in the SPG, and
by the finding that the WG spent more time in feeding in spring and winter. Assamese
macaques are flexible in their selection of food and activity budgets in relation to the
seasons.
Feeding was a major activity of the Assamese macaques in the Nagarjun forest.
Behavioural records showed that the proportions of behaviour spent feeding in As-
samese macaques in the WG and SPG were 0.55 and 0.37, respectively. Previous stud-
ies of wild Assamese macaques also showed that they spent most of their time feeding
and foraging: 40%, [Sarkar et al., 2012], 30% in males and 34% in females [Schulke et
al., 2011], and 43% [Chalise, 2010]. Despite the availability of garbage at the army
camp, the Assamese macaques in the SPG did not spend all of their feeding time eat-
ing human waste food. However, it is clear that the human food reduced the con-
sumption of natural food resources by these animals. As a result, overall feeding and
other activities of the group changed too. The dependency on human food and re-
duced feeding time might increase their time for other activities. However, the SPG
spent less time moving because they did not have to move or forage much to obtain
natural food. Because of the reductions in feeding and moving time, the SPG might
be able to allocate their time budget to resting and social activities. In contrast, the
WG had to move and forage over a large area to meet their food demand. Conse-
quently, these animals were less able to spend time resting or engaged in social be-
haviours. The same response to food availability was found in Barbary macaques in
Morocco [Alami et al., 2012] and in rhesus macaques in Bangladesh [Jaman and
Huffman, 2012].
Assamese macaques consumed a broad range of food items, but the major food
items were fruits for the WG and waste scraps for the SPG. Fruits accounted for 65.6%
of the diet of the WG. Although the SPG spent a considerable time (52.9%) eating
human food, they consumed many plant parts and other natural foods, so fruits ac-
counted for 22.5% of the total feeding records. For the WG and SPG, the consump-
tion of young leaves and mature leaves was only 3.5 and 7.6%, respectively, and 7.3
and 2%, respectively, but several studies in the highlands of Nepal, China, Bhutan,
and India concluded that Assamese macaques are primarily folivorous [Ahsan, 1994;
Srivastava, 1999; Chalise, 2003; Zhou et al . , 2011]. One recent study, by Chalise et al.
[2013a], in the Shivapuri forest of SNNP found that the primary food sources of As-
samese macaques between December and March were young and tender leaves, as
well as burgeoning twigs, followed by seeds, at 38.2, 35.3%, respectively. In this study,
we did not observe macaques in either group in early spring (March), when young
leaves are abundant, or during late spring, when some plants had fruits in the study
area. This could be a reason why macaques in this study apparently ate fewer young
leaves. However, the results of this study are like those of a study in Thailand [Schul-
ke et al., 2011] and a previous study in Nepal [Koirala and Chalise, 2014], in that the
diets of Assamese macaques comprised more fruits than leaves and flowers. Assamese
macaques ate more fruits when they were abundant [Zhou et al., 2011]. Similarly,
other macaques, such as crested black macaques ( M. nigra ) [O’Brien and Kinnaird,
1997] and Tonkean macaques ( M . tonkeana ) [Riley, 2007], are also frugivorous.
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A higher percentage of the diet of Assamese macaques was provided by tree spe-
cies [Zhou et al., 2011], consistent with the results of this study, which found that
more than 57% of food plants were trees. In relation to the high percentage of trees
as food plants, this study found that the Assamese macaques of SNNP were quite ar-
boreal. WG animals spent almost 99% of their time in trees, and this value was about
86% for SPG macaques. Wild Assamese macaques in Thailand did not use forest
strata uniformly; they were almost 90% arboreal. The SPG in this study spent about
5% less time in trees than macaques in Thailand, but the WG animals were more ar-
boreal than the Assamese macaques of Thailand. Unlike this study, Assamese ma-
caques in China spent more time on the ground, but the sympatric species, rhesus
macaques, used lower and middle tree canopies more frequently than Assamese ma-
caques [Zhou et al., 2014]. In this study site, rhesus macaques also inhabited the same
habitat. They seemed less arboreal than Assamese macaques, and they fed on those
plants on the floor that were not eaten by Assamese macaques [Koirala, unpubl. data].
Trees were the most common food resource, followed by shrubs and climbers. Only
few instances of herb species consumption were recorded in the study groups, con-
sistent with the arboreal habits of the species. Rhesus macaques of SNNP fed on 28
species of tree plants and 13 species of herbs [Bashyal, 2005], which is more than the
number of herb species consumed by Assamese macaques. Thus, Assamese and rhe-
sus macaques showed inter-species differences in their diets and habitat use, allowing
them to be sympatric. However, further investigation is needed to clarify the relation-
ships between them.
Common leopards, a predator of Assamese macaques, inhabit the Nagarjun for-
est. Security staff who stay in the Nagarjun forest have many records of direct encoun-
ters with leopards within the study area [Dambar Bista, pers. commun.]. During field
visits, B.P. recorded much evidence of macaques running up to the canopy, with
alarm calls, probably because a leopard was nearby [Koirala, unpubl. data]. When
domestic dogs at the army camp chased and threatened the macaques, they climbed
up into the trees, indicating that Assamese macaques have a defence mechanism
against predators. These facts suggest that Assamese macaques are arboreal to ensure
safety from predators and to avoid food competition with rhesus macaques. How-
ever, further studies are needed to examine the exact reasons behind their arboreal
behaviour.
The Assamese macaques in this study showed seasonal changes in their diets and
activity budgets. The macaques, especially in the WG, adapted to the availability of
natural food resources. However, we found no correlation between feeding times and
the availability of food in nature. Our estimates showed that fruit availability was
highest in winter, but unexpectedly, the WG animals spent the maximum time feed-
ing in the same season. This may be because Schima wallichii offers high fruit avail-
ability in winter and the fruits of this tree have a very small edible part. The macaques
need to break a hard pericarp to eat the small seed, resulting in them spending more
time in feeding and foraging activity in the winter.
Although the staff at the army camp did not usually hand food to the macaques,
the available waste influenced their behaviour and habituated them to the humans.
There is no cultivated land in the surrounding area, which is why there was no further
influence of provisioning. However, such provisioning outside protected areas may
possibly promote crop raiding and, in turn, human-macaque conflict. Additionally,
provisioning could transmit parasites between humans and macaques. Thus, we
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72 Koirala/Chalise/Katuwal/Gaire/Pandey/Ogawa
should be careful about provisioning for the conservation of Assamese macaques in
their natural and healthy habitat.
In conclusion, the dietary compositions and activity budgets of Assamese ma-
caques are affected by the availability of human food, which the Assamese macaques
in the SPG were highly dependent on. The macaques in the WG spent more time
feeding and moving compared with the macaques in the SPG. The macaques, espe-
cially in the WG, changed their diets and activity budgets seasonally and were highly
frugivorous, although they also ate a wide range of trees, herbs, shrubs and climbers,
as well as insects and their secretions. Assamese macaques in both groups were most-
ly arboreal.
Acknowledgements
We are very grateful to Primate Conservation Inc. USA and JSPS KAKENHI (grant No.
25440253 and 16K07539) for providing financial support for this research, and Idea Wild, USA,
for providing the required equipment. We are grateful to Lt. Col RC Khatry, commander of the
Bhairav Prasad battalion, Nagarjun, for granting permission and providing support for observa-
tions of macaques in the army camp area in the national park. We are indebted to and express
our sincere gratitude to Sujita Dhakal, Aruna Koirala, Yamuna Koirala, Chiring Lama, Chandra
Bahadur Budathoki, Kishor Tamang, Bhuban Paudyal, Purnaman Shrestha, Sabita Gurung, Sajan
Shrestha, and Bibash Shrestha, for their kind cooperation during the fieldwork. We also thank
Rita Chhetri from the National Herbarium and Plant Laboratories, Godawari, Lalitpur, Nepal,
for her kind help with the plant identification. We sincerely thank Buddi Sagar Poudel for his
guidance and much help during the manuscript preparation and data analysis.
Disclosure Statement
The authors declare that no conflicts of interest exist in relation to this work.
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