Beza Mahafaly Special Reserve: Long-Term Research on Lemurs in Southwestern Madagascar

  • Robert W. SussmanEmail author
  • Alison F. Richard
  • Joelisoa Ratsirarson
  • Michelle L. Sauther
  • Diane K. Brockman
  • Lisa Gould
  • Richard Lawler
  • Frank P. Cuozzo


The Beza Mahafaly Project in southwestern Madagascar was founded in 1975. It was established as a collaborative effort among the University of Madagascar (now University of Antananarivo), Washington University, Yale University, and the local communities for long-term training and research, biodiversity conservation, and socioeconomic development. Beza Mahafaly consists of two noncontiguous forest parcels separated by 10 km that became a protected area (Réserve Spéciale) in 1986: an 80-ha gallery forest and a 520-ha xerophytic spiny forest. The region has a diversity of habitats and a very diverse and highly endemic flora and fauna, including four species of lemurs found in or near the reserve. The ringtailed lemur (Lemur catta) and Verreaux’s sifaka (Propithecus verreauxi) have been the subject of our long-term research. In this chapter we highlight some of the results of this research. Our multidisciplinary studies illustrate the feasibility of collecting long-term data on careers of individual animals and of obtaining large samples on numerous animals, across numerous social groups, in relatively isolated breeding populations. Thus, we can provide insights into many of the demographic, socioecological, anthropogenic, and epidemiological factors that shape the local ringtailed lemur and sifaka population. Here we summarize how ringtailed lemur demographic structure is affected by climatic perturbations (drought); how aspects of general health (parasite loads and dental health) are directly related to habitat, dietary, and anthropogenic factors; how tight birth seasonality in sifaka can elicit stress responses in males associated with increased male aggression, group takeovers, and infanticide risk; how life history schedules are related to evolutionary responses to extreme climatic fluctuations; and how directional selection among sifaka males leads to longer, stronger legs, but not to increase in male body mass relative to females.


Mating Season Gallery Forest Birth Season Ringtailed Lemur Sportive Lemur 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We are grateful to the Government of Madagascar, Madagascar National Parks (MNP formerly ANGAP), the University of Antananarivo (The School of Agronomy (ESSA) especially the Forestry Department of this School (ESSA-Forêts) for permission to work at Beza Mahafaly Special Reserve. We especially appreciate the past and the present help, advice, and logistical support of our colleagues in Madagascar, notably Jo Ajimy, Krista Fish, James Loudon, Teague O’Mara, Jennifer Ness, Rafidisoa Tsiory, the late Rakotomanga Pothin, Ranaivoson Andrianasolo, Razakanirina Daniel, Ramanoelina Panja, Rasoarahona Jean, Rajoelison Gabrielle, Bruno Ramamonjisoa, Ranaivonasy Jeannin, Jessica Scott, and Youssouf Jacky. Our profound thanks go to the members of the BMSR Monitoring Team past and present, and in particular Enafa, Elahavelo, Emady Rigobert, Ellis Edidy, Efitiria, Eboroke Sylvain, Ranarivelo Ny Andry, Randrianarisoa Jeannicq, Ravelonjatovo Sylvia, Razanajaonarivaly Elyse, Ratsirarson Helian, and as well as to the Madagascar National Park team on the ground now led by Andry Randrianandrasana. Enafa’s skill with the blow gun is extraordinary, and together the BMSR Monitoring Team not only made it possible to capture so many animals safely but also, thereafter, to census and monitor them regularly.

We are grateful to our U.S. colleagues who continue to make important contributions to research on sifaka at BMSR, most notably the members of the Beza Mahafaly Sifaka Research Consortium (BMSRC), Marion Schwartz (Sifaka Database Manager), Patricia L. Whitten, Laurie R. Godfrey, and Robert E. Dewar. For veterinary assistance we thank David Miller, Martha Weber, Scott Larsen, Anneke Moresco, Mandala Hunter, Heather Culbertson, Kerry Sondgeroth, Katie Eckert, Rachel Mills, Angie Simai, Jessica Kurek, and Catherine Woods. Long-term support of the BMSR Monitoring Team has kindly been provided by the Liz Claiborne and Art Ortenberg Foundation in collaboration with Yale University and the University of Cambridge. Numerous funding agencies have generously supported our on-going research at BMSR, notably National Science Foundation, World Wildlife Fund, Margot Marsh Biodiversity Foundation, Schwartz Family Foundation Trust, St. Louis Zoo, Primate Conservation Inc., the International Primatological Society, the Indianapolis Zoo (Department of Science and Conservation), Tany Meva, MNP, the Wenner-Gran Foundation, National Sciences and Engineering Research Council of Canada, the University of North Dakota (SSAC; Faculty Research Seed Money Council; Arts, Humanities and Social Sciences Award Committee) and North Dakota EPSCoR, the American Society of Primatologists, the Lindbergh Fund, the John Ball Zoo Society, the National Geographic Society, the University of Colorado-Boulder (Council on Research and Creative Work, Innovative Grant Program), and Washington University. The electronic record of the primate specimens and their assigned Beza Mahafaly Osteological Collection (BMOC) numbers is available online through the University of Massachusetts, Amherst, Collections website (Brockman et al. 2008).

[]. Finally, we thank Peter Kappeler, David Watts, and an anonymous reviewer for their excellent comments on this paper.


  1. Brockman DK (1994) Reproduction and mating system of Verreaux’s sifaka, Propithecus verreauxi, at Beza Mahafaly, Madagascar. PhD thesis, Yale University, New HavenGoogle Scholar
  2. Brockman DK (1999) Reproductive behavior of female Propithecus verreauxi at Beza Mahafaly, Madagascar. Int J Primatol 20:375–398CrossRefGoogle Scholar
  3. Brockman DK, van Schaik CP (2005) Seasonality and reproductive function. In: Brockman DK, van Schaik CP (eds) Seasonality in primates: studies of living and extinct human and nonhuman primates. Cambridge University Press, Cambridge, pp 269–306CrossRefGoogle Scholar
  4. Brockman DK, Whitten PL (1996) Reproduction in free-ranging Propithecus verreauxi: estrus and the relationship between multiple partner matings and fertilization. Am J Phys Anthropol 100:57–69PubMedCrossRefGoogle Scholar
  5. Brockman DK, Whitten PL, Richard AF, Schneider A (1998) Reproduction in free-ranging male Propithecus verreauxi: the hormonal correlates of mating and aggression. Am J Phys Anthropol 105:137–151PubMedCrossRefGoogle Scholar
  6. Brockman DK, Whitten PL, Richard AF, Benander B (2001) Birth season testosterone levels in male Verreaux’s sifaka, Propithecus verreauxi: insights into socio-demographic factors mediating seasonal testicular function. Behav Ecol Sociobiol 49:117–127CrossRefGoogle Scholar
  7. Brockman DK, Godfrey LR, Dollar LJ, Ratsirarson J (2008) Evidence of invasive Felis silvestris predation on Propithecus verreauxi at Beza Mahafaly Special Reserve, Madagascar. Int J Primatol 29:135–152CrossRefGoogle Scholar
  8. Brockman DK, Cobden AK, Whitten PL (2009) Birth season glucocorticoids are related to the presence of infants in sifaka (Propithecus verreauxi). Proc R Soc Lond B 276:1855–1863CrossRefGoogle Scholar
  9. Crowley BE, Godfrey LR, Irwin MT (2011) A glance to the past: subfossils, stable isotopes, seed dispersal, and lemur species loss in southern Madagascar. Am J Primatol 73:25–37PubMedCrossRefGoogle Scholar
  10. Cuozzo FP, Sauther ML (2004) Tooth loss, survival, and resource use in wild ring-tailed lemurs (Lemur catta): implications for inferring conspecific care in fossil hominids. J Hum Evol 46:623–631PubMedCrossRefGoogle Scholar
  11. Cuozzo FP, Sauther ML (2006) Severe wear and tooth loss in wild ring-tailed lemurs (Lemur catta): a function of feeding ecology, dental structure, and individual life history. J Hum Evol 51:490–505PubMedCrossRefGoogle Scholar
  12. Dewar RE, Richard AF (2007) Evolution in the hypervariable environment of Madagascar. Proc Natl Acad Sci USA 104:13723–13727PubMedCrossRefGoogle Scholar
  13. Gentry AH (1993) Diversity and floristic composition of lowland tropical forest in Africa and South America. In: Goldblatt P (ed) Biological relationships between Africa and South America. Yale University Press, New Haven, pp 500–546Google Scholar
  14. Gould L (1997a) Intermale affiliative behavior in ringtailed lemurs (Lemur catta) at the Beza Mahafaly Reserve, Madagascar. Primates 38:15–30CrossRefGoogle Scholar
  15. Gould L (1997b) Affiliative relationships between adult males and immature group members in naturally occurring ringtailed lemurs (Lemur catta). Am J Phys Anthropol 103:163–171PubMedCrossRefGoogle Scholar
  16. Gould L (2006) Lemur catta ecology: what we know and what we need to know. In: Gould L, Sauther ML (eds) Lemurs: ecology and adaptation. Springer, New York, pp 255–274Google Scholar
  17. Gould L, Ziegler TE (2007) Variation in fecal testosterone levels, inter-male aggression, dominance rank and age during mating and post-mating periods in wild adult male ring-tailed lemurs (Lemur catta). Am J Primatol 69:1325–1339PubMedCrossRefGoogle Scholar
  18. Gould L, Sussman RW, Sauther ML (1999) Natural disasters and primate populations: the effects of a 2-year drought on a naturally occurring population of ring-tailed lemurs (Lemur catta) in southwestern Madagascar. Int J Primatol 20:69–84CrossRefGoogle Scholar
  19. Gould L, Sussman RW, Sauther ML (2003) Demographic and life-history patterns in a population of ring-tailed lemurs (Lemur catta) at Beza Mahafaly Reserve, Madagascar: a 15-year perspective. Am J Phys Anthropol 120:182–194PubMedCrossRefGoogle Scholar
  20. Gould L, Ziegler TE, Wittwer DJ (2005) Effects of reproductive and social variables on fecal glucocorticoid levels in a sample of adult male ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Reserve, Madagascar. Am J Primatol 67:5–23PubMedCrossRefGoogle Scholar
  21. Heckman KL, Rasoazanabary E, Machlin E, Godfrey LR, Yoder AD (2006) Incongruence between genetic and morphological diversity in Microcebus griseorufus of Beza Mahafaly. BMC Evol Biol 6:e98. doi: 10.1186/1471-2148-6-98 CrossRefGoogle Scholar
  22. Kappeler PM, Fichtel C (2011) A 15-year perspective on the social organization and life history of sifaka in Kirindy Forest. In: Kappeler PM (ed) Long-term field studies of primates. Springer, HeidelbergGoogle Scholar
  23. Lawler RR (2007) Fitness and extra-group reproduction in male Verreaux’s sifaka: an analysis of reproductive success from 1989–1999. Am J Phys Anthropol 132:267–277PubMedCrossRefGoogle Scholar
  24. Lawler RR (2009) Monogamy, male-male competition, and mechanisms of sexual dimorphism. J Hum Evol 57:321–325PubMedCrossRefGoogle Scholar
  25. Lawler RR, Richard AF, Riley MA (2003) Genetic population structure of the white sifaka (Propithecus verreauxi verreauxi) at Beza Mahafaly Special Reserve, southwest Madagascar (1992–2001). Mol Ecol 12:2307–2317PubMedCrossRefGoogle Scholar
  26. Lawler RR, Richard AF, Riley MA (2005) Intrasexual selection in Verreaux’s sifaka (Propithecus verreauxi verreauxi). J Hum Evol 48:259–277PubMedCrossRefGoogle Scholar
  27. Lawler RR, Caswell H, Richard AF, Ratsirarson J, Dewar RE, Schwartz M (2009) Demography of Verreaux’s sifaka in a stochastic rainfall environment. Oecologia 161:491–504PubMedCrossRefGoogle Scholar
  28. Loudon JE (2009) The parasite ecology and socioecology of ring-tailed lemurs (Lemur catta) and Verreaux’s sifaka (Propithecus verreauxi) inhabiting the Beza Mahafaly Special Reserve. PhD thesis, University of Colorado, BoulderGoogle Scholar
  29. Loudon JE, Sauther ML, Fish KD, Hunter-Ishikawa M, Jack IAY (2006) One reserve, three primates: applying a holistic approach to understand the interconnections among ring-tailed lemurs (Lemur catta), Verreaux’s sifaka (Propithecus verreauxi), and humans (Homo sapiens) at Beza Mahafaly Special Reserve, Madagascar. Ecol Environ Anthropol 2:54–74Google Scholar
  30. Loudon JE, Sponheimer M, Sauther ML, Cuozzo FP (2007) Intraspecific variation in hair δ13C and δ15N values of ring-tailed lemurs (Lemur catta) with known individual histories, behavior, and feeding ecology. Am J Phys Anthropol 133:978–985PubMedCrossRefGoogle Scholar
  31. Millette JB, Sauther ML, Cuozzo FP (2009) Behavioral responses to tooth loss in wild ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Special Reserve, Madagascar. Am J Phys Anthropol 140:120–134PubMedCrossRefGoogle Scholar
  32. Morris WF, Altmann J, Brockman DK, Cords M, Fedigan LM, Pusey AE, Stoinski TS, Bronikowski AM, Alberts SC, Strier KB (2011) Low demographic variability in wild primate populations: fitness impacts of variation, covariation, and serial correlation in vital rates. Am Nat 177:e14–28. doi: 10.1086/657443 PubMedCrossRefGoogle Scholar
  33. Rainwater TR, Sauther ML, Rainwater KAE, Mills RE, Cuozzo FP, Zhang B, McDaniel LN, Abel MT, Marsland EJ, Weber MA, Jack IAY, Platt SG, Cobb GP, Anderson TA (2009) Assessment of organochlorine pesticides and metals in ring-tailed lemurs (Lemur catta) at Beza Mahafaly Special Reserve, Madagascar. Am J Primatol 71:998–1010CrossRefGoogle Scholar
  34. Ratsirarson J (2003) Réserve Spéciale de Beza Mahafaly. In: Goodman SM, Benstead JP (eds) The natural history of Madagascar. University of Chicago Press, Chicago, pp 1520–1525Google Scholar
  35. Ratsirarson J (2008) La Réserve Spéciale de Beza Mahafaly. In: Goodman SM (ed) Paysages naturels et biodiversité de Madagascar. Muséum National d’Histoire Naturelle, Paris, pp 615–626Google Scholar
  36. Ratsirarson J, Randrianarisoa J, Ellis E, Emady RJ, Efitroarany RJ, Razanajaonarivalona EH, Richard AF (2001) Beza Mahafaly: écologie et réalités socio-économiques. Rech Dev B 18:1–104Google Scholar
  37. Richard AF, Rakotomanga P, Schwartz M (1991) Demography of Propithecus verreauxi at Beza Mahafaly, Madagascar: sex ratio, survival, and fertility, 1984–1988. Am J Phys Anthropol 84:307–322CrossRefGoogle Scholar
  38. Richard AF, Rakotomanga P, Schwartz M (1993) Dispersal by Propithecus verreauxi at Beza Mahafaly, Madagascar: 1984–1991. Am J Primatol 30:1–20CrossRefGoogle Scholar
  39. Richard AF, Dewar RE, Schwartz M, Ratsirarson J (2000) Mass change, environmental variability and female fertility in wild Propithecus verreauxi. J Hum Evol 39:381–391PubMedCrossRefGoogle Scholar
  40. Richard AF, Dewar RE, Schwartz M, Ratsirarson J (2002) Life in the slow lane? Demography and life histories of male and female sifaka (Propithecus verreauxi verreauxi). J Zool Lond 256:421–436CrossRefGoogle Scholar
  41. Sauther ML, Cuozzo FP (2008) Somatic variation in living, wild ring-tailed lemurs (Lemur catta). Folia Primatol 79:55–78PubMedCrossRefGoogle Scholar
  42. Sauther ML, Cuozzo FP (2009) The impact of fallback foods on wild ring-tailed lemur biology: a comparison of intact and anthropogenically disturbed habitats. Am J Phys Anthropol 140:671–686PubMedCrossRefGoogle Scholar
  43. Sauther ML, Sussman RW, Cuozzo F (2002) Dental and general health in a population of wild ring-tailed lemurs: a life history approach. Am J Phys Anthropol 117:122–132PubMedCrossRefGoogle Scholar
  44. Sauther ML, Fish KD, Cuozzo FP, Miller DS, Hunter-Ishikawa M, Culbertson H (2006) Patterns of health, disease, and behavior among wild ringtailed lemurs, Lemur catta: effects of habitat and sex. In: Jolly A, Sussman RW, Koyama N, Rasamimanana H (eds) Ringtailed lemur biology. Springer, New York, pp 313–331CrossRefGoogle Scholar
  45. Sponheimer M, Codron D, Passey BH, de Ruiter DJ, Cerling TE, Lee-Thorp JA (2009) Using carbon isotopes to track dietary change in modern, historical, and ancient primates. Am J Phys Anthropol 140:661–670PubMedCrossRefGoogle Scholar
  46. Steele E (1975) Needed: virtue and money. Defenders Wildlife 50:90Google Scholar
  47. Strier KB, Altmann J, Brockman DK, Bronikowski AM, Cords M, Fedigan LM, Lapp H, Liu X, Morris WF, Pusey AE, Stoinski TS, Alberts SC (2010) The Primate Life History Database: a unique shared ecological data resource. Methods Ecol Evol 1:199–211PubMedCrossRefGoogle Scholar
  48. Sussman RW (1991) Demography and social organization of free-ranging Lemur catta in the Beza Mahafaly Reserve, Madagascar. Am J Phys Anthropol 84:43–58CrossRefGoogle Scholar
  49. Sussman RW (1992) Male life history and intergroup mobility among ringtailed lemurs (Lemur catta). Int J Primatol 13:395–413CrossRefGoogle Scholar
  50. Sussman RW, Rakotozafy A (1994) Plant diversity and structural analysis of a tropical dry forest in southwestern Madagascar. Biotropica 26:241–254CrossRefGoogle Scholar
  51. Sussman RW, Ratsirarson J (2006) Beza Mahafaly Special Reserve: a research site in southwestern Madagascar. In: Jolly A, Sussman RW, Koyama N, Rasamimanana HR (eds) Ringtailed lemur biology. Springer, New York, pp 43–51Google Scholar
  52. Sussman RW, Green GM, Porton I, Andrianasolondraibe OL, Ratsirarson J (2003) A survey of the habitat of Lemur catta in southwestern and southern Madagascar. Primate Conserv 19:32–57Google Scholar
  53. Sussman RW, Sweeney S, Green GM, Porton I, Andrianasolondraibe OL, Ratsirarson J (2006) A preliminary estimate of Lemur catta population density using satellite imagery. In: Jolly A, Sussman RW, Koyama N, Rasamimanana HR (eds) Ringtailed lemur biology: Lemur catta in Madagascar. Springer, New York, pp 16–31CrossRefGoogle Scholar
  54. White F (1983) The vegetation of Africa: a descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. UNESCO (United Nations Educational, Scientific and Cultural Organization), ParisGoogle Scholar
  55. Wingfield JC, Hegner RE, Dufty AM Jr, Ball GF (1990) The “Challenge Hypothesis”: theoretical implications for patterns of testosterone secretion, mating systems, and breeding strategies. Am Nat 136:829–846CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2012

Authors and Affiliations

  • Robert W. Sussman
    • 1
    Email author
  • Alison F. Richard
    • 2
  • Joelisoa Ratsirarson
    • 3
  • Michelle L. Sauther
    • 4
  • Diane K. Brockman
    • 5
  • Lisa Gould
    • 6
  • Richard Lawler
    • 7
  • Frank P. Cuozzo
    • 8
  1. 1.Department of Anthropology and Environmental ScienceWashington UniversitySt. LouisUSA
  2. 2.Yale UniversityNew HavenUSA
  3. 3.School of Agronomy, Department of Water and Forestry (ESSA/Forêts)University of AntananarivoAntananarivoMadagascar
  4. 4.Department of AnthropologyUniversity of ColoradoBoulderUSA
  5. 5.Department of AnthropologyUniversity of North CarolinaCharlotteUSA
  6. 6.Department of AnthropologyUniversity Of VictoriaVictoriaCanada
  7. 7.Department of AnthropologyJames Madison UniversityHarrisonburgUSA
  8. 8.Department of AnthropologyUniversity of North DakotaGrand ForksUSA

Personalised recommendations