Chapter 3 Ecological Correlates of Diurnal Primates of the Western Ghats

INTRODUCTION The Western Ghats, a series of hills ranges, run parallel to the western coast of southwestern India from 21ºN to 8ºN (Pascal, 1988). Being close to the Arabian Sea, these hill ranges receive heavy southwest monsoons. Whereas the western slopes and the ridges contain primarily tropical evergreen rainforests, the eastern slopes, which are rain shadow, contain deciduous and scrub forests. The Ghats with a length of about 1600 kms from north to south and with an east to west width of 30-80 kms are a mosaic of heterogeneous habitat types. Although the human activity has been present in these hills since prehistoric times, the organized state sponsored forestry and non-forestry activities began about 200 years back (Chandran, 1997). Commercial plantations, tea and coffee cultivation, building of hydel dams and power generation etc., brought millions of people to these hills. As a result, the habitats, especially the regions under rainforests, have undergone drastic changes. The most conspicuous effect in these hills has been the fragmentation of rainforests. In most places, the rainforests fragments now stand isolated because of the gaps created by plantations of no-rainforest species and commercial crops. Despite such factors, the Western Ghats still harbor a diversity of plant and animal life because of which, the hills have been recognized as one of eight ‘hottest hotspots’ of biodiversity in the world (Myers et al., 2000). Due to the canopy contiguity and the availability of a variety of trees bearing fruits throughout the year, the rainforests of the Western Ghats are extremely rich in arboreal fauna. The most conspicuous of the arboreal mammals are four species of diurnal non-human primates including lion-tailed macaque (Macaca silenus), Nilgiri langur (Semnopithecus johnii), bonnet macaque (M. radiata) and Hanuman langur (S. entellus). Whereas lion-tailed macaque and Nilgiri langur are restricted to the Western Ghats, the other two species are widespread throughout southern India. The effect of rainforest fragmentation and habitat deterioration has been most drastic on lion-tailed macaque. Because of its highly selective feeding habits, limited range of occupancy (about 2500 km2), delayed sexual maturity, long inter birth interval, low population turnover and a small remaining wild population (about 3500 individuals), this species has been classified as endangered (IUCN, 2003). Further, the remaining population consists of 49 subpopulations isolated in rainforest fragments in 8 locations (Molur et al., 2003). Since forest fragments are also under intense human pressure, significant effects have been observed on demography, ranging patterns, feeding habits and reproductive rates of lion-tailed macaques (Kumar et al., 1995, Singh et al., 2001). Whereas quite a few studies are available on lion-tailed macaques, the information on the other species of primates in the Western Ghats is scanty (Sugiyama, 1968; Green and Minkowski, 1977; Kurup, 1978; Bhat, 1982; Karanth, 1985; Kumar et al., 1995; Singh et al., 1997a,b, 2000, 2002; Ramachandran and Joseph, 2001). A study of the distribution of primates in the Western Ghats has attained a special significance recently due to updating of primate taxonomy (Groves, 2001). Whereas lion-tailed macaque and Nilgiri langur are considered monotypic, and two subspecies of bonnet macaque (M. radiata radiata and M. radiata diluta) have already been described, confusion still prevails regarding Hanuman langur. Several subspecies of Hanuman langur present in the Western Ghats are now being classified as distinct species (Groves, 2001). However, hardly any field data are available on the distribution and status etc. from actual populations today and the classifications have been made largely on the basis of sample specimens collected decades back. It, therefore, becomes necessary not only to survey the primate populations in the Western Ghats afresh, but also to determine the ecological correlates of the distribution patterns and to assess the impact of anthropogenic factors on the remaining primate populations.

The present study focuses on the status and distribution of primates in the rainforests of Brahmagiri-Makut and Sirsi-Honnavara in the state of Karnataka. These areas were selected because very little information is available on primates in the rainforests of the Western Ghats in Karnataka, and these two localities represent two distinct ecozones. The two regions also differ in terms of anthropogenic factors. An additional interest in this study was to find out if any viable localities for the conservation of endangered lion-tailed macaque exist in the rainforests of Karnataka. STUDY AREA The study was carried out in two geographical regions of the Western Ghats including Brahmagiri-Makut, the southern end, and Sirsi-Honnavara, a northern region, of the Western Ghats respectively in the state of Karnataka (Fig. 3.1). Table 3.1 presents information on the locations, forest status and physical features of the two study areas.

Fig. 3.1. The Western Ghats and two study locations

The forests of Brahmagiri-Makut lie between 75º50’-76º3’E and 12º5’-12º13’N with an area of 324 km2 in Kodagu district (Fig. 3.2). The area receives both Southwest and Northeast monsoons with an average annual rainfall of about 6000 mm. The region includes three administrative forest ranges viz. Srimangala and Makut (Wildlife), which form Brahmagiri Wildlife Sanctuary, and

Makut (Reserve Forest). On the west of these hills is the state of Kerala with agricultural villages, on the east are coffee and cardamom plantations, on the north are wet evergreen forests and grassy mountain slopes, and on the south is a continuation of moist deciduous forests of the Rajiv Gandhi National Park. The forests of Brahmagiri-Makut are broadly classified as Medium Altitude Forest Type with Mesua ferrea-Palaquium ellipticum type of vegetation (Pascal, 1988). Cullenia exarillata, a species with a special importance for primates and found in abundance southwards in the Western Ghats, is absent in these forests. Table 3.1. Physical Features of the Two Study Regions Geographical location

Forest Division

Forest Range

Forest status

Altitude range (m asl)

Rainfall (mm)

Forest type

Srimangala PA 800-1608 6000 Grass hills, Shola, Evergreen and Moist-deciduous

Makut (WL)

PA 65-1110 “ Evergreen and Moist-deciduous

Brahmagiri-Makut 750 50’-760 3’E 1205’-12013’N

Virajpet

Makut (RF) RF 60-1744 “ Grass hills, Evergreen and Moist-deciduous

Janmane RF 275-540 5000 Evergreen and Moist-deciduous

Kyadagi RF 424-700 “ Evergreen and Moist-deciduous

Sirsi

Siddapura RF 401-542 “ Evergreen and Moist-deciduous

Kumta RF 134-545 “ Degraded rain forest and Moist-deciduous

Honnavara RF 98-470 “ Evergreen, Moist-deciduous and Degraded rain forest

Sirsi-Honnavara 74035’-74047’E 14015’-14025’N

Honnavara

Gersoppa RF 172-615 “ Evergreen, Moist-deciduous and Degraded rain forest

WL – Wildlife; PA – Protected Area; RF – Reserve Forest The forests of Sirsi-Honnavara lie between 74035’-74047’E and 14015’-14025’N in the district of Uttara Kannada. The selected study area with about 200 km2 of rainforest was between Sharavathy and Aghanashini rivers (Fig. 3.3). The official status of the forest is Reserve Forest with interspersed revenue lands. A number of villages with large areas under cultivation of commercial crops including areca nut (Areca catachu) and paddy (Oryza sativa) are scattered inside. The forest has been broadly classified as Low Altitude Rainforest by Pascal (1988) and as ‘West Coast Tropical Evergreen Forest’ by Champion and Seth (1968). Due to the human interference, the Dipterocarpus indicus-Diospyros condolleana-Diospyros oocarpa vegetation type found southwards has been replaced by Persea macrantha-Diospyros spp- Holigarna spp vegetation type (Pascal, 1988). It extends up to the plateau of Aghanashini river and forms the northern limit of the evergreen forests of the plains and low elevations. METHODS The study was conducted from November 2001 to January 2002 in Brahmagiri-Makut, and from February 2002 to April 2002 in Sirsi-Honnavara. I followed the methods of ‘total count’ for lion-tailed macaque and ‘encounter rate’ (NRC, 1981; Whitesides et al., 1988) to estimate the

distribution and abundance of other primate species. These methods have been reliably used in several earlier primate studies in the Western Ghats including Sringeri Forest Range (Singh et al., 2000), Silent Valley National Park (Ramachandran and Joseph, 2001) and Anaimalai hills (Singh et al., 1997a,b, 2002). The details of the effort employed are provided in Chapter 2. Since both study areas harbored tropical rainforests, it was assumed that neither the ‘visibility factor’ nor the ‘detectability factor’ would affect observations or bias the data to any significant extent. After sighting a group, sufficient time was spent to get the proper count of individuals. Individuals were classified as adult male, adult female and immature (including sub-adults, juveniles and infants), based on morphological characteristics (Roonwal and Mohnot, 1977; Kumar 1987; Harvey et al., 1991; Ramachandran and Joseph, 2001; Singh et al., 2002). The location and the demographic data allowed the groups to be differentiated and given distinct names for lion-tailed macaques. The abundance of a species is represented as encounter rate i.e. groups/km. The information on hunting practice and intensity was gathered through interviewing local people, forest officials, hunters, encounters with hunters, animal traps found in the forest etc. (Chapter 2). RESULTS All four species of diurnal primates of South India viz., bonnet macaque, lion-tailed macaque, Nilgiri langur and Hanuman langur, were found to occur in Brahmagiri-Makut. In Sirsi-Honnavara, Nilgiri langur was absent and the other three species were found to occur. Table 3.2 Distribution of Primates in Brahmagiri –Makut Species Location Forest type Altitude

range (m asl)

BM Kuppangal (1), Irpu (2), Evergreen forest, Moist deciduous

811-1308

BM, HL Naathangal (3), Makut (16) Evergreen forest, Moist deciduous

60-655

BM, LTM (?), NL

Sempole (7), Perumalumalai to Aabylu (8) Evergreen forest 755-1170

BM, LTM (?), HL

South of Aabylu (9), Mepalli (10), Nadakani (11), Pookala (12), Votekolli (13), Sollekolli (14), Kottarchipari (15), Arabithittu (17), Kulimakki (18), Matre (19), Kooratu (21)

Evergreen forest 60 -1744

BM, LTM, HL Udumbe (20) Evergreen forest 200-600 NL Munikal shola (4), Dharekadu (5) Evergreen forest, Montane

shola 1305-1400

None Sholas around Narimalai (6) Montane shola 1300-1400 BM-Bonnet Macaque, LTM-Lion tailed Macaque, NL-Nilgiri Langur, HL-Hanuman Langur, Numbers in parentheses indicate locations in Fig. 2. LTM(?) – Possible occurrence of lion-tailed macaque Table 3.2 and Fig. 3.2 present the distribution of primates in Brahmagiri-Makut. On the eastern slopes of the southern Srimangala Range where the forest was moist deciduous, only bonnet macaque was found to occur. At the southeastern tip of this Range with small patches of forests and coffee plantations, Hanuman langur and bonnet macaques were sympatric. At the higher elevation with thin shola* forests around Narimalai, all primate species were absent. However, in * A shola is a narrow patch of rainforests in hollows amidst grasslands at mountain tops in the Western Ghats

the similar type of sholas at Munikal and Dharekadu towards west, Nilgiri langur was the only primate species present. This population was found to be the northernmost population of Nilgiri langur in its distributional range in the entire Western Ghats. In the rainforests that continued towards northwest along the Kerala border at Sempole and Perumalumalai to Aabylu, Bonnet macaque and Nilgiri langur were sympatric. In the rainforest at northern Aabylu, Mepalli, Nadakani and Pookala of Srimangala Range, Votekolli, Sollekolli, Kottarchiparai of Makut (Wildlife) Range and Arabithittu, Kulimakki, Matre and Kooratu of Makut (Reserve Forest) Range, bonnet macaque and Hanuman langur were found to occur as sympatric species. Along with bonnet macaque and Hanuman langur, the lion-tailed macaque was sympatric at Udumbe of Makut (Reserve Forest) Range. Further details of the distribution of Hanuman langur are given in Fig. 3.2. Except one group each at Nathangal, Udumbe, Puliparai and Kulimakki, which were isolated stray groups, rest of the groups were located between Votekolli and Makut.

Fig. 3.2. Three administrative Ranges of Brahmagiri-Makut. Numbers indicate places mentioned in Table 3.2. RF – Reserve Forest; WL - Wildlife In Sirsi-Honnavara, north of Sharavathi river to Chiksuli in Kyadagi Range, bonnet macaque, lion-tailed macaque and Hanuman langur were found to be sympatric. In rest of the region, bonnet macaque and Hanuman langur were sympatric. Since lion-tailed macaque is an endangered species, a special attempt was made to locate all groups of this species. Appendix 3.1 presents the information on the distribution of the lion-tailed macaque in Sirsi-Honnavara. Each group was assigned an identity on the basis of location (Fig. 3.3) and demographic structure. In total, 32 groups of lion-tailed macaques were recorded. This population of lion-tailed macaque was found to be the northernmost population of its distributional range in the entire Western Ghats.

Table 3.3 Primate Groups at Different Altitudes in Brahmagiri-Makut and Sirsi-Honnavara

Brahmagiri-Makut Sirsi-Honnavara Altitude range (m asl)

BM HL NL Overall(%)

LTM BM HL Overall (%)

<200 3(17.65) 2(10.00) - (9.22) 1(3.13) 1(2.13) 5(3.31) (2.86) 201-400 7(41.18) 10(50.00) - (30.39) 1(3.13) 5(10.64) 19(12.58) (8.78) 401-500 - 3(15.00) - (5.00) 10(31.25) 24(51.06) 72(47.68) (43.33) 501-600 1(5.88) 4(20.00) - (8.63) 15(46.87) 15(31.91) 43(28.48) (35.75) 601-700 2(11.76) - - (3.92) 5(15.62) 2(4.26) 12(7.95) (9.28) 701-800 4(23.53) 1(5.00) 1(14.29) (14.27) - - - - 801-900 - - 1(14.29) (4.76) - - - - >900 - - 5(71.42) (23.81) - - - - Values in parentheses indicate percentages Table 3.3 presents information on distribution of primates at different altitudes. In Brahmagiri-Makut, the Nilgiri langur was found between 700-1400 m asl, though 71.41% of the groups were above 900m asl. On the other hand, 50% of the Hanuman langur and 41% of the bonnet macaque groups were found between 200-400 m asl. Since only one group of lion-tailed macaque was recorded at 430 m asl in this region, it is not listed in Table 3.3. In Sirsi-Honnavara, primate species were found between 150-700 m asl, though 79% of the groups of all species were found between 400-600 m asl.

Fig. 3.3. Locations of lion-tailed macaque groups in Sirsi-Honnavara. See Appendix 3.1. Table 3.4 presents the data on abundance of primates in the two study regions. Bonnet macaque and Hanuman langur were encountered in all the ranges. Nilgiri langur was encountered only in Srimangala. Lion-tailed macaque was encountered in Makut (RF) in Brahmagiri-Makut and in all the ranges except Janmane and Kumta in Sirsi-Honnavara. A total of 15 groups of Bonnet

Table 3.4 Primate Groups Encountered in Different Ranges of Brahmagiri- Makut and Sirsi- Honnavara

Bonnet macaque

Lion-tailed macaque

Nilgiri langur Hanuman langur

Study region

Range Km walked

No Groups/Km No Groups/Km No Groups/Km No Groups/Km Srimangala 80.2 3 0.037 0 - 9 0.112 1 0.012 Makut (WL)

38.5 5 0.129 0 - 0 - 14 0.363

Makut (RF)

119.3 7 0.058 1 0.008 0 - 6 0.050

Brahmagiri- Makut

Total 238 15 0.06 1 0.004 9 0.038 21 0.088 Janmane 20.3 5 0.25 0 - 0 - 22 1.10 Kyadagi 95.4 27 0.28 17 0.18 0 - 87 0.91 Siddapura 8.0 2 0.25 2 0.25 0 - 5 0.62 Kumta 6.5 2 0.31 0 - 0 - 8 1.23 Honnavara 19.4 5 0.25 3 0.15 0 - 22 1.13 Gersoppa 48.4 14 0.29 10 0.21 0 - 29 0.60

Sirsi- Honnavara

Total 198 55 0.28 32 0.16 0 - 173 0.87 macaque encountered provided an encounter rate of 0.06 groups/km. The encounter rate of bonnet macaque in Srimangala and Makut (RF) was 0.037 and 0.058 respectively, whereas it was higher in Makut (WL) (0.129groups/km). The encounter rate of Hanuman langur in Srimangala and Makut (RF) was 0.012 and 0.05 groups/km respectively, whereas it was considerably higher (0.363 groups/km) in Makut (WL). The overall abundance of bonnet macaque (0.28 groups/km) and Hanuman langur (0.87 groups/km) in Sirsi-Honnavara was higher than 0.06 and 0.088 groups/km respectively in Brahmagiri-Makut. Nine groups of Nilgiri langur occurred with an encounter rate of 0.038 groups/km only in evergreen forests bordering Kerala state and in the southwestern part of the Srimangala range. The lion-tailed macaque, with only 3 animals in a group, was encountered in Makut (RF) in Brahmagiri-Makut. In Kyadagi, Siddapura, Honnavara and Gersoppa ranges of Sirsi-Honnavara, 32 groups of lion-tailed macaque were recorded, providing an encounter rate of 0.16 groups/km. The encounter rates were almost even across all the ranges. In Sirsi-Honnavara, the abundance of Hanuman langur (0.87 groups/km) was higher than that of bonnet macaque (0.28 groups/km) and lion-tailed macaque (0.16groups/km). Table 3.5 presents information on group size of different primate species. I was not able to gather data on group size in lion-tailed macaque and bonnet macaque in Brahmagiri-Makut since they were very shy. The group size of lion-tailed macaque and bonnet macaque was 24.7 (N=20) (range 9-48) and 19.0 (N=5) (range 5-36) respectively in Sirsi-Honnavara. The group size of Nilgiri langur in Brahmagiri-Makut was 4.25 (N=4) (range 3-6). The group size of Hanuman langur was 6.5 (N=6) (range 3-9). The group size of Hanuman langur in Sirsi-Honnavara was 10.31 (N=13) (range 5-16). Extrapolating to 32 groups of lion-tailed macaques, the total population in Sirsi-Honnavara was estimated to be about 790 individuals. The group density and the animal density were 0.28 groups/km2 and 6.87 animals/ km2 respectively. The number of groups in each group size category of 11 to 20, 21 to 30 and 31 to 40 was 6, 6 and 5 respectively. Only two groups had less than 10 individuals and one group had more than 40 individuals. The age/sex composition was available only in nine groups (Table 3.6). The number of adult males per group varied from one to four, with a mean of 1.89 and median of 2. More than 50% of the groups had two adult males. Number of males in the group increased as the group size increased. Adult males constituted 8.86% of a group. The number of adult females per group varied from 3 to15, with a mean of 7.89 and median of 9. The adult females constituted 36.98% of a group. The

Table 3.5 Group Size in Different Species Brahmagiri-Makut Sirsi-Honnavara Animal No. of groups

No. of animals

Mean group size

No. of groups

No. of animals

Mean group size

BM - - - 5 95 19.0 LTM - - - 20 494 24.7 HL 6 39 6.5 13 134 10.31 NL 4 17 4.25 - - - mean adult sex ratio was 4.18 females per male. The immature class (sub adult, juvenile and infant) constituted 54.16% of a group, whereas infants alone constituted 17.71%. The adult female to immature ratio was 1:1.46 and the adult female to infant ratio was 1:0.48. The overall adult to immature ratio was 1:1.18. Table 3.6 Demography of Lion-tailed Macaque

Group Size Adult male

Adult female

Subadult and juvenile

Infant Total immature

Chiksuli 30 2 9 11 8 19 Kerekulivotekodlu 24 2 9 7 6 13 Hosathota 11 1 4 2 4 6 Samplevore-A 9 1 4 3 1 4 Salikanu 36 4 15 14 3 17 Hasolli 27 2 11 8 6 14 Kanukumbri 9 1 3 4 1 5 Suthlamane 31 2 10 16 3 19 Hukkali 15 2 6 5 2 7 Total 192 17 71 70 34 104 Mean 1.89 7.89 7.78 3.78 11.56 % of group 8.86 36.98 36.46 17.71 54.16

In Sirsi-Honnavara, villages were scattered throughout the rainforest and large tracts were cleared for planting commercial crops such as paddy and areca. However, the forest was still contiguous through hills slopes and valleys. Present information revealed that hunting pressure on primates was severe in Brahmagiri-Makut due to the preference for primate meat in almost all communities. On the other hand, the belief systems in Sirsi-Honnavara prevented most people from hunting primates for meat. The detailed data on interviews, trap sighting, encounters with hunters, hunting rate etc. is reported elsewhere (Kumara and Singh, 2004a). DISCUSSION One of the most outstanding observations during this study was the sighting of Nilgiri langur groups in Srimangala Range. Although, Nilgiri langur was recorded in Brahmagiri Hills in 1912-13 (Ryley and Shortridge, 1913), later reports indicated the presence of Nilgiri langur only upto Nilgiri Hills, quite south of Brahmagiri Hills (Hohmann and Sunderraj, 1990) or an ‘uncertain’ status in Brahmagiri Hills (Oates, 1979). The present population of Nilgiri langur in Srimangala, therefore, marks the ‘certain’ status of this species in Brahmagiri-Makut. Further, this is the present northernmost population of this species in the entire Western Ghats. In the Western Ghats south of Brahmagiri-Makut, Nilgiri langur is sympatric with lion-tailed macaque in the rainforests. However, the distribution of lion-tailed macaque continues northwards upto north of Sharavathy river. Hohmann and Sunderraj (1990) hypothesized that in the central Western Ghats, the Nilgiri langur was probably hunted for meat, and the habitat vacated by this species was invaded by Hanuman langur. Present data do not support this argument since all primates are

severely hunted in Brahmagiri-Makut and the hunting pressure on primates is very low in the northern areas of the Western Ghats in Karnataka where lion-tailed macaque is still present. I propose that the absence of Nilgiri langur north of Brahmagiri-Makut is due to ecological factors. As mentioned earlier, Brahmagiri-Makut marks a very specific ecotone with absence of Cullenia exarillata but rich in Lauraceae. “This forest type occupies the medium elevations of the Western Ghats from almost its southern extremity up to Brahmagiri Ghats which form a natural border between Kerala and Karnataka. This relief forms a barrier, north of which Cullenia is no longer found” (Pascal, 1988; p. 113). It is probably this ecotone demarcation that can explain the end of the Nilgiri langur’s distributional range in the Western Ghats. Such habitat and ecological correlation is further strengthened by the present data that even within Brahmagiri-Makut forests, the Nilgiri langur was found to occur only in the higher altitudes. On the other hand, the protection against hunting was maximum in areas at Votekolli and Makuta, which have high densities of other primates but are characterized by the absence of Nilgiri langur. A unique feature of this population of Nilgiri langur was a small group size (4.25 individuals) as compared to the group size of 5.89 and 11.00 in Silent Valley (Ramachandran and Joseph, 2001) and Anaimalai Hills (Singh et al., 1997a) respectively in the southern populations in the Western Ghats. Since lion-tailed macaque is an endangered species, a special attention has been paid in the recent years to the distribution, status and conservation of this species. The total wild population of about 3500 individuals is distributed into 49 subpopulations, many of which are single, isolated groups in forest fragments (Molur et al., 2003). There are only a few self-sustainable subpopulations where 10 to 15 groups of lion-tailed macaques have adjacent or overlapping home ranges forming one substantially large breeding population. The present study is a major step towards the conservation of this endangered species. I have described not only the northernmost population of lion-tailed macaque, but have also found a population of 32 groups in a single, contiguous tract of rainforest where not more than 6 groups were reported earlier (Bhat, 1982; Karanth, 1985). With the probable exception of Kalkad-Mundanthurai, the southernmost population of this species in the Western Ghats, Sirsi-Honnavara population is the single largest subpopulation of lion-tailed macaque in its entire distributional range. This subpopulation, therefore, deserves special attention for conservation of this species, which was only a few decades back thought to be at the brink of extinction. In Sirsi-Honnavara, the lion-tailed macaque population appears to be healthy as indicated by a large frequency of groups in the modal groups size, a one to about four adult male to adult female ratio and the presence of a large percentage of immature individuals in the group. All these factors show a healthy demography and breeding status. However, a few concerns regarding long term conservation and management of lion-tailed macaque in Sirsi-Honnavara must be pointed out. These forests have a status of only Reserved Forests and do not come under any legally notified Wildlife Sanctuary or National park. The protection status of the forests, therefore, is not optimum. Since agricultural villages are scattered throughout the region, construction of roads, buildings, encroachment into forest land and habitat deterioration appear to be imminent. It is not possible to shift the villages out of these forests, but if the area is declared as legally protected as a lion-tailed macaque reserve, it may arrest further encroachment and habitat disturbance. The lion-tailed macaque population may then remain a large and biologically self-sustainable single population for the long-term conservation of this species. A unique feature of this population is the relatively larger group size. In the Western Ghats, the group size in lion-tailed macaque appears to consistently increase from south to north: 16.30 in Anaimalai Hills (Singh et al., 2002), 19.64 in Silent valley (Ramachandran and Joseph, 2001), 20.11 in Sringeri (Singh et al., 2001) and 24.70 in Sirsi-Honnavara (present study). The ecological regulators of the distribution of primates in southern peninsular India are clearly discernible from the present study. The bonnet macaque and the Hanuman langur are widespread

species, and are found in a variety of habitat types. On the other hand, lion-tailed macaque and Nilgiri langur are endemic only to the wet forests of the Western Ghats. From the southern tip of the Western Ghats upto Brahmagiri-Makut, a conspicuous tree species is Cullenia exarillata. However, the dipterocarpus forests continue further north up to Sirsi-Honnavara. At Brahmagiri-Makut and at Sirsi-Honnavara, there are major ecological changes in the Western Ghats, and these ecotones interestingly also mark the northern ends of the distribution of Nilgiri langur and lion-tailed macaque respectively. Even within large ecological distributional ranges, micro level ecological factors could be observed related to the distribution of primate species. The lion-tailed macaque is frugivorous/insectivorous, the bonnet macaque is omnivorous, and the two langur species are primarily folivorous. In the present study areas, the two macaque species and either one of the langur species could be found to be sympatric. On the other hand, the two langur species were rarely sympatric. Whereas the Nilgiri langur was restricted to the higher altitude rainforests including the montane shola forests occupied exclusively by Nilgiri langur in Brahmagiri-Makut, the Hanuman langur was found at lower ranges, extended even into typical deciduous forests of the adjoining areas. In Sirsi-Honnavara, where Nilgiri langur was absent, Hanuman langur was present in most of the areas. This type of spatial separation has also been observed in other places in the Western Ghats where these two langur species occupy adjacent and connected forests but are rarely sympatric (Singh et al., 1997a). This type of separation could not only be a spatial separation to avoid food competition, but also a mechanism for breeding isolation. If these two species ever occur together, a possibility of hybrids is not ruled out (Chellam, 1985; Hohmann, 1988). A special mention needs to be made about Hanuman langur in Brahmagiri-Makut. The total area under study was about 324 km2. Except for three and one stray groups each towards north and south respectively, all groups of Hanuman langur were found in a small area between Votekolli and Makuta (Fig. 3.2). Groves (2001) in his recent taxonomic revision of primates, has classified Semnopithecus hypoleucos as a distinct species. He has mentioned a rather small distributional range of this species corresponding to exactly the same place where I found several groups of langurs clustered. If at all S. hypoleucos is a distinct species, it would be interesting to further explore its distributional range north- and southwards in the Western Ghats and adjoining eastern plateau forests. A study could also be initiated on the behavioral ecology of this population (species?) hitherto not studied. The Hanuman langur population in Sirsi-Honnavara is either S. dussumieri (Groves, 2001) or S. entellus achates (Molur et al., 2003) or even S. e. hypoleucos (D. Brandon-Jones – personal communication). In any case, a comparative ecological and behavioral study of Hanuman langur populations in these study regions deserves merit. The anthropogenic factors show remarkable effect on abundance of species. Such an effect was clearly visible in the present study. Because of the traditional practices and religious affinities, the people in Sirsi-Honnavara do not hunt primates for meat although these people live inside these forests. On the other hand, the people living around Brahmagiri-Makut on both eastern and western sides have a special preference for primate meat. Karanth (1985) estimated 10 groups of lion-tailed macaques in Brahmagiri-Makut whereas only one group was sighted during the present study. As a matter of fact, the forests of Brahmagiri-Makut that largely come under a Wildlife Sanctuary, are much less disturbed than the forests in Sirsi-Honnavara. Yet, the primate abundance in Sirsi-Honnavara was much higher than in Brahmagiri-Makut. Local hunting may result not only in significant overall reduction in the number of individuals of a species (including a total local extinction), it may also affect the local distribution pattern. In Brahmagiri-Makut, both bonnet macaque and hanuman langur were found with the higher abundance in Makut (Wildlife) as compared to their abundance in Srimangala and Makut (Reserve Forest). This was

due to the fact that because of the presence of large number of Forest Guards and officials at Makut (Wildlife), the protection in this range was higher than in the other two ranges. On the other hand, in Sirsi-Honnavara, all three species showed a remarkably even distribution in all ranges of their occurrence. This brings out a very important factor related to conservation of species in their natural habitats. A legal declaration of an area as a ‘Protected Area’ may accord a higher protection of the forest and animals by the Government, but the long term and sustainable conservation requires attitudinal changes in the local people. Education regarding the status and ecological significance of endangered and vulnerable species is a significant step in this direction. Appendix 3.1 Details of Lion-Tailed Macaque Groups in Sirsi-Honnavara Identity Name Location Altitude

m asl Group size

HV 1 Sannamanegudda 3 kms north-west of Nirgod, at the slope towards Hirebylu

417 ?

HV 2 Mavinamara savalu 4 kms west to Kodigadde & Nirgod at the slope towards Hirebylu

417 ?

HV3 Hirebylu Hirebylu path, 4kms from Kodigadde 247 ? KD1 Chiksuli 5 kms south of Beerlamakki 505 30 KD2 Krishnaghtta 2 kms south –east to Chiksuli, near

Kanumane 615 16

KD3 Kerekuli-votekodlu 2 kms east of Krishnaghatta near Kerekuli

550 24

KD4 Hosathota 3 kms south to Kerekuli, between Naginamane & Melinamane

430 11

KD5 Samplevore-A 3 kms south-east to Melinamane near Balemane

440 9

KD6 Samplevore-B “ 440 15 KD7 Salikanu 2 kms south-west to Melinamane

between Hegdegudda & Kodigadde 600 36

KD8 Dasur 1 km from Dasur towards Kudegodu 545 ? KD9 Kudegodu 5 kms from Dasur towards Kudegodu 446 ? KD10 Happaregoli 3 kms north- west from Galamavu 577 39 KD11 Kalegadde 5 kms west to Galamavu 545 18 KD12 Galamavu West to Galamavu village 616 14 KD13 Suthlamane 1km south-east to Suthlamane village 667 31 KD14 Doddaguddekanu 2 kms south to Suthlamane village 595 30 KD15 Kotegudda 3 kms from Singumane towards

Malemane 652 ?

KD16 Hukkali 1 km south to Hakkali village 568 15 KD17 Tormai Forests around Tormai village 658 28 SD1 Malemane 2 kms north-west to Malemane village,

at Guddeganapati 533 38

SD2 Katlekanu Forests around Katlekanu 530 ? Identity Name Location Altitude

m asl Group size

GR1 Hasolli South-west to Mugatihole, 5kms from Melinamane

458 27

GR2 Kanukumbri 5 kms from Melinamane, near Hasolli, south-east to Mugatihole

515 9

GR3 Kengikuli 3 kms south-east to Kodagi village 464 ?

towards Sasiguli GR4 Mahime 5 kms north-east to Mahime village

towards Sasiguli 270 ?

GR5 Sasiguli-1 Forests south to Sasiguli village 568 ? GR6 Sasiguli-2 3 kms south-east to Sasiguli village 540 ? GR7 Dundamavu-1 Forests south to Dundmavu village 488 23 GR8 Dundamavu-2 1 km north east to Dundmavu 568 ? GR9 Matnigadde 1 km north-east to Matnigadde 518 33 GR10 Votehalla Forests around Votehalla village 401 48