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Drastic population decline and conservation prospects of roadside dark-bellied bonnet macaques (Macaca radiata radiata) of southern India


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We carried out a survey on roadside dark-bellied bonnet macaques (Macaca radiata radiata) on the highways around the south Indian city of Mysore. The present survey was the fourth since 1989 on the same populations. We divided the habitats into intensive cultivation (IC), wet cultivation (WC), and scrub forests (SC). The number of groups has significantly reduced from 54 to 31 and the number of animals has declined from 1,207 to 697 from 1989 to 2009. This decline has been recorded only in the IC and WC areas, whereas the population in SC with places of Hindu worship has remained stable. Due to the loss of roadside Ficus trees over the years, the habitat of the monkeys has almost disappeared. Since bonnet macaque is not primarily a forest-dwelling species, the seemingly widespread primate may soon become 'threatened' if the non-forest populations continue to decline. Scrub forests in small hillocks housing Hindu temples remain the only prospective places for conservation of bonnet macaques.
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Drastic population decline and conservation prospects of roadside
dark-bellied bonnet macaques (Macaca radiata radiata)
of southern India
Mewa Singh Joseph J. Erinjery
Theethira S. Kavana Kuladeep Roy
Mridula Singh
Received: 26 July 2010 / Accepted: 16 January 2011 / Published online: 15 February 2011
ÓJapan Monkey Centre and Springer 2011
Abstract We carried out a survey on roadside dark-bel-
lied bonnet macaques (Macaca radiata radiata) on the
highways around the south Indian city of Mysore. The
present survey was the fourth since 1989 on the same
populations. We divided the habitats into intensive culti-
vation (IC), wet cultivation (WC), and scrub forests (SC).
The number of groups has significantly reduced from 54 to
31 and the number of animals has declined from 1,207 to
697 from 1989 to 2009. This decline has been recorded
only in the IC and WC areas, whereas the population in SC
with places of Hindu worship has remained stable. Due to
the loss of roadside Ficus trees over the years, the habitat of
the monkeys has almost disappeared. Since bonnet maca-
que is not primarily a forest-dwelling species, the seem-
ingly widespread primate may soon become ‘threatened’
if the non-forest populations continue to decline. Scrub
forests in small hillocks housing Hindu temples remain
the only prospective places for conservation of bonnet
Keywords Bonnet macaque Scrub forests
Conservation Hindu temples Population decline
Most of the primate populations have been declining dur-
ing the past few decades due to habitat loss, habitat frag-
mentation, and biotic pressures. As a result, a large number
of primate species are listed today as ‘threatened’ (IUCN
2010). Due to its diversity of habitat types, India, in
addition to a large number of other faunal species, is home
to 22 species of primates of which six are listed as
‘threatened’ and another six as ‘vulnerable’ (IUCN 2010).
Rhesus macaques, bonnet macaques, and Hanuman langurs
of the Indian subcontinent are considered ‘least concern’
due to their widespread distribution and large numbers.
Whereas the conservation biologists and wildlife managers
have paid serious attention to the conservation of ‘threa-
tened species’, little concern has been shown to the so-
called ‘least concern’ species. However, the numbers and
the distributional ranges of several such primate species in
India have drastically declined over the past few decades
(Kumara et al. 2009; Singh and Rao 2004; Southwick and
Siddiqi 1994a,b). It has been suggested that conservation
measures should be taken up for such species now before
these species also become ‘threatened’ (Kumara et al.
Periodic monitoring is the first step in assessing the
status of wild populations in order to devise the appropriate
conservation measures (Eudey 2008). Whereas several
surveys have been carried out on primate populations in
India, not a single large population has been monitored
over the long term. The bonnet macaque is an endemic
species to southern India ranging from the Godavari River
in the north to the peninsular tip of the south (Roonwal and
Mohnot 1977). The northern part of its range is inhabited
by the dark-bellied bonnet macaque (Macaca radiata
radiata) and the southern part of the range is inhabited by
M. Singh (&)J. J. Erinjery T. S. Kavana K. Roy
Biopsychology Laboratory, University of Mysore,
Mysore 570006, India
M. Singh
Maharaja’s College, University of Mysore,
Mysore 570006, India
Primates (2011) 52:149–154
DOI 10.1007/s10329-011-0234-x
the pale-bodied bonnet macaque (M. r. diluta) (Molur et al.
2003). Large populations of these two subspecies inhabit
human-inhabited landscapes and only a small portion of the
population inhabits forests and protected areas for wildlife
(Kumara et al. 2009). Over the past few decades, there has
been a rapid development in India in the areas of agricul-
tural expansion, concrete constructions, and widening of
highways. On one hand, these developmental activities
have drastically reduced the habitat of the bonnet maca-
ques, and on the other, people have increasingly become
intolerant of the macaques due to their crop-raiding
behavior. These changes are expected to influence the local
range and population size of the species.
For the past 20 years, we have been monitoring a pop-
ulation of dark-bellied bonnet macaques around the city of
Mysore within a radius of about 100 km. The previous
surveys were carried out in 1989 (D’Souza and Singh
1992), 1998 (Sharma 1998), 2003 (Singh and Rao 2004),
and the present survey was carried out in November to
December of 2009. In this article, we report the present
status of the bonnet macaque population in this region and
analyze the overall trends in this metapopulation and the
subpopulations. We also suggest possible conservation
measures for the non-forested bonnet macaques.
Study area
The study included eight highways around the south Indian
city of Mysore (12.18°N and 76.42°E), 11 adjacent roads
connecting the highways, and one scrub forest (SF) at
Chamundi Hill near the city (Fig. 1). We surveyed a total
of 1,052 km. The study area was divided into three habitat
Intensive cultivation
This is the region where the agricultural lands on roadsides
are irrigated through canals and wells and there is year-
round cultivation of sugarcane, vegetables, rice paddies,
cereals, etc. The dominant roadside vegetation included
sparse trees of Ficus bengalensis,F.religiosa,Tamarindus
indica, and Eucalyptus sp. The monkeys often raided crops
for food.
Wet cultivation
The agriculture in this habitat type was dependent on rains
during the monsoon months between June and October.
The cultivated crops included groundnut, cereals, and
beans. The roadside vegetation was sparse and included
Ficus bengalensis,F.religiosa,Tamarindus indica,Cocos
nucifera,Polyalthia longifolia, and Eucalyptus sp. In this
habitat as well, the monkeys often raided the crops for
Scrub forest
The Chamundi Hill range situated near the city of Mysore
consisted of thorny and SFs dominated by Acacia,Aza-
dirachta,Eucalyptus, and Zizyphus sp. This is a reserved
forest and there are no agricultural lands within the hill
range. A characteristic feature of the hill is the presence of
two places of Hindu worship, one at the bottom of the hill
and one at the top of the hill. These temples are regularly
visited by devotees, tourists, and monkeys, which, in
addition to the feeding in the forest, receive abundant
handouts of cooked food from visitors.
We carried out the study over a period of 2 months
between November and December of 2009. We traveled
the distance by foot and on a motorcycle/jeep at the speed
of \6 km/h. Since all the monkey groups were visible on
the roadsides, and the general location of each monkey
group has been known for the past two decades, we used
the method of ‘total counts’. The location map for these
groups was prepared in 1989 (D’Souza and Singh 1992)
Fig. 1 Map of the study area. Double lines show the roads that were
surveyed. 1Mysore, 2Antharsanthe, 3Begur, 4Handpost, 5Hangala,
6Nanjangud, 7Chamarajanagar, 8Yelandur, 9Kollegal,10 T. Narsipura,
11 Ramanagaram, 12 Kanakapura, 13 Periyapatna, 14 Anechaukur,
15 Chamundi Hills, 16 Gaddige, 17 Hunsur, 18 HD Kote, 19 Gundelpet,
20 HN Pura, 21 Channarayapatna, 22 Srirangapatna
150 Primates (2011) 52:149–154
and even during the present survey, we located a group
within 0.5 km of the 1989 location. If a group near an
earlier location was not sighted, a thorough search of the
surrounding area including the crop lands was made. Fur-
ther, the disappearance of the group was confirmed with the
villagers and the farmers in the area. After spotting a group,
we collected data on the number of animals in each age–
sex class. Due to the flat terrain, sparse trees, and clear
visibility in all habitat types, it was easy to see all the
animals. All the animals in a group of bonnet macaques
were mostly within a distance of about 50 m. Still, we
spent a few hours with each group and made repeated
counts to ensure that all animals were counted. The age–
sex classes included adult males (C5 years), adult females
(C3 years), sub-adult males (3–5 years), juvenile
(1–3 years), and infants (B1 year). Since the survey was
linear transects, we calculated the encounter rate as number
of groups/animals per kilometer. If the distance between
any two groups was more than 4 km, we considered these
groups as belonging to different subpopulations. We cal-
culated the occupied area as the total distance covered by a
subpopulation plus 4 km (adding 2 km on each side). We
calculated the intrinsic rate of natural increase (r)as
in which N
was the present number of animals,
was the number during the previous survey, e was the
base of natural logarithms, and twas the number of years
between two surveys. We compared the present data with
those from the previous surveys. All the previous surveys
were carried out employing the same methodology as in the
present survey as one of the authors (Mewa Singh) initiated
these surveys in 1989 and has since then been supervising
these studies with an active participation.
Overall population dynamics
Table 1presents the data on different sectors of survey,
habitat type, distance covered in each sector, and the
number of bonnet macaque groups and animals during the
four surveys. The number of groups has significantly
Table 1 Bonnet macaque populations in different transects in four different surveys
Sector Sector no.Habitat
covered (km)
Number of groups Number of animals
1989 1998 2003 2009
Mysore–Antharsanthe 1 WC 70 7 7 4 2 113 116 43 26
Handpost–Begur 2 WC 44 3 3 4 2 62 48 71 27
Mysore–Hangala 3 WC 75 11 10 10 5 192 157 170 132
Nanjangud–Chamarajanagar 4 WC 39 2 3 1 0 40 50 21 0
Chamarajanagar–Yelandur 5 WC 43 1 2 2 0 28 60 11 0
Kollegal–T. Narsipura 6 WC 67 9 8 6 7 257 169 125 102
Mysore–Ramanagaram 7 IC 91 7 8 8 1 133 103 168 34
Ramanagaram–Kanakapura 8 WC 28 0 1 1 2 0 12 16 19
Mysore–Kanakapura 9 WC 98 6 5 3 4 166 95 46 111
Mysore–Periyapatna 10 IC 42 1 1 0 0 6 6 0 0
Periyapatna–Nagarahole Road 11 IC 35 0 0 1 0 0 0 13 0
Chamundi Hills Road 12 SF 19 7 7 7 8 210 206 266 246
Kollegal Cross Road 13 WC 10 0 0 0 0 0 0 0 0
Mysore–Gaddige-Hunsur 14 IC 85 0 0 0 0 0 0 0 0
Gundelpet–Chamarajanagar 15 WC 34 0 0 0 0 0 0 0 0
Mysore–HN Pura 16 IC 92 0 0 0 0 0 0 0 0
HN Pura–Channarayapatna 17 IC 22 0 0 0 0 0 0 0 0
Mysore–Channarayapatna 18 IC 88 0 0 0 0 0 0 0 0
Srirangapatna–T. Narsipura 19 IC 35 0 0 0 0 0 0 0 0
T. Narsipura–Nanjangud 20 WC 35 0 0 0 0 0 0 0 0
Total 1,052 54 55 47 31 1,207 1,022 950 697
D’Souza and Singh (1992)
Sharma (1998)
Singh and Rao (2004)
Present study
Primates (2011) 52:149–154 151
reduced from 54 in 1989 to 31 in 2009 (v
=7.88, df =3,
p\0.05). Likewise, the number of animals has declined
from 1,207 in 1989 to 697 in 2009 (v
=138.08, df =3,
Population dynamics habitat-wise
The data on population status and dynamics during the four
surveys for each of the habitats are presented in Table 2.
Whereas the number of animals has significantly declined
in intensive cultivation (IC) (from 139 to 34) and wet
cultivation (WC) (from 858 to 417) (v
=96.61, df =3,
p\0.01 and v
=189.49, df =3, p\0.01, respectively),
the number has increased in SF (from 210 to 246)
=10.83, df =3, p\0.01). The present population in
WC is about 48.61% of the 1989 population (r=-0.07/
year). In IC, the present population is only 24.46% of the
1989 population (r=-0.04/year). The SF population has
recorded a growth with an r=0.01/year. Although in
2009, the mean group size in SF and WC was 30.75 and
18.95, respectively, the size did not differ significantly due
to large variations in group size in both habitats (t=1.93,
df =28, p=0.06). The most noticeable feature of groups
size is the significant decline in the number of groups in the
medium size range (11–30 animals) from 26 in 1989 to
only seven in 2009 in WC (v
=15.25, df =3, p\0.01)
as well as in the total population from 32 groups in 1989 to
12 groups in 2009 (v
=15.08, df =3, p\0.01). SF with
12.94 animal/km had the highest encounter rate followed
by WC with 0.76 animals/km and IC with 0.07 animals/
km. The area actually occupied by the macaques in the
survey region reduced from 25 to 4 km in IC, from 117 to
77.5 km in WC whereas it has remained the same in SF. In
IC, six subpopulations have now reduced to only one
subpopulation. In WC, 18 subpopulations in 1989 have
now become 17 subpopulations with only 48.61% of the
original population. The increased distance among the
subpopulations therefore indicates a high degree of popu-
lation fragmentation. All groups in SF have overlapping
home ranges making it a single subpopulation. Overall, the
number of animals per subpopulation has reduced from
48.3 in 1989 to 36.68 in 2009.
Age–sex ratios in habitat types
The data presented in Figs. 2,3, and 4show that the age–
sex ratios did not differ among the habitat types for adult
male:adult female (Kruskal–Wallis v
=1.19, df =2,
p=0.55), adult:immature (Kruskal–Wallis v
df =2, p=0.44) and adult female:immature (Kruskal–
Wallis v
=3.04, df =2, p=0.22). Although the pro-
portion of adult females recorded an increase during 1998
Table 2 Bonnet macaque populations in different habitat types during four surveys
Habitat type Year No. of
No. of
area (km)
Animals in
rOverall rNo. of sub-
(490 km)
1989 8 139 17.37 0.016 0.28 25 (5.10)
5.56 – 6 23.2
1998 10 121 12.10 0.020 0.25 31 (6.33) 3.90 -0.015 6 20.2
2003 9 181 20.11 0.018 0.37 29 (5.92) 6.24 0.080 6 30.2
2009 1 34 34.00 0.002 0.07 4 (0.81) 8.50 -0.280 -0.07 1 34.0
Wet cultivation
(543 km)
1989 39 858 22.00 0.072 1.58 117 (21.57) 7.33 18 47.7
1998 38 695 18.29 0.070 1.28 124 (22.84) 5.60 -0.023 21 33.1
2003 31 503 16.23 0.057 0.93 108 (19.89) 4.66 -0.065 24 21.0
2009 22 417 18.95 0.040 0.77 77.5 (14.27) 5.38 -0.030 -0.04 17 24.5
Scrub forest
(19 km)
1989 7 210 30.00 0.368 11.05 19 (100) 11.05 1 210.0
1998 7 206 29.43 0.368 10.84 19 (100) 10.84 -0.002 1 206.0
2003 7 266 38.00 0.368 14.00 19 (100) 14.00 0.051 1 266.0
2009 8 246 30.75 0.420 12.95 19 (100) 12.94 -0.010 0.01 1 246.0
(1,052 km)
1989 54 1,207 22.35 0.051 1.15 161 (15.34) 7.50 25 48.3
1998 55 1,022 18.58 0.052 0.97 174 (16.54) 5.87 -0.018 28 36.5
2003 47 950 20.21 0.045 0.90 156 (14.83) 6.09 -0.015 31 30.6
2009 31 697 22.49 0.029 0.66 100.5 (9.55) 6.93 -0.050 -0.03 19 36.7
km surveyed
Values in parentheses indicate percent area occupied
152 Primates (2011) 52:149–154
in all habitats and then steadily decreased, the difference
for the adult male:adult female ratio was not significant
among habitats over the surveys (Fig. 2) (Kruskal–Wallis
=6.18, df =3, p=0.10). The ratio of immature
individuals against adult females has declined over the
surveys with a sharper decline in IC, though the difference
among habitats for this ratio was non-significant (Fig. 3)
(Kruskal–Wallis v
=3.04, df =2, p=0.22). However, a
significant difference among surveys was observed for
adult:immature ratio (Fig. 4) (Kruskal–Wallis v
df =3, p=0.04) in which the immature ratio was 1:0.94,
1:0.71, 1:0.70, and 1:0.58 during 1989, 1998, 2003, and
2009 surveys, respectively. This indicated that the number
of immature individuals in relation to adults has been
continuously decreasing with a sharper decrease in IC.
The main results of the present study reveal that the pop-
ulation of roadside bonnet macaques, a so-called ‘least-
concern species’, has been drastically declining, and the
farmers whose crops are often raided by these monkeys are
not favorably disposed towards coexistence with the
monkeys. Except in the SF habitat, which also houses two
important places of Hindu worship in the Chamundi Hill,
the population of bonnet macaques has continually
declined over a period of 20 years as indicated by the four
surveys. Further, the results also showed that there has
been a continuous decline in the adult:immature ratio in the
groups from 1:0.94 in 1989 to only 1:0.58 in 2009. This
proportion of immature individuals is too small as com-
pared to the ratio of 1:1.14 for bonnet macaques in the
entire state of Karnataka, which includes several forest
habitats (Kumara et al. 2009). The total decline in the
population of bonnet macaques in the study area has been
due to the disappearance of several groups and not due to a
decrease in the group size. The results clearly indicate that
the bonnet macaques in their traditional habitats of road-
sides adjoining crop lands have no conservation future at
all. This is probably true of the other primates such as
rhesus macaques in northern India (Southwick and Siddiqi
During the past two decades or so, the Indian economy
has opened up as a more liberal free-market economy. This
has accelerated the pace of developmental activities that
includes the widening of highways. All the highways
included in our surveys had almost a continuous canopy of
banyan (Ficus bengalensis) trees some 30 years ago
(Mewa Singh, pers. obs.) and these fig trees provided an
important resource base for the monkeys and other arboreal
animals including squirrels and birds. A typical example in
the present study was that of Sector 7 (Mysore-Ramana-
garam Road), which links Mysore to the metropolitan city
of Bangalore. In the early 1960s, the entire stretch of
140 km between Mysore and Bangalore had an almost
continuous distribution of bonnet macaques (Paul Simonds,
personal information). In the year 2004, the government
started laying a four-lane highway on this road. Almost all
the roadside trees have been felled, which has left little
shelter or resource base for the monkeys. Similar road
widening has also been carried out in the other IC regions
since 2004. The last resort for the remaining monkeys in
these areas has been raiding crops for their food, which has
now become a daily affair. At the same time, the govern-
ment has provided more electricity to the farmers and the
once only monsoon fed crop lands are now irrigated by
electric bore wells. The cultivation, therefore, has become
more intensive and the farmers have increasingly become
intolerant of the crop-raiding monkeys. These farmers now
1989 1998 2003 200 9 1989 1998 2003 2009 1989 1998 2003 2009
Intensive cultivation Wet cultivation Scrub forest
Adul t ma le : Ad ult fem al e
Adult male Adult female
Fig. 2 Adult male:adult female ratio in different habitats over four
Adul t fem ale : I m matu re
Adult female Immature
1989 1998 2003 2009 1989 1998 2003 2009 1989 1998 2003 2009
Intensive cultivation Wet cultivation Scrub forest
Fig. 3 Adult female:immature ratio in different habitats over four
Adult : Immature
Adult Immature
1989 1998 2003 2009 1989 1998 2003 2009 1989 1998 2003 2009
Intensive cultivation Wet cultivation Scrub forest
Fig. 4 Adult:immature ratio in different habitats over four surveys
Primates (2011) 52:149–154 153
want the remaining monkeys trapped and released away
from their crops.
However, it may be noted that though the Indian farmer
has a negative attitude towards monkeys in a situation of
human–monkey conflict, he is not opposed to the conser-
vation of monkeys in areas where there is little such con-
flict. This is evident from the fact that the population in the
SFs of Chamundi Hill, which is also surrounded by crop
lands, has remained stable, and has even increased over the
years. We proposed earlier (Singh and Rao 2004) that since
India is dotted with small hillocks, many of which also
house Hindu temples, such places may be the only prospect
for conservation of non-forested primates. Sacred groves in
India and Africa have been observed to have a high degree
of protection and biodiversity (Bhagwat et al. 2005;
Mgumia and Oba 2003), and such places can also support
commensal primates. Since such species of primates have
very low densities in protected forests, a neglect of con-
servation of these non-forested populations may soon result
in these species also becoming threatened or undergoing
local extinction. This is probably true even for non-forest-
dwelling primate species in other countries such as
baboons and mangabeys in Africa (Hill 2000; Saj et al.
2001). In some regions of western Japan, the Japanese
macaques have already undergone local extinction
(Watanabe and Muroyama 2004). Muroyama and Eudey
(2004) advocate that theories and techniques that address
the needs of local people for relief from crop raiding by
macaques need to be developed for effective conservation
of such species.
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... Due to their proximity to humans, the bonnet monkey and the lion-tailed monkey groups are exposed to considerable human interferences (like teasing, chasing, etc.) and life-endangering hazards (like uncovered live wires, fast-moving traffic, etc.); deaths caused by the latter threats are not uncommon. These groups have also been part of long-term demographic and behavioral studies and hence, have been under regular monitoring (Cooper, Aureli, & Singh, 2004a, 2004b, 2007Kumara, Kumar, & Singh, 2010;Mangalam, Desai, & Singh, 2014Singh, Erinjery, Kavana, Roy, & Singh, 2011;Singh & Rao, 2004). ...
Existing models of attachment do not explain how death of offspring affects maternal behavior. Previous descriptions of maternal responsiveness to dead offspring in nonhuman anthropoids have not expounded the wide variation of deceased-infant carrying (DIC) behavior. Through the current study, we attempt to (a) identify determinants of DIC through a systematic survey across anthropoids, (b) quantitatively assess behavioral changes of mother during DIC, and (c) infer death perception of conspecifics. Firstly, we performed phylogenetic regression using duration of DIC as the dependent variable. Secondly, we undertook case studies of DIC in the bonnet monkey and the lion-tailed monkey through behavioral sampling. Results of phylogenetic Generalized Linear Mixed Model (Nspecies = 18; Ncases = 48) revealed a strong homology (H2 = 0.86). We also obtained a high intraspecific variation in DIC and found DIC to be affected by mother’s age, context of death, habitat condition, and degree of arboreality. We found bonnet mothers to carry their deceased offspring for 3.56 ± 4.03 SD days (N = 7) with diminished feeding, enhanced passivity, and social isolation during DIC and progressive decline in protection/attentiveness of corpse and attachment. Following Anderson (2016)’s framework of death perception, we interpreted repeated sensory investigation of corpses by mothers as comprehending causality of death, inanimate handling of corpse and its defense as comprehension of non-functionality, and a progressive disinterest of mothers in them as perceiving irreversibility of death. Lastly, we integrated DIC with mother-infant attachment theories and proposed a conceptual model characterizing DIC with causal determinants.
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Nonhuman individuals and groups, living in anthropogenic landscapes, often adopt adaptive foraging strategies, mediated by their day-today interactions with humans and their artefacts. Exploring such novel behavioral manifestations, especially in the Anthropocene, offers us insights into behavioral innovations and their transmission in such rapidly changing ecologies. In this study, employing field experiments, we investigated an example of human-induced, extractive foraging behavior-the extraction of liquid contents from plastic bottles-in a synurbic bonnet macaque Macaca radiata population. The main aims of the study were to examine the distribution, diversity, inter-individual variability and intra-individual flexibility of bottle-directed manipulative behaviors, and to explore the social and environmental factors driving this behavioral practice. We video-recorded the manipulation of partially filled plastic bottles and the extraction of liquid across four groups of bonnet macaques in southern India. Two socio-demographic factors-age class and group membership-and one environmental factor-food provisioning-were identified as major determinants of inter-individual variation in the performance of sophisticated manipulative techniques and in bottle-opening success. Our results also suggest that age-related physical maturation, experiential trial-and-error learning, and possibly social learning contributed to the acquisition of foraging competence in this task. These findings illuminate the mechanisms underlying inter-individual behavioral variability and intra-individual behavioral flexibility amongst free-ranging individuals of a cercopithecine primate species, traditionally known for its ecological adaptability and behavioral plasticity. Finally, this study documents how the presence of humans, their artefacts and their activities facilitate the development of certain behavioral traditions in free-ranging nonhuman populations, thus providing valuable insights into how human-alloprimate relations can be restructured within the increasingly resource-competitive environments of the Anthropocene.
Investigations into zoonotic disease outbreaks have been largely epidemiological and microbiological, with the primary focus being one of disease control and management. Increasingly though, the human–animal interface has proven to be an important driver for the acquisition and transmission of pathogens in humans, and this requires syncretic bio‐socio‐cultural enquiries into the origins of disease emergence, for more efficacious interventions. A biocultural lens is imperative for the examination of primate‐related zoonoses, for the human‐primate interface is broad and multitudinous, involving both physical and indirect interactions that occur due to shared spaces and ecologies. I use the case example of a viral zoonotic epidemic that is currently endemic to India, the Kysanaur Forest Disease, to show how biocultural anthropology provides a broad and integrative perspective into infectious disease ecology and presents new insights into the determinants of disease outbreaks. Drawing on insights from epidemiology, political ecology, primate behavioral ecology and ethnoprimatology, this paper demonstrates how human‐primate interactions and shared ecologies impact infectious disease spread between human and nonhuman primate groups. Bonnet macaque on a wire near Bengaluru city, India. Image Credit: Shaurabh Anand.
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A growing body of research focuses on how anthropogenic factors affect the behavior and ecology of primates and their ecosystems. Infrastructural development, such as roads, is an increasingly pervasive anthropogenic impact that destroys primate habitat, affects the distribution and dispersal of primates, and facilitates human–primate interactions. At our field site in Bantimurung-Bulusaraung National Park, Sulawesi, Indonesia, a major road bisects the habitat of the endangered moor macaque (Macaca maura). Beginning in 2015, we observed a behavioral shift by our main study group: they began spending more time along the road foraging in trash pits and waiting for provisions from vehicles. Our objective in this study was to examine how access to anthropogenic foods has affected the group’s ranging behavior by comparing ranging data collected before (2010–2011) and after the shift (2016–2017). In contrast to what we expected, home ranges were significantly larger and daily travel distance was significantly longer after the shift compared to before. As predicted, mean distance to the road decreased after the shift. These results likely reflect the irregular and spatially dispersed nature of provisioning at this site. The macaques appear to be attracted to the road because it presents opportunities to obtain palatable and energy-dense foods. Our results indicate that moor macaques are able to flexibly adjust their ranging behavior in response to anthropogenic impacts. However, given the risks of being in proximity to roads and humans, management of this emerging human–macaque interface is needed.
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Female reproductive success depends to a large extent on infants’ ability to survive to maturity. While most studies of female reproductive success have focused on the effects of individuals’ sociodemographic factors (e.g., age/parity, dominance rank) on offspring survival among wild primates living in less disturbed habitats, little research has focused on offspring survival in urban or periurban animals. Here we investigated sociodemographic and anthropogenic determinants of infant survival (up to 1 yr of age) in free-ranging bonnet macaques (Macaca radiata) living in a periurban environment in Southern India. We conducted the study from November 2016 to May 2018, on two groups of bonnet macaques at the Thenmala tourist site in the state of Kerala. Fifty infants were born across two birth seasons. Of these infants, 29.2% died or disappeared in 2017 and 26.9% died or disappeared in 2018. We found that infant survival was strongly influenced by the mother’s parity: infants of experienced mothers had a better chance of survival than those of first-time mothers. We also found that male infants were more likely to die than female infants. However, we found no effects of mothers’ dominance rank, or of frequency of mothers’ interactions with humans and time spent foraging on anthropogenic food, on infant survival. Our results, consistent with findings from other wild primate species, show that even in challenging human-impacted environments, experienced bonnet macaque mothers have greater success than inexperienced ones.
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With the uncontrolled expansion of anthropogenic modifications of the environment, wildlife species are forced to interact with humans, often leading to conflict situations that have detrimental effects for both wildlife and humans. Such interactions are escalating globally, making it crucial for us to devise strategies for both, the management of conflict and the conservation of these often-threatened species. We studied a case of potentially detrimental human-wildlife interactions between an endemic, habitat-specialist primate, the lion-tailed macaque Macaca silenus and resident human communities that has developed in recent years in the Western Ghats mountains of southern India. Primates provide useful model systems to understand the extent and nature of behavioural changes exhibited by wildlife in response to anthropogenic habitats with varying degrees of human influence. We documented behaviours, including foraging and intra-species social interactions, to examine the decisions made by the macaques as they exploited four human-modified habitats, which, for the purpose of this study, have been qualitatively characterised to include structural features of the habitat, type of food resources available and the presence of humans. Access to human-origin food, either cooked or packaged, acquired directly from homes or garbage pits, in the human-dominated habitat appeared to significantly reduce active foraging and searching for food, allowing them to engage in other behavioural activities, such as resting. Furthermore, patterns of reciprocated affiliation dissipated in certain human-dominated habitats, with individuals seeming to have adopted novel behavioural strategies, leading to altered social dynamics in the troop, possibly in response to provisioning. This study thus highlights the importance of understanding behavioural changes displayed by animals in response to human interactions; such knowledge could be crucial for the planning and implementation of management and conservation strategies for endangered species such as the lion-tailed macaque and possibly other wildlife in the increasingly anthropogenic landscapes of the tropical world.
Seed dispersal is an ecological process crucial for forest regeneration and recruitment. To date, most studies on frugivore seed dispersal have used the seed dispersal effectiveness framework and have documented seed-handling mechanisms, dispersal distances and the effect of seed handling on germination. In contrast, there has been no exploration of "disperser reliability" which is essential to determine if a frugivore is an effective disperser only in particular regions/years/seasons or across a range of spatio-temporal scales. In this paper, we propose a practical framework to assess the spatial reliability of frugivores as seed dispersers. We suggest that a frugivore genus would be a reliable disperser of certain plant families/genera if: (a) fruits of these plant families/genera are represented in the diets of most of the species of that frugivore, (b) these are consumed by the frugivore genus across different kinds of habitats, and (c) these fruits feature among the yearly staples and preferred fruits in the diets of the frugivore genus. Using this framework, we reviewed frugivory by the genus Macaca across Asia to assess its spatial reliability as seed dispersers. We found that the macaques dispersed the seeds of 11 plant families and five plant genera including at least 82 species across habitats. Differences in fruit consumption/preference between different groups of macaques were driven by variation in plant community composition across habitats. We posit that it is essential to maintain viable populations of macaques across their range and keep human interventions at a minimum to ensure that they continue to reliably disperse the seeds of a broad range of plant species in the Anthropocene. We further suggest that this framework be used for assessing the spatial reliability of other taxonomic groups as seed dispersers.
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Species conservation depends on the needs of the species concerned. For example, obligatory forest-dwelling and typically urban primate species require different conservation approaches. Here, I compare the ecology, life history, and behavior of two species of macaques in southern India; review what we know about the conservation challenges for each; and propose conservation measures. Lion-tailed macaques (Macaca silenus) are endemic to the rainforests of the Western Ghats and are endangered as a result of fragmentation of their habitats and low population growth. I recommend identification of contiguous forest habitats for viable lion-tailed macaque populations, improving the quality of their degraded habitats, and the linking of forest fragments with native vegetation as management measures for their conservation. In contrast, the bonnet macaque (M. radiata) is a habitat generalist endemic to southern India, but is losing its range to the relatively larger bodied and more aggressive rhesus macaque (M. mulatta). It is not a typically forest-dwelling species, and its populations are declining drastically in its traditional habitats that include Hindu temples, tourist spots, and roadsides with fruit-bearing trees and agricultural crops. The population has remained stable only where the habitat allows the macaques to forage in scrub jungle and includes a temple where the monkeys can obtain food from visitors. India is dotted with small hillocks with natural scrub or deciduous forest vegetation with one or two Hindu temples, and it appears that such habitats are the most suitable places for the long-term conservation of this species. These two case studies serve as examples of the different approaches needed to conserve forest-dwelling and urban primates, and with appropriate modifications, may inform the conservation of many other species.
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Primates inhabiting human-modified landscapes often exploit matrix habitat to supplement their diet with cultivated foods, at times resulting in economic losses and conflict with local people. Understanding human-nonhuman primate interactions and the attitudes and perceptions of local people towards crop feeding species are crucial to designing effective species-based management plans. Over a 12-month period, we used scan sampling to study the consumption of cultivated foods and matrix use patterns by two habituated groups of Bale monkeys (Chlorocebus djamdjamensis), Ethiopian-endemic bamboo specialists, in two forest fragments (Kokosa and Afursa) set amidst human settlements and farmland in the southern Ethiopian Highlands. Further, we conducted interviews with local people to document their attitudes and perceptions towards Bale monkeys at the two sites. We found that Bale monkeys at Kokosa, a more degraded habitat by most measures, consumed significantly more cultivated foods than their counterparts at Afursa. Moreover, Bale monkeys at Kokosa spent significantly more time in the matrix than in the forest habitat, while monkeys at Afursa spent significantly less time in the matrix than in the forest habitat. Not surprisingly, local people displayed a more negative attitude towards monkeys inhabiting Kokosa than those inhabiting Afursa. The differences in Bale monkey cultivated food consumption and matrix use patterns-as well as in local people's attitudes and perceptions towards Bale monkeys-between Kokosa and Afursa are probably associated with differences in habitat structure, degree of habitat alteration, and land-use practices between the sites. We conclude that to ensure long-term coexistence between Bale monkeys and local people in human-modified landscapes, it is vital to incorporate nearby matrix habitats into management plans and to work closely with local communities to develop effective nonlethal crop protection strategies, thereby reducing the likelihood of negative interactions between Bale monkeys and humans.
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Crop foraging or crop raiding concerns wildlife foraging and farmers' reactions and responses to it. To understand crop foraging and its value to wildlife or its implications for humans requires a cross-disciplinary approach that considers the behavior and ecology of wild animals engaging in this behavior; the types and levels of competition for resources between people and wildlife; people's perceptions of and attitudes toward wildlife, including animals that forage on crops; and discourse about animals and their behaviors and how these discourses can be used for expressing dissent and distress about other social conflicts. So, to understand and respond to conflicts about crop damage, we need to look beyond what people lose, i.e., crop loss and economic equivalence, and focusmore on what people say about wildlife and why they say it.
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Much has been written about insect damage to standing crops, but an area that has received little attention within agricultural development, conserva-tion, and primatological literature is that of primates and the potential damage they can cause to farmers' fields. This is likely to become an increasingly important issue for people interested in primates, as conservation projects adopt a more integrated approach to take account of local people's perspec-tives and needs. The aim of this paper is to examine the impact of crop raiding by primates, particularly baboons, on farmers living around the southern edge of the Budongo Forest Reserve, Uganda. I use data gathered during monthly farm surveys and informal discussion groups, along with time budget data, to demonstrate that 1) baboons can cause extensive damage to field crops, such as maize and cassava; 2) proximity of the farm to the forest edge and the presence or absence of neighboring farms affect the likelihood of any farm sustaining crop damage from baboons; and 3) in addition to the direct costs associated with crop losses attributed to baboon foraging activity, there are indirect costs of baboon crop raiding such as increased labor demands to protect crops from them and, occasionally, to replant crop stands badly damaged by baboons. These results have important implications for future primate conservation policy and practice.
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Although sacred groves arc, important for conservation in India, The landscape that surrounds them has a vital influence on biodiversity within them. Research has focused on tree diversity inside these forest patches. In a coffee-growing region of the Western Ghats, however landscape outside sacred groves is also tree covered because Planters have retained native trees to provide shade for coffee plants. We examined the diversity of trees, birds, and macrofungi at 58 sites-10 forest-reserve sites, 25 sacred groves, and 23 coffee plantations-in Kodagu district. We measured landscape composition and configuration around each site with a geographic information system. To identify factors associated with diversity we constructed multivariate models by using a decision-tree technique. The conventional measures of landscape such as patch size did not influence species richness. Distance of sacred groves front The forest reserve bad a weak influence. The measures of landscape structure (e.g., tree cover in the surroundings) and stand structure (e.g., variability in canopy height) contributed to The variation in species richness explained by multivariate models. We suggest that biodiversity present within sacred groves bets been influenced by native tree cover in the surrounding landscape. To conserve this biodiversity The integrity of the tree-covered landscape matrix will need to be conserved.
Population surveys of rhesus monkeys in north central India showed 88 % of the total population sampled in commensal or semi-commensal habitats in 1959-60, and 86 % in 1990-91. In the latter surveys, there was a significant shift from semi-commensal into commensal, and a minor increase in forest-dwelling, non-commensal populations. Over the 30-year span, group sizes increased in five habitat categories, whereas they declined in village and urban habitats despite a general increase in these populations. Rhesus populations in India are increasing again after substantial decline, and are adapting to human population pressure by increased commensalism.
Over 50 species of nonhuman primates live in 20 nations in Asia, interacting ecologically with over 3 billion people. The population status of different species of primates ranges from rare and endangered to abundant and relatively secure. The distribution, numbers, and trends of most species are not known, even for those such as Macaca fascicularis which are extensively used in biomedical research. Field surveys of rhesus populations since 1959 showed a serious 90% decline in population numbers in northern India throughout the 1960s and 1970s, followed by a partial recovery throughout the 1980s. Factors probably responsible for this recovery include a ban on trapping for export, increased agricultural production and economic development, increased conservation efforts, and perhaps a renewal of respect and tolerance for wildlife by the people of India. The history of rhesus populations in India can provide guidelines for conservation and management programs for other species. © 1994 Wiley-Liss, Inc.
The crab-eating or long-tailed macaque (Macaca fascicularis) of tropical Southeast Asia is a widespread but rapidly declining species. The threats to the species are manifold and include habitat loss and degradation that increasingly result in conflict with expanding human populations in both rural and urban landscapes, as well as trapping and trade for pharmaceutical testing, research, and development. The greatest threat from the trade is in the Indochinese region, especially Cambodia where in 2003–2004 macaques began to be harvested from the wild, ostensibly for captive breeding for export to China and to the USA and elsewhere. The lucrative operations, however, may serve to “launder” wild-caught monkeys and appear to have resulted in their disappearance even from legally protected areas. Much of the impetus for this trade appears to be biowarfare research in the USA, the country that is the world's largest user of primates. Macaca fascicularis is classified as of “Least Concern” in the IUCN/SSC 2008 Red List of Threatened Species. It is imperative that the conservation status of the species be reassessed and that the impact of trade on the species be assessed by the CITES Secretariat.
Sacred groves, some as old as several centuries, may have contributed to the preservation of some plant species in Tanzania. This has led conservationists to consider whether sacred groves could be used for in situ conservation. Eight sacred groves of the Ugunda chieftaincy of the Wanyamwezi in central Tanzania representing burial sites that varied from 6–300 years old were inventoried to compare woody species richness and taxonomic diversity with those of forest plots in a state managed Forest Reserve. Although they occupied a relatively small area the sacred groves had greater woody species richness and taxonomic diversity than the state managed Forest Reserve. The forest plots and the groves shared a species similarity index of 45%, suggesting that the sacred groves contributed to in situ conservation of the miombo woodland biodiversity. Some of the woody species were absent in the forest plots, also suggesting that groves served as a refuge for some species. An inventory of all existing sacred groves could provide important information on their role in in situ conservation. In order to promote them for community-based conservation of biodiversity the government should declare sacred groves as preservation sites, and incorporate them into modern conservation systems.
Forty-seven property owners in Entebbe, Uganda were questioned about vervet monkey activities on their property. Our main objective was to investigate the interactions between humans and vervet monkeys in an agricultural area adjacent to a forest zone. Other studies have reported that farms located within 300 m of a forested boundary probably incur the greatest risk of crop-raiding. Two other factors that may influence susceptibility to vervet crop-raiding were also examined: the types of crops grown and the types of direct preventative measures used. The effect of these two factors on vervet crop-raiding is not straightforward. However, the distance a property is located from the forest edge is an important factor influencing vervet crop-raiding. Surveyed gardens 200 m from the forest edge received significantly less crop-raiding than farms located 100 m or 50 m (P = 0.040, < α = 0.05).We suggest that the development of nonagricultural activities on land directly adjacent to forested areas may reduce vervet crop-raiding by deterring vervets from travelling greater distances from the forest edge due to increased obstacles or risks.
Whereas much attention is paid to the conservation and management of threatened species of primates, little work is reported on the species that are at lower risk. We report data on demography and population dynamics in commensal bonnet macaques (Macaca radiata radiata) sharing human habitats in Intensive Cultivation, Dry Cultivation and Scrub Forest zones around the city of Mysore, south India. We also compare the data with the previous data collected in 1989 and 1998 on the same groups by our research team. The population has suffered an overall decline of 21.3% (at an intrinsic rate (r) of –0.017) with Dry Cultivation habitat alone recording over 41% decline (r=–0.038) in 14 years. The Dry Cultivation habitat has also shown increased population fragmentation due to increased distance among groups, and decreased numbers of individuals per subpopulation. In Dry Cultivation areas, the number of small groups has been increasing, whereas the number of medium and large groups has been decreasing. We discuss such trends in the context of human-monkey conflict for shared resources. We propose that effective conservation strategies for lower risk species, especially if they are by-and-large commensal, must be planned now before they become threatened. We further propose that Scrub Forests with places of worship, and with the least human-monkey conflict but otherwise close to intensively used human habitats, are the most suitable and stable habitats for conservation of bonnet macaques.