Order Number 9401818

Ecology and conservation of the endangered lion-tailed macaque (Maeaea silcnus) in the landscape mosaic of the

Menon, Shaily A., Ph.D. The Ohio State University, 1993

Copyright ©1998 by Menon, Shaily A. All rights reserved.

UMI 300N. ZeebRd. Ann Aibor, MI 48106 ECOLOGY AND CONSERVATION OF THE ENDANGERED LION­ TAILED MACAQUE (MACACA SILENUS) IN THE LANDSCAPE MOSAIC OF THE WESTERN GHATS

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Shaily A. Menon, B.Sc., M.Sc., M.S.

******

The Ohio State University

1993

Dissertation Committee: Approved by F. E. Poirier, Dept, of Anthropology C. Davis, School of Natural Resources B. Anderson, Center for Mapping (Co-Advisors) T. Bookhout, Dept, of Zoology Interdisciplinary Program Copyright by Shaily A. Menon 1993 For Santosh and Arvind Menon With Love and Gratitude And For The Lion-tailed Macaques W ith Hope ACKNOWLEDGEMENTS

I express gratitude to Dr. Frank E. Poirier for his encouragement throughout my doctoral program and for patient tolerance of my 'independent streak.1 I am deeply indebted to Dr. Craig Davis for his support of the interdisciplinary nature of this project since its inception. H. S. Panwar and A. J. T. Johnsingh, Wildlife Institute of , Dehra Dun, provided invaluable administrative support as host-country collaborators. I gratefully acknowledge Dr. Ajith Kumar's guidance during field research, Dr. Ravi Chellam's help with preliminary field surveys, and R. Gopalan’s help with identification. The director of the Botanical Survey of India, Coimbatore, graciously allowed me access to their herbarium. I acknowledge the support of the Tamil Nadu Forest Department, the Chief Wildlife Warden of Tamil Nadu, and P. C. Tyagi (Wildlife Warden, Indira Gandhi Wildlife Sanctuary and National Park) and his family. Much of the success of the project is due to the dedication of my tracker, Thangavelu. I am grateful to Dr. David Cunningham, Center for Mapping, for assistance with image analysis and to Dr. Prem Goel and Cathie Hannon, Department of Statistics, for their advice on data analysis. This research received funding from the National Science Foundation, L. S. B. Leakey Foundation, World Wildlife Fund-US, and The Ohio State University Graduate School Alumni Research Award. It is my pleasure to acknowledge their support. I owe deep gratitude to K. K. and Sharada Menon for their gracious hospitality, S. Satish for his encouragement, and the people of Valparai for their warm acceptance of my presence in their midst. I am beholden to Lori Sheeran and James Stewart for their humor, empathy, and constructive input during the writing of this manuscript. For their inexorable confidence in me, I shall remain forever grateful to Catita Williams and David Butler. VITA

1986 ...... M.S. Animal Behavior and Ecology, University of South Carolina, Columbia. 1984 ...... M.Sc. Animal Physiology, Bombay University, Bombay, India. 1982 ...... B.Sc. Biology, St. Xavier’s College, Bombay University, Bombay, India.

PUBLICATIONS

Menon, S. 1992. Significance of Forest Fragments in the Landscape Mosaic of the Western Ghats. Smithsonian Symposium on Forest Remnants in the Tropical Landscape p. 36 (Abstract).

Menon, S. 1992. Conservation of the endangered lion-tailed macaque (M. silenus) in the landscape mosaic of the Western Ghats. American Journal of Primatology 27(1):47 (Abstract).

DeCoursey, P. and S. Menon. 1991. Circadian photo-entrainment in a nocturnal rodent: quantitative measurement of light-sampling activity. Animal Behavior 41:781 -785.

Menon, S., P. DeCoursey, and D. Bruce. 1988. Frequency modulation of a circadian pacemaker during photoentrainment. Physiological Zoology 61(2): 186-196.

Menon, S., Poirier, F.E., and W.R. Dukelow. History of the American Society of Primatologists (ASP) and International Primatological Society (IPS). In “History o f American Anthropology ’ F. Spencer ed. Garland Press. In press.

FIELDS OF STUDY

Major Field: Interdisciplinary Program Studies in Conservation Biology, Ecology, Primatology TABLE OF CONTENTS

ACKNOWLEDGEMENTS...... Hi

VITA ...... v

TABLES OF CONTENTS...... vi

LIST OF TABLES...... viii

LIST OF FIGURES...... ix

CHAPTER PAGE

I. IN TRODUCTION...... 1

II. LITERATURE REVIEW ...... 6

Anthropogenic Disturbance in the Western Ghats and its Effects on Rainforest Vegetation...... 6 Nonhuman Primate Survival in Disturbed Forests ...... 14 Behavioral and Life-History Characteristics of the Lion-tailed M acaque ...... 17

III. STUDY SITES AND METHODS...... 27

Study S ite ...... 27 M eth o d s...... 37 Vegetation Structure and Composition ...... 37 Activity Pattern and Feeding Ecology of the Lion-tailed Macaque ...... 39

IV. RESULTS AND DISCUSSION...... 45

Vegetation Structure and Composition...... 45 Regeneration of Vegetation...... 55 Habitat Use by the Lion-tailed Macaques at Puthuthotam Cardamom Forest...... 62 Activity Patterns of Lion-tailed Macaques at Puthuthotam Cardamom Forest...... 65 Diet and Feeding Ecology of the Lion-tailed Macaques at Puthuthotam Cardamom Forest ...... 73

V. CONCLUSIONS AND RECOMMENDATIONS...... 83

APPENDICES...... 93 A. Plant Species at Puthuthotam Cardamom Forest and Varagaliar B. Notes on Methodology C. Spatial Analyses

REFERENCES CITED...... 112 LIST O F TABLES

TABLE...... PAGE

1. General characteristics of primary and pioneer plant species...... 13 2. List of fauna observed at Puthuthotam Cardamom Forest ...... 34 3. Summary of instantaneous scan sampling data collected at Puthuthotam Cardamom Forest...... 43 4. Data recorded for each individual in scans ...... 43 5. Instantaneous scan sampling codes and definitions ...... 44 6. Comparison of vegetation characteristics of three forests in the Western G hats...... 47 7. Number of seedlings, saplings, and of plant species in 20 plots of 100 m^ each at Puthuthotam Cardamom Forest ...... 56 8. Number of species in 10 frequently disturbed plots and 10 less frequently disturbed plots of 100 each at Puthuthotam Cardamom Forest 59 9. Time budgets of lion-tailed macaque groups in three forests ...... 66 10. List of plant species in the diet of lion-tailed macaques at Puthuthotam Cardamom Forest...... 75 11. Plant species at Puthuthotam Cardamom Forest and Varagaliar ranked according to percent of lion-tailed macaque diet ...... 78 LIST OF FIGURES

FIGURES PAGE

I. Rainforest distribution along the Western Ghats ...... 28 2 Some rainforest fragments in and around the Anaimalai Sanctuary ...... 30 3. Monthly rainfall in Puthuthotam Cardamom Forest and Varagaliar ....35 4. Schematic diagram of Puthotottam Cardamom Forest ...... 36 5. Species-area curve for three 400 x 5 m transects at Puthuthotam Cardamom Forest...... 46 6. Vertical profile of trees at Puthuthotam Cardamom Forest ...... 49 7. Girth distribution of in a 400 x 5 m transect at Puthuthotam Cardamom Forest...... 51 8. Girth distribution of plants in a 400 x 5 m transect at Varagaliar ...... 51 9. Girth distribution of plants in a 40 x 40 m plot at Puthuthotam Cardamom Forest...... 52 10. Girth distribution of plants in a 40 x 40 m plot at Vargaliar ...... 52 II. Number of trees (girth ^1 0 cm) per species in a 400 x 5 m transect at Puthuthotam Cardamom Forest ...... 54 12, Number of trees (girth S 10 cm) per species in a 40 x 40 m plot at Puthuthotam Cardamom Forest ...... 54 FIGURES PAGE

13. Regeneration of 12 plant species in 20 plots of 100 m2 each at Puthuthotam Cardamom Forest ...... 58 14. Population structure of two climax species at Puthuthotam Cardamom Forest and Silent Valley...... 60 15. Percent of daytime spent by lion-tailed macaques in difFerent canopy levels at Puthuthotam Cardamom Forest ...... 62 16. Diurnal variation in use of different canopy levels by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 63 17. Percent of daytime spent in different activities by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 65 18. Diurnal variation in daytime spent in difFerent activities by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 66 19. Monthly variation in the percent of daytime spent resting by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 70 20. Monthly variation in the percent of daytime spent ranging by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 71 21. Monthly variation in the percent of daytime spent foraging by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 71 22. Monthly variation in the percent of daytime spent feeding by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 72 23. Monthly variation in the percent of daytime spent in other activities by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 72 FIGURES PAGE

24. Monthly variation in percent of feeding time spent on plant and animal foods by lion-tailed macaques at Puthuthotam Cardam om Forest ...... 73 25. Variation in the percent of daytime spent foraging on difFerent surfaces by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 74 26. Monthly variation in the use of 11 key plant species by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 79 27 (a-g). Comparison between monthly availability and use of excelsa, Artocarpus spp. F icusspp., Litsea oleiodes, peltata, Coffea arabica, and Diospyros spp. by lion-tailed macaques at Puthuthotam Cardamom Forest ...... 81 28. Representation of age-sex classes in the scan samples of behavior...... 99 29. Representation of age-sex classes in the nearest neighbor dataset for each animal ...... 99 30. Flowchart for spatial analyses...... 103 31. Overlay of 3 GIS files: roads, rivers, and parcels...... 107 32. Mean spectral values of 100 classes in TM bands 3 and 4 ...... 108 33. Cluster.GIS containing 7 landcover classes ...... 109 34. Final map with five land-use classes in a portion of the Anaimalai S anctuary ...... 110 C H A PT ER I

INTRODUCTION

The survival of most nonhuman primate species is inextricably tied to tropical rainforest conservation and management because as many as 80-90% of living nonhuman primate species are specialized to inhabit tropical, moist, broadleaf forests (Napier and Napier 1967). Formulation of management strategies for threatened primate populations must increasingly rely on data assessing the extent of change in vegetation due to disturbances and how effectively primate species adjust to these changes. The present research discusses rainforest fragmentation in the Western Ghats, south India, and its effect on vegetation and the feeding ecology of an isolated group of the endangered lion-tailed macaque. Wild populations of lion-tailed macaques are restricted to tropical rainforests in the Western Ghats of South India (IUCN 1976, Green and Minkowski 1977, Kurup 1978, Kumar 1985). Surveys in the late 1970s

I 2 indicated that lion-tailed macaque populations were confined to large forests, such as those in the Silent Valley (176 km^) and the Agastyamalai Hills (753 km^). Such forests were highlighted as the only viable sites for the long-term conservation of lion-tailed macaques (Green and Minkowski 1977, Henry et al. 1984). However, in 1988, Kumar (personal communication) conducted a systematic survey of the Western Ghats and located several groups of lion-tailed macaques in small isolated patches of forest, some as small as 35 ha. Data extrapolated from Ali (1985) and Kumar (1987, and personal communication) indicate that approximately 50% of the wild lion-tailed macaque habitat is distributed in relatively small, discontiguous, forest patches. Most of these forest patches are 0.5 km^ to 20 km^ in area and less than 30% of the wild lion-tailed macaque population is distributed in forests larger than 100 km2 in area (Kumar 1987). Isolation oflion-tailed macaque populations has resulted from large-scale clearcutting of forests for tea, cofFee, eucalyptus, and cinchona cultivation. Forest patches are subjected to ongoing disturbance through selective logging, firewood collection, and cardamom cultivation in the undergrowth. Given the heterogeneity of the landscape and increasing human population densities and demands in the Western Ghats, it is unreasonable to expect that large tracts of prime forest will be preserved in total isolation from human contact in the future. Since we are faced with an increasingly patchy juxtaposition of degraded forests, protected forests, monoculture plantations, and human habitation, it becomes critical to examine the status of the vegetation and wildlife in forest fragments. 3

The objeccives of this study are as follows: 1. To determine the differences in plant species composition, structure, and regeneration of tropical vegetation in a disturbed rainforest fragment compared with large, protected forests. A frequently disturbed and selectively logged forest is expected to have a lower species diversity and larger gaps in girth classes than a protected forest. The hypothesis is tested that species are adapted for regeneration under the most common disturbance regime (Denslow 1980). Thus, pioneer species are expected to be present in greater proportions in a frequently disturbed forest than in a protected forest. Moreover, vegetation in frequently disturbed areas within forest fragments should have a larger proportion of pioneer species than climax species. Large-seeded fruit forms a major component of the diets of many frugivores, such as the lion-tailed macaque. Since most pioneer species are not producers of large-seeded fruit, disturbance is expected to foster invasion of vegetation that does not contribute to the frugivore diet. 2. To determine differences in the activity patterns and feeding ecology of lion­ tailed macaques in an isolated forest fragment compared with those in protected habitats. Results from several studies on responses of other primate species to habitat disturbance have indicated that primate species in disturbed, and therefore marginal, habitats spend less time feeding than those in protected, unlogged, or higher quality habitats (Johns 1986, Marsh 1981a, Oates 1977). The hypothesis is tested that lion-tailed macaques in a disturbed forest fragment would be expected to spend less time feeding than those in protected forests. Lion-tailed macaques in a degraded forest should spend more time on the ground and in lower and middle levels of the canopy than lion-tailed macaques in large protected forests. Lion-tailed 4

macaques in a disturbed forest fragment are expected to have higher levels of infant mortality. What impact could altered abundances and distribution of food sources be expected to have on survival of lion-tailed macaques in a disturbed forest fragment? Vegetation studies in logged forests reveal that logging not only reduces total plant food abundance but also alters the distribution and relative availability of difFerent food types (Johns 1983a). Primate species inhabiting disturbed forests respond to altered resource conditions by making several dietary modifications. Since plant species diversity is expected to be lower in a degraded forest, lion-tailed macaques in such forests would be forced to rely on a smaller number of food species. Marginal diet items, such as , which are difficult for most macaques to digest, are expected to form a larger proportion of the diets of animals occupying a degraded forest patch. Lion-tailed macaques in a marginal habitat adjoining agricultural lands are expected to supplement their diet with cultivated crops such as coffee. 3. To propose alternatives for lion-tailed macaque conservation and the management of their fragmented rainforest habitat in the Western Ghats in light of data obtained in this study and in previous studies in protected forests. What are the anthropogenic pressures acting upon rainforest fragments in the Western Ghats and how might human needs be reconciled with wildlife conservation?

The data obtained from this research, along with their analysis and significance, are presented as follows. Chapter II contains a review of literature relevant to anthropogenic disturbances in the rainforests of the Western Ghats, the effects of such disturbances on rainforest vegetation, and an overview of 5 studies on the survival of nonhuman primates in disturbed forests and the life- history characteristics of the lion-tailed macaque. The study site and methodology are described in Chapter III. Chapter IV contains results and discussion on the vegetation characteristics of a forest fragment compared with those in a protected forest, regeneration patterns within a disturbed, isolated fragment, and on the feeding ecology and activity patterns of lion-tailed macaques in an isolated forest fragment. In Chapter V, I summarize my conclusions and discuss recommendations for the management of isolated forest fragments and lion-tailed macaque groups within them. C H A PT E R II

LITERATURE REVIEW

In this chapter I present background information on anthropogenic disturbance in the Western Ghats and its effects on rainforest vegetation, the survival of primate species inhabiting disturbed forests, and life-history characteristics of the endangered lion-tailed macaque.

Anthropogenic Disturbance in the Western Ghats and its Effects on Rainforest Vegetation

Paleobotanical evidence indicates that the climate in the late Cretaceous and early Tertiary was humid and tropical throughout peninsular India (Meher-Homji, 1974, 1978). The climate became drier as the landmass drifted northward and rainforests began to regress from the central and eastern parts of

6 7 the peninsula. However, the west coast of India continued to have a wet climate and a short dry season ofless than six months. The Western Ghats thus became a refugium for the humid plant genera of the wet evergreen forests (Meher- Homji 1979). Palynological data indicate that the climate of the southern state of Kerala has remained the same since the Miocene (Ramanujan and Rao 1977). Subsequent changes in rainforest distribution along the Western Ghats have been a direct result of human interference. Most of the initial exploitation of these rainforests by local human populations was low in intensity and only the more accessible regions were cleared for shifting cultivation (Pascal 1988). In the late 18th and early 19th century, however, rainforest exploitation became extensive and systematic with large scale, selective deforestation. Commonly exploited tree species included Dipterocarpus indicus (Dipterocarpaceae) andPalaquium ellipticum (Sapotaceae) for plywood, Calophyllum elatum (Clusiaceae) for aircraft manufacture during World War I and for building masts, Mesua ferrea (Guttiferae) for railroad ties, and Artocarpus species (Moraceae) for construction of boats and furniture (Stracey 1959, Pascal 1988). Large areas of prime forest were cleared by private owners for tea, coffee, and cardamom plantations in the 1800s. For example, the evergreen forests of ‘South Coimbatore’ occupied approximately 20,000 ha in a continuous belt between 450 and 1,500 m above sea level. However, half of this forested area was ‘dereserved’ for tea or cinchona cultivation (Stracey 1959). Pascal (1988) discusses both the present distribution of the difFerent rainforest types in the Western Ghats and their probable extent before extensive human encroachment. Rainforests along the Western Ghats occur mainly in the three southern states of Karnataka, Kerala, and Tamil Nadu. In Kerala, 8 dense evergreen vegetation dominated by the Dipterocarpaceae family may have extended all along the coast to altitudes of 700 m in the erstwhile region of Travancore (from the southern end of the Western Ghats to the Palghat Gap). High human population densities in the coastal and lower elevations of Travancore resulted in widespread destruction of these forests, except in extremely inaccessible areas. The indicator species that characterizes this forest formation, Dipterocarpus bourdillonii, is fast disappearing; Pascal (1988) observed only 15 remnant trees in the southern regions of the Western Ghats. Rainforests in the state of Karnataka are also dominated by Dipterocarp trees. DifFerent associations of these forests occur at present from 14° 15' to 11° 30' N but once may have extended to 10° 30' N. Dipterocarp rainforests are low altitude forests that do not extend above elevations of 700 m. Most of these forests in Karnataka are fragmented and degraded, with lower tree density and species diversity than the original forests. Construction of numerous reservoirs and dams such as those at Varahi, Chakra, Savehaklu, and Kalinadi has hastened this process (Pascal 1988). At medium elevations (700 -1400 m), Dipterocarp vegetation is replaced by the Cullenia-Mesua-Palaquium forest type which is the best preserved, in extent and quality, of the medium elevation forests in the Western Ghats (Pascal 1988). It occurs from the southern end of the Western Ghats to the border between Kerala and Karnataka in the north. Extensive forests of this type occur in the Nilgiri region (Silent Valley and Attapadi, Kerala), the Anaimalai region (Indira Gandhi Wildlife Sanctuary and National Park, Tamil Nadu), and the Agastyamalai Hills (Kalakkad-Mundanthurai Tiger Reserve, Tamil Nadu). Although these are protected forests, they are subjected to varying levels of disturbance. These disturbances include clearing of large areas 9 for teak, eucalyptus, tea, and cinchona plantations, inundation of large forests for the construction of dams and reservoirs, and the building of roads and canals through forests. Large areas of prime forest were cleared by private owners for tea, coffee, and cardamom plantations in the 1800s, These disturbances have caused the present-day landscape mosaic of the Western Ghats to be dotted with isolated forest fragments ranging in size from 35 ha to greater than 10 km^. The smaller fragments are essentially ‘islands’ surrounded by monoculture plantations and human habitation. Several of the isolated private forest fragments in the Western Ghats appear, to the casual observer, to be stands of primary forest and are often included as undisturbed forests in statistics compiled by forest officials (Green and Minkowski 1977). However, they are, in reality, non-regenerating clumps of trees functioning as shade providers for cardamom cultivation in the undergrowth. The main study site of this research was one such forest fragment. I summarize, below, the procedure involved in cardamom cultivation in wet evergreen forest fragments, as described by Green and Minkowski (1977) and Rivals and Mansour (1974). Elettaria cardamomum is a 2 m tall leafy plant cultivated for its seed pod which is used as a condiment. A perennial herbaceous monocot of Zingiberaceae, the ginger family, it grows wild in the wet evergreen forests of the Western Ghats and is also cultivated as a commercial plant. Cardamom plants require high levels of humidity, and therefore thrive in the understory of wet evergreen forests. Before cardamom cultivation begins all herbs, shrubs, small trees, saplings, and lianas are removed from the forest. Mature trees forming the topmost canopy are thinned and usually about 85% of them are left standing for shade. 10

Forest regeneration is prevented by regular and systematic weeding of all undergrowth in cardamom fields. The absence of regeneration, thinning of the forest, and replacement of natural undergrowth with cardamom plants modifies the forest in many ways including diminishing its ability to conserve soil and water. Cardamom leaves have low transpiration rates of 1.6 mg/g/min versus an average for rainforest trees of 8.9 mg/g/min (Walter 1971). The soil is vulnerable to erosion and windfall losses of canopy trees is increased (Green and Minkowski 1977). Small second-growth trees such as Erythrina indica* Clerodendron spp., M acaranga spp., orM allotus spp. that are good for shading the cardamom are introduced by managers of the plantation or are encouraged to grow naturally in gaps. When a cardamom field is finally abandoned, the ensuing regenerated forest is of a different composition than normally regenerating wet evergreen forests due to altered soil characteristics, influence of excess light penetration, and paucity of seeds from the few remaining large trees. An entire succession sequence must occur before the climax formation is once again attained, a process which could take 100 years (Green and Minkowski 1977). The history of cardamom cultivation is summarized by Rivals and Mansour (1974). Cardamom cultivation began on a large scale in 1800 in response to increased demands for its export. Approximately 65,000 ha of forests in the Western Ghats were dedicated to cardamom cultivation. In 1966, India was the largest producer of cardamom (4000 dried pods) followed by (1500 dried pods) and Guatemala (500 dried pods). About half of India’s cardamom production was exported. In the past several years, cardamom production in India has been erratic and has declined sharply due to adverse climatic conditions and parasitic 11 diseases. Infestation of the bases and bracts of the inflorescence by the thrips, Taeniopsis cardamomi, and a viral mosaic disease considerably weaken and hence reduce yields of infected plants. Other parasites affecting cardamom yield include borers of the stem, fruit, and rhizomes and caterpillars that feed on the leaves. Several years of strong measures against the parasites would be required to return to high cardamom yields (Rivals and Mansour 1987). Several pioneer species occur naturally at low densities in the rainforests of the Western Ghats. These include, () and Olea dioica (Oleaceae). Changes in the microclimate, as a result of fragmentation, and creation of large gaps in the forest canopy, as a result of selective logging, encourage the proliferation of naturally occurring pioneer species. Furthermore, species usually absent or rare in evergreen forests are introduced into these forests because the altered conditions are favorable for their growth (Pascal 1988). An example of a species that establishes itself in forest fragments in the Western Ghats is Oroxylum indicum (Bignoniaceae), characteristic of moist deciduous forests. Wet evergreen forests are distributed predominantly among developing countries in the tropics and are subject to large-scale anthropogenic disturbances. Alteration of these forests is estimated at a minimum rate of 11 million ha (Whitmore 1980, 1990) to 13 million ha per year (Ehrlich and Ehrlich 1986). These figures translate to a loss or change in 1.2 to 1.4% of the total forest cover annually. Since the beginning of this century, approximately 62% of the total area of tropical rainforest has been logged or converted for non-forest use (Conservation International 1990). If this rate of disturbance continues, most of the world’s tropical forests will be reduced to logged or otherwise disturbed patches by the end of this century. 12

Research on forest fragmentation through the late 1970s and early 1980s focused mainly on biogeographical considerations related to species-area curves, size and shape of reserves, and the value of several small versus single large reserves (Diamond 1975, 1976, May 1975, Simberloff and Abele 1976, 1982, SimberlofF 1988). These discussions have little practical value to the manager confronted with the issues of the usefulness of forest fragments and their management (Saunders et al. 1987, Saunders et al. 1991). Saunders et al. (1991) describe fragmentation of the landscape as creating several patches of natural vegetation of difFerent sizes within a heterogenous matrix of vegetation and/or land use. Fragmentation brings about physical change in the microclimate through radiation, wind, and water fluxes, and biotic change which depends on factors such as time since isolation and distance between remnants (Saunders et al. 1991). Subsequent to fragmentation, a patch can be expected to lose a proportion of originally present species and to gain a complement of invading species (Verner 1986, Murphy 1989). Continued recruitment of native plant species will depend on the distance of the fragment from other areas of native vegetation and on the extent and nature of gaps in the forests produced by the disturbance. As indicated by the intermediate disturbance hypothesis (Connell 1978), natural disturbances are not uncommon in tropical rainforests and frequent disturbances of intermediate intensities may, in fact, account for the maintenance of high levels of species diversity in tropical forests. The gap paradigm (Watts 1947, Whitmore 1975, 1978, 1982, 1989, Brokaw 1985, Hubbell and Foster 1986, Swaine and Whitmore 1988) was formulated to explain forest dynamics as driven by gap formation in the forest through natural disturbances. According to this paradigm, tree species respond to gaps 13 in one of two major ways. One group of species colonizes large gaps; seeds of these species germinate only in the open under high light intensities. Such species are referred to as pioneer species (Swaine and Whitmore 1988). The second group, non-pioneer, primary, or climax species, are shade-tolerant and grow in small gaps under the mature canopy of primary forest trees or under a canopy of pioneers colonizing a large gap. Table 1 lists general characteristics of pioneer and climax species. Hubbell and Foster (1986) describe a third group of “late secondary species”, most of whose characteristics resemble those of pioneer species except that they have higher wood densities, and more complex architecture, and are longer lived than most pioneer species. Whitmore (1988), however, argues that such a classification confuses matters and refers to them as long-lived, pioneer species.

Table 1. General characteristics of primary and pioneer plant species

Characteristics Primary species Pioneer species Germination Shade/sun and shade High/specific light intensities Shade tolerance High Low Photosynthetic rate Low High Growth rate Low High Wood density Moderate-high Low Seed dormancy Brief-none Long Seed size and no. Large and few Small and many Variable and diverse Wide dissemination Branching Complex Simple Life-span Long Short Examples Mesna ferrea Macaranga peltata Cullenia excelsa Clerodendron viscosum

From Hubbell and Foster (1986) 14

This interpretation of a dichotomous tree species response to gaps is somewhat simplistic and neglects variability in other aspects of the environment (Canham 1989, Leiberman et al. 1989, Poulson and Platt 1989). Nevertheless, as Whitmore (1988) suggests, the gap paradigm is a useful general description of forest dynamics. Intensive studies of forest dynamics for more than 20 years suggest that the gap paradigm is applicable to forests at all latitudes (Whitmore 1988). The cycle of gap, building, and mature phases of forest growth, under the gap paradigm, can be summarized briefly as follows. Natural disturbances, such as treefalls, create openings in forest canopies. Seedlings within these gaps grow into an immature forest of saplings which subsequently develops into mature trees. Thus, forests can be described as spatial mosaics of vegetation in different structural phases. Varying intensities of disturbances generate gaps of different sizes. Variation in gap size contributes to the heterogeneity o f species composition within a forest because the sequence of forest growth in a large gap is as follows: only pioneer seedlings germinate initially, grow into saplings, and then into a mature forest of pioneers. When individual pioneer trees die they create small gaps in the canopy. Seedlings of non-pioneer species flourish in the shady conditions of these small gaps and grow to maturity, eventually taking over the small gaps. Pioneer species never have the opportunity to grow to adulthood as long as gaps remain small.

Nonhuman Primate Survival in Disturbed Forests

Several resesarchers have developed models predicting fragmentation effects on faunal species (Dobkin and Wilcox 1984, Gotfryd and Hansell 1984, 15

Haila 1984, McLellan et al. 1984, Murphy and Wilcox 1984, Temple 1984, Urban and Shugart 1984, Forman and Gordon 1986, Rolstad 1991, Hanski and Gilpin 1991). The majority of these studies, however, have focused on birds in temperate regions. Increasing fragmentation and altered plant species composition in forest fragments have direct repercussions for many primate species. Information available through 1980 revealed that of the extant primates, only 21 species (14%) are generalists that can exploit both open and wooded habitats, 131 species (86%) live only in forested areas, and a minimum of 32 species (21%) are completely restricted either to primary forests or to one specific habitat type (Wolfheim 1983: 735). A few relatively recent studies have examined the response of primates to habitat disturbance in Southeast Asia, Africa, and the neotropics (Poirier 1968, 1969, 1971, Southwick and Cadigan 1972, Wilson and Wilson 1975, 1976, Marsh and Wilson 1981, Ayres 1983, Johns 1983a, b, 1985a, b, c, 1986, Skorupa 1986). A study of five primate species in a Malaysian forest before and after selective logging revealed that survival is influenced by the extent of logging, the ability to survive the period of high mortality immediately following logging, and the ability to adjust to the changed conditions of the regenerating forest (Johns 1983a, b). Specialist feeders, or species with a low dietary diversity, such as proboscis monkeys ( Nasalis larvatus) and orangutans ( Pongo pygmaeus), experienced higher mortality after selective logging of their forest habitat (Saltier and MacKenzie 1981, Wilson and Wilson 1975). On the other hand, generalist feeders, such as the crab-eating macaque (Macaca fascicularis), the pig-tailed macaque (M. nemestrina), the banded-leaf monkey (Presbytis 16

melalophus), the dusky leaf monkey (P. obscura), the common gibbon (Hylobates lar) and C. mitis, did not experience severe negative effects in selectively logged forests (Wilson and Wilson 1976, Marsh and Wilson 1981, Johns 1983a, b, and Skorupa 1986), Data from existing literature on primate survival in disturbed habitats were used to examine how survival correlated with the following three variables: body weight, frugivorous or folivorous diet type, and dietary diversity (Johns and Skorupa 1987). Dietary diversity was measured by the percent of the primate's diet made up by the 5 most frequently used food species. The analysis revealed that vulnerability of primates to habitat disturbance increased with increasing reliance on a frugivorous diet and, to a somewhat lesser degree, with increasing body weight. When the effects of diet type and body weight were controlled, dietary diversity showed no correlation with survival ratios. The reason for increased vulnerability with frugivory may be that the abundance and availability of fruit is more affected by disturbance than that of leaves. Moreover, regeneration of foliage is likely to occur before reappearance of fruit resources. Johns and Skorupa (1987) conclude that the response of primates to selective logging and to other forms of moderate habitat disturbance appears to be primarily a reaction to changed abundances and distribution of different food types. In Johns’ (1986) study in Malaysia, less fruit and flowers were available in logged forests so both banded leaf monkeys and white-handed gibbons fed more on young leaves. Leaf monkeys ate twice, and gibbons three times, the normal quantity of leaves. Feeding on less-nutritious foods places certain constraints on an animal. A gibbon that is forced to eat leaves must spend up to 80% of the day resting, to conserve energy. Before logging, the distribution of four representative taxa of food species in the study site (Johns 1986) did not 17 differ significantly from random. Following logging, however, the habitat became a mosaic of cleared patches. Both the gibbon and the leaf monkey showed a marked increase in the use of the lower to middle canopy over the upper canopy. Johns (1983) and Johns and Skorupa (1987) note that in southeast Asia, the majority of commercial timbers are not primate food species and food trees are lost only through incidental damage during logging. This factor along with the ability of southeast Asian primates in disturbed forests to change their diet and feeding behavior accounts for their survival in logged forests (Berenstain 1986, Johns 1986, Johns and Skorupa 1987). In Africa and South America, however, commercial timbers are frequently also important primate food trees and lost food sources in such forests are not compensated for by colonizing tree species, which are rarely useful to primates as food sources (Johns 1983). This is the major factor accounting for the inability of the neotropical primate, Chiropotes s. satanus, to survive in heavily logged forests (Ayres 1981, Johns 1985a).

Behavioral and Life-historv Characteristics of The Lion-tailed Macaque

The lion-tailed macaque (Macaca silenus) is endemic to the Western Ghats and is a prime example of rainforest specialization among primates. In this section, I present a brief overview of the literature concerning the evolutionary history, habitat, diet, annual ranges, competitors, predators, vocalization, reproductive biology, and present status of lion-tailed macaques in the wild. 18

Lion-tailed macaques are the only obligate rainforest dwelling macaques. Green and Minkowski (1977) state that the diet of lion-tailed macaques can be obtained only from the diverse flora and fauna of an undisturbed, mature rainforest, suggesting that lion-tailed macaques evolved in, and are adapted to, the rainforest habitat. While other macaques are semi-terrestrial to varying extents, wild lion-tailed macaques typically spend less than 1% of their time on the ground and are considered to be the only exclusively arboreal macaque (Southwick et al. 1970). Most other macaques such as Japanese (M. juscata), rhesus {M. mulatto), long-tailed (M. fascicularis), and bonnet macaques (M. radiata) are characterized by greater behavioral and ecological flexibility than are lion-tailed macaques. For example, M. fascicularis inhabits mangroves, nonriverine primary forests, riverine secondary forests, slash-and-burn fields, and other human rural and urban environments, such as temple sanctuaries (Wheatley 1980) and on secondary islands where they were introduced by humans (Poirier and Smith 1975, Poirier and Farslow 1984). M. mulatto has adapted to environments vastly different from the forest habitat (believed to be its original habitat), in villages, towns, cities, railway stations, roadsides, canals, and temples (Teas et al. 1980). According to Teas et al. (1980), the adaptability o f M . m ulatto is expressed primarily in behavior and, unlike lion-tailed macaques, rhesus macaques are sufficiently plastic in their behavior to accommodate close association with humans, Fooden (1975, 1976, 1980), Nozawa et al. (1977), and Eudey (1980) suggest that macaques belonging to the silenus-sylvanus group are archaic members of the based on certain primitive morphological characteristics and the disjunct distribution of extant species. From analyses of such morphological features as tail length and skull structure, Fooden (1975) 19 considers M . silenus to be most similar to the ancestral M acaca population which began the Asian dispersal of the genus. He argues that the silenus subgroup originated in peninsular India and dispersed northward to continental and insular Southeast Asia. This dispersal occurred when evergreen forests were present throughout peninsular India. Subsequent desiccation, associated with Pleistocene glacials, resulted in the disappearance of rainforests in eastern and central India and led to a disjunction of the Indian and Southeast Asian populations. M . silenus may have been restricted subsequently within its rainforest refugium due to competition from the rapidly expanding and more ecologically flexible M . radiata in peninsular India (Eudey 1980). A different view of phylogenetic relationships within the genus M acaca arises from immunological assays and electrophoresis of serum proteins performed by Cronin et al. (1980). These data indicate that sylvanus is the more archaic group and that silenus evolved more recently. However, the variability of electrophoretic integrity in the study, due to various reasons including extremely small sample sizes restricted to captive animals, makes the placements of sylvanus and silenus, in particular, rather inconclusive (Cronin et al. 1980). Detailed studies of molecular data from feral populations are required to establish the systematics of the genus Macaco. Lion-tailed macaque habitat is typically composed of lofty, dense, evergreen trees, 30-50m tall, with dense canopies (Green and Minkowski 1977). These are either the dipterocarp forests of the northern Western Ghats or the more southern forests in which Cullenia is abundant. D om inant tree associations of forests in which lion-tailed macaques occur are: Cullenia - Mesua - Palaquium, Cullenia - Calophyllum, Poecihneuron - Cullenia, Palaquium - Mangifera, Dipterocarpus - Mesua - Palaquium, and Persea - H oligam a - 20

Diospyros (Green and Minkowski 1977, Kumar 1987, Ali 1985). Food sources of lion-tailed macaques occur mainly in the upper canopy layers, but the flowers and fruit of vines, small trees and shrubs also are eaten. In the southern forests, C ullenia and Artocarpus are the main food sources (Green and Minkowski 1977). Cullenia is absent in the dipterocarp forests of Karnataka; the main lion-tailed macaque food sources there are Artocarpus, Mangifera, Eugenia, Ficus, and Caryota urens (Bhat 1984). Johnson (1980) observed lion­ tailed macaques eating fruit of Cojfea arabica, a cultivated species, in abandoned coffee plantations adjoining forest patches. Lion-tailed macaques show a higher level of carnivory than most macaques; they have been observed feeding on snails, insects (adults, pupae, and larvae), lizards, tree frogs, bird eggs and nestlings, and giant Malabar squirrel nestlings (Green and Minkowski 1977, Kumar 1987, and Hohmann: personal communication). In one of the earliest studies on lion-tailed macaques, Sugiyama (1968) observed a troop for 2 months during which they ranged over an area of approximately 2 km2. Kumar (1985, 1987) reported smaller annual ranges of 1 km^ in forests of the Anaimalai region. For lion-tailed macaque groups at Kalakkad-Mundanthurai, Green and Minkowski (1977) observed an annual home range of 5 km2. Home ranges of adjacent troops overlapped at the edges but not in a core area of about 3 km 2. Green and Minkowski offer the following explanation for the large home ranges. Despite monthly variation in the diet of lion-tailed macaques, Cullenia and Artocarpus are the most important year-round foods; thus troops are forced to range widely to exploit scattered aggregations or rare, isolated, individual trees for fruit constituting the bulk of their diet in a given month. According to Kumar (1985), the difference between the rainforests in Anaimalai and Kalakkad-Mundanthurai may explain 21 this discrepancy in annual range size. A part of the forest in which lion-tailed macaques occurred at Kalakkad-Mundanthurai was under cardamom cultivation. Cardamom plantations often are lumped under the category of undisturbed forest. In reality, they are equivalent to clear-felled forests because in the absence of regeneration the mature forest will not be replaced (Kumar 1985). The lion-tailed macaque diet at Kalakkad-Mundanthurai came from vines (12.3% of the total diet), small trees and shrubs (23.1% of the total diet), and big trees (27.7% of the total diet). In the part of the monkey’s range that is undergrown with cardamom, the mature stand of trees is thinned or selectively logged and all vines, seedlings, saplings, and young trees are cleared on a regular basis. Since these areas have reduced plant food and invertebrate density, the monkeys would need a greater area to obtain their food (Kumar 1985). In some areas, cardamom plantations serve as a secondary ranging area outside primary forests for lion-tailed macaque groups that may otherwise face extreme limitations of spatial and food resources (Kurup 1978). Feral female lion-tailed macaques give birth to their first young at 5 years of age and subsequently have only 1-2 more offspring in their lifetimes; male lion-tailed macaques first breed at 8 years (Green and Minkowski 1977). According to Kumar (1987), age at first birth for females in the wild was about 6.6 years and interbirth intervals were 3.6 years on average. In captivity, female lion-tailed macaques reach sexual maturity at about 4 years, males at 5 years. Males require multiple mounts to achieve ejaculation, and copulation is primarily a phenomenon of the follicular phase of the female’s cycle in captive populations. Lindburg et al. (1989) report that the interbirth interval for lactating females in captivity averages just over 17 months with no evidence of 22 birth seasonality. Kumar and Kurup (1985b), however, report two birth seasons per year in wild and captive lion-tailed macaques. The main frugivorous competitors of lion-tailed macaques are the giant Malabar squirrel, giant flying squirrel, Indian flying fox, toddy cat, Jerdon's imperial pigeon, great pied hornbill, Nilgiri langurs, hanuman langurs (in the northern ranges of the Western Ghats), and bonnet macaques (Bhat 1984, Green and Minkowski 1977). Green and Minkowski (1977) have observed lion-tailed macaques displaying alarm and mobbing reaction to leopards, wild dogs, pythons, and eagles, all of which are known or potential predators. Hohmann and Herzog (1985) identified 17 distinct vocal patterns in captive and wild lion-tailed macaques which they classified into 3 categories: (1) Situation-specific, (2) Interaction-specific, and (3) Individual-specific. ‘Loud’ calls made by the adult male when he is separated from the group and ‘warning’ calls made by adults and subadults of both sexes are both examples of situation-specific calls. During field observations by Hohmann and Herzog (1985)» lion-tailed macaques were observed to emit ‘warning’ calls on 3 occasions when unidentified birds of prey flew low overhead. In each instance, the group members reacted by fleeing to lower, denser vegetation for cover, and all subsequent locomotion and vocalization ceased until the danger was past. Other situation-specific calls include the female copulation call, the male copulation call, and several infant calls. Examples of interaction-specific calls are the ‘grunt’ call used to threaten conspecifics, the ‘rattle’ call used to threaten other species, and the ‘squeak’, ‘scream’, ‘shriek’, and ‘twit’ calls — all used during agonistic interactions. 23

Individual-specific calls lack a clear context. An example is the ‘whoo’ call made by males and females of all ages, often in sequence. The ‘whoo’ call is uttered at low frequencies when resting and feeding and at higher frequencies during the beginning of locomotion. Infant ‘whoo’ calls keep wandering infants in contact with their mothers. No other macaque species is reported to have a vocal pattern in their repertoire similar to the ‘loud’ call of male lion-tailed macaques. Loud vocalizations common in gibbons, howlers, and colobines were thought to be absent in baboons and macaques. Lion-tailed macaque ‘loud’ calls exhibit the acoustic conditions of long-distance sound transmission in forest habitats. These include a repetition of units and phrases, absence of transitional forms to other vocal patterns, emission from high places above the ground, stereotyped structure, broadcasting over a broad range of frequencies, and concentration of the main energy at low frequencies. Hohmann and Herzog (1985) suggest two reasons why the ‘loud’ vocalization is present in male lion-tailed macaques: high levels of arboreality and their social organization. In a dense arboreal habitat, which is not conducive to visual communication, the loud call enables the adult male to maintain vocal contact with the rest of the group. Most terrestrial/semi­ terrestrial macaques live in large multi-male groups whereas lion-tailed macaques typically live in small groups with 1-2 adult males. Hohmann and Herzog (1985) claim that individual adult lion-tailed macaque males can wander away from a group without fear of competition from other males and hence need a ‘loud’ call to maintain group cohesiveness from long distances in the canopy. Kumar and Kurup (1985a) state that the single adult male in their study group frequently wandered away from the group, maintaining an almost constant vigil for the presence of neighboring troops by climbing tall trees and 24 scanning the area around him. In light of the occurrence of intertroop copulations during encounters between troops, the male’s behaviors of active vigilance, herding of troop members, and inducing them to retreat when faced with a dominant troop, suggests an effort by the adult male to maintain exclusivity of reproductive resources (Kumar and Kurup 1985a). Thus, Hohmann and Herzog’s (1985) claim that the adult male has no fear of competition is perhaps not justified. However, their explanation that the ‘loud’ call may serve to maintain group cohesion still holds true for a male who has to leave the group to look out for neighboring groups. Lion-tailed macaques are limited to rainforests in the three states of Karnataka, Kerala, and Tamil Nadu. Estimates of the total number of lion­ tailed macaques in the wild have ranged from as few as 60 to as many as several thousand. When the lion-tailed macaques were brought into the limelight in the early 1980s, in connection with the debate over the proposed Silent Valley hydroelectric project, only 60 individuals were reported to exist in the wild (Ali 1985). Green and Minkowsi (1977) estimated the total lion-tailed macaque population in the wild at about 400 individuals, Kurup (1978) projected the total lion-tailed macaque population at 55 groups or 825 individuals (at an average group size of 15), and Ali (1982) put the estimate at 1000 individuals. The general consensus at the Lion-tailed Macaque Symposium, held in 1982 at Baltimore, based on a review of information gathered after 1975, was that the wild population of lion-tailed macaques was “in no case over 2000 animals” (Kurup 1988). But Karanth (1985) estimated 200 lion-tailed macaque groups (approximately 3000 individuals) in the state of Karnataka alone. This figure relied heavily on sighting reports of lion-tailed macaque groups by local people and was almost certainly an overestimate (Kurup 1988). Moreover, Bhat 25

(1984) had previously indicated that only a few groups still survive in Karnataka in evergreen forests of North Kanara and Shimoga at altiudes between 500 and 700 m. Kurup (1988) ascribes discrepancies in estimates of the wild lion-tailed macaque population to the facts that remote, inaccessible, dense, evergreen forests make censusing difficult and that most estimates are derived from extensive surveys at gross scales. H e states, furthermore, that lion-tailed macaque ranging patterns change substantially in habitats of different types and qualities which further confounds attempts at arriving at population estimates. For example, a group in a dense forest might range over a 100 ha area while one in a poorer habitat might have a home range of 15-20 km^ (Kurup 1988). Subsequent to 1982, efforts at censusing lion-tailed macaque groups focused on intensive surveys in smaller segments of the animal’s habitat. In the latest, and perhaps the most reliable, revision Kurup (1988) estimates a total lion-tailed macaque population of 1760 individuals distributed as follows in the three states: Kerala (900 individuals), Tamil Nadu (360 individuals), and Karnataka (500 individuals). Roughly half of this total population (approximately 850 individuals) is estimated to be distributed in isolated forest fragments (Kumar 1987). The reintroduction of captive lion-tailed macaques into the wild has been a much-debated issue. Lindburg and Gledhill (1992) have reviewed the status of the captive population of lion-tailed macaques in North America, Europe, Asia, and Africa. A steady rise in captive births beginning in the 1950s, which continued until recently, has brought the captive population to 176 individuals. About 56% of these are considered surplus in light of a 1991 decision by the North American Species Survival Plan (SSP) of temporarily 26 scaling back the captive population to improve its genetic diveristy. Thus, as Lindburg and Gledhill (1992) point out, unlike concerns of some other endangered species, management of captive lion-tailed macaques faces novel problems related to steady-state propogation. The authors recognize that reintroduction of surplus captive individuals to the wild is not an easily resolved issue nor is it popular among conservationists who consider the financial resources more effectively spent on protection of existing lion-tailed macaque habitat. Lindburg and Gledhill (1992) make the case and state several reasons why it is unrealistic for scientists in foreign countries and critics of zoos in the U.S. to presume that monies allocated to zoos for captive breeding programs can be easily reallocated for conservation efforts in the wild. Most captive breeding programs, however, do consider reintroduction as a final measure of the program’s success. According to the authors, therefore, ‘experimental’ reintroduction of captive lion-tailed macaques should be encouraged before a crisis situation develops in the wild, first, to gain knowledge that will be useful in a future time of need, and second, as an avenue for involving zoos more directly in saving wild habitat in the future. CHAPTER III

STUDY SITE AND METHODS

S tu d y S ite

The research described here was conducted in the rainforests of the Western Ghats, south India. The Western Ghats were uplifted during the Tertiary when the Indian Plate collided with Eurasia, causing high levels of tectonic activity and a series of elevations near the west coast of India (Pascal 1988). The relief is 1600 km long and extends from 8° N (the southern tip of India) to 21° N (Fig. 1). Its continuity is broken by the 30 km wide Palghat Gap (10° 30' N) and the narrow Shencottah Pass (9° N). The highest point in this chain of hills is Anaimudi peak at 2695 m. Violent regressive erosions etched deep valleys in the steep western slopes of the Ghats. Most of the major rivers flow east along the gradual eastern slopes (Pascal 1988).

27 28

Karnataka Forest! fDipterxarpxeae)

Nilglri Region ^.iCullenla-MesiuhPalapuluml

P algh atC ap ^ ^ n a lm a l a i Region Rainforest Distribution lOJltenUbMetua-Palaquiuml n n Before 18th Century (Low Altitude)

Before 18th Century IMedlum Altitude) Cardamom Hills (DipterocarpaceaeJ

Agastyamalai Region ^ ICullenlH-MesLta-Palaquiumj

100 km

78 -

Fig 1. Rainforest distribution along the Western Ghats. (Adapted from Kurup 1988 and Pascal 1988) 29

The formation of the Himalayas and the Western Ghats established the current monsoon patterns in India (Meher-Homji 1979). The monsoons are seasonal winds that blow from the Indian Ocean toward the Asian continent in summer and in the opposite direction in winter. The winter winds are dry whereas the summer winds are extremely moisture-laden and are the source of torrential monsoon rains (Pascal 1988). The intense, fast-travelling monsoon front arrives from the south-west. It reaches the Tropic of Cancer within two weeks of its arrival at the southern tip of the Indian peninsula, thus covering 15 degrees of latitude in approximately as many days (Pascal 1988). The north- south Western Ghats block this advancing front. Thus, the west coast of southern India has a wetter climate than the east coast and central India. The wet period exceeds six months annually in many regions along the west coast. Precipitation is highest both in a 80-120 km band parallel to the west coast and along the western relief of the Ghats. The average annual rainfall in these regions is more than 3000 mm and usually occurs in two peaks: one in June- July and a second, lower one in October-November. Rainfall and altitude gradients are two major factors determining vegetation distribution in the Western Ghats. The steep west-east precipitation gradient along the Western Ghats determines the distibution of rainforests. Evergreen forests require annual rainfall levels greater than 2000 mm; these levels occur mainly along the western relief. The annual rainfall drops to less than 1000 mm towards the eastern regions of the Ghats, a precipitation level insufficient to support evergreen formations. Areas with average annual rainfall between 1500 and 2000 mm have moist-deciduous forest formations and those with less than 1500 mm have dry-deciduous formations. The rainforest distribution also follows altitude gradients. Evergreen formations typically 30 occur at altitudes between 600 m and 1200 m. Tropical montane forests grow between 1200 m and 2000 m. At altitudes above 2000 m wind and frost conditions restrict tree growth to small pockets in sheltered valleys whereas exposed areas have grassland vegetation. The local term, ‘shola’, for these small forest pockets is often used to represent any wet evergreen forest.

vo

Sanctuary Boundary Rainforest Fragments

Fig. 2. Some rainforest fragments in and around the Anaimalai Sanctuary. (Adapted from Kumar 1987) 31

The Anaimalai Wildlife Sanctuary (Fig. 2) was established in the Anaimalai region of the Western Ghats in 1972. It is located at 76° 49.3' - 77° 21.4' E and 10°13.2' - 10° 33.3' N and covers an area of 987 km^. In the late 1980s an area of 330 km^ was raised to the status of a National Park, and the sanctuary was renamed the Indira Gandhi Wildlife Sanctuary and National Park. Most of the sanctuary is hilly, and altitudes range from 350 to 2500 m above sea level. Several tea and coffee monoculture plantations lie adjacent to the sanctuary. Isolated patches of rainforests are interspersed between these private plantations. The main study site, Puthuthotam Cardamom Forest (PCF), is a 65 ha disturbed forest patch surrounded on three sides by tea and coffee plantations and on one side by a heavily used road. PCF is located at 10° 20' N and 76° 58' E. It is a private forest and is underplanted (the forest floor is cleared for planting while several mature trees are left standing to provide the shade and humidity necessary) with cardamom and coffee. Additionally, it has an ongoing history of selective logging. PCF is home to a single lion-tailed macaque group of 43 individuals. I collected data in 1990-91 on feeding ecology, activity, and ranging patterns of the study group, and the phenology, floristic structure and composition, and regeneration of vegetation at PCF. The second study site is the protected forest of Varagaliar (VG) located within the sanctuary. VG, together with several adjacent forests, has a total area of approximately 16 km^. These forests contain approximately ten groups of lion-tailed macaques. Data on vegetation were collected in VG in 1990-91. Lion-tailed macaque populations in VG were the subjects of studies on feeding ecology in 1979-80 and 1983-84 by Kumar (1987). 32

Rainfall data for the duration of the study period (1990-91) at PCF were obtained from a rain gauge monitored at the office of the Public Works Department at Valparai. Rainfall records at Valparai for -several previous years were obtained from the same office. Kumar (1987) has comparable data on rainfall at VG during 1983-84. Rainfall at PCF starts around March and reaches its highest levels in June-July during most years. Subsequently, the levels decrease steadily until December or show a second lower peak between September and November (Fig. 3). Both PCF and VG show the effects of the south-west monsoon winds and the north-east monsoon winds. Annual rainfall in PCF (2735 mm - average of six years) is on average approximately twice as high as that at VG (1343 mm - average of two years, from Kumar 1987). The sanctuary contains a wide range of vegetation types corresponding to the altitude gradient. These include scrub and dry thorn forests, dry deciduous forests, moist deciduous forests, wet evergreen forests, montane forests, and grasslands. The wet evergreen forest at VG is located at 650 m and appears to be a transition between the low-alti tude Dipterocarp forests and medium-altitude Cullenta-Mesua-Palaquium forests (following classification by Pascal 1988). The forest fragment at PCF is located at 1085 m and corresponds to the Cullenia-Mesua-Palaquium type of wet evergreen forest. Eight hundred of the 2000 plant species found in South India are present in the sanctuary. Several of these are rare or endangered species such as Diospyros niiagarica, Combretum ovalifolium, and Gnetum ula. The sanctuary contains a representative set of the animal species occurring in the Western Ghats. Reptiles found within the sanctuary include pythons, cobras, vipers, monitor lizards, flying lizards, tortoises, and crocodiles. 33

Avifauna characteristic of the sanctuary include drongos, orioles, thrushes, doves, flowerpeckers, woodpeckers, parakeets, eagles, and hornbills. Mammals include tigers, leopards, elephants, gaur, sambhar, spotted deer, barking deer, mouse deer, wild boar, wild dogs, sloth bears, pangolins, porcupines, Nilgiri tahr, civets, otters, Malabar giant squirrels, and flying squirrels. There are five non-human primate species in the sanctuary: the slender loris ( Loris tardigradus), common langur (Presbytis entellus), (P. johnii), bonnet macaque (Macaca radiata), and lion-tailed macaque (M. silenus). Surveys of several isolated forest fragments, within the sanctuary and in areas surrounding it, by the author and by Kumar (personal communication) indicate that several animals inhabit these fragments or use them as ’stops’ in their ranging or migration routes. Wildlife observed within isolated fragments includes lion-tailed macaques, Nilgiri langurs, barking deer, mouse deer, wild boar, giant squirrels, Great Indian Hornbills, and numerous other birds and reptiles. These fragments serve as rest stops to leopards, tigers, wild dogs, and elephant herds moving between forests. Table 2 lists the fauna I observed at Puthuthotam Cardamom Forest. 34

Table 2. List of fauna observed at Puthuthotam Cardamom Forest

Mammals Birds

Common Name (Species) Common Name (Family)

Tiger (Panthera tigris) Hornbills (Bucerotidae) Leopard (Panthera pardus) Thrushes (Muscicapidae) Elephant (Elephas maximus) Flycatchers (Muscicapidae) Bison (Bosgaurus) Woodpeckers (Picidae) Sambhar (Cervus unicolor) Mynas (Stum idae) Barking deer (Muntjacus muntjac) Barbets (Capitonidae) Mouse deer (Tragulus meminna) Owls (Strigidae) Lion-tailed macaque (Macaca silenus) Drongos (Dicruridae) W ild dog (Cuon alpinus) Hawks (Accipitridae) Wild boar (Sus scrofa) Kingfishers (Alcedinidae) Porcupine (Hystrix indica) Jungle fowl (Phasianidae) Giant squirrel (Ratufa indica) 35

Fig. 3. Monthly rainfall in mm at Puthuthotam Cardamom Forest and Varagaliar. (Source: PCF data for 1982-91 from Public Works Department office, Valparai; VG data for 1982-83 from Kumar 1987) 36

Tea plantation

Streams

Roads

Coffee Nursery

iSiV* Frequently Disturbed Areas - Vegetation / Transects

Tea plantation |~ | Vegetation Plot

Fig. 4. Schematic diagram of Puthuthotam Cardamom Forest. Major landmarks and vegetation transects and plots are indicated. 37

Methods

The main study site, Puthuthotam Cardamom Forest (PCF), has a total area of 65 ha A map of PCF was prepared with the help of a pre-existing estate map. Direct measurements were made to establish locations of all major trails, vegetation transect, and other landmarks (Fig. 4). The total area of Varagaliar (VG), the second study site, is 16 km2 (1600 ha), approximately 25 times that of PCF. An area of 100 ha, which includes the home range of one lion-tailed macaque group (Kumar 1987, and personal observation) was demarcated for vegetation studies at VG.

Vegetation Structure and Composition

Data on vegetation structure and composition were collected at PCF along three 400 x 5 m transects and in one 40 x 40 m plot. Vegetation data at VG were collected along a 400 x 5 m transect and one 40 x 40 m plot. The choice of sample plots of 40 x 40 m made the data from this research comparable with those collected by Pascal (1988) in several undisturbed forests in the Western Ghats. I numbered and tagged all trees and lianas with gbh (girth at breast height) 2: 10 cm in the vegetation transects and plots at PCF and VG. I recorded the following data for each tagged plant: species and family, gbh, height (m), and presence or absence of epiphytes on trees. I collected leaf specimens, and flowers and fruit when available, poisoned them with formalin, pressed them between sheets of grey blotting paper tied between wooden planks, and dried them over low heat. I mounted the treated specimens on 38 herbarium sheets and identified the plant species at the BSI (Botanical Survey of India) herbarium, Coimbatore. If conclusive identification was not possible because fruit or flower samples were lacking, I noted the local name of the plant. Height of trees was calculated with the aid of a clinometer. Where use of the clinometer was obstructed by other trees, height was visually estimated within 10 m classes. Tree density per ha was calculated from the number of trees in the sample plots. The Shannon-Weiner index (H 0 was used to calculate plant species diversity:

H' = - 2 ( « / N)\og (t»/ N) (l) m i 1 where, ni = number of individuals of the species V, s = number of species in the plot, and N = total number of individuals in the plot.

I diagrammed a profile of 5 x 40 m to illustrate the vertical structure of the forest at PCF (Fig. 7). This profile is comparable with those drawn by Pascal (1988) for protected forests. Workers at the Puthuthotam Estate regularly and systematically cleared all undergrowth in PCF, as part of the cardamom cultivation protocol, until about 5 years before this study. To examine the extent and nature of regeneration of the natural vegetation, I recorded the number and species of seedlings (gbh <10 cm), saplings (gbh 10 to 30 cm), young trees (gbh 31 to 50 cm), and mature trees (gbh > 50 cm) in 20 randomly placed 10 x 10 m plots. 39

Riverine areas were defined as areas lying within 100 m of a river or stream. Frequently disturbed areas were arbitrarily defined as areas within 100 m of the fragment boundary or of a major trail extending across the fragment and frequented by humans. Plots were randomly selected from a numbered grid. Data were collected in the first 5 plots that occurred in each of the following areas: frequently disturbed riverine, less frequently disturbed riverine, frequently disturbed non-riverine, and less frequently disturbed non-riverine, respectively.

Activity Pattern and Feeding Ecology of Lion-tailed Macaques at Puthuthotam Cardamom Forest

I collected data in 1990-91 on the feeding ecology and activity patterns of a feral lion-tailed macaque group confined to Puthuthotam Cardamom Forest (PCF). The study group consisted of 43 individuals of the following age- sex composition: 2 adult males, 12 adult females, 13 subadults, 10 juveniles, and 6 infants. Groups in undisturbed forests are much smaller (12-20 animals per group) but have a similar age-sex composition as the group at PCF. Adult males are usually greater than 10 years old, and are identifiable by their large body size, large canines, and well-demarcated manes and tail tufts. Adult male lion-tailed macaques have an average body length of 55 cm and an average body weight of 8 kg (Green and Minkowski 1977). Adult females are at least 6- 7 years old, usually multiparous, and are identifiable by their smaller body size and elongated nipples. Both adult males and females have survived beyond the age of 30 years in captivity (Lindburg et al. 1989), but average life expectancy in the wild is probably shorter. Subadult females are approximately 5-6 years 40 old, subadult males are 5-10 years old, and juveniles are 1-5 years old; none of these individuals has well-developed tail tufts. The entire group was well habituated, and 9 monkeys were individually identifiable beyond general age- sex class. The study area was divided into numbered plots by overlaying a transparent 50 x 50 m grid on the base map. Ranging patterns were estimated by marking on the map the location of the troop during each scan. If more than 70% of the group was visible, the group center was marked. When the individuals were widely dispersed in loosely defined subgroups, as was often true, the subgroup center was plotted. The instantaneous scan sampling method (Altmann 1974) was used to collect intensive behavioral and ecological data on the animals in the study group. The study group at PCF was observed for 6 days every month for 12 months starting from September 1990. Data were recorded on several activity and diet parameters for all individuals observable within a period of 5 minutes. This procedure was repeated every 15 minutes. Each 5 minute period of data collection is henceforth referred to as a ‘scan* and data pertaining to a single individual within a 5 minute scan is referred to as a ‘record’. A total of 14,171 records were made in 659 hours of observations (Table 3). Data collected for each record are listed in Table 4. The codes used for recording data and the definitions of different activity categories are listed in Table 5. Unusual occurrences were recorded ad libitum during 10 minute scan intervals between routine 5 minute scans. Observations began on most days at dawn before the troop left its sleeping trees and continued for 12 hours each day. Heavy rain delayed or disrupted observations during several days in June, July, and August. 41

The instantaneous scan sampling method allows standardized comparisons between months within a study and between studies conducted by different researchers in different study sites (Marsh 1981b). In this study, 6 days of monthly scanning and 5 minute scans were selected to obtain data comparable with previous research on lion-tailed macaques in protected forests (Kumar 1987, Kurup and Kumar 1993). A primary use of the instantaneous scan sampling method is to create an activity time budget by estimating the amount or percent of time that individuals within a group devote to various activities (Altmann 1974). The percent of daytime spent on an activity by the group is defined as the sum of records of animals engaged in the activity in a day expressed as a percentage of the total number o f animals recorded in that day:

P — n t Af(lOO) (2) ti a ' ' where, Pa = % of daytime spent by the group on activity la\

na = number of records in a day when the activity was V and N = total number of records in a day.

Phenological data on 102 trees of 20 tree species provide an estimate of plant food availability each month. Selection of tree species for phenological sampling was based on preliminary observations of lion-tailed macaque diet at PCF and prior knowledge of lion-tailed macaque plant foods obtained from the literature and local people. Six representative trees or shrubs of each species were selected and marked. However, for 6 of the 20 species only 1 to 4 individuals were located. Each month all the marked trees were visited on the 42 day before observations of the lion-tailed macaque study group began. The presence of flush (new leaves), flowers, raw fruit, and ripe fruit was noted for each tree in the phenological sample. The amount of flowers or fruit on each tree in the sample was not quantified. The density (d z) of each species at PCF was calculated from vegetation data in transects and plots. The monthly availability of fruit from each plant species relative to other species in the phenological sample was estimated with the following formula (Goel personal communication):

(3) where, A i = percent availability of fruit and flower of species i, = number of trees of species i fruiting in the sample, di = density of species i at PCF, ». - number o f trees of species i in the phenological sample, and k - total number of tree species in the phenlogical sample.

The formula provides a crude estimate of plant food availability because the sample sizes are small and only presence/absence data were collected. A i estimates the importance of the fruit or flowers of species i relative to the other species fruiting or flowering in the sample. Use of plant species by the lion­ tailed macaques was estimated by the percent of daytime spent feeding on a particular species out of the total time spent feeding in a day. Data from the scan samples were analyzed with the Kruskal-Wallis one­ way analysis of variance to test for variation between months in activities, diet items, plant species, and foraging surface. 43

Table 3: Summary of instantaneous scan sampling data collected at Puthuthotam Cardamom Forest

No. of No, of Total Mean scans Month days hours records per day September 6 65.5 769 44.3 October 6 51.25 573 34.3 November 6 72 1180 48 December 6 68 1321 46.3 January 6 72 1594 48 February 6 71 1778 47.3 March 6 72 1913 47 April 6 72 1786 47 May 6 65 1887 43.2 June 3 19.25 498 26 July 2 5 165 10.5 August 4 26 707 26.3 Total 63 659 14,171 468.2

Table 4: Data recorded for each individual in a scan

1 Date 2 Time 3 Weather conditions (clear, cloudy, heavy rain, and windy) 4 Plot number (corresponding to a numbered grid) 5 Individual identification 6 Nearest neighbor (within a 10 m radius around the individual) 7 Height of individual in the canopy 8 Height of canopy above the individual 9 Activity of the individual 10 Diet type (if the activity is feeding) 11 Plant species (if the activity is feeding) 12 Foraging surface (if the activity is foraging) 13 Presence of other frugivores in the plot 14 Comments 44

Table 5: Instantaneous scan sampling codes and definitions

Weather Individual I.D. 1 Clear 1 Adult female 2 Cloudy 2 Adult male 3 Light rain 3 Subadult 4 Misty 4 Juvenile 5 Heavy rain 5 Adult female with infant 6 Windy Height in canopv (meters) Foraging Surface 0 0 1 Dried leaves 1 1 to 10 2 Tree bark 2 11 to 20 3 Tree holes and other snags 3 21 to 30 4 Shrubs 4 > 3 1 5 Ground 6 Tree canopy 7 Epiphytes Activity 1 Resting (includes sleep and inactivity; excludes grooming) 2 Resting and grooming 3 Ranging (directional travel by an individual or entire group) 4 Foraging (includes scouting for food sources and local travel within a tree clump) 5 Feeding (actual manipulation of food item or intake of food) 6 Sexual activity (includes sniffing, examination of estrous swelling, and mounting) 7 Agonistic activity (chasing or gestural/vocal behaviors in an agonistic context) 8 Play (chasing in a non-agonistic context, hanging from branches etc.) 9 Predator alarm (gestural or vocal response to potential1 predator) Diet/Food Source 1 Leaf 10 Epiphyte/liana 2 Flower 11 Invertebrate 3 12 Bird egg 4 Fruit flesh 13 Bird nestling 5 Seed 14 Giant Malabar squirrel 6 Resin 15 Water 7 Mushroom 16 Pith 8 Moss/lichen 17 Bat 9 Honey/honeycomb 18 Other CH A PTER IV

RESULTS AND DISCUSSION

Vegetation Structure and Composition

Forest types of different species composition occur along an altitude and latitude gradient within the wet evergreen forests of the Western Ghats and are often named after the most abundant species within them (Pascal 1988). Puthuthotam Cardamom Forest lies at an altitude of 1085 m and corresponds to the Cullenia exceba - Mesua ferrea - Palaquium ellipticum type of forest. This vegetation type occurs between latitudes 8° 50' and 10° 50' N and at medium altitudes (600-1400 m). The forest at Varagaliar is at an altitude of 650 m and corresponds to a transition between the Cullenia exceba - Mesua ferrea - Palaquium ellipticum type and the Dipterocarpus bourdillonii - Dipterocarpus indicus - Anacolosa densiflora type, which occurs between latitudes 8° 50' and 10° 50' N and at low elevations (0-650 m).

45 46

The species-area curve for the transects at Puthuthotam Cardamom Forest has a gradual slope that reaches a plateau at a cumulative species number of 45 (Fig. 5). However, the species list for Puthuthotam Cardamom Forest (Appendix 1) which includes all identified species in the transects (0.6 ha), plot (0.16 ha), regeneration plots (0.2 ha), and those observed to be eaten by the lion-tailed macaque group, contains more than 70 species.

50 45 40 g 35 •S 30 I 25 1 20 ■* 15 1

0 1 5 913 17 21 25 29 33 3741 45 49 63 57 Cumulative area (x100 sq.m.)

Fig. 5. Species-area curve for three 400 x 5 m transects at Puthuthotam Cardamom Forest. Only trees with girth ^ 10 cm are included. 47

Table 6. Comparison of vegetation characteristics of three forests in the Western Ghats.

PCFVG ARF a Sample plot size (ha) 0.16 0.16 0.2 Study area (ha) 65 100 > Total number of individuals (gbh S 10 cm) 130 170 303 Tree density (/ha) 812 1063 1520 Number of species (gbh ^10 cm) 31 41 32 Species diversity (H') 4.0 4.8 4.0 aAttapadi Reserve Forest data from Pascal (1988).

Data collected in the sample plots at Puthuthotam Cardamom Forest and Varagaliar are summarized in Table 6. Data obtained by Pascal (1988) at Attapadi Reserve Forest (ARF) are included for comparative purposes. Attapadi Reserve Forest, a protected forest adjacent to the Silent Valley Forest, lies at an altitude of 900 m, and is representative of the Cullenia excelsa - Mesua ferrea - Palaquium ellipticum forest type. Attapadi Reserve Forest does not appear to have been exploited in the past but has been subjected to recent fellings in the undergrowth by the local human population (Pascal 1988). Varagaliar, on the other hand, is recovering from damage caused by selective logging up to about 50 years ago (Hohmann and Sunderraj 1987) and Puthuthotam Cardamom Forest has a history of selective logging and undergrowth clearing. 48

As expected, tree density is lower at Puthuthotam Cardamom Forest than at Varagaliar and Attapadi Reserve Forest. The species diversity index, H 't is identical for Puthuthotam Cardamom Forest and Attapadi Reserve Forest, but the species composition of the two forests is different. Puthuthotam Cardamom Forest has been invaded by pioneer species, such as Clerodendron viscosum, as a result of fragmentation and disturbance. The higher species diversity index at Varagaliar may be because it is a low altitude forest (Kumar personal communication). The vertical profile at Puthuthotam Cardamom Forest (Fig. 6) reveals a large gap in the top and middle canopy layers. There were 2 trees (11.1% of the total trees in the plot) in the height class between 20 and 30 m and 1 tree (5.5%) between 10 and 20 m. A lower, more continuous layer consists of 15 plants with heights between 0 and 10 m and girths between 10 and 36 cm. Emergent trees, which reach heights of 40 m or above, are absent in the profile at Puthuthotam Cardamom Forest. In a 5 x 50 m profile at Attapadi Reserve Forest, Pascal (1988) observed 2 emergents (5.4% of the total trees in the plot), 3 trees (8.1%) in the 20 to 30 m, 13 trees (35.1%) in the 10 to 20 m, and 19 plants (51.4%) in the 0 to 10 m height classes, respectively. The paucity of trees in the middle layer of the canopy seen in the vertical profile at Puthuthotam Cardamom Forest is confirmed by girth distribution graphs. A uniform L-shaped girth distribution is indicative of an abundance of younger individuals and gradually declining numbers of older individuals with Fig. 6. Vertical profile of trees at Puthuthotam Cardamom Forest. Area sampled = 5 x 40 m.

VS

i 50 larger girths. Uniform L-shaped girth distributions are obtained in healthy, regenerating forests. Pascal (1988) obtained L-shaped girth distributions in his 2000 sample plot in the Attapadi Reserve Forest. The girth distributions in the plots at Puthuthotam Cardamom Forest and Varagaliar do not show a uniform L-shaped pattern (Figs. 7 and 8). The girth distribution in the transects at Puthuthotam Cardamom Forest and Varagaliar deviate similarly from the L-shaped pattern (Figs. 9 and 10). Gaps in the girth distribution indicate that selective logging has taken place in the forest. Past selective logging of trees at Varagaliar is reflected by a low representation of trees with girths between 20 and 80 cm. The girth distribution at Puthuthotam Cardamom Forest is more markedly disrupted because of systematic clearing of undergrowth in the past and ongoing selective logging. The numbers of individuals of different species in a transect and a plot at Puthuthotam Cardamom Forest are shown in Figs. 11 and 12. In both, the largest number of plants belong to pioneer, invading, or cultivated species. For example, in the transect (Fig. 11), two pioneer species,Clerodendron viscosum, and Macaranga peltata, constituted 18.3% (20 individuals) and 11% (12 individuals) of the total species in the transect. Mesopsis emini, an introduced species, comprised 13.8% (15 individuals) of total species. Mesua ferrea, a light tolerant climax evergreen species (Pascal 1988), formed 12.8% (14 individuals) of the total species in the transect. Each of the other species in the transect formed between 0.9% (1 individuals) and 5.5% (6 individuals) of the total species. Similarly, in the plot (Fig. 12), Cojfea arabica, a cultivated species, and Maesa perotettiana, a pioneer species, constituted 24.6% (32 individuals) and 14.6% (19 individuals) of the total species in the plot. 51

A

GBH

Fig. 7. Girth distribution of plants in a 400 x 5 m transect at Puthuthotam Cardamom Forest. Only plants with girths 10 cm are included.

35

ra il 00 GBH

Fig. 8. Girth distribution of plants in a 400 x 5 m transect at Varagaliar. Only plants with girth S 10 cm are included. 52

too

*

z 20

10 30 SO 70 80 110 130 ISO 170 190 210 230 250 270 200 310 330 350 GBH

Fig. 9. Girth distribution of plants in a 40 x 40 m plot at Puthuthotam Cardamom Forest. Only plants with girths 10 cm are included.

60 ■

50 ■

40 ■

30 ■

10 • F l r i n n r i n fli-i

110 130 150 170 400 GBH

Fig. 10. Girth distribution of plants in a 40 x 40 m plot at Varagaliar. Only plants with girth £ 10 cm are included. 53

Graphs for relative density of plants of different species in the transect and plot at Varagaliar are not shown because species at Varagaliar were incompletely identified. An evergreen species, inconclusively identified as a Syzigium spp. (local name “arinaval”), constituted 14.8% (21 plants) of the total species in the transect. All other species in the transect formed between 0.7% (1 plant) and 7.7% (11 plants) of the total species in the transect. Another evergreen species, (local name “poovathi”) probably belonging to the Dipterocarp family, formed 10.6% (18 plants) of the total species in the plot at Varagaliar. All other species in the plot constituted between 0.6% (1 plant) and 8.8% (15 plants) of the total. Similarly, in the plot at Attapadi Reserve Forest (Pascal 1988), three climax evergreen species, Aglaia anamallayana , Cullenia excelsa, and Palaquium ellipticum, formed 21.4%, 17.1%, and 9.5% of the total species. The pioneer species Clerodendron viscosum and Macaranga peltata were absent in plots at Attapadi Reserve Forest and Varagaliar. 54

25 -

A s. Of

z

Species

Fig. 11. Number of trees (girth £ 10 cm) per species in a 400 x 5 m transect at Puthuthotam Cardamom Forest. The numbers in parentheses represent the relative densities of each species in percentages.

35

30

25

20 t O I Q si 5? w _ w* sL si to ^ C. | 2 | Si g g C. w m 1 ! 3 49 S' I 1 W to II 10 S 3 8 49 E 1 | I itfi- 0 3 2 2 2 5 I |-|j'jto-q:oa.5gog^0agaiStill a §1 1 i f 1111 0 Innnnnnnnnnnnn^

Species

Fig. 12. Number of trees (girth £ 10 cm) per species in a 40 x 40 m plot at Puthuthotam Cardamom Forest. The numbers in parentheses represent the relative densities of each species in percentages. 55

Regeneration of Vegetation

The numbers of individuals of different species in the 20 plots (100 m^ each) at Puthuthotam Cardamom Forest are listed according to three girth classes in Table 7. An unidentified climax evergreen species (local name “sholapoovathi”) has the largest number of seedlings (944 individuals). This species appears to be more light-tolerant than most climax evergreen species. The pioneer species, Clerodendron viscosum, has 918 seedlings and the introduced species, Cojfea arabica, is third with 795 seedlings. Abundances of 12 plant species at Puthuthotam Cardamom Forest are plotted in Fig. 13. The first 10 species were frequent contributors to the diet of the lion-tailed macaque study group. The data on diet use were obtained from scan sampling of behavior and are described later in this chapter. The two most frequent contributors to the lion-tailed macaque diet, Cullenia exceba and Artocarpus spp., occur at low densities in Puthuthotam Cardamom Forest and did not show a stable population structure. Litsea oleiodes, a climax species, and Macaranga peltata, a pioneer species, have more stable population structures. Two of the key diet species, Oroxylum indicum and Semecarpus travancorica, were missing in the sample plots. Syzigium lac turn, Diospyros sylvatica, Erythrina subum bram, and Mesua ferrea are missing representatives in the 11 to 30 cm girth class. Mesua ferrea is included in this graph because of its importance as the preferred ‘sleeping tree’ of the lion-tailed macaques at Puthuthotam Cardamom Forest. The last species in the graph is the pioneer species, Clerodendron viscosum. It did not contribute to the lion-tailed macaque diet but constituted the highest proportion of all seedlings and saplings. Table 7. Number of seedlings, saplings, and trees of plant species in 20 plots 100 m2 each at Puthuthotam Cardamom Forest.

Ltm Girth (cm) Species Dieta Type 0 to 10 11 to 30 >30 "sholapoovathi" C 944 2 2 "sholathadacha " > C 15 3 0 Actinodaphne bourdillonii c 65 4 0 Aphanamixis pofystachya * c 1 0 0 Artocarpus spp. * c I 1 5 Bhesa indica c 0 0 2 Canarium strictum c 8 1 0 Cardamomum elettaria * I 116 0 0 Cinchona spp. I? 21 4 2 Cinnamomum malabatrum c 7 1 0 Clerodendron viscosum p 918 88 1 Coffea arabica * I 795 10 0 Croton reticulattts c 152 1 1 Cryptocarya spp. c 49 0 1 Cullenia excelsa * c 35 0 0 Datura stramomium p 35 0 0 Diospyros sylvatica * c 98 1 1 Elaeocarpus spp. * c 0 0 1 Erythrina subumbrans * p 2 1 2 Euvodia Ittmur-ankenda p 1 0 0 * Ficus spp. c 29 5 2 Flacourtia montana * c 2 0 1 Glochiodion ellipticum * c 1 0 0 Lantana camara * p 1 0 0 Limonia spp. c 25 0 0 * Litsea deccanensis c 170 14 3 Litsea oleiodes * c 123 7 1 Macaranga peltata * p 43 6 2 Maesa perotettiana ♦ p 189 14 0 Table 7. (contd.) Number of seedlings, saplings, and trees of plant species in plots of 100 each at Puthuthotam Cardamom Forest.

Ltm Girth (cm) Species Diet* Type 0 to 10 11 to 30 >30 * tetracocctu P 4 0 0 Mangifera indica * C 69 1 4 Mastixia arborea C 3 0 1 * Mesopsis emini I 24 2 0 * Mesua ferrea C 171 0 7 Myristica attennata * C 0 0 1 Neolitsea zeylaniea * C 7 1 0 Oka dioica P 67 1 1 Oroxylum indiaim * C 0 0 0 Persea macarantha c 32 6 0 Phyllanthus spp. c 57 10 0 * Semecarpus travancorica c 0 0 0 Stemonerats tetrandms * p 1 0 0 Sterculia guttata p 11 1 0 * Syzigium laetum c 3 0 1 * Syzigium spp. c 15 3 0 Toona ciliata c 22 1 0 * Vepris biloaiiaris c 4 0 0

^ h e asterix indicates those species Forming part of the lion-tailed macaque diet at PCF.

Key for Species Type: C: Climax forest species native to the forest type P: Pioneer species native to the forest type I: Introduced or cultivated species 58

Fig. 13. Regeneration of 12 plant species in 20 plots of 100 m2 each at Puthuthotam Cardamom Forest. Girths are divided into 3 classes of 0 to 10 cm, 11 to 30 cm, and >30 cm. The Y axis indicates the proportion o f each girth class represented by the 12 species.

To test the prediction that most species in an area will be adapted to the prevalent gap disturbance regime (Denslow 1980), I compared the abundance of 28 known climax evergreen species and 12 pioneer species found in 10 plots in ‘less frequently disturbed’ areas with their abundance in 10 plots in ‘frequently disturbed’ areas at Puthuthotam Cardamom Forest (Table 8). The assumption is made that frequently disturbed areas have larger openings or gaps and are exposed to higher temperatures and light intensities than less frequently disturbed areas. Although the 100 m cut-off point from forest edges and major trails is somewhat arbitrary, it was based on observations that human activity was most often encountered in these areas. Eighteen of the 28 climax species had greater abundances in less frequently disturbed plots and 8 of the 12 pioneer species had greater abundances in frequently disturbed plots. The 59

Table 8. Numbers of species in 10 frequently disturbed plots and 10 less frequently disturbed plots of 100 m2 each at Puthuthotam Cardamom Forest. Plants of all girths are included.

Climax species LFDa FD Pioneer Species LFDa FDb "shola poovathi" 781 167 Clerodendron viscosum 486 521 Mesua ferrea 130 48 Maesa perottetiana 58 145 Litsea oleiodes 96 35 Olea dioica 18 51 Litsea deccanensis 89 98 Macaranga peltata 11 40 Mangifera indica 64 10 Datura stramomium 0 35 Croton reticulatus 63 91 Erythrina subumbrans 1 4 Actinodaphne bonrdillonii 53 16 Mallotus tetracoccm 2 2 Cryptocarya spp. 45 5 Euvodia htmttr-ankenda 0 1 Diospyros spp. 42 58 Lantana camara 0 1 Persea macarantha 35 3 Stemonerctts tetrandus 1 0 Limonia spp. 24 1 Oroxylum indicum 0 0 Cullenia excelsa 13 22 Sterculia guttata 12 0 Canarittm strictum 9 1 naval ” 8 10 Neolitsea zeylanica 8 0 Cinnamomum malabatrum 5 3 Poona ciliata 5 17 Vepris bilocukris 4 0 Artocarpus spp. 3 4 Flacourtia montana 3 0 Syzigium laetum 3 1 Mastixia arborea 1 3 Bhesa indica 1 1 Elaeocarpus spp. 1 0 Glochidion ellipticum 1 0 Myristica attenuata 1 0 “shola thadacha “ 1 0 Aphanamixis polystachya 0 1 aLFD: plots in less frequently disturbed areas defined as areas lying beyond 100m of the forest edge or major forest trails. ^FL): plots in frequently disturbed areas defined as areas within 100m of the forest edge or major forest trails. 60 abundance of pioneer species was significantly higher than that of climax species in frequently disturbed areas (Analysis o f Variance, p < 0.05). Two climax species, Mesua spp. and Cullenia exceba, show more stable population structures in the protected Attapadi Reserve Forest (Fig. 14a) than at Puthuthotam Cardamom Forest (Fig. 14b), Mesua seedlings appear to survive better than Cullenia seedlings in disturbed conditions. However, both species are logged for timber and have no saplings or young trees and disproportionately few mature trees at Puthuthotam Cardamom Forest.

B0 30 160 60 360 1 390 1490

Mesua spp, Cullenia excelsa (a)

35 12.5

855 175

Mesua ferrea Cullenia excelsa

(b)

Fig. 14. Population structure of two climax tree species at (a) Attapadi Reserve Forest and (b) Puthuthotam Cardamom Forest. The numbers in the boxes correspond, from bottom to top, to the numbers of seedlings, saplings, young trees, and mature trees, respectively. Data for Attapadi Reserve Forest are from Manilal et al. (1989). 61

Future species composition of forest fragments depends on the disturbance regime prevalent in the fragment and its effect on climax species regeneration. Selective logging and other forms o f habitat destruction alter species composition in wet evergreen forests by increasing soil dessication and penetration of light to the forest floor (Green and Minkowski 1977). Shade- tolerant trees, such as Cullenia and Palaquium, do not regenerate as abundantly as in forests prior to logging (Aiyar 1932, Kadambi 1954). Selective logging is often accompanied by introduction of commercially desirable species resulting in gradual enrichment of some species and impoverishment of others (Pascal 1988). Examples of species that have been introduced in selectively logged forests for their commercial value include Mesua ferrea, Palaquium ellipticum, Artocarpus species. Some climax rainforest trees that can better tolerate microclimate fluctuations of light and moisture survive better in large gaps. Such species, as listed by Pascal (1988), include Artocarpus spp. (Moraceae), Canarium strictum (Burseraceae), Cinnamomum spp. (Lauraceae), Dimocarpus longan (Sapindaceae), Ficus spp.(Moraceae), Elaecarpus serratus (Elaeocarpaceae), Holigamia arnottiana (Anacardiaceae), Mangifera indica (Anacardiaceae), Myristica attenuata (Myristicaceae), Persea macarantha (Lauraceae), and Mesua ferrea (Guttiferae). Tolerance of Mesua ferrea to disturbance is exhibited at Puthuthotam Cardamom Forest (Fig. 14b) by the high proportion o f seedlings. Unfortunately, however, Mesua ferrea is a highly desirable timber species and very few adult trees remain in the forest fragment. 62

Habitat Use by the Lion-tailed Macaques at Puthuthotam Cardamom Forest

The lion-tailed macaque study group at Puthuthotam Cardamom Forest spent 4.9% of its time on the ground. Although Puthuthotam Cardamom Forest was a degraded forest with large gaps between trees, the group spent the largest proportion of its time (51.9%) in the topmost level of the canopy. Of the remaining time, 26.5% and 16.7% were spent on the middle and lowest level of the canopy, respectively (Fig. 15). The difference in time spent in different canopy levels is statistically significant (Kruskal-Wallis test, p < 0.0001). However, the group did not exhibit significant variation in the percent of daytime spent in different canopy levels during different parts of the

Ground Lowest Topmost Canopy level

Fig. 15. Percent of daytime spent by lion-tailed macaques in different canopy levels at Puthuthotam Cardamom Forest. The canopy levels on the x- axis correspond to heights of 0 m (ground), 1-10 m (lowest), 11-30 m (middle), and >30 m (topmost) in the canopy. Within each level, the height of the canopy above the individual is represented as shown in the legend. 63

50.0 45.0 40.0 35.0 '------0600to 1000 hours 30.0 25.0 20.0 1400 to 1800 hours 15.0 10.0 5.0 0.0 1 to 10 11 to 20 21 to 30 >31 Height in meters

Fig. 16. Diurnal variation in use of different canopy levels by lion-tailed macaques at Puthuthotam Cardamom Forest.

day (Fig. 16) except for heights in the canopy between 21 and 30 m. The group spent most time in this height class during early morning hours and least time during mid-day (Kruskal-Wallis test, p< 0.05). O f the time spent on the ground by the study group, 4.4% was spent in open areas without tree cover, such as the road and large clearings within the forest. Activities performed on the ground included ranging, foraging, feeding, playing, and drinking water from streams. The lion-tailed macaques crossed clearings in the forest by coming to the ground. They also crossed the road to raid Artocarpus trees growing in the coffee plantation on the other side of the road. Human presence in the coffee plantation was sometimes an indirect and 64 often a direct deterrent, in the form of gesturing and stone-throwing, which prevented them from spending much rime in the coffee plantation. Although lion-tailed macaque individuals migrate between groups within interconnected forests, they do not typically cross large gaps. They are very infrequently seen outside their rainforest habitat except where forest destruction is extensive. In more than 1700 hours of observations, Green and Minkowski (1977) did not observe lion-tailed macaques crossing adjoining tea or coffee plantations to the nearest forest patch even if the plantation was the shortest path between patches. O f the time spent on the ground, only 0.5% was spent under trees on the forest floor. This indicates that if trees or shrubs were available the group preferred to stay off the ground. Individuals were observed to jump across large gaps in the canopy even when it required significant effort. I observed three instances of lion-tailed macaques falling from the canopy at Puthuthotam Cardamom Forest during the study period. The first two individuals were juveniles and were observed to fall while clearing gaps in the canopy. Both these individuals survived their falls. The third instance involved the death of an infant. Kumar (1987) did not record any instances of falls resulting in deaths in 6 years of observations on lion-tailed macaques in the protected forest of Varagaliar. Johns (1981) notes that infant mortality is high during logging and immediately following logging. Almost all the primate infants previously present in his study groups disappeared by the time logging was completed. 65

Activity Patterns of Lion-tailed Macaques at Puthuthotam Cardamom Forest

The lion-tailed macaque group at Puthuthotam Cardamom Forest showed significant variation in time spent in different activities (Fig. 17, K-W test, p < 0.001). The largest proportion of time each day was spent ranging (median = 33.8%). The second most common activity was foraging (median = 22.8%). Feeding and resting took up about a fifth of the total time each (median values 17.1% and 16.4%, respectively). Time spent resting was significantly lower during early morning hours (from 0600-1000 hours) than during the remainder of the day (Kruskal-Wallis test, p < 0.01). Differences in time spent ranging, foraging, feeding, and in ‘other’ activities (Fig. 18) were not statistically significant at different times of the day. Other activities include sexual activity, agonistic activity, play, and predator alarm. Activity categories are defined in Table 5.

60 O Min ■ Mean ° Max ♦ Median

40 ■

20 ■

Resting Ranging Foraging Feeding Other

Fig. 17. Percent of daytime spent in different activities by lion-tailed macaques at Puthuthotam Cardamom Forest. 66

40 0600-1000 hours

30 1400-1800 hours

20

Resting RangingForaging Feeding Other

Fig. 18. Diurnal variation in daytime spent in different activities by lion-tailed macaques at Puthuthotam Cardamom Forest.

Table 9. Time budgets of lion-tailed macaques in three forests. Values represent average percent of daytime spent in different activities by lion-tailed macaques in each forest.

Activity PCF V G a VG/ASt> Resting 16.0 6.6 27.8 Foraging 23.7 36.3 26.7 Feeding 17.9 39.6 27.8 Ranging 34.0 13.9 15.0 O ther 8.4 3.7 3.5

a Data from Kumar (1987) for lion-tailed macaques at VG b Combined data for lion-tailed macaques at VG and AS from Kurup and Kumar (1993) 67

Table 9 compares the time budgets of lion-tailed macaques at Puthuthotam Cardamom Forest with those obtained by Kumar (1987) and Kurup and Kumar (1993). Kumar’s (1987) data are for lion-tailed macaques at Varagaliar. Kurup and Kumar (1993) combined data for groups at Varagaliar and AS (Anaikunthi Shola), both protected forests in the Anaimalai Sanctuary. Some of the differences in the time budgets in Kumar (1987) and Kurup and Kumar (1993) are due to the way in which activity categories are described in the two studies. Kumar (1987) defines 'resting’ as inactivity or sleeping, ‘ranging’ or ‘travel’ as movement that excludes ‘foraging’, ‘foraging’ as visual exploration for food, including pauses of inactivity, and ‘feeding’ as actual intake of food and manipulation of potential food sources. Kurup and Kumar (1993), on the other hand, define ‘feeding’ as actual intake of food items only, ‘foraging’ as active search for and selection of food items before eating, ‘resting’ as inactive sitting or standing, and ‘ranging’ or ‘moving’ as movements from one canopy level to another or over greater distances. Kumar’s (1987) data tends to overestimate ‘foraging’, by including rest pauses, and ‘feeding*, by including some foraging activities. Kurup and Kumar’s (1993) definitions of activity categories are much closer to those used in the present study. Lion-tailed macaques at Puthuthotam Cardamom Forest spend less time feeding and resting and more time ranging and foraging. Lion-tailed macaques at VG/AS spend approximately equal amounts of daytime resting, foraging, and feeding and less time ranging. Studies on other nonhuman primates also show that groups inhabiting disturbed habitats tend to spend less time feeding than those in more protected 68 habitats. Marsh (1981a) and Oates (1977) found that colobus monkeys living in marginal habitats in Africa (Tana and Chobe, respectively) spent less time feeding and more time resting than groups in an apparently better habitat (Kibale Forest Reserve). Oates (1977) suggests that the increased resting in marginal habitats may be explained by reduced energy requirements for thermoregulation because the climate at Chobe is much warmer than at Kibale. The same holds true for Tana and Kibale in Marsh’s (1981a) study. The climate at Puthuthotam Cardamom Forest is considerably cooler, especially during winter months, than at Varagaliar and Anaikunthi Shola. Thus, thermoregulation is a plausible explanation lor the increased ranging instead of resting at Puthuthotam Cardamom Forest. However, it can be ruled out because the average amount of daytime spent ranging at Puthuthotam Cardamom Forest did not show significant difference during different months (Fig. 20). Marsh (1981a) proposed that the number of species in the diet may be another equally plausible explanation for decreased amounts of time spent feeding in marginal habitats. Marsh’s (1981b) study at Tana recorded 22 species in the diet of red colobus monkeys and a study by Struhsaker (1975) recorded 56 species in red colobus diet at Kibale. In Oates’ (1977) study at Chobe, colobus monkeys fed on only 7 species compared with 42 species at Kibale. In this study, the Puthuthotam Cardamom Forest group fed on 42 species whereas the group at Varagaliar (Kumar 1987) fed on 81 species. Thus, low dietary diversity and decreased amount of time spent feeding are characteristics that are shared by the colobus monkeys at Tana and Chobe and by lion-tailed macaques at Puthuthotam Cardamom Forest. However, Marsh’s 69

(1981a) reasoning that low feeding times are explained by greater problems of detoxifying monotonous diets does not necessarily apply to frugivores like lion- tailed macaques. Moreover, there is a difference in how the remainder of the time is spent by the Puthuthotam Cardamom Forest study group. Lion-tailed macaques at Varagaliar and VG/AS spent more time resting than ranging as did the colobus monkeys in the protected habitat at Kibale and the marginal habitats of Tana and Chobe, whereas the lion-tailed macaques at Puthuthotam Cardamom Forest spend more time ranging than resting. There are several reasons that might account for increased ranging at Puthuthotam Cardamom Forest: 1. Puthuthotam Cardamom Forest has a very high human presence and the monkeys are often disturbed by the close proximity of twig pickers, people cutting wood from fallen trees, and people walking along the major forest trails. 2. As seen in the vegetation analyses, Puthuthotam Cardamom Forest is an extremely degraded habitat, possibly much more so than Tana or Chobe, and thus requires long searches for food items. Moreover, colobus monkeys are predominantly folivorous, and leaves, in general, require a shorter search time than food items constituting an omnivorous diet. 3. The unusually large size of the lion-tailed macaque group at Puthuthotam Cardamom Forest may be the most important factor compelling the group to be constantly on the move to ensure optimal foraging and to fulfil dietary requirements of all the individuals in the group. These characteristics, high human presence, extent of habitat degradation, and size of the group, are not seasonal in nature and were constants throughout the year. Their explanatory power is reinforced by the fact that of all the activities, ranging is the only one that did not vary 70 significantly with month (Fig. 20). Figs. 19 through 23 show the monthly variation in the average percent of daytime spent resting, ranging, foraging, feeding, and in other activities by the lion-tailed macaques at Puthuthotam Cardamom Forest. The mean, median, minimum, and maximum percent of daytime spent in the activity are plotted for each month.

45 * Mean 0 Min a Max 0 Median 40

35 30

20

15 10

0 JF A M JJS AM 0 N D

Fig. 19. Monthly variation in the percent of daytime spent resting by lion­ tailed macaques at Puthuthotam Cardamom Forest. Kruskal-Wallis test, p < 0.01.

A 71

* Mean 0 Min 0 Max V Median

30 25

Fig. 20. Monthly variation in the percent of daytime spent ranging by lion­ tailed macaques at Puthuthotam Cardamom Forest. Kruskal-Wallis test, no significant difference between months.

GO • Mean 0 Min a Max O Median SO

40

20

Fig. 21. Monthly variation in the percent of daytime spent foraging by lion­ tailed macaques at Puthuthotam Cardamom Forest. Kruskal-Wallis test, p < 0.001. 72

• Mean ° Min o Max • Median

45 40 35 30

|I 2025 15 10 5 0 FJ MJAMJ A 5 0 N D

Fig. 22. Monthly variation in the percent of daytime spent feeding by lion­ tailed macaques at Puthuthotam Cardamom Forest. Kruskal-Wallis test, p < 0.001.

■ Mean o Min □ Max V Median

30 i

25 ■

20 ' i >

10 ■

Fig. 23. Monthly variation in the percent of daytime spent in other activities by lion-tailed macaques at Puthuthotam Cardamom Forest, Kruskal- Wallis test, p < 0.001, 73

Diet and Feeding Ecology of the Lion-tailed Macaques at Puthuthotam Cardamom Forest

Plant foods formed a significantly greater portion of the annual diet of lion-tailed macaques than did animal foods (Fig. 24, Kruskal-Wallis test, p < 0.05). Invertebrates such as insects, grubs, and caterpillars formed only 3.5% of the annual diet of lion-tailed macaques at Puthuthotam Cardamom Forest. Of the various plant parts in the lion-tailed macaque diet at Puthuthotam Cardamom Forest, fruit flesh constituted 34.5%, fruit flesh and seeds (mainly Artocarpus spp. whose flesh and seeds were consumed at the same time) constituted 29.9%, flowers (mainly of Cullenia excelsd), constituted 18.4%, seeds constituted 12.6%, pith (mainly of Cardamom elettaria and “agil”) constituted 1.3%, and leaves constituted 0.5%.

100 00 so 70 ------P lan t 60 SO ------Invertebrate

30 20 10 0 F M A M

Fig. 24. Monthly variation in the percent of feeding time spent on plant and animal foods by lion-tailed macaques at Puthuthotam Cardamom Forest. 74

There is a significant difference in the time spent foraging on different foraging surfaces (Fig, 25, Kruskal-Wallis test, p < 0.01). Lion-tailed macaque diet is predominantly frugivorous and they do most of their foraging (51.8%) in the canopy foliage. Considerable amounts of time were also spent foraging for animal prey, mainly insects and grubs, in large, dried and curled up Macaranga \czves (7.6%); by prying off loose pieces of tree bark (11.5%); and in tree holes and snags (14.9%). I observed an adult female lion-tailed macaque catch and eat two bats out of several that flew out of a tree hole after she put her hand in it. This is the first time that lion-tailed macaques have been observed to eat bats. Other animal food items eaten by lion-tailed macaques but not observed at Puthuthotam Cardamom Forest, include bird eggs, bird nestlings, and giant Malabar squirrel nestlings.

90 80- 70- 0 Median 60- ■ Mean

40 ' « Min 30-

2 0' ° Max 10 ■

f 5? B 'l

Fig. 25. Variation in percent of daytime spent foraging on different surfaces by lion-tailed macaques at Puthuthotam Cardamom Forest. 75

Table 10. List of plant species in the diet oflion-tailed macaques at Puthuthotam Cardamom Forest.

Speciesa Part eaten Anacardiaceae Semecarpus travancorica hypocarp Bignoniaceae Oroxylum indicum mesocarp Bbenaceae Diospyros sylvatica seed Elaeocarpaceac Elaeocarpus munronii mesocarp Elaeocarpus spp. mesocarp Euphorbiaceae Macaranga peltata (=roxburghii) flower/mesocarp (=albns) flower Flacourtiaceae /Bixaceae Flacourtia montana mesocarp Guttiferae Mesua ferrea mesocarp, shoots Icacinaceae ?Stemoncrcus (=Gomphandra) tetrandrus (-polymorpha) Lauraceae L. deccanensis mesocarp L. insignis mesocarp L. oleiodes mesocarp Neolitsea zeylanica mesocarp Loranthaceae Loranthtu tomentus mesocarp (=excelsa) seed/flower Moraceae Artocarpus heterophyllus (-integrifolia) seed/mesocarp A. hirsutus seed/mesocarp Ficus glaberrima mesocarp F. glomerata mesocarp 76

Table 10 (contd.) List of plant foods in lion-tailed macaque diet at PCF.

Speciesa Part eaten F hispida mesocarp F. macrocarpa mesocarp F. microcarpa (=retnsa) mesocarp F. nervosa mesocarp F. ?tjakela mesocarp F. ?travancorica mesocarp Mvristicaceae Myristica (- Knema) attenuata seed coat Myrsinaceae Maesa perrotettiana mesocarp Mvrtaceae Syzigium (=Eugenia) lacttim (=lacta) mesocarp Papilionaceae Erythrina subumbrans (~lithosperma) nectar Rubiaceae Coffea arabica mesocarp Rutaceae Vepris bilocttlaris mesocarp/gum Sapotattftt Mimusops elengi mesocarp Palaqtiium elliptiam mesocarp Sterculiaceae Sterculia guttata seed Lantana camara mesocarp Zingiberaceae Elettaria cardamomum pith Mushrooms Lichens Family unidentified Mesopsis emini mesocarp ’aghil' pith 'naval' inconclusively identified species are preceded by a ? 77

The plant species contributing to the lion-tailed macaque diet at Puthuthotam Cardamom Forest are listed in Table 10. Lion-tailed macaques at Varagaliar fed on 81 plant species in one year (1982-83 data from Kumar 1987) whereas the group at Puthuthotam Cardamom Forest fed on only 42 species in one year. O f these, 11 species constituted 90.73% of the annual diet at Puthuthotam Cardamom Forest and 20 species comprised 90.66% of the diet at Varagaliar (Table 11). At Puthuthotam Cardamom Forest, Cullenia excelsa and Artocarpus spp. each formed more than 20% of the annual diet. In contrast, no single species contributed more than 18.5% to annual lion-tailed macaque diet at Varagaliar. Green and Minkowski (1977) also note that Cullenia fruit formed a major component of lion-tailed macaque diet in their study site at Kalakkad-Mundanthurai but they do not quantify its proportion in the diet. Cullenia is not abundant at Varagaliar, as it is a transitional forest, and contributed only 0.4% to lion-tailed macaque diet. The biggest contributors to the annual lion-tailed macaque diet at Varagaliar were Bischofia javanica (18.5%) and Diospyros microphyUa (15.2%). Among the species listed in Table 11, only Artocarpus spp., Ficus spp, andSemecarpus travancorica were common to both sites. However, their contributions to the lion-tailed macaque diets in both sites were different. Artocarpus and Ficus spp. contributed to a much greater extent to the diet at Puthuthotam Cardamom Forest (23.92% and 14.26%, respectively) than at Varagaliar (6.7% and 9.21%, respectively). Interestingly, 2 of the 11 key species in the lion-tailed macaque diet at Puthuthotam Cardamom Forest are not climax evergreen species. Macaranga peltata, a pioneer species, constituted 3.69% and Coffea arabica, a cultivated species, constituted 5.69% of the annual diet. 78

Table 11, Plant species at Puthuthotam Cardamom Forest and Varagaliar ranked according to percent of lion-tailed macaque diet. Species names present in both sites are indicated by bold type.

PCF VGA

Rank Speciesb Percent Species& Percent o f Se t CttmS ofSet Cum.c I Cullenia excelsa 24,77 24.77 Bischofia javanica 18.5 18,5 2 Artocarpus spp. 23.92 48.69 Diospyros microphylla 15.2 33.7 3 Ficus spp. 14.26 62.95 Ficus spp. 9.21 42.91 4 Litsea oleiodes 8.23 71.18 Artocarpus spp. 6.7 49.61 5 Coffea arabica 5.69 76.87 Mangifera indica 5.7 55.31 6 Macaranga peltata 3.69 80.56 Aglaia roxburghiana 4.6 59.91 7 Qroxylum indiatm 3.33 83.89 Bambusa arundanaceae 4.3 64.21 8 Syzigittm lactttm 2.8 86.69 Salmalia malabarica 4.0 68.21 9 Diospyros sylvatica 1.51 88.2 Dillenia pentagyna 3.9 72.11 10 Semecarpus travancorica 1.42 89.62 Syzigittm cumini 3.6 75.11 11 Erythrina subumbrans 1.11 90.73d Diospyros montana 3.5 78.61 12 Mushrooms, lichens 1.85 80.46 13 Glycosmis mauritania 1.8 82.26 14 Dalbergia sissoides 1.8 84.06 15 Vepris bilocttlaris 1.4 85.46 16 Nephelmm longana 1.2 86.66 17 Saccopetalum tomentosum 1.1 87.76 18 Semecarpus travancorica 1.0 88.76 19 Caesalpinea bondtti 1.0 89.76 20 Chilocarptts atroviridis 0.9 90.66d a VG data for 1982-83 from Kumar, 1987 (Appendix: Table 11.2) b Artocarpus and Fiats species are grouped together for PCF and VG c Cumulative percent d 31 and 61 species, respectively, make up the remaining 9% for PCF and VG 79

1 Cultma D Artentrpm 1 Ftna 1 Uttt* 0Coffra EH Mnearmifii §Omqtum

0Sjzi/jum B Dwify m B Smrcdrpus B Eryihriitd B Othtr

100

80 I 80 1 40 S. 20 0 J FMAMJJ ASOND

Month

Fig. 26. Monthly variation in the use (estimated by the percent of daytime spent feeding on a particular species out of total feeding time) of 11 key plant species by lion-tailed macaques at Puthuthotam Cardamom Forest.

There is significant monthly variation in the use of different plant species by lion-tailed macaques at Puthuthotam Cardamom Forest (Fig. 26, Kruskal- Wallis test, p < 0.0001). Figs. 27 (a - g) compare the use in the lion-tailed macaque diet of Cullenia excelsay Artocarpus spp., Ficus spp., Litsea oleiodes, Coffea arabica, Macaranga peltata, and Diospyros sylvatica, respectively, with their availability at Puthuthotam Cardamom Forest. Graphs for Oroxylum indicum and Semecarpus travancorica are not provided because they were absent in the vegetation sample plots and transects at Puthuthotam Cardamom Forest. The density value used in the formula for availability could, therefore, not be calculated for these two species. The Cullenia flowering season usually begins in 80

October and lasts through January (Pascal 1988 and phenological data in this study). During these months, Cullenia flowers formed a significant proportion of the lion-tailed macaque diet at Puthuthotam Cardamom Forest. Cullenia fruit were eaten in proportion to their availability in August and September, but not in March, April, and May. Since Cullenia fruit are covered with spines and remain green most of the time, the degree of ripeness was difficult for me to estimate. However, Cullenia fruit fall is highest around July (Pascal 1988). Cullenia fruit are probably raw in March, April, and May and were, therefore, not the preferred food in these months. Cullenia, Artocarpus spp., and Ficus spp. were the only species that contributed to the lion-tailed macaque diet for more than 8 months in the year at Puthuthotam Cardamom Forest. In April, May, and June, Litsea fruits were available and were one of the preferred foods of the lion-tailed macaques in those 3 months, Artocarpus was a preferred food in most of the months when it was available except in May when a marked preference was shown for Litsea fruit. Ficus was consumed in small amounts almost year round, but it became the major component (72%) of the lion­ tailed macaque diet in September. This was because Cullenia, Artocarpus, and Litsea fruit were not available in September. The other main contributors to the diet in September were Cullenia flowers. Fruit of Cullenia, Artocarpus, Litsea, and Ficus, were absent in October, November, and December. During these three months, Cullenia flowers, and the fruit of Diospyros spp. and of the pioneer species, Macaranga peltata, were the main contributors to the lion­ tailed macaque diet. Fruit of Diospyros spp., Macaranga peltata, and Coffea arabica were not eaten in proportion to their availability, but appear to be consumed when preferred food sources were at a premium. 81

70 -|

60 '

BO •

40 •

20 •

J= l

(a)

90 -I 60 ' 70 ■

S O •

30 •

20 •

(b)

80 70 ■ 60 •

40 - " PitH t in £>*rt 20 -

(c)

Fig. 27. Comparison between monthly availability and use of Cullenia excelsa, Artocarpusspp., F icuss pp.,Litsea oleiodes, Macaranga peltata, Coffea arabica, and Diospyros spp. by lion-tailed macaques at Puthuthotam Cardamom Forest. 82

Fie. 27 contd.

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o M AJAM JJF $ O N

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70

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CONCLUSIONS AND RECOMMENDATIONS

The factor most inimical to the survival of the lion-tailed macaque is not so much their small total number but, rather, their dispersion in small, isolated, and disturbed rainforest fragments in the Western Ghats, India. The total lion­ tailed macaque population in the wild is estimated at 1760 individuals (Kurup 1988); approximately half of these inhabit isolated forest fragments (Kurup 1987). The main objectives of this study were to determine disturbance effects on the vegetation in a rainforest fragment and on the ecology of an isolated lion-tailed macaque group, and to recommend strategies for the management of lion-tailed macaque groups in habitat fragments. Intensive data on plant species composition, forest structure, and the ecology of lion-tailed macaques in a disturbed forest fragment were not available before this study. Management decisions ideally require long-term data on vegetation and lion-tailed macaque ecology from several forest

83 84

fragments. Unfortunately, this option is not available for isolated groups of lion-tailed macaques because the destruction of forest fragments in the Western Ghats is inevitable and, in some cases, imminent. For example, subsequent to this study approximately 800 trees have fallen following permission to selectively log 332 trees at Puthuthotam Cardamom Forest (Ahimaaz 1992 and personal communication). Conservation biologists are often forced to make decisions based on incomplete data; not doing so could result in the irrevocable loss of a natural resource (Diamond 1986). During the study, I made extensive surveys to locate rainforest fragments in the Anaimalai region. Three of four isolated fragments I visited are < 100 ha in size, are located at altitudes of approximately 1000 m, and contain resident populations of lion-tailed macaques. All three are surrounded by tea or coffee plantations and have ongoing anthropogenic disturbances. The fourth fragment is at an altitude of approximately 1200 m and is connected to a larger, protected forest. It has no resident lion-tailed macaques but is sometimes visited by groups inhabiting the protected forest (Kumar: personal communication). The flora, fauna, and human pressures appear to be very similar in privately owned forest fragments in the Anaimalai region and conclusions reached from this study in Puthuthotam Cardamom Forest can be extrapolated to them until further data become available. Vegetation data obtained in this study clearly indicate that frequent disturbance in forest fragments creates conditions conducive to invasion by and proliferation of pioneer species. These pioneer species are absent or occur at low densities in large, protected forests. Disturbance effects on forest structure include interrupted girth class distribution and large gaps in the middle and top canopy layers. Primary or climax species in the forest fragment did not have 85 stable population structures typically found in protected forests. Regeneration of primary species will continue to be discouraged because disturbance is ongoing, frequent, and of high intensity in most isolated fragments. Eventually, primary species can be expected to become locally extinct. For example, only four mature individuals of the species Palaquium ellipticum, one of the three species that the forest type is named after, could be located in the entire fragment of Puthuthotam Cardamom Forest! Changes in forest structure and plant species composition have direct and indirect effects on lion-tailed macaques inhabiting a forest fragment. Ripe, sugar-rich fruits form a major component of the lion-tailed macaque diet in protected and disturbed habitats. However, because they rely heavily on a few fruiting species, lion-tailed macaques in forest fragments are susceptible to ‘ecological crunches’ (Wiens 1977) which reduce the carrying capacity of the habitat. Such crunches are known to result from poor fruiting due to failed rainfall or from reduced food availability immediately following selective logging. Of the four tree genera that formed 70% of the lion-tailed macaque diet at Puthuthotam Cardamom Forest, Cullenia and Artocarpus are desirable timber species. During certain months, up to 60% of the lion-tailed macaque diet comprised either Cullenia or Artocarpus. Ficus is an important food source for lion-tailed macaques because it was available almost all year round and contributed significantly to their diet during months when few other species are fruiting. Another important food source at Puthuthotam Cardamom Forest is Litsea, which forms a significant proportion of the lion-tailed macaque diet during the three months in which it fruits. The abundance of such keystone plant resources determine, in part, the carrying capacity of the environment for most of its animal biomass (Leighton and Leighton 1983, Terborgh 1983). It 86 might be possible to enhance or maintain the carrying capacity of a degraded forest by management actions that increase the abundance of such plant resources. Terborgh (1986) suggests that as long as desirable densities of keystone plant resources are maintained the rest of the forest could be managed for timber production without reducing the carrying capacity of the forest. He cautions, however, that at present we know niether how to manipulate the species composition of tropical forests on a commercial scale nor what the long­ term consequences of such manipulations might be. Moreover, knowledge of the biology of the animal is crucial because a tree species might be important to the animal even if it does not contribute significantly to the diet. For example, Mestta ferrea, the ironwood tree, a heavily logged timber species, is a tall tree with a wide canopy and is the preferred ‘sleeping tree1 of the lion-tailed macaques at Puthuthotam Cardamom Forest. Lion-tailed macaques in the disturbed forest fragment have modified their diets and behaviors. They spend less time feeding, eat leaves and flowers to a greater extent, include fruit and flowers of pioneer and cultivated species in their diets, and spend more time in lower and middle levels of the canopy than do lion-taiied macaques in protected forests. The plasticity in the diet and behavior of primates, which has been emphasized by many studies, ensures immediate survival of primates in logged or otherwise disturbed forests. However, stresses exerted on the animals during critical periods of low food production may have negative implications for the population structure in the future. Dependence on marginal and low energy foods and an increased need to spend time ranging to locate food sources in disturbed forest fragments could adversely affect birth rates, life spans, and mortality rates of the lion-tailed macaque groups. Increased infant and juvenile mortality can be inferred from 87 the high incidence of accidental falls observed in this study. Increased predation risk may be expected due to large gaps in the canopy and the increased need to use the ground and lower canopy levels in the disturbed forest fragment. Lion-tailed macaque groups in large, protected forests fission when they reach the critical size of approximately 24 to 30 animals. Wild lion-tailed macaques have average groups of 15 to 20 individuals. The study group at Puthuthotam Cardamom Forest did not fission despite reaching the large size of 43 individuals, because the forest fragment is not large enough to accommodate two separate groups and the nearest forest is several kilometers of intervening plantations and human habitation away. Large group sizes have been associated with decreasing birth rates in lion-tailed macaques (Kumar 1987). Furthermore, high adult female to male ratios in larger groups cause increased female sexual competition, in the form of sexual harassment during mating, resulting in suppression of reproduction by ovulating females (Kumar 1987). Thus, reproductive rates are likely to decrease as the group size of lion­ tailed macaques in forest fragments increases. A significant proportion of rainforest vegetation and wildlife in the Western Ghats occur in isolated fragments outside the network of protected areas. What are the implications of habitat fragments and isolated ‘metapopulations’ (Levins 1970) of wildlife in the greater agricultural, natural, and urban landscape mosaic of the Western Ghats? Puthuthotam Cardamom Forest contains a large resident group of lion-tailed macaques and a nesting pair of Great pied horn bills (Buceros bicomis), both endangered species, and several other representatives of the local flora and fauna (See Table 2). Animals with large home ranges such as wild elephants, wild dogs, tigers, and leopards frequently pass through the forest fragment as they travel between larger forests. 88

Loss of forest fragments could affect home ranges of elephants, tigers, and leopards possibly resulting in increased conflict with humans. It is unlikely that the opposite, i.e. decrease in conflict upon loss of fragments, will occur because carnivores do not prefer forested habitats over grasslands or plantations, elephants are found more often in grasslands, and bison prefer plantations (Balakrishnan and Easa 1986). Thus, animals with large home ranges can be expected to continue crossing plantations even if forest fragments are lost. Forest fragments may, in fact, serve as refugia for wandering carnivores and herds of elephants thereby reducing their contact with humans. Besides their importance as habitat for wildlife, forest fragments offer several direct and indirect benefits to the local human population. They provide commercial benefits to the private owners through the cardamom crop and as sources of timber. These fragments are the only sources of free firewood for the dense population of plantation workers living in their immediate vicinity. Indirect benefits of forest fragments to humans include temperature regulation, water conservation, and prevention of soil erosion. India's forest resources, like those in many other tropical countries, will become increasingly fragmented under pressures of expanding human populations and their demands. In such a milieu, conservation efforts must be broadened to include involvement by local people. Conservation goals should include satisfaction of basic human needs for fuelwood and sources of livelihood together with protection of wildlife and its habitat. The protection and management of already existing forest fragments in the Western Ghats is an attractive alternative both for biodiversity conservation and for use by local humans. 89

Management alternatives for isolated lion-tailed macaques and their fragmented habitats can be broadly categorized into the following three possibilities: (1). no action, (2). abandoning all fragments and translocating the lion-tailed macaques, and (3). full protection and intensive management of all forest fragments. Consequences of the decision to take no action would include loss of most forest fragments and approximately half of the wild lion-tailed macaque population possibly by the end of this century. The decision to abandon all fragments has the advantage of attempting to mitigate the anticipated loss of isolated lion-tailed macaque groups. This decision involves identification of protected forests that would support lion-tailed macaques and translocation of lion-tailed macaque groups from isolated fragments to such forests. This alternative faces constraints such as the dearth of suitable habitat for translocation and mortality risks inherent in the process of capture and translocation and during the critical period immediately following transfer. The third option involves protection of fragments and active management of vegetation and lion-tailed macaques in the fragments. Protection and management of all isolated fragments is probably an unrealistic expectation because of financial constraints that would limit the level of protection and management efforts. A combination of the second and third options will maximize investment of limited financial resources into those fragments most likely to survive and support resident wildlife populations including the lion-tailed macaque. I recommend the following steps to meet this objective: 1. A team of ecologists, anthropologists, and local wildlife managers should be formed to rapidly locate isolated forest fragments and assess their viability. The following criteria should be used to determine fragment viability: 90

density of keystone plant resources of lion-tailed macaques to estimate carrying capacity of the forest fragment, plant and animal species diversity in the fragment, extent of soil degradation to assess regeneration conditions for vegetation, fragment size and edge-area ratios, and importance of the fragment for local fuelwood needs. The forest fragments should be ranked according to priority for conservation efforts. 2. Those forest fragments that are determined to be viable should be purchased/acquired immediately by the state governments or made available for purchase by philanthropic, conservation, or other non-governmental organizations. 3. Buffer zones of fast-growing native plant species should be planted along fragment edges. Coupled with regulated distribution of fuelwood, these buffer zones would serve the dual purpose of reducing damage along fragment edges and providing the local people with a much-needed source of fuel. This would win the support of local people for voluntary protection of the fragment. 4. Lion-tailed macaques in fragments that are determined to be non-viable should be captured and translocated to protected forests. Lion-tailed macaques within viable fragments should be periodically monitored and management actions taken as the need arises. For example, the death of the adult male lion-tailed macaque in the absence of suitable replacement would have serious implications for the single-male, multi-female lion-tailed macaque groups. Furthermore, occasional transfer of males in or out of isolated groups may be required to maintain genetic diversity. Recent advances in genetic testing on hair and fecal samples should be attempted to 91

determine the amount of inbreeding depression in isolated lion-tailed macaque groups. 5. Regeneration of desirable climax plant species should be closely monitored. Presence of a species in a forest remnant is no guarantee of its continued existence without successful reproduction and recruitment; managers therefore need to examine the age structure of species in forest fragments to identify vulnerable species to be targeted for special management (Saunders et al. 1991). Some keystone plant species for the lion-tailed macaques may have few or no regenerating seedlings and would have to be reintroduced into the fragment. 6. Research stations should be established in some fragments where local and international researchers can do academic and management oriented research into forest fragment and wildlife conservation. Data on long-term responses of wildlife and vegetation to different logging practices are sorely lacking. Any logging operations in forest fragments should have a research component added to collect such data. The importance of accurate maps and potential uses of GIS should be realized and employed in research and effective implementation of management plans. Income generated from charges to research scientists using the facilities and information generated will both contribute to future conservation of the forest fragments. 7. The infrastructure necessary for ecotourism should be developed for some fragments such as Puthuthotam Cardamom Forest. Puthuthotam estate has been in decline for the past few years and in essence is a no longer a productive proposition (Ahimaaz 1992). The plight of estate workers is bad; several of them have been unemployed for several years and receive no compensation from the management. Ecotourism offers the following 92

benefits: generation of revenue that can be put into conservation efforts; increased local, national, and international awareness; and provision of alternative sources of income to locals several of whom are severely underpaid or unemployed. Uncontrolled tourism does have a negative side where it becomes the agent of disturbance and damage to wildlife and wild habitats and precautionary measures will have to be taken to prevent these.

O f all the endangered non-human primate species in India, the lion-tailed macaque is the only one that occurs nowhere else in the wild and it should be declared a national treasure. APPENDIX A

Plant Species in vegetation sample at Puthuthotam Cardamom Forest

Anacardiaceae Elaeagnaeceae Holigarna arnottiana Elaeagnus conferta Semecarpus travancorica Euphorbiaceae M angifera indica ?Brynia spp. Aristolochiaceae Antidesma menasu Thottea siliquosa Croton reticulatns Bignonaceae Croton spp. Oroxylum indictim Glocbidion ellipticum Burseraceae Macaranga peltata (=roxburghii) Canarium strictum Mailotus tetracoccus (=albus) Calastracaceae Phyllanthus spp. Bhesa (=Kurrima) indica Flacourtiaceae/Bixaceae Combretaceae Flacourtia montana ?Unidentified Gesneriaceae Cornaceae Aescbynanthus perrottetii Mastixia arborea G uttiferae Ebenaceae Mesua ferrea Diospyros sylvatica Icacinaceae D. spp. ?Stemonercu$ (= Gomphandra) Elaeocarpaceae tetrandnts (=polymorpha) Elaeocarpus m unronii E. spp.

93 94

Lauraceae Mvristicaceae IBeilschmieda wightii M yristica (= Knema) attenuata Actinodaphne bourdillonii Myrsinaceae Cinnamomttm malabatrum Ardesia sonchifolia (=villosa) Cryptocarya bourdillonii Maesa perrotettiana C. lawsonii Mvrtaceae C. ?beddomei Syzigium (=Eugenia) lactum (=lacta) Litseacoriaceae Oleaceae L. deccanensis Olea dioica L, insignis Papilionaceae L. oleiodes Erythrina subumbrans (=lithosperma) Neolitsea zeylanica Rubiaceae Persea (= Machiltts) macarantha Cinchona spp. Loranthaceae Coffeaarabica Loranthus tomentus Rutaceae Malvaceae Euodia Itimur-ankenda(= roxburghiana) Cullenia exarillata (=excelsa) Limonia ?acidissima Meliaceae Vepris bilocularis Aglaia (=Lansium) anamalayana Sapindaceae Aphanamixis (=A moora) polystachya Dimocarpus (= Nephelitim) longon (=longana) Toona (=Cedrela) ciliata (-toona) Sapotaceae Trichilia (=Heynea) Chrysophyllum lanceo latum connaroides (=trijuga) Mimusops elengi Moraceae Palaquitim ellipticum Artocarpus beterophyllus (=integrifolia) Solanaceae A. hirsutus Datura stramonium Ficus beddomei Sterculiaceae F.glaberrima Sterctdia guttata F.glomerata Verbenaceae F. hispida Clerodendron viscosum F. macrocarpa Lantana camara F. microcarpa ( -retusa) Zingiberaceae F. nervosa Elettaria cardamomum F. ?tjakela Unidentified F, ?travancorica Mesopsis emini 'aghil* 'naval' mushrooms, lichens 95

Plant Species in vegetation sample at Varagaliar

Anacardiaceae Moraceae Holigamagrahami Ficus lacor (=infectoria) Annonawa? Ficus spp. Unonapannosa Myristicaceae Bignonaceae Myristica (- Knema) attenuata Oroxylum indicum Oleaceae Burseraceae Oleadioca Canariumstrictum Rutaceae Dipterocarpaceae Vepris bilocularis Dipterocarpus indictts Sapotaceae Ebenaceae Palaquium ellipticum Diospyros microphylla Sapindaceae Diospyros sylvatica Dimocarpus (= Nephelium) longon (=longana) Euphorbiaceae Filicinm decipiens Agrostistachis spp. Verbenaceae Baccaurea courtallemis Vitex pubescent Glochidion ellipticum Unidentified Ostodeszeylanica 'sembakam' Antidesma menasu 'ari naval' Flacourtiaceae/Bixaceae 'nakliamaram’ Flacourtia montana 'agni-pela' Gnetaceae 'tondu-puliyam' Gnetiim ula 'vella-kungalium' Guttiferae ’nadunari’ Calophyllum elatum (=tomentosnm) 'sholavengai' Lauraceae 'atta-parta' Actinodaphne bourdillonii 'vatta-kanni' Cinnamomum malabatrum ‘poovachi1 Cryptocarya spp. 'karavappilla' Melastomataceae 'vakkanaimaram' ?Memycylon depressum 'vilpadarai' Malvaceae 'maravacti' Salmalia malabaricum 'inallagai-inaram' Meliaceae ’manjakad' Trichilia ( =Heynea) cannaroides (=trijuga) 10 unidentified species for which no local names were available APPENDIX B

Notes on Methodology

I present, here, a brief discussion on the advantages and limitations of the instantaneous scan method of data collection used in this study. An instantaneous scan should ideally record behavior of all visible group members at the exact moment in time. In practice, the behaviors of the visible members of a group are recorded within a very short period of time and the result is assumed to approach a simultaneous sample of all the observed individuals (Altmann 1974). To keep sampling time brief and consistent, the behavioral and ecological categories that are recorded are simple, coded, clearly defined, and easily distinguishable from other categories. Each scanned individual is observed for a brief and consistent amount of time (Altmann 1974). The instantaneous scan sampling method provides a ‘snapshot’ of the activities and location of various individuals of the group that are within the observer’s field of vision. Since these snapshots are ideally taken systematically at even intervals throughout the day, they provide a statistically valid sample of the entire day’s activity.

96 97

There are, however, many limitations and sources of bias associated with this manner of data collection. Comparisons of time budgets between studies and even between two study groups within the same study are meant to be gross measures of behavioral differences (cf. Marsh 1981a). Five or six days form only a small portion of each month. Therefore, variation between observation periods at monthly intervals is only an approximation of variation between months (Marsh 1981b). The apparently arbitrary choice of a month as a sampling interval is legitimate because it enables comparison of time budgets and behavioral variables between wet and dry seasons without m aking a p rio ri assumptions about the significance of different amounts of rain (Marsh 1981b). In order to draw inferences from variation between observation periods, appropriate tests of heterogeneity of variations, such as analysis of variance, should be first conducted between and within observation periods (Marsh 1981b). This is not always possible because certain kinds of behavioral data cannot be collected as continuous variables. Moreover, data obtained from repeated observations of the same individuals or groups has been open to the criticism of being dependent data (Marsh 1981b). However, Kolmes (1984) counters this criticism by arguing that although scan-sampling records behaviors performed by the same group of individuals, the behaviors themselves are sequentially independent. Sources of bias in the instantaneous scan sampling method include differential representation of various age-sex classes in scans, differential visibility of individuals engaged in different kinds of activities, and the association between dispersion and activity (Clutton-Brock 1977). Animals engaged in activities involving motion are more easily noticed by the 98 observer. Intragroup dispersion is greater during activities like ranging and lower during activities such as feeding and resting. These two biases are in opposite directions and therefore self-correcting to some unknown extent (Kumar 1987). One effect of these two biases can be to increase the total number of animals recorded in a scan when the group is predominantly engaged in particular activities. This can be compensated by expressing the number of animals engaged in each activity category in a scan as a proportion of the total number of animals recorded in that scan. The averages of these percentages are used to calculate overall estimates (Clutton-Brock 1974). Thus, scans with few numbers of records are given equal weight to scans with relatively large numbers of records. However, the study group at Puthuthotam Cardamom Forest, was unusually large. There were 43 individuals in the study group in this study whereas the study group at Varagaliar (Kumar 1987) started at 12 in 1979 and reached the size of 24 during 1983-84. Therefore, in this study, scans with larger numbers of records were more representative of the group's behavior than those with few records. Therefore, the number of animals engaged in an activity in a scan was not divided by the number of animals recorded in that scan i.e. scans were not equally weighted. Although visibility of animals was easier at Puthuthotam Cardamom Forest due to the level of habitat degradation, the study group was considerably dispersed at certain times. Additionally, Puthuthotam Cardamom Forest had twice the amount of rainfall as Vargaliar making observations difficult or at times preventing them. The months of June, July, and August are represented by fewer scans and total records than the other 9 months in this study. However, time budgets and 99 activity patterns do not change significantly if these 3 months are excluded from the analyses.

Representation of Age-sex Classes

10 H Expected 4 0 . —. □ Observed 3 5 f 3 0 | 25 % 20 | 15 S 10 IB I ill lB~i Unidentified Adult males All Adult Subadults Juveniles Adult Adult females females females with without infants infants

Fig 28. Representation of age-sex classes in the scan samples of behavior

NN

I Expected 40 35 ^ □ Observed 30 25

20 ■ 15 10

UnidentifiedJtihik Adult males All Adult Subadults Juveniles Adult Adult females females females with without infants infants

Fig. 29. Representation of age-sex classes in the nearest neighbor data set for each scan animal. 100

An analysis of age-sex classes in the instantaneous scan sample reveals that not all classes were represented as expected i.e. the number of records of different age-sex classes in the sample were not in proportion to their representation in the study group (Fig. 28). The same trends in deviation from expected representation is seen in the nearest neighbor data set (Fig. 29). In both data sets, adult males, adult females (with and without infants combined), and subadults do not deviate significantly from expected values. Juveniles, however, are under-represented in both data sets. Also, when adult females are divided into those with and without infants, the former are underrepresented and the latter over-represented in the data sets. One explanation is that at least a few of the unidentified individuals belong to the juvenile category. Adult females with infants tended to lag behind the main group and could, therefore, have been underrepresented in the samples. Furthermore, since I could individually identify only four of the six adult females with infants, a female with infant could have been recorded as a female without infant when she was seen away from her infant. However, these deviations in representation of age-sex classes probably do not alter the conclusions about time budgets and feeding ecology of the study group at Puthuthotam Cardamom Forest. A PPEN D IX C

Spatial Analyses

Analysis of spatial data was carried out for a section of the Anaimalai Wildlife Sanctuary including areas of human habitation in the plains to the north of the sanctuary and private monoculture tea and coffee plantations to the south. The forest fragment at PCF and the protected forests at VG lie within this larger area. The following sources of spatial data were used: 1. Survey of Land Records Maps (1:50,000) ofValparai Taluk, Coimbatore district, 1987 and Pollachi Taluk, Coimbatore district, 1977 (Year of first publication 1876). 2. Landsat Thematic Mapper data (Bands 1, 2, 3, and 4) from February 25, 1990, covering an area from 76° 50' to 77° E and 10° 20' to 10° 30' N. 3. Crude map of rainforest fragment distribution in the Anaimalai region from Kumar (1987 and personal communication). 4. Knowledge of the study area from surveys and a 2 year intensive field study of the vegetation and distribution and behavior of an endangered prim ate.

101 102

Map rectifications and image processing were performed with ERDAS Version 7.4 Image Processing Software. A flowchart of the sequence of steps taken in image processing is presented in Fig. 30. The map projections were most probably Polyconic. However, since this information did not appear on all the maps, the coordinates of the maps were converted from Latitude-Longitude to a UTM projection. Eight known points on the maps were digitized in digitizer inches. A GCP (Ground Control Point) file was created for the 8 digitized points. This file contained 2 sets of coordinates for the same location: the data file coordinates in Latitude-Longitude and the map coordinates corresponding to the new grid on which the maps were being projected. The coordinate transformation command (COORDN) computed a transformation matrix with an error tolerance of 30 m RMS. The H transform option in the DIG UTIL function allowed conversion of the coordinates of the DIG files to UTM map coordinates. Next, DIGPOL was used to digitize 3 map layers, roads, rivers, and land parcels, and GRD PO L was used to convert the digitized files to GIS files. A composite map containing roads, rivers, reservoirs, and land parcel information was created with the OVERLAY command. The TM scene was obtained in 4 pieces each in a floppy disk. To facilitate processing, I produced a single composite image by digitally mosaicing together the 4 pieces using the STITCH command in ERDAS. This image was then rectified and georeferenced to the UTM coordinates of the digitized maps. Thirty points scattered around the map were identified and a GCP file of their map and file coordinates was created. A second order, nearest neighbor rectification with a 20 m maximum error tolerance was 103

iR o a d i

Survey o f G rdpol Land records Roads R oads.1GIS^- (W a te r Map, 1:50,000 M ap w ith I Urban Area D ig u til D igpo! G rd p o l p — O verlay Land- Recode ] D ig UTM Rivers Rivers.,gi4- H ie.G IS H Transform C o o rd s. | Private Estates 1 (Jrdpo^i UTM Projection Land t'arcelf.Gll G C P File ^ P arw li —|Anaimalai SanctuarJ

Parambikulam Sane]

W ater

Urban Unvegctated Areas LandiatTM 90 Floppy Disk 1 Unvegetated Areas In Sanctuaries LandsatTM 90 Floppy D isk 2 Stitch Recode ■|TM .LAN| -^TM .GISj *''^ ltr | Cluitcr.CjISl1 Vegetation in Urban and Private Areas N Rectify LandsatTM 90 Floppy D isk ;t Vegetation in Urban, Private and Sanctuary Areas LandsatTM 90 Floppy D isk 4 Vegaution Mainly in Sanctuaries

Isolated Points

| W ater

Unvegetated Areas | Landuse.GIS —

Cluster.GIS |— | Natural Vegetation (Fammbikulain SanctuarjJ

L' | Natural Vegetation (Anaimalai Sanctuary) |

Fig. 30. Flowchart for spatial analysis. 104 used. Rectification was done to correct spatial distortions in the image and to link the image to a specific coordinate system. Using the same coordinate system for the image and the maps allows these data to be compared and analyzed in later steps. Band ratioing was done to better delineate vegetation and general topographical features in the scene. Ratio images involve the division of digital values in one spectral band by the corresponding values in another band. A ratioed image effectively compensates for the brightness variation caused by varying topography {such as vegetation in sunlit versus shadowed sides of a ridge) and emphasizes the color content of the data (Lillesand and Kiefer 1987). The following ratios were performed on the image: 1. T M 3/T M 4 2. TM1+TM2+TM3+TM4 Certain mathematical combinations of spectral values in different bands are found to be sensitive indicators of the presence and condition of green vegetation and are referred to as vegetation indices (Lillesand and Kiefer 1987). One such index is the Transformed Vegetation Index (TVI) given by the equation: TVI = [{(TM4-TM3)/(TM4+TM3)} + 0.5]0*5 * 100. The maps obtained from band ratioing and the TVI were useful in delineating natural vegetation. They did not assist in further classification of the natural vegetation into different types and were, therefore, not used in subsequent steps. The next step was to classify the satellite image to produce a thematic map o f various landcover classes. Classification is usually done by spectral pattern recognition and may be supervised or unsupervised. Supervised 105 classification involves identification of several training sets for different land-use or vegetation types based on ground-truthing or aerial photographs to confirm the acccuracy of the training sets. Aerial photographs were not available for the area, and detailed gound-truthing was beyond the scope of the project because of the time and expense involved in a return to the field. Therefore, unsupervised image classification was performed. Unsupervised classification employs a clustering algorithm to separate the scene into the specified number of pattern classes. The CLUSTR command, in ERDAS, for sequential unsupervised clustering using the minimum-distance classifier, was used to obtain 100 land-cover classes. Following the method employed by Zeff and Merry (1993), for each of the 100 clusters the spectral means in TM Band 4 (near infra-red band ranging from 0.76 to 0.9 pm) were plotted against those in TM Band 3 (red band ranging from 0.63 to 0.69). These two bands produce the most contrast between land-cover features (Jensen 1986). Points belonging to the same land-cover type are expected to be close to each other in such a plot. Each point on the plot was examined against the backdrop of the original TM image. Based on knowledge of the study area and the spatial context of the 100 classes, the points on the plot were lumped to form distinct classes representing major land-cover types in the study area. A GIS file, Cluster, was created containing 7 classes. The map of roads, rivers, and parcels was recoded into a new GIS file, Land-Use. The MATRIX command was carried out using the Cluster file as the column file and the Land-Use file as the row file. The MATRIX procedure in ERDAS is used to combine 2 GIS files to produce a new one. The new file contains information about how the class values from the 106

original GIS files overlap. A unique class value is assigned in the new map for each coincidence of input class values from the original files. The map obtained from the MATRIX procedure was recoded into 5 classes. The SCAN command was used to smooth out the different classes and produce the final map. A circular scanning window of 5 pixel radius was used together with the “Majority” option. The SCAN procedure in ERDAS is used to filter a CIS file. Every pixel is analyzed spatially according to the pixels surrounding it. The “Majority” option assigns to each pixel the class value that represents majority of the class values in the scanning window. This option operates like a low frequency filter to clean up a “salt and pepper” GIS file (ERDAS Field Guide 1990). An overlay of the roads, rivers, and parcels files produced the Land- Use.GIS file (Fig. 31) containing the following 6 classes: 1. Roads, 2. Water, 3. Urban areas, 4. Private estates, 5. Anaimalai sanctuary, and 6. the adjoining Parambikulam sanctuary. The plot of mean spectral values of 100 unsupervised classes in TM band 3 vs. TM band 4 is provided in Fig. 32. A new file, Cluster.GIS, was generated from the clumped classes in Fig. 33 and recoded to contain the following 7 classes: 1. Water, 2. Urban unvegetated areas, 3. Unvegetated areas within the sanctuary, 4. Vegetation occurring in urban areas and private estates, 5. Vegetation occurring in urban areas, private estates, and sanctuary, 6. Vegetation occurring mainly within the sanctuary, and 7. Isolated points not corresponding to any specific land- cover class. The MATRIX command produced a new file on which the SCAN operation was performed producing the final file, Map.GIS (Fig. 34). Map.GIS was recoded into the following 5 classes: 1. Roads, 2. Water, 107

| | Water | | Private Estates jParambikulam Sanctuary Urban Areas (Anaimalai Sanctuary |\]Roads

Fig. 31. Overlay of 3 GIS files: roads, rivers, and parcels. Scale: 1 inch = 4 km. 108

120 -i Isolated Poinu

100 - Vegetation in Urban, Private, and Sanctuary Altai 80 - ji

Vegetation in Sanctuaries

Isolated Points

20 ■ W ater

10 20 30 40 7060 80 90

Fig. 32. Mean spectral values of 100 classes in TM bands 3 and 4.

3. Unvegetated areas, 4. Cultivated vegetation, and 5. Natural vegetation. The category natural vegetation includes all non-cultivated vegetation in urban areas, private estates, and in protected areas. One of the most useful ecological applications of Landsat data is in the production of accurate and up-to-date vegetation maps which are acutely lacking for most tropical forests and several temperate forests. This can be done by unsupervised image classification, supervised classification, or by visual interpretation of the satellite imagery. The success of image classification depends on whether different land-use and land-cover types have distinctive spectral signatures (Scott et al. 1993). No classification system devised is equally suitable to all applications even within one subdiscipline, such as wildlife management (Parker 1977). Although supervised classification has been successfully applied in mapping urban and agricultural features, it has proved generally unsuccessful in mapping 109

□ W ater Vegetation _ _ I(Anaimalai Sanctuary) Vegetation □ Cultivated Vegetation (Parambikulam Sanctuary) Unvegetated Areas " \ | Roads

Fig. 33. Cluster.GIS containing 7 landcover classes. The white rectangle at the bottom-center of the map is an area of faulty data in the original TM image which was masked. 110

Natural Vegetation ^"(A naim alai Sanctuary)

Fig. 34. Final map with Five land-use classes in a portion of the Anaimalai Sanctuary. The white rectangle at the bottom-center of the map is an area of faulty data in the original TM image which was masked. Ill natural vegetation due to the inherent spectral heterogeneity in natural vegetation (Scott et al. 1993). Visual interpretation of satellite data involves delineation of polygon boundaries on printed false-color composites of the image based on perceived differences in image tone, texture, and context. However, although polygon boundaries can often be discerned with high accuracy, polygon labeling has to be done with the aid of recent aerial photographs, reliable land-use and land-cover maps, or field studies (Scott et al. 1993). Due to the unavailability of good topographic maps and aerial photographs, and absence of ground-truthing, further classification of the natural vegetation landcover in the study area into different types of vegetation was not possible. Such classifications are facilitated by soil, slope, and recent topographic maps of the region which are either not available or are not easily accessible. In a status report on the management of national parks and sanctuaries in India, Kothari et al. (1989) note that 38 of the 43 national parks and 146 of the 191 sanctuaries answering their questionnaire reported the availability of a map for their area or sent a map along with the questionnaire. However, a large proportion of the maps had only the barest details, such as sanctuary or park boundary, streams, and rivers. Furthermore, there were few copies of the maps available which restricted their availability to visitors and researchers (Kothari et al. 1989). Local wildlife staff members lacked access to Survey of India topographic sheets relevant to their area. The absence, or inaccessibility, of good topographic and thematic maps is a stumbling block in the development and effective implementation of management plans for the region. REFERENCES CITED

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