Journal of Threatened Taxa | | 26 August 2021 | 13(9): 19352–19357


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

#7521 | Received 13 June 2021 | Finally accepted 11 August 2021



The captive population of the Lion-tailed Macaque Macaca silenus (Linnaeus, 1758). The future of an endangered primate under human care


Nilofer Begum 1, Werner Kaumanns 2, Alexander Sliwa 3 & Mewa Singh 4


1,2 LTM-Research and Conservation, Eschenweg 5, 37130, Gleichen, Germany.

1 Department of Biology, Chemistry, Pharmacy, Freie Universität Berlin, Berlin, Germany.

3 Zoologischer Garten Köln, Riehler Straße 173, 50735 Köln, Germany.

4 Biopsychology Laboratory, Institution of Excellence, Vijnana Bhavan, University of Mysore, Mysuru, Karnataka 570006, India.

1, 2, 3, 4 (corresponding author)




Abstract: For conservation breeding, the endangered Lion-tailed Macaques have been maintained in North America under SSP since 1983 and in Europe under EEP since 1989. Based on a growing interest to support the species long-term survival, the SSP population increased considerably during the first few years of the programme but due to space problems and resulting birth control measures, it has drastically declined to small numbers and a non- breeding status at present. The EEP population continually increased till 2012, but due to the lack of spaces and birth control practises, it has gradually declined since then. It is emphasised that the knowledge gained from field studies on Lion-tailed Macaques in India and its incorporation for captive management under EEP has helped develop appropriate management strategies. Captive propagation of the Lion-tailed Macaque in India, the habitat country, can profit from the successes and drawbacks of the long-term management experiences of SSP and EEP.


Keywords: Captive breeding, SSP, EEP, Indian captive population, meta population management.




Editor: Anonymity requested.            Date of publication: 26 August 2021 (online & print)


Citation: Begum, N., W. Kaumanns, A. Sliwa & M. Singh (2021). The captive population of the Lion-tailed Macaque Macaca silenus (Linnaeus, 1758). The future of an endangered primate under human care. Journal of Threatened Taxa 13(9): 19352–19357.


Copyright: © Begum et al. 2021. Creative Commons Attribution 4.0 International License.  JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by providing adequate credit to the author(s) and the source of publication.


Funding: SERB India and Wels Zoo Austria.


Competing interests: The authors declare no competing interests.


Acknowledgements: Mewa Singh acknowledges the Science and Engineering Research Board, Government of India for the award of a SERB Distinguished Fellowship (Award Number: SB/S9/YSCP/SERB-DF/2018(1). Nilofer Begum acknowledges Tiergarten Wels for supporting her project and LTM conservation in general.




For most of its history, the captive population of the Lion-tailed Macaque (LTM) was mainly constituted by the North American and the European subpopulations and by a number of other small subpopulations (e.g., India and Japan). Figure 1 and Figure 2 provide an overview on its development, births, imports, and losses. They, like other data used for this paper, are based on the last edition of the international studbook for the LTM (Sliwa & Begum 2019). The North American breeding programme (Species Survival Plan, SSP) for the LTM was established in 1983 with 163 individuals in about 30 zoos (Gledhill 1985). The European programme (European Endangered Species Programme, EEP) was established in 1989, comprising 89 individuals in 12 institutions. Currently, the latter comprises 322 individuals in 44 institutions. The EEP was coordinated by Dr. Werner Kaumanns (German Primate Center; since 2000 curator of primates at Cologne Zoo) till his retirement in 2006. Dr. Alexander Sliwa, Cologne Zoo, is the coordinator since then. The European population grew slowly but steadily to a size of 338 individuals in 2012 but decreased to a current size of 322 individuals in 2018 (Figure 1). The number of births decreased drastically since 2011 (global- Figure 2, European- Figure 3, for more information see below). The American SSP population with its first coordinator and (International) studbook keeper Laurence Gledhill had its peak size and productivity in the decade after the start of the SSP, with about 269 individuals in 1988. Currently, there are only 31 individuals living (Sarno 2018). The reasons for the decrease were space problems, widely executed birth control measures in the 1990s, ageing, and possibly loss of interest (Lindburg 2001; Ness 2011, 2013). The Indian captive population currently comprises 51 individuals including 16 wild-born macaques. The Japanese Lion-tailed Macaque subpopulation has 76 individuals; other smaller stocks comprise 36 individuals totally.

The current global population comprises 516 individuals in 98 zoos. The wild population of the LTM at present is estimated to be about 4,000 individuals, distributed in 47 isolated subpopulations at seven locations (Singh et al. 2020), with less than 2,500 mature individuals in about 200 groups. The current captive population in 98 groups, therefore, constitutes about 11% of the global population.

The breeding programmes for the LTM always acted with a perspective on the species in the wild. The establishment of the SSP for the LTM was realised assuming that at that time only about 1,000 LTMs were left in the wild (see Hill 1971). To establish a reserve in zoos was intended. Contacts and cooperation between American and Indian institutions were realised (including financial support for field studies). American scientists and especially Dr. Donald Lindburg, San Diego Zoological Society, contributed important studies both with reference to the biology of the species and its captive propagation (e.g., Lindburg et al. 1989; Lindburg & Gledhill 1992; Lindburg & Harvey 1996).

Almost since its establishment, the European LTM population was managed  in contact with Indian wildlife biologists. Results from their studies on the wild population in its natural habitat (Western Ghats, southern India) were integrated. Since 1998 (till 2004) the annual reports for the captive population also reported on the status and other relevant aspects of the wild population. This was based on a close (ongoing) cooperation of the first EEP coordinator with Dr. Mewa Singh and Dr. Ajith Kumar. Prof. Mewa Singh, University Mysore, leading Indian primatologist, and wildlife biologist visited Germany to work on LTM matters with Dr. Werner Kaumanns since the 1990s more than 25 times. Mainly due to Mewa Singh and his working group, the conservation of the LTM became and is still   an important issue in India. In addition to grants from major Indian sources, some of the studies were financially supported by German Primate Center, Volkswagen Foundation, various American and European zoos, and private persons. Due to this work, the current status of the species and conservation needs are well known, and the Lion-tailed Macaque is one of the best-studied macaque species, both in the wild and in captivity (for an overview see Singh et al. 2009 and Kaumanns et al. 2013). In situ and ex situ studies resulted in a large number of publications that cover aspects of husbandry and management, conservation and especially many aspects of the species biology. A number of Prof. Singh’s students were involved in Lion-tailed Macaque studies and will continue working for the conservation of the species. Efforts to save the LTM in India got much support through the Fifth International LTM Conference in 1999, that was organised by Mewa Singh at the University of Mysore and supported by the EEP coordinator. Two volumes (58, 59) of German Primate Center’s Primate Report (Schwibbe et al. 2000, 2001) report on the results of the conference. These reports provide an overview on the status of in situ and ex situ research and captive propagation efforts for the species.

The contact with Indian colleagues, the involvement in field studies on a number of aspects of the species biology, and the resulting knowledge, significantly influenced the management of the EEP population. From the beginning of the EEP’s existence, the importance of behavioural and especially social aspects, breeding patterns and aspects of life histories were emphasised. According to Singh et al. (2006), especially considering the reproductive system and social system of the species, is the key to the conservation of the species. EEP policy strongly went for this. Although close cooperation between EEP and SSP was initiated during the Third International LTM Conference (1990) in San Diego, the programmes developed differently. In the SSP population, birth control on a large scale, based on a strict genetic management was carried out from about 1988 onwards (Lindburg & Gledhill 1992; Lindburg et al. 1997; Lindburg 2001). Figure 1 demonstrates the effects on the development of the global historical population. It also shows the latter’s “recovery” (2001--–2011) and a new decline from 2012 onwards. This results from a strong decrease in the number of births (Figure 2). This decrease is induced by the development of the European population (Figure 3). Birth control has been carried out there, too, to deal with space problems. Under these pressurising conditions, the EEP long-term management plan edited in 2016 (Sliwa et al. 2016) recommends further birth control measures on a large scale.

Birth control on a large scale over long periods of time to control population size, however, can have enormous risks for the survival of a population. The example of the SSP population and a number of relevant studies (Kaumanns et al. 2013; Penfold et al. 2014; Kaumanns & Singh 2015; Kaumanns et al. 2020) demonstrate possible negative consequences and elaborate ways to stop negative trends.

We are afraid that under the conditions given, the EEP population’s and therefore the global captive population’s, long-term survival is threatened – given the trend in population development continues and no serious changes in management are initiated soon. The ‘Endangered’ status of the LTM in the wild (Singh et al. 2020) with increasing fragmentation of its range of distribution and habitat destruction, strongly recommend, to continue with preserving a reserve in zoos, especially in India. Measures to stabilise the European and thus the global captive population, and new steps towards achieving its long-term survival are urgently required in order to prevent a loss of reproductive potential, like it happened in the SSP population. The European population is the only captive population that is still large and potentially productive enough to be developed further as a reserve. It seems, that space problems and other infrastructural limitations currently hinder to achieve this goal. EEP participants should consider whether all means to allow more breeding again are really exhausted or whether stopping birth control or more moderate schedules are possible, at least. It is suggested that more should be done to preserve the population’s breeding potential, size, and structure, with the goal to send European LTMs back to other regions and  especially to their country of origin.

Zoos in India keep a small LTM population with a number of potential founders. Many zoos however report breeding problems. According to the last edition of the international studbook, totally 51 animals are kept in 10 Indian zoos, six of which keep less than three animals each. There are two zoos with more individuals – Chennai (n= 20) and Trivandrum (n= 10). These group sizes come close to group sizes in the wild. Historically, Chennai Zoo contributed to more than one-third (n= 64) of the captive births in India (n= 185) and between 2003 and 2018, it contributed to 75% (n= 45) of births in Indian zoos (n= 60) in this period. This might be due to an accumulation of husbandry know-how, personnel experience, and constancy in the management system. Delhi Zoo played an important role in the past by contributing to 49 births, many of them in the 1970s–80s. Judging from the experiences in the European breeding programme, successful breeding requires allowing groups to grow undisturbed, to larger sizes of around 20 individuals with differentiated demographic structures that allow the females to live permanently in their natal groups and to maintain strong social bonds (female-bonded system; see Kumar 1987). This would allow intergenerational overlap and to acquire the necessary social and cognitive competence to interact properly in a complex  social system and to raise offspring (Kaumanns et al. 2013). According to field observations, only the males are the mobile elements of the Lion-tailed Macaque social system (Kumar 1987; Kumar et al. 2001). Under captive conditions only males should be transferred between groups (for details see Kaumanns et al. 2006). More information derived from the studies in the wild (e.g., Kumar 1987; Krishnamani & Kumar 2000; Umapathy & Kumar 2000; Sharma 2002; Sushma 2004; Singh et al. 2006) is available to be used in designing keeping systems for the species. It refers to the species’ arboreal life, selective and individualised foraging on diverse plant and animal species, seasonal variations in diet, large time spent in foraging and exploration, maintenance of large interindividual distances, low reproductive turnover, and a number of special features of the reproductive system. Many aspects have been emphasised for the management of the species in the international breeding programmes. Their consideration would also support successful breeding in the country of origin especially with its advantage of natural living conditions, availability of native food plants and large open-air enclosures.

The Indian zoo community is interested in building up a larger, more productive population in cooperation with the European Breeding Programme (Govindhaswamy Umapathy, pers. comm. 03.viii.2020). This constellation provides a chance to develop perspectives and solutions for problems on both sides. A cooperation could provide spaces for Lion-tailed Macaques from European zoos. Even more importantly, a larger and productive Indian population supported both in terms of animals and know-how from Europe could serve as an interface between the captive and the wild populations. It could be used for a number of conservation purposes – including providing animals for reintroductions in the long run. The establishment of an “Indo-European Lion-tailed Macaque reserve population” would require careful planning. An integrated (One-Plan) approach needs to be developed that aims at the integration of the know-how on the species and the conservation-oriented research interests as provided by the above-mentioned Indian scientists and their institutions. It furthermore should aim at the development of the infrastructural conditions in selected Indian zoos as required for an appropriate management and husbandry aiming at conservation breeding (for a more elaborated outline on this topic see Singh et al. 2012). Research institutions, selected Indian zoos in the range states of the species (like Chennai, Trivandrum, and Mysore) and the EEP should cooperate closely. A small board of experts from these institutions should be established to guide and supervise the project. Previous attempts to establish a breeding programme for the LTM in India and to transfer breeding groups from the USA and from Europe did not work out well due to bureaucratic issues and difficulties with local competence and motivation (see also Krishnakumar & Manimozhi 2000; Singh et al. 2009). The proposed new approach should be designed such that corresponding problems are minimised. It is of particular importance to ‘institutionalise’ captive propagation of the LTM in its country of origin  more strongly. It should include to choose a competent coordinator who permanently overlooks and organises the work above the level of individual zoos and is supported by the Central Zoo Authority of India. A successfully carried out project would also serve as a model for other species and co-operations. It could help to establish Indian zoos as important partners in metapopulation management programmes especially concerning endemic Indian species like the LTM. It is important to note thereby, that time is running out for the development and establishment of international metapopulation management programmes (see Macdonald & Hofer 2011; Powell et al. 2019). They are needed to overcome the sustainability problems threatening many captive populations. Many of them are shrinking for instance due to breeding problems. In terms of climate, available space, and other resources, a number of zoos in India could establish very good keeping systems for the LTM. As elaborated by Singh et al. (2012), conservation breeding in Indian zoos, however, still requires a serious change in professional attitudes, training opportunities and infrastructural requirements. The future of the global captive population of the LTM, for instance, may depend on progress there.

Many zoos and many dedicated people in several countries worked for the survival of the LTM in the wild and for the establishment of a reserve population under human care over many decades. They achieved a lot. Currently, much of what has been achieved with the captive population is at risk. To allow a development ending with a captive LTM population without much breeding and thus with a low conservation potential would be against professional standards and simply sad.


For figures - - click here





Gledhill, L.G. (1985). Analysis of statistics submitted to the Lion-tailed Macaque studbook - North American zoos, pp. 237–238. In: Heltne, P.G. (ed.). The Lion-tailed Macaque: Status and Conservation. Alan R. Liss, New York, 411 pp.

Hill, C.A. (1971). Zoo’s help for a rare monkey. Oryx 11(1): 35–38.

Kaumanns, W., E. Krebs & M. Singh (2006). An endangered species in captivity: Husbandry and management of the Lion-tailed Macaque (Macaca silenus). MyScience 1: 43–71.

Kaumanns, W., M. Singh & A. Sliwa (2013). Captive propagation of threatened primates–the example of the Lion-tailed Macaque Macaca silenus. Journal of Threatened Taxa 5(14): 4825–4839.  

Kaumanns, W. & M. Singh (2015). Towards more individual-based and fitness- oriented captive mammal population management. Current Science 109(6): 1121–1129.  

Kaumanns, W., N. Begum & H. Hofer (2020). “Animals are designed for breeding”: Captive population management needs a new perspective. Journal of Zoo and Aquarium Research 8(2): 76–85.

Krishnakumar, N. & A. Manimozhi (2000). What ails captive breeding programmes in India -Lion-tailed Macaque (LTM) - A case study. Zoos’ Print 15(5): 3–4.

Krishnamani, R. & A. Kumar (2000). Phyto-ecology of the Lion-tailed Macaque (Macaca silenus) habitats in Karnataka, India: Floristic structure and density of food trees. Primate Report 58: 27–56.

Kumar, A. (1987). The ecology and population dynamics of the Lion-tailed Macaque (Macaca silenus) in South India. PhD Thesis. University of Cambridge, 174pp.

Kumar, M.A., M. Singh, H.N. Kumara, A.K. Sharma & C. Bertsch (2001). Male migration in Lion-tailed Macaques. Primate Report 59: 5–18.

Lindburg, D.G., A.M. Lyles & N.M. Czekala (1989). Status and reproductive potential of Lion-tailed Macaques in captivity. Zoo Biology Supplement 1: 5–16.

Lindburg, D.G. & L.G. Gledhill (1992). Captive breeding and conservation of Lion- tailed Macaques. Endangered Species Update 10: 1–4.

Lindburg, D.G. & N.C. Harvey (1996). Reproductive biology of captive Lion-tailed Macaques, pp. 318–341. In: Fa, J.E. & D.G. Lindburg (eds.). Evolution and Ecology of Macaque Societies. Cambridge University Press, Cambridge, 597pp.

Lindburg, D.G., J. Iaderosa & L. Gledhill (1997). Steady-state propagation of captive Lion-tailed Macaques in North American zoos: A conservation strategy, pp. 131–150. In: Walli, J (ed.). Primate Conservation: The Role of Zoological Parks. American Society of Primatologists, Vol.1., Chicago, 252 pp.

Lindburg, D.G. (2001). A century of involvement with Lion-tailed Macaques in North America. Primate Report 59: 51–64.

Macdonald, A.A. & H. Hofer (2011). Editorial: Research in zoos. International Zoo Yearbook 45: 1–6.

Ness, T. (2011). North American Studbook for Lion-tailed Macaques (Macaca silenus). Minnesota Zoological Gardens, Minnesota, 131 pp.

Ness, T. (2013). North American Studbook for Lion-tailed Macaques (Macaca silenus). Minnesota Zoological Gardens, Minnesota, 136 pp.

Penfold, L.M., D. Powell, K. Traylor-Holzer & C.S. Asa (2014). ‘Use it or lose it’: characterization, implications, and mitigation of female infertility in captive wildlife. Zoo Biology 33(1): 20–28.

Powell, D.M., C.L. Dorsey & L.J. Faust (2019). Advancing the science behind animal program sustainability: An overview of the special issue. Zoo Biology 38(1): 5–11.

Sarno, A. (2018). North American Studbook for Lion-tailed Macaques (Macaca silenus). Kansas City Zoo, Kansas City, 175 pp.

Schwibbe, M., M. Singh, W. Kaumanns & C. Knogge (Eds.) (2000). Proceedings of the Fifth International Symposium on Lion-tailed Macaque, Mysore, India, Part 1. Primate Report 58. German Primate Center, Goettingen, 95pp.

Schwibbe, M., M. Singh, W. Kaumanns & C. Knogge (Eds.) (2001). Proceedings of the Fifth International Symposium on Lion-tailed Macaque, Mysore, India, Part 2. Primate Report 59. German Primate Center, Goettingen, 108 pp.

Sharma, A.K. (2002). A study of reproductive behaviour of Lion-tailed Macaque (Macaca silenus) in the rainforests of Western Ghats. PhD Thesis. Department of Zoology, University of Mysore, 102 pp.

Singh, M., A.K. Sharma, E. Krebs & W. Kaumanns (2006). Reproductive biology of Lion-tailed Macaque (Macaca silenus): An important key to the conservation of an endangered species. Current Science 90(6): 804–811.

Singh, M., W. Kaumanns, M. Singh, H.S. Sushma & S. Molur (2009). The Lion- tailed Macaque Macaca silenus (Primates: Cercopihecidae): conservation history and status of a flagship species of the tropical rainforests of the Western Ghats, India. Journal of Threatened Taxa 1(3): 151–157.

Singh, M., W. Kaumanns & G. Umapathy (2012). Conservation-oriented captive breeding of primates in India: Is there a perspective? Current Science 103(12): 1399–1400.

Singh, M., A. Kumar & H.N. Kumara (2020). Macaca silenus. The IUCN Red List of Threatened Species 2020: e.T12559A17951402. Downloaded on 21 August 2021.

Sliwa, A., K. Schad & E. Fienieg (2016). Long-term management plan for the Lion- tailed Macaque (Macaca silenus) European Endangered Species Programme (EEP). Zoologischer Garten Köln, Köln, Germany and European Association of Zoos and Aquaria (EAZA), Amsterdam, Netherlands, 60 pp.

Sliwa, A. & N. Begum (2019). International Studbook 2018 for the Lion-tailed Macaque Macaca silenus Linnaeus, 1758, edition 2018, Vol. 9. Zoologischer Garten Köln, Cologne, v+188 pp.

Sushma, H.S. (2004). Resource utilization and niche separation in sympatric rainforest arboreal mammals. PhD Thesis. University of Mysore, 150 pp.

Umapathy, G. & A. Kumar (2000). Impacts of the habitat fragmentation on time budget and feeding ecology of Lion-tailed Macaques (Macaca silenus) in rainforest fragments of Anamalai Hills, South India. Primate Report 58: 67–82.