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Review Journal of Threatened Taxa A history of primatology in (In memory of Professor Sheo Dan Singh) – Mewa Singh, Mridula Singh, Honnavalli N. Kumara, Dilip Chetry & Santanu Building evidence for conservaton globally Mahato, Pp. 16715–16735 DOI: htps://doi.org/10.11609/jot.6524.12.13.16715-16735 www.threatenedtaxa.org ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

Communicatons Review University campuses can contribute to wildlife conservaton in urbanizing regions: a case study from Nigeria – Iliyasu Simon, Jennifer Che & Lynne R. Baker, Pp. 16736–16741 DOI: htps://doi.org/10.11609/jot.6316.12.13.16736-16741 A history of primatology in India (In memory of Professor Sheo Dan Singh) Killer Whale Orcinus orca (Linnaeus, 1758) (Mammalia: Cetartodactyla: Delphinidae) predaton on Sperm Whales Physeter macrocephalus Linnaeus, 1758 (Mammalia: Cetartodactyla: Physeteridae) in the , Mewa Singh, Mridula Singh, Honnavalli N. Kumara, Dilip Chetry & Santanu Mahato Sri Lanka – Ranil P. Nanayakkara, Andrew Suton, Philip Hoare & Thomas A. Jeferson, Pp. 26 September 2020 | Vol. 12 | No. 13 | Pages: 16715–16735 16742–16751 DOI: htps://doi.org/10.11609/jot.5006.12.13.16742-16751 DOI: 10.11609/jot.6524.12.13.16715-16735

The Critcally Endangered White-rumped Vulture Gyps bengalensis in Sigur Plateau, , India: Populaton, breeding ecology, and threats – Arockianathan Samson & Balasundaram Ramakrishnan, Pp. 16752–16763 DOI: htps://doi.org/10.11609/jot.3034.12.13.16752-16763

Avifauna of Saurashtra University Campus, Rajkot, , India – Varsha Trivedi & Sanjay Vaghela, Pp. 16764–16774 DOI: htps://doi.org/10.11609/jot.5113.12.13.16764-16774

Five new species of trap-door spiders (Araneae: Mygalomorphae: Idiopidae) from India – Manju Siliwal, Rajshekhar Hippargi, Archana Yadav & Dolly Kumar, Pp. 16775– 16794 DOI: htps://doi.org/10.11609/jot.5434.12.13.16775-16794

Rapid mult-taxa assessment around Dhamapur Lake (Sindhudurg, , India) using citzen science reveals signifcant odonate records – Neha Mujumdar, Dataprasad Sawant, Amila Sumanapala, Parag Rangnekar & Pankaj Koparde, Pp. 16795–16818 DOI: htps://doi.org/10.11609/jot.6028.12.13.16795-16818

Commercially and medicinally signifcant aquatc macrophytes: potental for improving livelihood security of indigenous communites in northern , India – Shailendra Raut, Nishikant Gupta, Mark Everard & Indu Shekhar Singh, Pp. 16819– 16830 DOI: htps://doi.org/10.11609/jot.5640.12.13.16819-16830

Leaf nutrients of two Cycas L. species contrast among in situ and ex situ locatons – Thomas E. Marler & Anders J. Lindström, Pp. 16831–16839 For Focus, Scope, Aims, Policies, and Guidelines visit htps://threatenedtaxa.org/index.php/JoTT/about/editorialPolicies#custom-0 DOI: htps://doi.org/10.11609/jot.6205.12.13.16831-16839 For Artcle Submission Guidelines, visit htps://threatenedtaxa.org/index.php/JoTT/about/submissions#onlineSubmissions Contributon to the Macromycetes of , India: 69–73 For Policies against Scientfc Misconduct, visit htps://threatenedtaxa.org/index.php/JoTT/about/editorialPolicies#custom-2 – Diptosh Das, Prakash Pradhan, Debal Ray, Anirban Roy & Krishnendu Acharya, Pp. For reprints, contact 16840–16853 DOI: htps://doi.org/10.11609/jot.4896.12.13.16840-16853

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Journal of Threatened Taxa | www.threatenedtaxa.org | 26 September 2020 | 12(13): 16715–16735 ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) PLATINUM OPEN ACCESS DOI: htps://doi.org/10.11609/jot.6524.12.13.16715-16735

#6524 | Received 04 August 2020 | Final received 07 September 2020 | Finally accepted 17 September 2020

R e v A history of primatology in India i e (In memory of Professor Sheo Dan Singh) w

Mewa Singh 1 , Mridula Singh 2 , Honnavalli N. Kumara 3 , Dilip Chetry 4 & Santanu Mahato 5

1,5 Biopsychology Laboratory, Insttuton of Excellence, Vijnana Bhavan, University of Mysore, Manasagangothri, Mysuru, 570006, India. 1 Zoo Outreach Organizaton, No. 12, Thiruvannamalai Nagar, Saravanampat - Kalapat Road, Saravanampat, , 641035, India. 2 Department of Psychology, Maharaja’s College, University of Mysore, Mysuru, Karnataka 570005, India. 3 Salim Ali Centre for Ornithology and Natural History, Anaikat P.O., Coimbatore, Tamil Nadu 641108, India. 4 Aaranyak, 13 Tayab Ali Byelane, Guwahat, 781028, India. 1 [email protected] (corresponding author), 2 [email protected], 3 [email protected], 4 [email protected], 5 [email protected]

Abstract: India harbors a wide diversity of with 24 species that include lorises, , langurs and gibbons. Systematc research on the primates in India started about 60 years ago. In order to develop a historical perspectve, we recognize three broad phases of research: largely natural history and base line research, primarily behavioral ecology research, and increasingly queston and hypothesis-driven research. We describe the old and the recent primate research in the country and suggest research areas for the future.

Keywords: Gibbon, Indian primates, langur, loris, .

Editor: Anonymity requested. Date of publicaton: 26 September 2020 (online & print)

Citaton: Singh, M., M. Singh, H.N. Kumara, D. Chetry & S. Mahato (2020). A history of primatology in India (In memory of Professor Sheo Dan Singh). Journal of Threatened Taxa 12(13): 16715–16735. htps://doi.org/10.11609/jot.6524.12.13.16715-16735

Copyright: © Singh et al. 2020. Creatve Commons Atributon 4.0 Internatonal License. JoTT allows unrestricted use, reproducton, and distributon of this artcle in any medium by providing adequate credit to the author(s) and the source of publicaton.

Funding: SERB Distnguished Fellowship (Award Number: SB/S9/YSCP/SERB-DF/2018(1)).

Competng interests: The authors declare no competng interests.

Author details: Mewa Singh is a renowned wildlife biologist at University of Mysore with over 45 years of research experience on Indian wildlife, especially primates. His interests are ecology, behavior and conservaton management. He has trained a generaton of students who are now excelling in many insttutons all over the world. Mridula Singh is a PhD in animal behavior and teaches psychology at Maharaja’s College, University of Mysore. She carried out a long-term feld study on maternal parental investment paterns in relaton to dominance hierarchy in Lion-tailed Macaques in the Anamalai Hills. H.N. Kumara is principal scientst at Salim Ali Centre for Ornithology and Natural History. His research interests encompass a large variety of taxa and topics in ecology, behavior and conservaton. He initated the frst long-term studies on the Nicobar Long-tailed Macaques. Dilip Chetry is a senior biologist with Aaranyak and carries out conservaton research and acton for wildlife, especially primates, in northeastern India. He also trains people in ecological research through various workshops and training programs. Santanu Mahato is a researcher at the University of Mysore and will be startng long-term studies on primates in the Anamalai Hills soon.

Author contributon: Mewa Singh conceptualized the artcle and other authors contributed to literature search and helped in artcle preparaton.

Acknowledgements: Mewa Singh acknowledges the Science and Engineering Research Board, Government of India for the award of a SERB Distnguished Fellowship (Award Number: SB/S9/YSCP/SERB-DF/2018(1)) which also supports Santanu Mahato. The authors acknowledge the assistance of Sayantan Das in preparing the fgure in the artcle.

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INTRODUCTION India to develop a historical perspectve. These include: a largely, natural history and baseline research; primarily India is home to at least 24 species of nonhuman behavioral ecology research; and increasingly queston primates that include two species of lorises, 10 species and hypothesis driven research. It must, however, be of langurs, 10 species of macaques, and two species of made clear that these are not strictly discrete phases as small apes, with several of them being threatened or all types of research have been carried out at all tmes, vulnerable (Table 1) (Molur et al. 2003). Several species but it is only a working tme frame to trace the research including the Slender Loris, , and perspectves and developments. Further, since there has Assamese Macaque also have distnct subspecies making not been any insttuton in the country with the primary India very rich in primate taxa. Due to this diversity, focus on primate research and conservaton, most research in primatology in India has made prominent research has been individual initatves scatered across strides. Although in the past, many researchers came diferent insttutons. In this review, therefore, we will to India from other countries and carried out pioneering primarily refer to Ph.D. theses and only few references, studies on diferent primate taxa, most of the published and website(s) wherever available, of each researcher research on Indian primates today is by Indian scientsts, since the total number of references on Indian primates unlike in many other habitat countries. At a conference number in thousands. This will help readers to fnd organized by the Associaton of Indian Primatologists, other publicatons of the same researchers. To our a group recently established by young primatology knowledge, we have included all Ph.D. theses writen researchers, at Bengaluru in 2019, one of us (Mewa Singh) on primates in India, either by an Indian or by foreign delivered the inaugural address on the development of researchers. We clarify that this artcle is not a critcal primatology in India, and was surprised to know that review of the contributons of Indian primatology, which many young scholars had not even heard the names of remains another topic to work on, but a repository of many of the pioneers. This fact motvated us to prepare informaton on the subject. It is, however, more than this artcle so that the historical informaton is made an annotated bibliography as the contributons of all available at one source before such valuable informaton the scientsts are discussed briefy. A brief tmeline of becomes inaccessible or is forgoten. This will also crucial developments in Indian primatology is illustrated provide a historical perspectve for the development of in Figure 1. primate research in India. Primates have been used in a wide variety of research in India. Classical paleontological and evolutonary studies have been carried out by A LARGELY NATURAL HISTORY AND BASELINE RESEARCH S.R.K. Chopra and Ashok Sahni at Panjab University, Chandigarh. Research on primate physiology has been Although a few short artcles on Indian primates in carried out by N.K. Lohia at University of , their natural habitats were published earlier (McCann Jaipur, by N.R. Moudgal, M.R.N. Prasad, N. Ravindranath 1933; Nolte 1955), it was only in the late 1950s and early and Polani B. Sheshagiri at the Indian Insttute of 1960s that systematc and relatvely long-term studies Science, Bengaluru, and by scientsts at the Natonal were taken up. The main emphasis in those studies Insttute for Research in Reproductve Health, Mumbai. was on the natural history of the study species, and At the Indian Insttute of Science, Aditya Murthy studies these descriptve studies provided a baseline for further neurophysiology of primates and Praveen Karanth works research. The researchers, therefore, focused on the on primate . In additon, primates have also species rather than on specifc research questons. The been employed for biomedical research at the Central pioneers of that era are Charles H. Southwick, Phyllis Drug Research Insttute, Lucknow, Natonal Insttute Jay Dolhinow, Paul E. Simonds, Yukimaru Sugiyama, and of Nutriton, Hyderabad, Natonal Insttute of Virology, Sheo Dan Singh. Pune, Post Graduate Insttute of Medical Educaton and Research, Chandigarh, Natonal Insttute of Immunology, Charles H. Southwick and All India Insttute of Medical Sciences, New Delhi. Although short descriptons of the behavor of Rhesus In this artcle, however, we will focus only on research Macaques and Hanuman Langurs were published before on primates in their natural habitats concerning ecology 1959 (McCann 1933; Nolte 1955), the frst long-term and behavior. feld study on the distributon, populaton status, ecology As it could probably be true for primatology globally, and behavior was undertaken by Charles H. Southwick we recognize three phases of primatology research in (https://neurotree.org/neurotree/publications.php?p

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id=56382&searchstring=&showflter=all). Southwick conducted his studies from 1959 to 60 in Utar Pradesh and in 1962 in West Bengal. The results of the surveys conducted on the Rhesus Macaque populatons in 280 villages, 30 towns, and 200 Hindu temples revealed that 16% villages, 69% towns and 6% Hindu temples had Rhesus Macaques (Southwick et al. 1961a), and 406, 37, 84 and seven groups of macaques were located along roadsides, canal banks, railroads, and forest areas, respectvely (Southwick et al. 1961b). The frst detailed study on the ecology and behavior of the Rhesus Macaque was published in 1965 (Southwick et al. 1965). The group size was 15.1 individuals with an average of 3.5, 6.9, 3.8, and 0.9 adult males, adult females, infants, and juveniles, respectvely. The study found a clear inter-group and within-group hierarchy and peaceful to violent inter-individual relatonships. This study also laid foundatons for further research in ecology and behavior of Rhesus Macaques. Southwick maintained his research collaboratons with Indian primatologists and contnued to publish on populaton trends in Rhesus Macaques untl 2001 (Malik et al. 1984; Southwick & Siddiqi 1966, 1977, 1988, 2001).

Phyllis Jay Dolhinow Though there were reports about the taxonomy of Hanuman Langurs, the frst comprehensive feld study on the demography, ecology and behavior of this species was published in 1965 by Phyllis Jay (Jay 1965, later known as Dolhinow, P.J.). Jay conducted an 18-month feld study between October 1958 and April 1960 at Orcha, a small village close to a forest in , and Kaukori, a village among agricultural lands in Utar Pradesh. The two habitats of langurs varied with respect to availability of forest, food items of langurs, human density and human interactons, interactons with other primates and forest dwelling including predators. The groups were largely bisexual with 1.5 to 2 adult females per male, but there were also all-male groups. The groups remained relatvely stable. Actvity peaked during dawn and dusk. The home ranges of the groups averaged from 1 to 3 square miles. The social interactons were usually peaceful and relaxed. There were no overt dominance interactons and actvites such as grooming could occur for over fve hours per day. The infants and juveniles were found to undergo a fairly long process of socializaton. Jay also provided a detailed ethogram of langurs describing a multtude of gestures and vocalizatons used by various age-sex classes. Dolhinow also carried out a survey on

Figure 1. Illustraton of the developments in Indian primatology with solid grey lines depictng initaton of feld research by seminal primatologists pre-1980 and establishment of independent research research independent of establishment and pre-1980 primatologists seminal by research feld of initaton depictng lines grey solid with primatology Indian in developments the of Illustraton 1. Figure groups focussed on primatology post-1990. Doted red lines mark organizaton of crucial meetngs, symposia and conferences and the same decade. belong to doted when they identcal on the tmeline) are mark individual year Colour designs of dots (that red projects. arrows trace the period of internatonal collaboratve Rhesus Macaques from 1964 to 1965 covering 9,510

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Table 1. Primate species in India with their IUCN and Indian Wildlife (Protecton) Act status.

Conservaton status

Common name Species IUCN IW(P)A

Lorises

1 Slender Loris Loris lydekkerianus EN I*

2 Slow Loris Nyctcebus bengalensis EN I

Langurs

3 Himalayan Grey Langur Semnopithecus ajax EN II

4 Bengal Hanuman Langur Semnopithecus entellus LC II

5 Lesser Hill Langur Semnopithecus hector NT II

6 Dark-legged Malabar Langur Semnopithecus hypoleucos LC II

7 Coromandal Grey Langur Semnopithecus priam NT II

8 Central Himalayan Langur Semnopithecus schistaceus LC II

9 Semnopithecus johnii VU I

10 Capped Langur Trachypithecus pileatus VU I

11 Golden Langur Trachypithecus geei EN I

12 Phayre’s Leaf Monkey Trachypithecus phayrei EN I

Macaques

13 Rhesus Macaque Macaca mulata LC II

14 Assamese Macaque Macaca assamensis NT II

15 Stump-tailed Macaque Macaca arctoides VU II

16 Pig-tailed Macaque Macaca leonina VU II

17 Arunachal Macaque Macaca munzala EN Not listed

18 White-cheeked Macaque Macaca luecongenys Unknown Not listed

19 Macaca thibetana NT Not listed

20 Lion-tailed Macaque Macaca silenus EN I

21 Bonnet Macaque Macaca radiata VU II

22 Nicobar Long-tailed Macaque Macaca fascicularis umbrosus VU I

Gibbons

23 Eastern Hoolock Gibbon Hoolock leuconedys VU I

24 Western Hoolock Gibbon Hoolock hoolock EN I

*Schedule types under IW(P)A. Lower schedule types have higher protecton than higher schedule types.

kilometers in rural and forest areas, and found one spending much tme in grooming and play. Hierarchies group per 11.6km in forests and 32.8km in rural areas within the group were found to be subtle. Females (Dolhinow & Lindburg 1978). indulged in promiscuous matng with extreme male tolerance. Groups had overlapping home ranges. Paul E. Simonds Simonds later visited India in 1963, 1970, and 1975 for Afer a short-term study by Nolte (1955), the frst subsequent research. Simonds (1965) became a base detailed study on the behavior of Bonnet Macaques line for subsequent studies on Bonnet Macaques. was published by Simonds (1965). Paul Simonds (https://anthropology.uoregon.edu/profile/simonds/) Yukimaru Sugiyama conducted his study on Bonnet Macaque between A contemporary of Phyllis Jay Dolhinow was October 1961 and June 1962 on the Mysore-Ooty Yukimaru Sugiyama (htps://www.researchgate.net/ road near Bandipur forests. The group size of Bonnet profle/Yukimaru_Sugiyama) from Kyoto University, Macaques varied from >10 to 40–60 individuals. Kyoto who conducted studies on Hanuman Langurs Simonds described the macaques as gregarious and and Bonnet Macaques in Dharwad from April 1961 to

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April 1963. Dr. D. Miyadi, Kyoto University, Kyoto was and in peer-groups of six monkeys each. There were the project director and the Indian collaborator was Dr. also mother-infant pairs in these experiments. The idea M.D. Parthasarathy from Central College, Bangalore. behind these experiments was to study the contributon Sugiyama reported the social structure of one-male of mothers, peers, and the group on the behavioral groups and instances of infantcide in Hanuman Langurs development of infant monkeys. Infants raised in peer for the frst tme and proposed that such a pathological groups when separated from others showed disturbed behavior could be due to high populaton densites, and behaviors including ‘protest’ and ‘despair’ (Singh 1977). could occur to induce estrus in females (Sugiyama 1964, When these infants were exposed to strangers, they 1965, 1966). The process involved oustng the dominant reacted aggressively just like the Rhesus Macaques in male and other males of a langur group by an outside their natural groups (Singh 1980). S.D. Singh passed male who took over the group, and the ousted males away on 26 June 1979 when he was less than 50 years formed an all-male group. He also carried out short- old. The students trained by him contnued their work term studies on Lion-tailed Macaques (Sugiyama 1968) and three of them, Waheeda Khan, Kiran Lata, and and Bonnet Macaques (Sugiyama 1971) during the Suraj Bhan, submited their theses under B.S. Gupta afore-mentoned period. Later, Sugiyama made several at Meerut. Waheeda Khan (Khan 1980) studied the visits to India in 1972–73 to study Himalayan langurs efect of urban environment on social, emotonal, and (Sugiyama 1976) and Rhesus Macaques, and in 1976, curiosity behaviors in Rhesus Macaques at diferent 1978, and 1997 to study Hanuman Langurs (Sugiyama age levels. The urban monkeys were more aggressive 1984). whereas the forest monkeys were shy and tmid. Urban monkeys also needed high levels of stmulaton Sheo Dan Singh to satate their curiosity. The same urban and forest When these feld studies were going on in the 1960s, monkeys were studied by Kiran Lata (Lata 1980) for there was also experimental work carried out on Rhesus development of cognitve behaviour. Environmental Macaques at the Department of Psychology, Punjab enrichment showed no efect on delayed response University, Chandigarh by Sheo Dan Singh that resulted learning, object discriminaton learning set, and oddity in some exceptonal fndings which unfortunately have discriminaton learning; however, the forest monkeys not found much menton in later literature on Indian were slower to adapt to experimental situatons than primates. Singh captured monkeys from urban and the urban monkeys. The above two studies were carried forest environments and studied behavioral diferences out in the Primate Laboratory at Meerut University. A between them under laboratory conditons. When feld study carried out by Suraj Bhan (Bhan 1983) on presented with novel stmuli, the urban monkeys were the development of behaviour in Rhesus Macaque far more actve and manipulated presented objects more infants found developmental trends in play and social than the forest monkeys (Singh & Manocha 1966) and interactons at diferent age blocks, with male infants were more responsive to stmuli of higher complexity and juveniles being more playful than females. N.K. (Singh 1968). Urban monkeys were more aggressive than Chandel (Chandel 1981), though trained at Meerut, forest monkeys whereas forest monkeys groomed more submited his thesis at the University of Rajasthan, ofen than urban monkeys (Singh 1969). The urban and reported that paterns of afliatve behaviors in adult forest monkeys, however, did not difer in their problem- and adolescent Rhesus Macaques changed with age and solving capabilites (Singh 1969) indicatng that whereas matng season. Rajbir Singh (Singh 1981), who also went the selecton pressures of an urban environment have to Rajasthan from Meerut, reported that in free-ranging shaped many new adaptve paterns in urban monkeys, Rhesus Macaques, the social interactons among group the basic intelligence does not seem to be afected. S.D. members are not random but paterned by age, sex, and Singh was invited to head the Psychology Department relatve dominance in the social hierarchy. at the newly established university at Meerut in 1971. Two other students of S.D. Singh, Raghubir Singh At Meerut, he contnued laboratory experiments on Pirta and Mewa Singh, made their careers in primatology Rhesus Macaques primarily on selectve lobectomy and established many milestones in Indian primate and learning, and also started experimental studies on research later. Pirta did his Ph.D. at Kashi Vidyapith these macaques in their natural habitats in Siwalik Hills University, his post-doctoral research at Utkal University, in the Asarori Range forests near Dehradun. The Rhesus and later worked in University tll his Macaque infants were separated from their mothers retrement. Mewa Singh did his Ph.D. at the University shortly afer birth and were reared in social isolaton of Mysore and contnues to work there as Professor (for

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Life) (The works of Pirta and Singh are discussed in later on many Indian primates, and Poirier advanced the sectons). research to fner details of ecology and behavior. He published several artcles in various journals from 1966 to 1970 but a detailed single report on the Nilgiri Langur PRIMARILY BEHAVIORAL ECOLOGY RESEARCH was published in 1970 (Poirier 1970). The Nilgiri Langurs are distributed in the southern Indian states of , This was the phase when baseline data were available Tamil Nadu, and Karnataka. The home ranges of Nilgiri on some Indian primates and researchers could now Langur groups overlapped with Bonnet and Lion-tailed look at the ecology and behavior of their study species in macaques and occasionally with Hanuman Langurs, greater detail. The approach stll was primarily species- and varied from ¼ to 1 square mile. The average group oriented. size was 8–9 individuals in bisexual groups, and solitary and uni-sex male groups were also observed. The main Donald G. Lindburg food of Nilgiri Langurs were leaves, fowers, buds, seeds, D.G. Lindburg came to India to study Rhesus Macaques and bark or stem. Poirier also reported other behaviors in 1964. Of the 21 months that the study lasted, the including social communicaton, dominance hierarchies, frst nine months were devoted to a feld survey on grooming, and play in detail. This report, therefore, Rhesus Macaques under the leadership of Phyllis Jay provided an excellent background for analytcal studies Dolhinow and during the remaining 12 months, Lindburg later. carried out an ecological and behavioral study of Rhesus Macaque at Asarori Forest, Siwalik Hills, at the Forest Suraj Mal Mohnot Research Insttute, near Dehradun (Lindburg 1971). Though there were a few Indian primatologists Prior to this study, only a few experimental studies including M.A. Beg, M.R. Siddiqi, M.D. Parthsarathy, and were carried out on the behavior of Rhesus Macaques H. Rahman who carried out feld work with the previously- by S.D. Singh (cited earlier) and Lindburg’s was the frst mentoned foreign scientsts, the frst independent account of Rhesus Macaque behavior from its natural feld study in India was undertaken by S.M. Mohnot at habitats. The Rhesus Macaque is the most widespread Jodhpur University. Mohnot started studying Hanuman non-human primate from 200E in Afghanistan to 1200E Langurs near Jodhpur in 1965, obtained his Ph.D. in 1974 in China, and from 180N in India to 400N in China. The and became a faculty member at the Department of habitat occupied by the species included Sal Shorea Zoology at Jodhpur University. In the years since then, robusta dominated deciduous forest and the area was Mohnot has carried out many long-term studies on heavily grazed by domestc bufaloes. The mean group primates, guided several Ph.D. students, hosted many size was about 32 though it varied signifcantly from foreign scholars, and made Indian primatology known to one habitat type to another. The groups were bisexual the world over. The duo of Roonwal and Mohnot also with adult male to female rato being about 1:3 and the published the frst comprehensive book on primates reproductve rate was about 90%. The home range of a of South Asia (Roonwal & Mohnot 1977). The langur large group was approximated to 15km2. The Rhesus populaton near Jodhpur is unique in many ways, it is Macaque is largely frugivorous but occasionally feeds an isolated populaton since hundreds of years with on insects and grasshoppers. Lindburg also provided no distributon of any other primates up to at least a detailed informaton on intergroup interactons, social hundred kilometers. The rocky and open nature of the behavior, and social organizaton and his artcle became region allows close day-long observatons throughout the general reference for later research on the ecology the year (Mohnot 1984). The basic social system, social and behavior of Rhesus Macaques. Lindburg contnued changes, and infant-killing were reported in the Jodhpur his associaton with Indian primatologists (Singh et al. langurs (Mohnot 1971; Makwana 1979a). (More details 1999) and also carried out conservaton breeding of on Mohnot’s work will come later along with his students Lion-tailed Macaques at San Diego Zoo, San Diego, USA. and collaborators).

Frank E. Poirier Steven Green, Karen Minkowski, John F. Oates, Rauf Frank E. Poirier (htps://anthropology.osu.edu/ Ali, and Jayashree Ratnam people/poirier.1) studied Nilgiri Langurs in the Nilgiri Hills Steven Green and Karen Minkowski (htp://www. between September 1965 and August 1966. This was momentum2.miami.edu/donors_steven_green.html) the tme when base line studies were already published from Rockefeller University collaborated with the

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Bombay Natural History Society to carry out a long feld individual fed at a given tme, and social rank afected study lastng 19 months from September 1973 to April spatal distributon and the feeding rate of an individual. 1975 on the Lion-tailed Macaque at Ashambu Hills in Tamil Nadu. Except for a short term study by Sugiyama Ajith Kumar (1968) and Karr (1973), nothing was published about Ajith Kumar (htps://www.ncbs.res.in/mscprogram/ the ecology and behavior of Lion-tailed Macaques tll program-management) carried out a six-year-long study 1977 (Green & Minkowski 1977). Surveys by Green & on the ecology and populaton dynamics of Lion-tailed Minkowski revealed that there were about 405 Lion- Macaques in the Anamalai Hills and received his Ph.D. tailed Macaques in the wild in the states of Karnataka, degree from University of Cambridge in 1987 (Kumar Tamil Nadu, and Kerala. The Lion-tailed Macaque is an 1987). He examined variatons in the demographic obligate rainforest dweller and primarily arboreal. The parameters and correlates of ecology and behavior. He average group size is about 15 individuals. Sexual maturity reported delayed sexual maturity in females, long inter- in females is atained at about fve years of age and very birth intervals, low populaton turnovers but high infant few infants are produced by a female in her lifetme. survival rates. These macaques fed on fruits and insects. The primarily fruit diet of the species is supplemented Larger groups had larger ranges than smaller groups. The with insects. Green & Minkowski also initated long- low birth rate was explained to be due to high matng term conservaton plans for the endangered Lion-tailed interference resultng in suppression of concepton Macaque. Green ofered a post-doctoral fellowship to in many females. This study site in the Anamalai Hills John F. Oates (htps://anthropology.commons.gc.cuny. became one of the most important centers for wildlife edu/john-f-oates/) to contnue research on the Lion- research in subsequent years. tailed Macaques. The conservaton initatves taken up by Green & Oates fnally resulted in a large area in Kalakad-Mundanthurai being maintained as a reserve INCREASINGLY QUESTION-ORIENTED AND HYPOTHESIS- primarily for the Lion-tailed Macaques (Sekhsaria 2015). DRIVEN RESEARCH Oates, however, shifed his focus to the Nilgiri Langur and commented on its geographical distributon and Although several researchers contnued to focus status (Oates 1979), and reported that most food items on partcular species, primate research, from the mid of the Nilgiri Langur had low fbre and condensed tannin 1970s, became increasingly queston-oriented and content but the staple food items were highly digestble hypothesis-driven. Therefore, while discussing this (Oates et al. 1980). As John Oates took over from Steven phase of development of primatology research in India, Green, Rauf Ali (htps://www.feralindia.org/user/78) we will not focus on the research by any individual but came to Kalakad-Mundanthurai to take over from John by several insttutons where primate research was Oates. Ali was registered for Ph.D. at the University of conducted or is stll underway. Bristol With John Crook (Ali 1981) and reported for the frst tme that in the subspecies of Bonnet Macaques Jai Narayan Vyas University, Jodhpur that he was studying at Agastyamalai, it was the females S.M. Mohnot (Mohnot 1974) established a major that emigrated and dispersed between the groups, primate research program at Jodhpur where he trained a behavior not known in the other subspecies of the a large number of Indian students and hosted several Bonnet Macaque or most other macaque species. Ali researchers from abroad. The unique Jodhpur populaton also observed that at the tme of intergroup encounters, of Hanuman Langurs became a major focus of ecological there was also heightened within-group aggression that research. Sarah Blafer Hrdy (Hrdy 1975) (htp://www. he explained as the group males trying to prevent the citrona.com/hrdy), a graduate student of Irvin DeVore females from crossing over to the opposite group. Years and tutored by E.O. Wilson and Robert Trivers arrived in later, following Rauf Ali, Jayashree Ratnam (htps:// Jodhpur in 1971 and S.M. Mohnot advised her to study www.ncbs.res.in/mscprogram/program-management) Hanuman Langurs at Mount Abu in Rajasthan. Infantcide arrived at the Mundanthurai forests to study Bonnet in langurs, reported by Sugiyama and Mohnot, was just Macaques with a more pointed queston about the becoming a hot topic in primatology and it was generally foraging decisions in a social forager (Ratnam 2002). being characterized as a ‘social pathology’ occurring Ratnam analyzed foraging decisions in response to primarily due to overcrowding. Hrdy observed many experimentally-induced variatons in ecological and cases of usurping control of diferent groups of langurs social variables. Social dominance determined which and instances of infantcide. The females who lost infants

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gave birth within 6–8 months afer the death of their their natal groups and weaned sons follow their fathers infants whereas the normal inter-birth interval was up to afer oustng. Emigratng males have a higher mortality about 27 months. Infantcide could also occur in places than females resultng in female-biased sex ratos. G. where there was no overcrowding. Hrdy, therefore, Agoramoorthy (Agoramoorthy 1987) observed several proposed that infant killing was not a social pathology cases of abortons in Hanuman Langurs happening but a male reproductve strategy to garner reproductve probably due to stress afer male take-over of a group access to females. In the years to come, sexual selecton and conjectured that an aborton may be a female as an explanaton for infantcide became the main reproductve strategy to avoid infantcide afer the birth focus of research on Hanuman Langurs. In 1977, the of the infant. Since the inter-birth interval in females Indo-German project, conceptualized by Christan whose infants were killed by the new males was shorter Vogel, was launched at Jodhpur to study interrelatons than other females, it appears that the male achieves between individual life-histories and troop histories higher reproductve success by infantcide. Hanuman (Vogel 1988). This project contnued for many years and Langurs are primarily folivorous but were also observed sponsored several German researchers who worked on to spend considerable tme feeding on insects, especially Jodhpur langurs. These langurs feed on 35 species of during the monsoon season, which supported the plants, and a unique foraging ‘inventon’ is feeding on ‘energy/nutrient maximizaton’ hypothesis (Srivastava the ‘waternut’ Trapa natans recently introduced to the 1989). Carola Borries (Borries 1989) (htps://www. region. The main food items of a large number of troops stonybrook.edu/commcms/anthropology/faculty-and- were Prosopis julifora and the food provided by humans staf/borries.php), a student of Christan Vogel, and (Winkler 1981). The females reached menarche at the hosted by S.M. Mohnot, studied competton in female age of 29 months, cycled for 24 days, had gestaton langurs of Jodhpur from October 1984 to January 1986. of about 200 days, and an inter-birth interval of 15.3 She found that the young females occupied high ranks, months. About 80% infants survived for six months and the hierarchical positon of a female declined but only 35.9% completed two years of life (Winkler with age. Reproductve success was rank related that et al. 1984). Volker Sommer (Sommer 1985) (htps:// declined signifcantly from high, over middle, to low www.ucl.ac.uk/anthropology/people/academic-and- ranking females (Borries et al. 1991). In Kumbalgarh teaching-staf/volker-sommer), a student of Christan Wildlife Sanctuary, there were about 69% bisexual and Vogel at Gotngen, and hosted by S.M. Mohnot studied about 31% all-male band groups of Hanuman Langurs, the Jodhpur langurs for his Ph.D. between October and their social organizaton was the same as in the 1981 and December 1982, and again as a post-doctoral Jodhpur langurs (Chhangani 2000) (htps://mgsubikaner. research fellow in 1986–1987. He witnessed and ac.in/wp-content/uploads/2015/10/1396502908- also inferred many cases of infantcide and discussed 2014Faculty_MemberDrAnilKumarChhangani.pdf). D.S. the male reproductve strategy with respect to three Rajpurohit (Rajpurohit 2005) determined dominance conditons, postulatng that the infantcidal male is not hierarchy and its role in social organizaton in Hanuman the father of the killed infants, but actually sires the langurs, and G. Sharma (Sharma 2007) reported subsequent ofspring, and infantcide shortens inter- details of paternal care in these langurs. In 1994, S.M. birth interval, and found that these conditons were Mohnot launched a mega research project (Indo-US largely met (Sommer & Mohnot 1985). Primate Project) for a period of fve years, which was Makwana (1979b) studied rhesus monkeys in Asarori, further extended by three years. Most of the research Siwaliks, and reported group size to vary from 6 to 90 with under this project was in northeastern India where a male to female rato of 1:2.7. Home range size varied Mohnot collaborated with P.C. Bhatachrjee at Gauhat with group size, and the daily range was about 1803 m. University, Guwahat. Since most researchers were from P.R. Ojha (Ojha 1979) observed that the tail carriage of the the northeast, we will have a separate secton for the Rhesus Macaque alpha male with the basal half carried research on northeastern Indian primates. upright, distal part leaning forward and the tp forming Hanuman Langurs being widespread have also been a loop is unique and is not shown by any other member studied in diferent habitats in India. The Himalayas are in the group. The upright posture is usually assumed a diferent ecological niche and the Himalayan Hanuman in tense situatons such as intergroup interactons or Langurs, though studied at diferent alttudes in Indian alarms. L.S. Rajpurohit (Rajpurohit 1987) (htps://www. and Nepalese Himalayas, exhibit traits such as mult- researchgate.net/profle/Lal_Rajpurohit2) reported male groups which are diferent from the langurs in the that the ousted males did not achieve residency in Indian plains that are largely uni-male (Bishop 1975).

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Hanuman Langurs in are organized between the two hands (Mangalam et al. 2014, 2016). in mostly uni-male groups with an adult sex rato of Sindhu Radhakrishna (Radhakrishna 2001) (htps:// 1:7.9. In Kanha, an atack by a male band on the uni- www.nias.res.in/professor/sindhu-radhakrishna) carried male study group, resultng in killing of three infants, out a 21-month-long study on the Slender Loris in the and the tming of takeover with respect to the birth forests of Dindigul, Tamil Nadu. Most of the aggressive season, supported the ‘sexual selecton’ and not the interactons among individuals were observed in the ‘social pathology’ hypothesis (Newton 1984). At contexts of territorial defense and matng; individuals Hill ranges of Siwalik Hills, 75% groups of Hanuman communicated through chemical and vocal signals, and Langurs were uni-male with a sex rato of 1:3.2, and immigratons were frequently seen in the study area these langurs shifed home ranges seasonally (Bala (Radhakrishna & Singh 2002). Females were seen to 2013). In Bidar District of Karnataka, there were 142, be in estrus largely in October–December and births 55, and 28 uni-male, mult-male and all-male groups, occurred in April–June; gestaton lasted for 5.5 months respectvely, with a total populaton of 6,384 individuals and there was an inter-birth interval of seven months (Patl 2019). In the six study groups, the populaton size (Radhakrishna & Singh 2004). Anantha Krishna Sharma of 199 in 2012 increased to 270 in 2014. (Sharma 2002) (htps://www.researchgate.net/profle/ Ananthakrishna_Sharma) studied reproductve biology University of Delhi and Vatavaran of the Lion-tailed Macaques in the Western Ghats and Several studies were carried out on primates at reported that there were two birth peaks in January to the Department of Anthropology, University of Delhi April and in September to December, which remained by Praveen K. Seth. Though most of his work was on constant over the years and across forest fragments of primate anatomy (which is beyond the scope of this diferent sizes (Sharma et al. 2006). In contrast, births artcle), Iqbal Malik (htp://www.vatavaran.org/) and he in captve Lion-tailed Macaques occurred throughout carried out studies on populaton trends and behavior the year. H.S. Sushma (Sushma 2004) (htps://www. of Rhesus Macaques. The surveys conducted between sacon.in/adjunct-scientst/) studied sympatric Lion- 1975 and 1980 revealed that in several localites, the tailed Macaques, Nilgiri Langurs, Bonnet Macaques, and populaton of Rhesus Macaques increased by 66–79%, Indian Giant Squirrels and found that both macaque mainly due to human-provisioning (Seth & Seth 1983a; species had relatvely narrow food niche breadth and Malik et al. 1984). The populatons, however, showed high niche overlap resultng in co-occurrence of these a declining trend in the later years (Malik 1989). Malik species only in a few months of the year. On the other was also involved in translocatons of urban Rhesus hand, Nilgiri Langurs had a low niche overlap with Lion- Macaques and found that most translocated monkeys tailed Macaques and both langurs and squirrels had appeared to exhibit normal behavior (Imam et al. 2000). broad niches resultng in tolerance and co-occurrence of all three species (Sushma & Singh 2006). B.A. Krishna University of Mysore, Mysuru (Krishna 2007) studied maternal and parental investment Mewa Singh (htp://uni-mysore.ac.in/psychology/ paterns in Lion-tailed Macaques and observed that faculty/dr-mewa-singh) started research on primates weaning occurs from fve and a half months to about at the University of Mysore in 1975 and has studied nine months of age of the infant (Krishna et al. 2008) Bonnet, Rhesus, Lion-tailed, and Long-tailed macaques, and the weaning paterns indicated that the dominant Hanuman and Nilgiri langurs, and Slender Loris. The females are positvely biased towards their male group has carried out extensive studies on the ecology, ofspring (Singh et al. 2007). Kuladeep Roy (Roy 2011) behavior, and conservaton management, both in situ (https://www.researchgate.net/profile/Kuladeep_ and ex situ, on primates with over 150 publicatons Roy) also studied sympatric primates in a diferent (see htps://uni-mysore.ac.in/psychology/faculty/dr- forest and found that though there was a high niche mewa-singh). Long-term feld studies were initated overlap between Lion-tailed and Bonnet macaques for on Slender Loris (Singh et al. 1999, 2000; Kumara et about two months, it occurred at the tme of resource al. 2006) and other primates (Kumara & Singh 2004) in abundance of the shared resources, and as a result, various habitats. Populaton trends in Bonnet Macaques both of these macaques species and the Hanuman have been monitored in and around Mysuru for over 30 Langur which is a folivore, unlike frugivore macaques, years showing a nearly 70% decline in the populaton could co-exist throughout the year (Singh & Roy 2011). (Erinjery et al. 2017a). Bonnet Macaques showed Kavana T.S. (Kavana 2015) and Joseph J. Erinjery (Erinjery laterality in various actvites as a division of labor 2016) (htps://www.researchgate.net/profle/Joseph_J_

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Erinjery) studied individual diferences indicatng Park, Karnataka, the study now covering a period of two behavioral variability in macaques (Bonnet and Lion- decades. The Bonnet Macaques usually live in bisexual tailed) and langurs (Nilgiri and Hanuman), respectvely. groups with each group having several adult males and Behavioral variability was higher in macaques than adult females. Sinha, however, found that in Bandipur, in langurs (Erinjery et al. 2017b). The ideal gas model 11 of the 21 groups of Bonnet Macaques were uni-male- approach showed that primates in the Western Ghats do mult-female groups (Sinha et al. 2005). The origin and not form actve associatons with each other (Erinjery et evoluton of this unique social organizaton in the Bonnet al. 2015). In context to langurs, Hanuman Langurs are Macaque has been explained through a research- more social than Nilgiri Langurs as the later are more based model illustratng the following process (Sinha folivorous and require to spend much tme in restng to & Mukhopadhyay 2013). In deciduous forests, food digest leaves (Kavana et al. 2015). Prerna Singh (Singh becomes scarce and patchy in the dry season, and at the 2017) observed abandonment of infants by Bonnet same tme, tourists passing through the forest ofer food Macaque females afer a group split, and reported that which is rich and clumped. These two factors induce weaning may also occur without aggressive responses intense competton among females in a mult-female- from mothers. Mathew Cooper from University mult-male group, leading to group fssion, resultng in of Georgia studied Assamese Macaques under the small groups of females. In such a situaton, it becomes supervision of Irvin Bernstein in 1997–98 for his Ph.D. easier for a single male to monopolize reproducton, (Cooper 1998) and worked as a post doc in 1999–2001 and consequently, the evoluton of uni-male social at the University of Mysore on dominance style and organizaton takes place. relatonship quality in Bonnet Macaques (Cooper et al. Sinha (Sinha et al. 2005) and his group also described 2004a,b). Under the supervision of Cliford Jolly, Lisa a new macaque species, the Arunachal Macaque Macaca Schloterhausen (Schloterhausen 2000) (htps://as.nyu. munzala from the forests of . Four edu/departments/anthropology/graduate/alumni/ students have carried out their Ph.D. research with Sinha. doctoral-alumni/schloterhausen-lisa.html) from New Rishi Kumar (Kumar 2012) (htps://www.researchgate. York University, New York in afliaton with the University net/profle/Rishi_Kumar24) reported the distributon, of Mysore reported that wild Bonnet Macaques moved demography, and behavioral ecology of mixed groups and fed on more food items but engaged less in social of Bonnet and Rhesus macaques in peninsular India actvites than the commensal bonnets. Anna Nekaris and observed that Rhesus Macaques were extending (Nekaris 2000) (htps://www.brookes.ac.uk/templates/ their range into the range of Bonnet Macaques, due pages/staf.aspx?wid=&op=full&uid=p0029554) from perhaps to human interventon (Kumar et al. 2011, Washington University, Missouri under the supervision 2013). Mayukh Chaterjee (Chaterjee 2013) (htps:// of David Tab Rasmussen, afliated with the University www.wt.org.in/members/mayukh-chaterjee/) studied of Mysore and observed that the Slender Loris was behavioral fexibility and social diversity in Bonnet exclusively faunivorous feeding on insects and small Macaques with special reference to uni-male groups. vertebrates, and distributed their actvity budget evenly Ecological diferences can cause intra-specifc and even between foraging and travelling. Cornelia Bertsch intra-populaton diferences in social systems which, from Austria collaborated with the research group and in turn, produce fexibility and variability in behavioral studied Lion-tailed Macaques from 1998 to 2003 in the strategies such as grooming and proximity. Narayan . Sharma (Sharma 2013) (htps://www.researchgate. net/profle/Narayan_Sharma3) studied efects of Natonal Insttute of Advanced Studies, Bengaluru habitat fragmentaton on primate populatons in Upper At NIAS, primate research was initated by Anindya Brahmaputra Valley where at least one species was lost in Sinha (htps://www.nias.res.in/professor/anindya- the last 30 years. Many, largely habitat specialist species, sinha) and joined later by Sindhu Radhakrishna. The have shown decline in their populatons whereas a more group has been focusing on primate ecology, behavior, commensal and generalist Rhesus Macaque has become conservaton, and cogniton. Afer Simonds (1965), more abundant. Geographical distance, dissimilarity Sinha (2001) provided the frst detailed descripton in spatal features and anthropogenic factors afected of distributon, ecology, and behavior of the Bonnet turnover in primate species compositon. Shreejata Macaque. Inarguably the best long-term feld study Gupta (Gupta 2016) (htps://www.researchgate.net/ on behavior of any primate species in India is Sinha’s profle/Shreejata_Gupta2) found that Bonnet Macaques research on Bonnet Macaques in Bandipur Natonal produce fexible and intentonal gestures with context

16724 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 September 2020 | 12(13): 16715–16735 J TT History of primatology in India Singh et al. variability. Gestures such as those used in play decrease there. Untl the early 1980s, virtually nothing was known with age and others such as those related to aggression about these primates in northeastern India. Anwaruddin and afliaton appear at the adult stage. Females had a Choudhury (Choudhury 1989) (htps://en.wikipedia. higher repertoire of signals than males. Anindya Sinha org/wiki/Anwaruddin_Choudhury) was the frst person also hosted Avant Mallapur (htps://www.linkedin. to begin primate research in northeastern India in 1986 com/in/avant-mallapur-b9b2a615), a Ph.D. student when he reported that the Pig-tailed and the Stump- from the University of Edinburgh, Edinburgh (Mallapur tailed macaques were restricted by the Brahmaputra 2005). Mallapur studied the efect of visitor presence River towards west of their range. He also discovered on behavior of captve Lion-tailed Macaques and found a new populaton of macaques which he described as a that the visitor presence might adversely afect welfare subspecies of Macaca thibetana but the same has now of causing an increase in abnormal and other been described as a new species with the name Macaca social behaviors (Mallapur et al. 2005). munzala. Sindhu Radhakrishna joined NIAS in 2002 and In one of the earlier studies conducted in 1987–1991 contnued surveys on lorises in northeastern India and on the Golden Langurs (Duta 1992), it was found that Kerala. In , the Bengal Slow Loris was found they were confned to the northwestern belt of Assam. in only two of the 16 sites surveyed and the habitat was The group had 7–13 individuals and shifed the home found to be severely afected by anthropogenic actvites range seasonally. They fed on fowers, blossoms, (Radhakrishna et al. 2010). In , the encounter leaves and fruits of 34 plant species. Atul Kumar Gupta rate of the Slow Loris was 0.22 individuals (Swapna et al. (Gupta 1997) (htp://tdu.edu.in/governance/akgupta/) 2008). The Slender Loris was found in 22 of the 36 forest observed that due to contnuing loss of forests, primate ranges surveyed in Kerala with an abundance ranging populatons in northeastern Indian region were being from 0.02 to 1.44 individuals/km (Radhakrishna et al. adversely afected. 2011). She is also tracing human-monkey relatonships Although a few feld studies were carried out in the in literature and folklore (Radhakrishna 2018). Asmita 1980s and 90s, it was in 1994 that long-term studies Sengupta (Sengupta 2015) (htps://www.atree.org/ on several primate species in northeastern India were users/dr-asmita-sengupta) studied seed dispersal as an initated under the Indo-US Primate Project with ecosystem functon by Rhesus Macaques in Buxa Tiger S.M. Mohnot of J.N.V. University, Jodhpur and P.C. Reserve in 2012–2014. Macaques dispersed 84% of the Bhatacharjee of Guwahat University as the principal 49 species they fed on. The macaque-handled seeds investgators. Prabal Sarkar (Sarkar 2000) (htps://www. were largely deposited outside the tree canopy, were ustm.ac.in/zoology-list-of-faculties/dr-prabal-sarkar/) undamaged and had enhanced germinaton. Provisioning reported that in Assamese Macaques, a provisioned reduced the daily range of macaques and resulted in group spent more tme on clumped resources than lesser seed dispersal than in unprovisioned macaque forest groups. As a result, the provisioned monkeys group. Shaurabh Anand (Anand 2019) (htps://www. were more aggressive towards each other, and to researchgate.net/profle/Shaurabh_Anand3) studied reduce tension, also indulged more in grooming than perceptons about human-Rhesus Macaque confict in the forest monkeys. Jayanta Das (Das 2002) found Himachal Pradesh. Whereas higher level forest ofcers Gibbons in three and fve reserved forests in Silchar and proposed strict rules for management of negatve Karimganj divisions respectvely with a median group interactons, the lower level ofcers agreed more with size of three. In Borajan where the forest damage was the farmers who actually sufered losses. Forest ofcers extensive, Gibbons were forced to move on ground to believed that sterilizaton and culling of monkeys were cross between food patches and they showed many the efectve interacton resoluton measures, the vigilance behaviors while walking on open grounds. Dilip farmers found these inefectve and against cultural and Chetry (Chetry 2002) (htps://www.researchgate.net/ religious beliefs. Therefore, there appears to be higher profle/Dilip_Chetry2) recorded alttudinal distributon human-human confict than human-animal negatve range of Stump-tailed Macaques from 50–1,300 m in interactons (Anand & Radhakrishna 2020). northeastern India with group size ranging from 2–63 individuals. Home range size was 336–587 ha. These Northeastern India and Gauhat University monkeys fed on 200 food plants. Stump-tailed Macaques Northeastern India is an extremely rich region for are terrestrial and non-seasonal breeders. Habitat primate diversity with 13 of the 27 Indian primate species loss, fragmentaton, jhum cultvaton, and traditonal including loris, macaques, langurs, and small apes found huntng were determined as the major threats. Rekha

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Medhi (Medhi 2004) (htps://www.researchgate.net/ (>100km2) forest patches, respectvely. Likewise, the profle/Rekha_Chetry) studied the behaviour of a semi- group size was 2.5, 3.29, and 3.9 in small, medium, provisioned introduced troop of Golden Langur outside and large fragments. There were fewer young in small its distributon range. Grooming played an important fragments than in medium and large fragments. role in maintaining inter-individual relatonship. She also reported development of neonate, neonate behaviour, North Eastern Hill University, Shillong intra-troop social dynamics and development of inert G.S. Solanki (Solanki 1987) and Awadhesh Kumar individual relatonships. Jihosuo Biswas (Biswas 2004) (Kumar 2006) (htps://nerist.ac.in/forestry/facultes/ (htps://www.researchgate.net/profle/Jihosuo_Biswas) awadhesh) studied the behavior of Capped Langurs in found that in Golden Langurs, the home ranges of the Pakhui Wildlife Sanctuary in Arunachal Pradesh and two study groups were 25 and 58 ha. Home range size found that the langurs spent 54% tme restng and 36% correlated with group size and there was an overlap in feeding. Age-sex classes did not difer in their tme- between ranges of diferent groups. There was a actvity budgets. Most actvites occurred between the bimodal feeding peak in the mornings and evenings. heights of 10 and 15m. Langurs fed on 52 plant species, These langurs spent about 44.75% of their tme restng, mostly of the family Moraceae, and 68% of their diet probably to digest leaves which are their primary food comprised leaves. Huntng for meat was the major items ingested. Joydeep Bose (Bose 2005) reported that threat for these langurs. Phayre’s Leaf Monkey is found only in southern Assam, Tripura, and . The home range size correlated Mizoram University, Aizawl with group size and varied from 2.7 to 17.6 ha with a day Zothansiama (Zothansiama 2013) (htps://mzu.edu. range of 304 to 592 m. Time-actvity budgets difered in/department-of-zoology/) found that Stump-tailed between groups in plantatons and forests. Leaves Macaques were scarcely distributed to the south bank (54%), followed by shoots (23.2%) and fruits, fowers, of Brahmaputra River. In a captve group of Stump- and seeds were ingested. Dhiraj Kumar Borah (Borah tailed Macaques, copulatons in females correlated with 2010) studied ecology and behavior of Capped Langurs sexual atractveness indicated by the bright red genital and found that though the tme-actvity budgets did skin color. Males ofen indulged in homosexual behavior not difer, the home range was larger and the day tme perhaps to regulate aggression. The forest study group length of movement was smaller in the undisturbed fed on fruits, leaves, shoots, and fowers of 36 plant habitat than in the disturbed habitat. Langurs spent species. Pallab Deb (Deb 2015) studied the Western most tme restng, followed by feeding, and grooming Hoolock Gibbon in southern Assam and Mizoram and was the predominant social actvity. Nabajit Das (Das found 51 individuals in 16 family groups with a mean 2013) (htps://www.researchgate.net/profle/Nabajit_ group size of 3.1. Tree density varied among diferent Das) found Bengal Slow Lorises in seven protected areas habitats. Gibbons used tree from 5 to 30 m, though most and two reserved forests in Assam with a density ranging of their actvites were between the heights of 6 and 25 from 2.78 to 9.21 animals/km2. Lorises spent 42.59% of m. They fed on fruits, fowers, leaves and petoles of their tme restng, followed by 27.46% in locomoton, 32 plant species. Most of the habitats of Gibbons were and 21.28% in feeding. They used 44 trees and 12 lianas under anthropogenic pressures. and the food consisted of 80.93% plant exudates and only 2.32% animal mater. The average home range size North Orissa University, Baripada was 15.60ha. Raju Das (Das 2012) (htps://www.researchgate.net/ profle/Raju_Das2) found that in the Chirang Reserved Forest, Golden Langurs fed on 91 plant species of trees PRIMATE STUDIES FROM THE NORTH-EAST IN and climbers. The spectrum of food plant species in OTHER UNIVERSITIES the diet of langurs appeared to be determined by the foristc compositon of the habitat. University of Cambridge Kashmira Kakat (Kakat 2005) (htps://www. Assam University, Silchar researchgate.net/profile/Kashmira_Kakati/research) Mofdul Islam (Islam 2015) (htps://www. found that in Hoolock Gibbons, the encounter rates researchgate.net/profle/Mofdul_Islam) studied were 0.09 animal/km, 0.23 animal/km, and 0.58 animal/ Hoolock Gibbons in the Inner-Line Reserved Forest in km in small (<5km2), medium (20–30 km2), and large Assam and observed that gibbons mainly fed on Ficus,

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Artocarpus, Dysoxylum, Gmelina, and Syzygium. Gibbon studied groups of Lion-tailed Macaques in 1989–91 populaton positvely correlated with canopy cover and in a degraded forest fragment near and in an tree density. The dramatcally changing land cover due undisturbed forest at Varagaliyar and reported that to reasons such as shifing cultvaton is a major threat to the monkeys in fragments spent more tme in ranging the gibbons in the region. than restng or feeding, and were more terrestrial than the macaques in undisturbed forest. R. Krishnamani North Eastern Regional Insttute of Science and (Krishnamani 2002) (htps://www.researchgate.net/ Technology, Nirjuli, India profle/Krishnamani_Ramanathan) reported that of the Kuldip Sarma (Sarma 2015) studied the Eastern 190 woody plant species in the study area, Lion-tailed Hoolock Gibbon in Lower Dibang Valley and Lohit Macaques used 74 as food trees. The relatve density districts of Arunachal Pradesh and found 77 groups and of these species being 57.1% indicated that the habitats three solitary individuals with a mean group size of 3.04. in Karnataka could support a good populaton of Lion- He estmated the presence of 6,953 groups of gibbons tailed Macaques. H.N. Kumara (htps://www.sacon.in/ with 21,710 individuals in the state. Groups in protected division/conservaton-biology/#fs-tabbed-12) joined forests had larger home ranges than in unprotected the insttute in 2010 and contnued studies on primates. forests. Surveys were carried out on Lion-tailed Macaques, Nilgiri Langurs, and Slender Loris in the unexplored forests, and Tezpur University, Tezpur several conservaton actons were initated, resultng in Mrigankhi Borah (Borah 2016) found that Hoolock the formaton of Sharavathi Lion-tailed Macaque Wildlife Gibbons in Hoolongapar fed on 54 plant species in which Sanctuary. K. Santhosh (Santhosh 2017) (htps://www. Ailanthus, Ichnocarpus, Trichosanthes, and Ficus were researchgate.net/profle/K_Santhosh2) reported that in the most preferred species. Fruits comprised 51.14% of the Lion-tailed Macaque in its northernmost populaton the diet. Vatca lanceaefolia and Artocarpus chaplasha in the Western Ghats, the mean monthly day path length grew and survived beter in the gaps than in understory correlated positvely with the number of fruitng trees, whereas Artocarpus chaplasha grew beter in the and tree density and fruit tree density also correlated understory. positvely with habitat use (Santhosh et al. 2015). Kumara initated long-term studies on the Nicobar Long- tailed Macaque (NLTM). This island populaton surveyed SOME OTHER CENTERS OF PRIMATE RESEARCH earlier (Umapathy et al. 2003) sufered a heavy loss to the 2004 Indian Ocean tsunami, however, in the following Salim Ali Center for Ornithology and Natural History, decade or so, the populaton of these macaques has Coimbatore not only recovered but increased (Velankar et al. 2016). Ajith Kumar initated a study to explore the impact Arijit Pal (Pal 2018) (htps://www.researchgate.net/ of habitat fragmentaton on various taxa including profle/Arijit_Pal4) found that in NLTM, the number of primates. G. Umapathy (Umapathy 1998) (htps://www. births correlated with the monthly rainfall, and about ccmb.res.in/People/Research-Group/G-Umapathy) 71% of the births occurred in the rainy season (Pal et al. found that in forest fragments, tree density and canopy 2018a). These macaques showed six extractve foraging height best predicted occurrence of primates, and it behaviors, including some being tool-aided, and teeth is suggested that arrestng further degradaton, and fossing (Pal et al. 2018b). Partha Sarathi Mishra (Mishra improving quality, of habitats along with retenton of 2020) (htps://www.researchgate.net/profle/Partha_ jackfruit orchards may help primate conservaton in Mishra3) found reciprocity, though not equitable, in the fragmented landscape (Umapathy & Kumar 2000). grooming among NLTM males, but no rank-related An Indo-German Project funded by the Volkswagen diferentaton of afliaton, indicatng lack of social Foundaton with Werner Kaumanns from German bonds in this subspecies (Mishra et al. 2020a). Females Primate Centre, Gotngen and Ajith Kumar from SACON directed grooming towards high ranking males, though as Principal Investgators was launched to further dominance hierarchy was less steep than expected in study the efect of forest fragmentaton. Though not Macaca fascicularis (Mishra et al. 2020b). based at SACON, the efect of forest fragmentaton on the behavior of Lion-tailed Macaques was also University of Rajasthan, Jaipur studied by Shaily Menon (Menon 1993), a student of It was found that mult-male Hanuman Langur Frank Poirier from The Ohio State University. Menon groups split only afer a takeover by a male from an all-

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male band, and the split resulted in formaton of two Kerala Forest Research Insttute, Peechi uni-male groups (Mathur & Manohar 1990). In another K.K. Ramachandran (htp://www.kfri.res.in/whois. study, Mathur & Manohar (1992) demonstrated that asp?ID=10&sub=1) initated primate research at KFRI, populaton density and not habitat disturbance was primarily on Lion-tailed Macaques and Nilgiri Langurs correlated with takeover in Hanuman Langurs. Reena and supervised Ph.D. research of two students. Gigi K. Mathur (htps://www.researchgate.net/profle/Reena_ Joseph (Joseph 1999a) (htps://nirmalacollege.ac.in/ Mathur2) at the University of Rajasthan also hosted department-faculty/dr-gigi-k-joseph/) reported the Linda D. Wolfe who arrived from the USA in 1984 on presence of 14 groups of Lion-tailed macaques and an NSF grant and studied feeding ecology of Rhesus 85 groups of Nilgiri Langurs in Silent Valley. The food Macaques. The monkeys in a town group foraged more, of Lion-tailed Macaques comprised 91% plant and 9% moved less, engaged in fewer aggressive acts, and slept animal mater. The average group size of Nilgiri Langurs less during the day than the monkeys in a forest/temple was 5.89. R. Suganthasakthivel (Suganthasakthivel complex (Wolfe 1992). 2011) (htps://www.researchgate.net/profle/R_ Suganthasakthivel) found a total of 13 groups with an Himachal Pradesh University, Shimla average group size of 15.4 of Lion-tailed Macaques and R.S. Pirta (htps://www.researchgate.net/profle/ 23 groups of Nilgiri Langurs with an average group size of Raghubir_Pirta) carried out his Ph.D. research at 6.5 in the Nelliampathy forests. The relatve abundances Kashi Vidyapith University, Varanasi where he studied of diurnal arboreal mammals ranged between 1.2 and cooperatve behavior in Rhesus Macaques and reported 12.6 individuals per 10km. that adult females were the main groomers, and grooming duraton increased during the matng season; Centre for Cellular and Molecular Biology, Hyderabad play was largely restricted to infants and juveniles (Pirta Afer his Ph.D. from SACON and post-doc from 1983). In cooperatve atacks, adult females were University of Mysore, G. Umapathy joined Laboratory help-givers in both forest and urban groups, though for the Conservaton of (LaCONES) cooperatve atacks were more among the later (Pirta at CCMB, Hyderabad and contnued his studies on 1984). Human disturbance caused more detrimental efects of habitat fragmentaton on demography, genetc efects on Hanuman Langurs who were largely restricted status, hormones, and endoparasites in primates in to broad-leaf forests than Rhesus Macaques who ofen the Anamalai Hills. Sustained habitat fragmentaton visited towns too (Ross et al. 1993). Anita Chauhan and exposure to human and livestock increased the (Chauhan 2009) (htps://www.researchgate.net/profle/ prevalence of endoparasite in Lion-tailed Macaques Anita_Chauhan5) reported that the Rhesus Macaques (Hussain et al. 2013) and Nilgiri Langurs (Tiwari et al. subsistng in temple areas interacted more aggressively 2017). M.S. Ram (Ram 2018) (htps://www.researchgate. with humans than the macaques in a marketplace net/profle/Ram_M_S2) constructed phylogeography and the Hanuman Langurs, due possibly to the heavy of the Lion-tailed Macaque across its range and found dependence of the temple macaques on human that these macaques in forest fragments had depleted resources. Santosh Kumar Sahoo (Sahoo 1993) carried mitochondrial diversity and that the Palghat gap in out a study from 1988 to 1991 in Shimla and found that the Western Ghats separated their populatons into the Rhesus Macaques engaged in agonistc interactons two almost 2.11 million year ago (Ram et al. 2015). In more than the Hanuman Langurs, and most of these the Endangered Golden Langur in Assam, though the agonistc interactons occurred during human feeding. genetc diversity was high, populatons in smaller forest Pramod Kumar (Kumar 1992) studied Rhesus Macaques fragments showed lower nucleotde diversity than in and Hanuman Langurs in Shimla between 1987 and larger fragments (Ram et al. 2015). 1991 and found that populatons of both species increased 26.61% and 25.86%, respectvely, during the Aligarh Muslim University, Aligarh study period. Rhesus Macaques were more urban than Ekwal Imam (Imam 2000) (htps://www.researchgate. langurs and the adult sex rato in the two species was net/profle/Ekwal_Imam) monitored populatons of 1:2.3 and 1:5.6, respectvely. As Shimla gets very cold in Rhesus Macaque near Aligarh. In the study region, winter, there were signifcant seasonal variatons in the 963 monkeys in 1993 increased to 1337 in 1995 with behavior of both species. a birth rate of 0.81 infant/female/year and high infant survivorship. These observatons agree with general trends in populaton increase in Rhesus Macaques in

16728 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 September 2020 | 12(13): 16715–16735 J TT History of primatology in India Singh et al. several places in 1980s and 1990s. Bharathiyar University, Coimbatore Shanthala Kumar (Kumar 2019) (htps://www. University of California, Davis researchgate.net/profle/Shanthala_Kumar) reported Uma Ramakrishnan (Ramakrishnan 1999) found that the relocaton of Bonnet Macaques from urban that in the wild bonnet macaques, discriminaton of areas to a wild habitat facilitated the spread of alien predators, and ant-predatory responses, develop endoparasites to Lion-tailed Macaques where these ontogenetcally as juveniles, subadults and adults parasites were otherwise absent. Even on an isolated respond to the stmuli diferently. island system, the increased use of human-dominated spaces by Nicobar long-tailed Macaques has resulted in University of Burdwan, Bardhaman an increased endoparasite richness and prevalence. Sangita Mitra (Mitra 2000) studied behavior of Assamese Macaques near Darjeeling during 1996 and 1999 in broadleaf forests up to 2,500m, though the FUTURE macaques were found mostly between 200 and 500 m. The adult sex rato was 1:2.88 and the ranges varied Natonal survey of primates from 0.6 to 3.5 km2. There have been some atempts in the past to census primates in diferent regions of the country. The University of Calicut, Kozhikode earliest surveys were on the Rhesus Macaques in certain Arboreal mammals with special reference to the parts of northern India initated by Charles Southwick Nilgiri Langur were studied in the high ranges in Kerala (described earlier in this artcle). Zoological Survey of (Joseph 1999b). A surprising fnding was that Nilgiri India undertook surveys in northern India under the Langurs were observed to feed on fruits (50%) more than leadership of R.P. Mukherjee (Mukherjee & Mukherjee on leaves (30% and fowers (20%). More carbohydrates 1972), in southern India by G.U. Kurup (Kurup 1984), and containing food were used by langurs in summer and in the northeastern India by J.R.B. Alfred (Alfred & Sat more protein and lipid were consumed during rainy 1990; Alfred et al. 2004) and some others at localized season and winter. places the details of which are available with ZSI at htp://faunaofndia.nic.in/. Primates in northeastern Maharaja Ganga Singh University, Bikaner India were also surveyed under the Indo-US Primate Rhesus Macaques in the Aravali Hills were actve Project (Srivastava 1999). Ullas Karanth (Karanth 1985) during mornings and evenings in summer, but more surveyed Lion-tailed Macaques in the state of Karnataka. actve during the day in winter (Sharma 2009). Most of these surveys are now outdated and were largely localized. It is only recently that a systematc Indian Insttute of Science, Bengaluru survey on Rhesus Macaques has been carried out in Though taxonomy research on primates has been Himachal Pradesh (Kumara & Singh 2020). There have going on in the Indian Insttute of Science for long, it been several status assessments of primates by Zoo is only recently that an ecological study by Mehreen Outreach Organizaton including PHVAs for the Lion- Khaleel (Khaleel 2020) (htps://www.researchgate.net/ tailed Macaque (Kumar et al. 1995), primates of South profle/Mehreen_Khaleel) has shown that Hanuman Asia (Molur et al. 2003) and Hoolock Gibbon (Molur et Langurs in Kashmir Himalaya are distributed wider al. 2005). There is, therefore, an urgent need to take up than previously thought. These langurs show seasonal a natonwide survey of primates in India. variatons in home ranges and diet in accordance with the energy maximizing strategy. Populaton monitoring Populaton dynamics can be studied if a populaton University of Madras, Chennai is monitored for decades. Most research studies are R. Sasi (Sasi 2018) provided the status and distributon of two to three year duratons, and hence cannot bring patern of lion-tailed macaque and slender loris for the out the stochastcity of primate populatons. A lesson less explored regions including Parambikulam landscape is learnt from the monitoring of Bonnet Macaque and Megamalai-Srivilliputhur-Tirunelveli hills in the populatons around the city of Mysore for over 30 years southern Western Ghats. now by Mewa Singh and his team which has shown a ~70% decline in these macaques (Erinjery et al. 2017a). We need to identfy populatons that are representatves

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of each species in diferent habitats and monitor them Conservaton breeding at least once in three years for several decades to Several species of primates in India are threatened. comprehensively know the trends. An expert group of primatologists and wildlife managers should decide which of these species require Taxonomic studies conservaton breeding in order to develop sustainable Several species of primates in India are classifed captve populatons for restocking or rehabilitaton, if into subspecies. In order to prioritze conservaton, it required in the future. An initatve has already been is necessary that genetc studies are carried out on taken for conservaton breeding of Lion-tailed Macaques all subspecies to determine whether there are more but it has not yielded desired results. Perspectves for distnct species. For example, the Hanuman Langur systematc management on the basis of analysis of was considered as only one species but it is now shown conservaton breeding programs in various countries that there are many distnct species of this langur have been provided to aide planning (Kaumanns et al. (Ashalakshmi et al. 2015). Morphologically similar 2005, 2013, 2020). populatons of the Lion-tailed Macaque north and south of the Palghat gap have been genetcally diferentated Natonal Primate Research Centre for the past 2.1 million years (Ram et al. 2015). There is a long pending demand of primatologists to have a Natonal Primate Research Centre that should Ecological and behavioral research with long term feld be managed both by primate biologists and wildlife studies managers, and funded by the Ministry of Environment, There are many subspecies of few primate species in Forest and Climate Change and Ministry of Science and India for which even baseline ecological and behavioral Technology, Government of India. This Centre would informaton are not yet available. There is therefore a not only initate new research but will also coordinate need to not only study these subspecies, but also all among various insttutons and governmental agencies species across varied habitats. There are hardly any long- in the country that are engaged in primate research term feld studies carried out in India. The one carried out and management. The Primate Centre could also host by Anindya Sinha and his group on Bonnet Macaques at a Natonal Primate Museum to showcase primate taxa, Bandipur brought out excellent results about evoluton educate people and build awareness for their protecton of sociality (Sinha & Mukhopadhyay 2013). A study on and conservaton. the Himalayan Grey Langur near Chamba by the Wildlife Informaton Liaison Development / Zoo Outreach A Platorm for Interactons Organizaton has been going on since 2012 (htps:// A Primatological Society of India was established www.speciesconservation.org/case-studies-projects/ in the late 1970s but it became inactve in the 1980s. himalayan-grey-langur/9744) though the results are yet Since then, there has not been a single platorm for to be published. Indian primatologists to meet and exchange informaton except a CAMP meetng conducted by Zoo Outreach Conservaton and management Organizaton (Molur et al. 2003). An enthusiastc group Many species of primates, because of their threatened of Indian primatologists have established a new platorm or vulnerable status, require actve interventon for named, Associaton of Indian Primatologists (htps:// in situ management. Wildlife Trust of India has been www.indianprimates.org/) in 2019 and conducted its carrying out rehabilitaton of Eastern Hoolock Gibbon frst natonal workshop at Indian Insttute of Science, from small and isolated forest fragments to large forest Bengaluru in November 2019. This platorm needs to complexes (Roy et al. 2015). Since diferent species be strengthened. inhabit diferent ecological setngs and experience anthropogenic pressures, Singh (2019), for example, Internatonal Cooperaton has provided approaches to management of urban- Only three important internatonal meetngs have and forest-dwelling species. Likewise, appropriate been held in India: the 7th Congress of the Internatonal management strategies need to be synthesised through Primatological Society at Bengaluru in January 1979, dialogues between primate biologists and wildlife Internatonal Symposium on Primates at Jodhpur managers. University in February 1982, and the 7th Asian Primate Symposium at Gauhat University in February 2020. The proceedings of the Bengaluru and Jodhpur meetngs led

16730 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 September 2020 | 12(13): 16715–16735 J TT History of primatology in India Singh et al. to seminal publicatons (Seth & Seth, 1983b, 1989a,b; reproductve success in free-ranging female Hanuman Langurs Roonwal et al. 1984, respectvely). We encourage the (Presbyts entellus). Internatonal Journal of Primatology 12: 231– 257. htps://doi.org/10.1007/BF02547586 research groups in India to become more actve in Bose, J. (2005). An ecobehavioural study of Phayre’s Leaf Monkey developing internatonal cooperaton so that expertse Trachypithecus phayrei (Blyth, 1847). PhD Thesis. Gauhat University, Guwahat. from diferent countries could be integrated in research Chandel, N.K. (1981). Afliaton, atenton and spacng mechanisms and conservaton. A successful example of such a among free-reanging Rhesus monkeys (Macaca mulata). PhD cooperaton has been the conduct of an internatonal Thesis, University of Rajasthan, Jaipur. Chaterjee, M. (2013). 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American Journal of Primatology 73: 335–346. Sarkar, P. (2000). Ecology and dynamics of social relatonships of htps://doi.org/10.1002/ajp.20900 Assamese macaque: Macaca assamensis (McClelland, 1839). PhD Singh, M., A. Udhayan, M.A. Kumar, H.N. Kumara & D.G. Lindburg Thesis. Gauhat University, Guwahat. (1999). Status survey of Slender loris Loristardigradus lydekkerianus Sarma, K. (2015). Studies on populaton status, behavioural and in Dindigul, Tamil Nadu, India. Oryx 33: 30–36. htps://doi. habitat ecology of Eastern Hoolock gibbon (Hoolock leuconedys) org/10.1046/j.1365-3008.1999.00033.x in Arunachal Pradesh, India. PhD Thesis. North Eastern Regional Singh, P. (2017). Parental investment paterns in relaton to dominance Insttute of Science and Technology (Deemed University), Nirjuli. status in Bonnet macaque Macaca Radiata. PhD Thesis. University Sasi, R. (2018). Geographical mapping and ecological impact of lion- of Mysore, Mysuru. tailed macaque and slender loris in Southern India. PhD Thesis. Singh, R. (1981). Study of behavioral paterns of age sex classes during University of Madras, Chennai. transitonal periods in free ranging Rhesus Monkeys (Macaca Schloterhausen, L. (2000). Town monkeys, country monkeys: A mulata). PhD Thesis. University of Rajasthan, Jaipur. socioecological comparison of a human commensal and wild group Singh, S.D. (1968). Efect of urban environment on visual curiosity of Bonnet macaques (Macaca radiata). PhD Thesis. New York behavior in Rhesus monkeys. Psychonomic Science 11: 83–84. University, New York. Singh, S.D. (1969). Urban monkeys. Scientfc American 221: 108–115. Sekhsaria, P. (2015). KMTR’s one-mile corridor. Sanctuary Asia 45–49. Singh, S.D. (1977). Efects of infant-infant separaton of young monkeys Sengupta, A. (2015). Seed dispersal by Rhesus macaques (Macaca in a free-ranging natural environment. Primates 18: 205–214. mulata). PhD Thesis. Manipal Academy of Higher Educaton, htps://doi.org/10.1007/BF02382959 Manipal. Singh, S.D. (1980). Xenophobic reactons of free-ranging Rhesus infant Seth, P.K. & S. Seth. (1983a). Brief report: Populaton dynamics of groups raised in natural habitat. Primates 21: 492–497. htps://doi.

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org/10.1007/BF02373837 Sugiyama, Y. (1966). An artfcial social change in a Hanuman Singh, S.D. & S.N. Manocha (1966). Reactons of the Rhesus Monkey langur troop (Presbyts entellus). Primates 7: 41–72. htps://doi. and the langur in novel situatons. Primates 7: 259–262. htps://doi. org/10.1007/BF01730557 org/10.1007/BF01730791 Sugiyama, Y. (1968). The ecology of the Lion-tailed macaque (Macaca Sinha, A. (2001). Report on the Monkey in the Town’s Commons: A silenus (Linnaeus)): A pilot study. Journal of Bombay Natural History Natural History of the Indian Bonnet Macaque. Report to Natonal Society 65: 283–292. Insttute of Advanced Studies, Bengaluru. Sugiyama, Y. (1971). Characteristcs of the social life of Bonnet Sinha, A., A. Data, M.D. Madhusudan & A. Mishra (2005). Macaca macaques (Macaca radiata). Primates 12: 247-266. htps://doi. munzala: A new species from Western Arunachal Pradesh, org/10.1007/BF01730414 Northeastern India. Internatonal Journal of Primatology 26: 977- Sugiyama, Y. (1976). Characteristcs of the ecology of the Himalayan 989. htps://doi.org/10.1007/s10764-005-5333-3 langurs. Journal of Human Evoluton 5: 249–277. htps://doi. Sinha, A. & K. Mukhopadhyay (2013). The monkey in the town’s org/10.1016/0047-2484(76)90029-4 commons, Revisited: An anthropogenic history of the Indian Bonnet Sugiyama, Y. (1984). Some aspects of infantcide and intermale Macaque, pp187–208. In: Radhakrishna, S., M.A. Hufman & A. competton among langurs, Presbyts entellus, at Dharwar, Sinha (Eds.), The Macaque Connecton: Cooperaton and Confict India. Primates 25: 423–432. htps://doi.org/10.1007/BF02381665 Between Humans and Macaques. Springer, xii+256pp. Sushma, M.S. (2004). Resource utlizaton and niche separaton in Sinha, A., K. Mukhopadhyay, A. Data-Roy & S. Ram (2005). Ecology sympatric rainforest arboreal mammals. PhD Thesis. University of proposes, behaviour disposes: ecological variability in social Mysore, Mysuru. organizaton and male behavioural strategies among wild Bonnet Sushma, H.S. & M. Singh (2006). Resource parttoning and interspecifc macaques. Current Science 89: 1166–1179. interactons among sympatric rain forest arboreal mammals of the Solanki, G.S. (1987). Ecology and behavoiur of Capped Langur in Western Ghats, India. Behavioral Ecology 17: 479–490. htps://doi. Pakhui Wildlife Sanctuary, India. PhD Thesis. Saurashtra University, org/10.1093/beheco/arj058 Rajkot. Swapna, N., A. Gupta & S. Radhakrishna (2008). Distributon survey Sommer, V. (1985). Weibliche und männliche reproduktonssstrategien of Bengal Slow loris Nyctcebus bengalensis in Tripura, northeastern der Hanuman-Languren (Presbyts entellus) von Jodhpur, Rajasthan, India. Asian Primates Journal 1: 37–40. India. PhD Thesis. University of Götngen, Gotngen. Thierry, B., M. Singh & W. Kaumanns (Eds.). (2004). Macaque Sommer, V. & S.M. Mohnot (1985). New observatons on infantcides Societes: A Model for the Study of Social Organizaton. Cambridge among Hanuman langurs (Presbyts entellus) near Jodhpur University Press, 418pp. (Rajasthan/India). Behavioral Ecology and Sociobiology 16: 245– Tiwari, S., M. Reddy, M. Pradheeps, G.S. Sreenivasamurthy & G. 248. htps://doi.org/10.1007/BF00310987 Umapathy (2017). Prevalence and co-occurrence of gastrointestnal Southwick, C.H., M.A. Beg & M.R. Siddiqi (1961a). A populaton survey parasites in Nilgiri Langurs (Trachypithecus johnii) of fragmented of Rhesus monkeys in villages, towns and temples of northern landscape in Anamalai Hills., Western Ghats, India. Current Science India. Ecology 42: 538–547. 113: 2194–2200. Southwick, C.H., M.A. Beg & M.R. Siddiqi (1961b). A populaton survey Umapathy, G. (1998). Impacts of habitat fragmentaton on the of Rhesus monkeys in northern India: II. Transportaton routes and arboreal mammals in the wet evergreen forests of the Anamalai forest areas. Ecology 42: 698–710. Hills in the Western Ghats, South India. PhD Thesis. Bharathiar Southwick, C.H., M.A. Beg & M.R. Siddiqi (1965). Rhesus monkeys University, Coimbatore. in North India, pp111–159. In: I. DeVore (Eds.), Holt, Rinehart & Umapathy, G. & A. Kumar (2000). The occurrence of arboreal Winston (eds.). Primate Behavior: Field Studies of Monkeys and mammals in the rain forest fragments in the Anamalai Hills, Apes, New York, 654pp. south India. Biological Conservaton 92: 311–319. htps://doi. Southwick, C.H. & M.R. Siddiqi (1966). Populaton changes of Rhesus org/10.1016/S0006-3207(99)00097-X monkeys (Macaca mulata) in India, 1959 to 1965. Primates 7: 303– Umapathy, G., M. Singh & S.M. Mohnot (2003). Status and 314. htps://doi.org/10.1007/BF01730644 distributon of Macaca fascicularis umbrosa in the Nicobar Islands, Southwick, C.H. & M.F. Siddiqi (1977). 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Journal of Threatened Taxa | www.threatenedtaxa.org | 26 September 2020 | 12(13): 16715–16735 16735 PLATINUM The Journal of Threatened Taxa (JoTT) is dedicated to building evidence for conservaton globally by publishing peer-reviewed artcles online every month at a reasonably rapid rate at www.threatenedtaxa.org. OPEN ACCESS All artcles published in JoTT are registered under Creatve Commons Atributon 4.0 Internatonal License unless otherwise mentoned. JoTT allows allows unrestricted use, reproducton, and distributon of artcles in any medium by providing adequate credit to the author(s) and the source of publicaton.

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

September 2020 | Vol. 12 | No. 13 | Pages: 16715–16926 Date of Publicaton: 26 September 2020 (Online & Print) www.threatenedtaxa.org DOI: 10.11609/jot.2020.12.13.16715-16926

Review A checklist of buterfy fauna of Bankura Town, West Bengal, India – Ananya Nayak, Pp. 16868–16878 A history of primatology in India (In memory of Professor Sheo Dan Singh) – Mewa Singh, Mridula Singh, Honnavalli N. Kumara, Dilip Chetry & Santanu A diversity of spiders (Arachnida: Araneae) from a cashew ecosystem in Kerala, Mahato, Pp. 16715–16735 India – Mamparambath Subramanian Smitha & Ambalaparambil V. Sudhikumar, Pp. 16879–16884 Communicatons Clinical and pathological fndings in a Dwarf Red Brocket Mazama rufna University campuses can contribute to wildlife conservaton in urbanizing (Mammalia: Cetartodactyla: Cervidae) atacked by dogs regions: a case study from Nigeria – Eduardo Alfonso Díaz, Gustavo Donoso, Carolina Sáenz, Ivete Dueñas & – Iliyasu Simon, Jennifer Che & Lynne R. Baker, Pp. 16736–16741 Francisco Cabrera, Pp. 16885–16890

Killer Whale Orcinus orca (Linnaeus, 1758) (Mammalia: Cetartodactyla: Indigenous uses and traditonal practces of endemic and threatened Chilgoza Delphinidae) predaton on Sperm Whales Physeter macrocephalus Linnaeus, Pine Pinus gerardiana Wall. ex D.Don by tribal communites in Kinnaur District, 1758 (Mammalia: Cetartodactyla: Physeteridae) in the Gulf of Mannar, Himachal Pradesh, northwestern Himalaya Sri Lanka – Swaran Lata, P.S. Negi, S.S. Samant, M.K. Seth & Varsha, Pp. 16891–16899 – Ranil P. Nanayakkara, Andrew Suton, Philip Hoare ­& Thomas A. Jeferson,­ Pp. 16742–16751 Notes The Critcally Endangered White-rumped Vulture Gyps bengalensis in Sigur Plateau, Western Ghats, India: Populaton, breeding ecology, and Range extension and frst confrmed record of the Flightless Anomalure threats Zenkerella insignis (Matschie, 1898) (Mammalia: Rodenta: Anomaluridae) in – Arockianathan Samson & Balasundaram Ramakrishnan, Pp. 16752–16763 Nigeria – Dolapo Oluwafemi Adejumo, Taiye Adeniyi Adeyanju & Temidayo Esther Avifauna of Saurashtra University Campus, Rajkot, Gujarat, India Adeyanju, Pp. 16900–16903 – Varsha Trivedi & Sanjay Vaghela, Pp. 16764–16774 Power lines as a threat to a canopy predator: electrocuted Harpy Eagle in Five new species of trap-door spiders (Araneae: Mygalomorphae: Idiopidae) southwestern Brazilian Amazon from India – Almério Câmara Gusmão, Danilo Degra, Odair Diogo da Silva, Lucas Simão – Manju Siliwal, Rajshekhar Hippargi, Archana Yadav & Dolly Kumar, de Souza, Angélica Vilas Boas da Frota, Carlos Augusto Tuyama, Maria Cristna Pp. 16775–16794 Tuyama, Thatane Martns da Costa, Ana Paula Dalbem, Adrian A. Barnet, Francisca Helena Aguiar-Silva & Manoel dos Santos Filho, Pp. 16904–16908 Rapid mult-taxa assessment around Dhamapur Lake (Sindhudurg, Maharashtra, India) using citzen science reveals signifcant odonate records First record of the Assam Leaf Turtle Cyclemys gemeli (Fritz et al. 2008) (Reptlia: – Neha Mujumdar, Dataprasad Sawant, Amila Sumanapala, Parag Rangnekar & Testudines: Geoemydidae) from the Darjeeling- Himalaya, India Pankaj Koparde, Pp. 16795–16818 – Aditya Pradhan, Niran Chetri & Saibal Sengupta, Pp. 16909–16911

Commercially and medicinally signifcant aquatc macrophytes: potental for Breeding biology of Malabar Tree Toad Pedostbes tuberculosus (Anura: improving livelihood security of indigenous communites in northern Bihar, Bufonidae) from Castle Rock, Karnataka, India India – Deepak Deshpande & Nikhil Gaitonde, Pp. 16912–16915 – Shailendra Raut, Nishikant Gupta, Mark Everard & Indu Shekhar Singh, Pp. 16819–16830 First record of Ourapteryx dierli Inoue, 1994 (Lepidoptera: Geometridae: Ennominae) from India Leaf nutrients of two Cycas L. species contrast among in situ and ex situ – Sanjay Sondhi, Dipendra Nath Basu & Krushnamegh Kunte, Pp. 16916–16919 locatons – Thomas E. Marler & Anders J. Lindström, Pp. 16831–16839 Notes on a communal roostng of two oakblues (Lepidoptera: Lycaenidae: Arhopala) and the Common Emigrant (Pieridae: Catopsilia pomona) buterfies Contributon to the Macromycetes of West Bengal, India: 69–73 in Utarakhand, India – Diptosh Das, Prakash Pradhan, Debal Ray, Anirban Roy & Krishnendu Acharya, – Sohom Seal, Debanjan Sarkar, Agnish Kumar Das & Ankush Chowdhury, Pp. 16840–16853 Pp. 16920–16923

First report of mango leaf gall midge Procontarinia robusta Li, Bu & Zhang Short Communicatons (Diptera: Cecidomyiidae) from India – Duraikannu Vasanthakumar, Senthilkumar Palanisamy & Radheshyam Murlidhar A new species of Platylestes Selys (Odonata: Zygoptera: Lestdae) from the Sharma, Pp. 16924–16926 coastal area of Kannur District, Kerala, India – K.G. Emiliyamma, Muhamed Jafer Palot & C. Charesh, Pp. 16854–16860

A frst complete documentaton of the early stages of Hampson’s Hedge Blue Acytolepis lilacea lilacea Hampson, 1889 (Lepidoptera: Lycaenidae) from Publisher & Host Western Ghats, Kerala, India – V.K. Chandrasekharan & Muhamed Jafer Palot, Pp. 16861–16867

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