Current research activities:

My research team is involved in several different areas of investigation. At present, most of my data are collected at the Buffalo Zoo. We have used a number of different data collection systems, but at present we are using an Excel app (for tablet or phone) for data collection. The primary purpose of these observations is to establish a “baseline” of normal behavior for the gorilla group. This allows us to then systematically examine the effect of environmental or social changes on the gorilla group; for example, the addition or removal of an individual, a diet change, or a new type of enrichment. This also provides a rich source of data for a variety of applied research questions.

Training observers takes a considerable amount of time, and it is not until you are trained, pass reliability tests, and “graduate” to the research team that you are truly making a contribution to the project. It’s likely to take you several weeks at mimimum to learn the skills you need to collect data. Then you’ll need to practice, on your own or with another researcher, until you are completely comfortable with the procedure. At this point, you may take an observer reliability test and formally begin to collect “real” data. Some students feel ready to take their reliability test after a few weeks, others practice for a whole semester. Either way is fine.

Once you have “passed”, you’ll be scheduled for 1-2 observation sessions a week at the zoo. Sessions last about 2 hours, and are conducted in the “AM” (between 10 and 12:30) or the “PM” (between 1 and 4). This means the schedule is pretty flexible. Students can observe singly or in pairs.

The second type of research that I do involves hormonal assessments. We collect fecal samples (well, zookeepers collect them, then send them to me). This may sound messy–and it is–but it allows us to explore hormonal changes that may affect, or be affected by, behavior. Current studies involve changes in reproductive hormones in white-cheeked gibbon females as they mature, and measures of stress hormones in gorillas in response to a visual barrier. This work requires great attention to detail in the lab, and a very good ability to follow directions. We use assay kits, and students who are interested practice the methods before actually running assays. Non-invasive endocrinological assessment pairs very well with observational behavioral data.

Once students have mastered skills, they may develop their own research projects (typically during their senior year). To date, “Team Ape” has expanded well beyond apes. We have completed studies on octopus, vampire bats, addax, horses, elephants, giraffe, Humboldt penguins, lions, tigers, and polar bears.

“Team Ape” has been a highly productive research team, because we work together and contribute to our long-term data base.

Team Ape with Ape 4-14

Team Ape members (L-R) Chase LaDue ’14, Sydney Cheroff ’18 and Liam Kelly ’15, with “Sidney” gorilla at the Buffalo Zoo

Members of Team Ape, past and present, at the Animal Behavior Society Meeting, 2017

Members of Team Ape, past and present, at the Animal Behavior Society Meeting, 2017

End-of-year party 2016, with seniors: Sara, Charles, and Megan

End-of-year party 2016, with seniors: Sara, Charles, and Megan

team ape 2015-16

Team Ape end-of-year party, December 2016

Domenic Romanello '18, Sydney Chertoff '18, and Margret Halfdanardottir '18 represent Team Ape in South Africa, 2017

Domenic Romanello ’18, Sydney Chertoff ’18, and Margret Halfdanardottir ’18 represent Team Ape in South Africa, 2017

Sydney destroying a defunct palm pilot, 2016

Sydney destroying a defunct palm pilot, 2016


Team Ape members Sydney Chertoff ’18 and Kelsey Trumpp ’15 model “gibbon jackets” at the education outreach fair, Animal Behavior Society meeting, Anchorage, AK 2015

The end of the palm pilots!

The end of the palm pilots!

Group photo

Team Ape members Matthew LeFauve ’14, Chase LaDue ’14 and Liam Kelly ’15 with Professor Jeanne Altmann at the 2014 Animal Behavior Society meeting in Princeton, NJ

Margret H, Magda L, and Sara S, end of year party, 2016

Margret H, Magda L, and Sara S, end of year party, 2016



Faith Burns, Ray Kleinfelder, Adrienne Rothenberg, Gary Steele
2010-2011 Faith Burns, Kelly Fox, Ray Kleinfelder, Chase LaDue, Matthew LeFauve, Alissa Mitchell, Heather Paye, Adrienne Rothenberg
2011-2012 Tyler Carver, Alex Hofner, Chase LaDue, Matthew LeFauve, Macy Madden,  Heather Paye, Lindsey Perkes-Smith, Madeline Prush, Lindsey Robbins, Claire Taberski, Kelsey Trumpp
2012-2013 Tyler Carver, Heather Desorcie, Mackenzie Green, Liam Kelly, Chase LaDue, Matthew LeFauve, Macy Madden, Heather Paye, Lindsey Perkes-Smith, Madeline Prush, Charles Ritzler, Claire Taberski, Kelsey Trumpp, Chelsa Wlodarczyk
2013-2014 Mallory Abel, Sydney Chertoff, Chase LaDue, Heather Desorcie, Mackenzie Green, Liam Kelly,  Matthew LeFauve, Macy Madden, Lauren McGee, Charles Ritzler, Claire Taberski,  Kelsey Trumpp, Chelsa Wlodarczyk, Nick Woodard
2014-2015 Mallory Abel, Tara Benczkowski,  Sydney Chertoff, Heather Desorcie, Olivia Hoffman, Liam Kelly, Megan Kozlowski, Lauren McGee, Eli Musik-Kotlowski, Charles Ritzler,  Sara Sperber, Rosemary Trahan, Kelsey Trumpp, Nick Woodard

Taryn Bansmer, Koryndee Bowlsby, Sydney Chertoff, Jon-Ellyn Cullen, Heather Desorcie, Margret Halfdanardottir, Olivia Hoffman, Anna Kidder,  Megan Kowlowski, Magda Lenczewski, Eli Musik-Kotlowski, Hanna Ridge, Charles Ritzler, Caeley Robinson, Brittany Rohacs, Domenic Romanello, Macy Rounds, Sara Sperber, Heather Spoor, Kari Tuite, London Wolff


Taryn Bansmer, Koryndee Bowlsby, Amy Bucklaew, Sydney Chertoff, Lexie Haley, Margret Halfdanardottir, Anna Kidder,  Magda Lenczewski, Eli Musik-Kotlowski, Hannah Ridge, Caeley Robinson, Domenic Romanello, Macy Rounds, Nicole Socie, Kari Tuite

 2017-2018 Taryn Bansmer, Adam Bator, Amy Bucklaew, Sydney Chertoff, Misa Gamble, Erin Guntrum, Lexie Haley, Margret Halfdanardottir, Megan Miller, Sera Muscoreil,  Joshua Pajek, Hannah Ridge, Abigail Robinson, Caeley Robinson, Domenic Romanello, Emie Tanner, Kari Tuite



Taryn Bansmer, Adam Bator, Amy Bucklaew, Misa Gamble, Erin Guntrum, Lexie Haley, Darnell Hester, Megan Miller, Joshua Pajek,  Abigail Robinson, Caeley Robinson, Emie Tanner, Kari Tuite



Adam Bator, Maddie Blackwell, Amy Bucklaew, Misa Gamble, Erin Guntrum, Lexie Haley, Kayla Kitchen, Megan Leight, Angie Liriano, Erica Meehan, Megan Miller, Joshua Pajek, Caeley Robinson, Clarice Scarpace, Alisa Slawinski, Emie Tanner, Hunter Wood

Publications (with links to PDF’s when available; for other reprints, please email me)


Early papers (1995 and earlier)

Weinberg, S. and Candland, D. 1981. “Stone-grooming” in Macaca fuscata. Am. J. Primatol. 1:465-468.

Weinberg, S. 1987. Possible cleaning symbiosis between Pica pica and Odocoileus hemionus. Southwestern Natur. 32:138-139.

Margulis, S. 1989. Introduction of a male colobus to an existing all-male group. Proceedings, 15th National Conference, American Association of Zoo Keepers, 31-37.

Margulis, S.W. 1993. Mate choice in rocky mountain mule deer (Odocoileus hemionus hemionus) bucks: is there a preference for does without fawns? Ethol. Ecol. Evol., 5:115-119.

Margulis, S.W., Altmann, J., and Ober, C. 1993. Sex-biased lactational duration in a human population and its reproductive costs. Behav. Ecol. Sociobiol., 32:41-45.

Margulis, S.W., Abbott, D.H., and Saltzman, W. 1995. Behavioral and hormonal changes in female naked mole-rats following removal of the breeding female from a colony. . Hormones and Behavior, 29:227-247.

Margulis, S.W., Chin, J., Warneke, M., Dubach, J., and Lindgren, V. 1995. The Y translocation of Callimico goeldii. Int’l. J. Primatol., 16:145-155.


Margulis. S. W. and Altmann, J. 1997. Behavioural risk factors in the reproduction of inbred and outbred oldfield mice., Animal Behaviour, 54:397-408.

Margulis, S. W. 1997. Inbreeding-based bias in parental responsiveness to litters of oldfield mice. Behav. Ecol. Sociobiol., 41:177-184.

Margulis, S. W. 1997. Linking observation and conservation: baboon research and conservation at Brookfield Zoo. Bison, 11:10-19.

Margulis, S. W. 1998. Differential effects of inbreeding at juvenile and adult life history stages in Peromyscus polionotus J. Mamm. 79:326-336.

Margulis, S. W. 1998. Relationships among parental inbreeding, parental behaviour, and offspring viability in oldfield mice.. Anim. Behav. 55:427-438.

Margulis, S. W., and Altmann, J. 1998. Linking observation and conservation: using baboons to teach zoo visitors about behavior and conservation. Pp. 125-128 in: Proceedings, 1998 AZA Annual Conference, Tulsa, OK.

Margulis, S. W., Havlik, M., and Ward, D. 2000 Behavior Matters, Connections curriculum unit, Brookfield Zoo and the Center for Learning Technologies in Urban Schools.

2001 – 2010

Margulis, S. W., Whitham, J., and Ogorzalek, K. 2001. Social interactions among female gorillas before and after the introduction of a new silverback. Pp. 79-82 in: Proceedings, The Apes: Challenges for the 21st Century, Chicago Zoological Society.

Margulis, S. W. and Walsh, A. 2001. Establishing a Keeper-Based Behavioral Monitoring Program: A Top-Down/ Bottom-Up Approach. Pp. 55-62 in: Proceedings, 2001 AZA Annual Conference. St. Louis, MO

Margulis, S.W. and Walsh, A. 2002. The effects of inbreeding on testicular sperm concentration in Peromyscus polionotus.. Reprod. Fertil. Dev., 14:63-67.

Koeninger Ryan, K., Lacy, R. C. and Margulis, S. W. 2002. Impacts of kinship and inbreeding on components of fitness. Reproduction and Integrated Conservation Science (W. V. Holt, A. R. Pickard, J. C. Rodger, D. E. Wildt, eds). Cambridge University Press.

Margulis, S.W., Whitham, J. C., and Ogorzalek, K. 2003. Silverback male presence and group stability in gorillas (Gorilla gorilla gorilla). Folia Primatologica, 74: 92-96.

Margulis, S.W., and Weber, T. 2003. Keepers and behavioral research: fostering the connection. Animal Keepers’ Forum, 30:285-287.

Margulis, S.W., Hoyos, C. and Anderson, M. 2003. The effect of felid activity on zoo visitor interest. Zoo Biology, 22:587-599.

Atsalis, S., Margulis, S.W., Bellem, A. and Wielebnowski, N. 2004. Sexual behavior and hormonal estrus cycles in captive aged lowland gorillas (Gorilla gorilla). American Journal of Primatology, 62: 123-132.

Margulis, S.W. 2004. Classroom activities in behavior. Pp. 521-528 in: Encyclopedia of Animal Behavior (M.Bekoff, ed.), Greenwood publishing company.

Margulis, S.W., Nabong, M., Alaks, G., Walsh, A., and Lacy, R.C. 2005. The role of early experience on subsequent parental behavior and reproductive success in oldfield mice (Peromyscus polionotus). Animal Behaviour, 69: 627-634.

Margulis, S. W. 2005. The collection of behavioral data. Pp. 23-26 in: Learning the Skills of Research: Animal Behavior Exercises in Laboratory and Field (E. Jacob, and M. Hodge, eds.), Instructor’s Resource CD to accompany Animal Behavior: An Evolutionary Approach (8th edition), J. Alcock. McGraw-Hill.

Margulis, S. W. 2005. Candid Camera: comparing and contrasting observation methods. Pp. 18-in: Learning the Skills of Research: Animal Behavior Exercises in Laboratory and Field (E. Jacob, and M. Hodge, eds.). Instructor’s Resource CD to accompany Animal Behavior: An Evolutionary Approach (8th edition), J. Alcock McGraw-Hill.

Atsalis, S., Kasnicka, C., Margulis, S., McGee, J., and Pruett-Jones, M. 2005. EthoTrak, lessons learned from electronic behavioral data monitoring. Proceedings, Association of Zoos and Aquariums Annual Conference (available electronically on website), 2005.

Margulis, S.W., Rafacz, M., and Jacobs, B. 2006. The effectiveness of environmental enrichment: Lessons learned and rules of thumb. In: Proceedings, 2005 International Conference on Environmental Enrichment, Columbia University, New York (N. Clum, S. Silver, P. Thomas, eds). Wildlife Conservation Society, NY.

Atsalis, S, and Margulis, S.W. 2006. Sexual and hormonal cycles in geriatric western lowland gorillas.. Int J Primatol. 27: 1663-1687.

Atsalis, S., Walsh, A., Pruett-Jones, M., Margulis, S., Apura, Z., and Oakley, T. 2006. EthoTrak: Electronic Behavioral Data Monitoring-A Manual. E- publication, Brookfield Zoo, 2006.

Margulis, S.W., Atsalis, S., Bellem, A., and Wielebnowski, N.  2007.  Assessment of reproductive behavior and hormonal cycles in geriatric western lowland gorillas. Zoo Biology , 26: 117-139 .

Sayer, E., Whitham, J., and Margulis, S.W. 2007. Who needs a forelimb anyway? Locomotor, postural and manipulative behavior in a one-armed gibbon.. Zoo Biology, 26:216-222.

Margulis, S.W. and Westhus, E. 2008. Evaluation of different observational sampling regimes for use in zoological parks. Applied Animal Behaviour Science, 110: 363-376.

Atsalis, S., Margulis, S.W., and Hof, P.R. (eds). 2008. Primate Reproductive Aging: Cross-taxon Perspectives. Karger Press.

Atsalis, S. and Margulis, S.W. 2008. Perimenopause and menopause-documenting life changes in aging female gorillas. Pp. 119-146 in: Atsalis, S., Margulis, S.W., and Hof, P.R. (eds). Primate Reproductive Aging: Cross-taxon Perspectives. Karger Press.

Atsalis, S. and Margulis, S.W. 2008. Primate reproductive aging: from lemurs to humans. Pp.186-194 in: Atsalis, S., Margulis, S.W., and Hof, P.R. (eds). Primate Reproductive Aging: Cross-taxon Perspectives. Karger Press.

Gentry, L., and Margulis, S.W. 2008. Behavioral effects of introducing pied tamarin (Saguinus bicolor) to black howler monkey (Alouatta caraya) and white-faced saki (Pithecia pithecia) in a zoological park.. American Journal of Primatology, 70: 505-509.

Margulis, S.W. and Pruett-Jones, M. 2008. Integrating science and husbandry: less is more. Pp. 25-38 in: The Well-being of Animals in Zoo and Aquarium Sponsored Research: Putting Best Practices Forward (T Bettinger and J Bielitzki, eds.). Scientists Center for Animal Welfare, Greenbelt, MD.

Margulis, S.W. 2008. Book review: Gorilla Society: Conflict, Compromise, and Cooperation Between the Sexes. American Journal of Physical Anthropology, 137: 241.

Watters, J.V., Margulis, S.W., and Atsalis, S. 2009. Behavioral monitoring in zoos and aquariums: a tool for guiding husbandry and directing research. Zoo Biol. 28: 35-48.

Rafacz, M., Margulis, S., and Santymire, R. 2009. Comparing the role of androgens in paternal care between gibbon species. Gibbon Journal, 5: 74-80.

Margulis, S.W. 2010. Methodology: sources of measurement error, reliability and validity. Encyclopedia of Animal Behavior (M. Breed and J. Moore, eds), pp 424-428. Elsevier, UK.


Fuller, G., Margulis, S., and Santymire, R. 2011. The effectiveness of indigestible markers for identifying individual animal feces and their prevalence of use in North American zoos. Zoo Biology, 30: 379-398.

Margulis, S.W., Burns, F., and Rothenberg, A. 2011. Sex ratio bias in managed populations of Hylobatids. Folia Primatologica, 211: 224-235.

Sarfaty, A. Margulis, S.W., Atsalis, S., and Sarfaty, A. 2011. Effects of combination birth control on estrous behavior in captive Western lowland gorillas, Gorilla gorilla gorilla. Zoo Biology, 31: 350-361.

Margulis, S.W., Steele, G.R., and Kleinfelder, R. E. III. 2012. Use of buckets as tools by Western lowland gorillas. Zoo Biology, 31: 260-266.

Rafacz, M.L., Margulis, S.W., and Santymire, R.M. 2012. Hormonal correlates of paternal care differences in the Hylobatidae. American Journal of Primatology, 74: 247-260.

Segurel, L.Thompson, E.E., Flutre, T., Lovstad, J.,Venkat, A., Margulis, S., Moyse, J., Ross, S.,Gamble, K., Sella, G., Ober, C. and Przeworski, M. 2012. Blood ties: ABO is a trans-species polymorphism in primates. PNAS. 109: 18493–18498.

Rafacz, M.L., Margulis, S.W., and Santymire, R.M. 2013. Hormonal and behavioral patterns of reproduction in female hylobatids. Animal Reproduction Science, 137: 103-112.

Robbins, L. and Margulis, S.W. 2014. The effects of auditory enrichment on gorillas. Zoo Biology. 33: 197-203.

LaDue, C., Madden, M., Perkes-Smith, L., and Margulis, S. 2014. Behavioral changes associated with pregnancy and infant development in captive gorillas. Animal Keepers’ Forum, 41:80-83.

LaDue, CA, Scott, NL, Margulis, SW. 2014. A Survey of musth among captive male elephants in North America: updated results and implications for management. Journal of the Elephant Managers Association.25: 18-24.

LeFauve, MK, and Margulis, SW. 2015. Functionality in tool use in Western lowland gorillas (Gorilla gorilla gorilla). Animal Behavior and Cognition.2: 96-104.

Harl, H., Stevens, L., Margulis, S., and Petersen, J. 2016. Gibbon aggression during introductions: an international survey. Journal of Applied Animal Welfare Science. DOI:   10.1080/10888705.2015.1130631

Robbins, L. and Margulis, S.W. 2016. Music for the birds: effects of auditory enrichment on captive bird species. Zoo Biology.35: 29-34.

Margulis, S.W., Volle, K.A., LaDue, C.A., Atsalis, S. 2017. What necropsy reports can tell us about menopausal and age-related changes in Western lowland gorillas. Journal of Zoo and Aquarium Research. 5: 11-15.

Margulis, S.W. 2017. Methodology: sources of measurement error, reliability and validity. Reference Module in the Life Sciences. Elsevier, UK.

Margulis, S.W. 2017. Zoos as venues for research: changes in focus, changes in perception. Chapter in edited volume, Increasing Legal Equality for Zoo Animals: Justice on the Ark (J. Donahue, ed).

Braude, S., Margulis, SW, Broder, D. 2017. The study of animal behavior provides valuable opportunities for original science fair projects: recommendations from The Animal Behavior Society, Education Committee. American Biology Teacher, 79: 438-441.

Lenczewski, M., Halfdanardottir, M., Margulis, SW. 2017. Now you see me (now you  don’t): a visual barrier study on a zoo-housed group of Western lowland gorillas. Anthropology: Bachelors to Dissertation (University at Buffalo), 4: 77-91.

Chertoff, S., Margulis, SW, and Rodgers, J. 2018. Visual processing of faces in juvenile Western lowland gorillas without the use of training or reinforcement: a pilot study.  Animal Behavior and Cognition, 5: 292-299.

Margulis, SW, and Romanello, D. 2019. Data coding, measurement error, and reliability. In: Encyclopedia of Animal Behavior, 2nd Edition. Pp 447-454.

LaBarge, LR, Hill, RA, Berman, CM, Margulis, SW, and Allan, ATL. 2019. Anthropogenic influences on primate antipredator behavior and implications for research and conservation. American Journal of Primatology. DOI: 10.1002/ajp.23087.

LaBarge, LR, Allan, ATL, Berman, CM, Margulis, SW, and Hill, RA. 2020. Reactive and pre-emptive spatial cohesion in a social primate. Animal Behaviour, 163: 115-126.