Dr Graham L Banes

About Me

I am an Associate Scientist in the Wisconsin National Primate Research Center at the University of Wisconsin - Madison. My research combines field, genomic and computational approaches to the study and conservation of non-human primates, focusing on the effects of human intervention in otherwise natural evolutionary processes. My laboratory hosts post-doctoral scientists, graduate students and numerous undergraduates, and integrates research, teaching and outreach with the University's wider community. Orang-utans (Pongo spp.) are our primary taxa of focus.

I originally trained in Zoology at the University of Aberdeen in Scotland, earning a Bachelor's degree (B.Sc. Hons, 2008) following study at the University of British Columbia (2006-2007) in Vancouver, Canada. For my honours thesis, I investigated the effects of forest disturbance on wild Bornean orang-utans and conducted line-transect surveys of orang-utan nests to re-determine population density in Tanjung Puting National Park, Indonesia. I then read for a Master's degree (M.Phil. by Research, 2010) in Biological Anthropological Science at Darwin College, University of Cambridge, England, where I inferred the subspecies composition of reintroduced orang-utans at Camp Leakey, in the same national park, and assessed their mitochondrial DNA diversity. I then completed a Ph.D. in Biological Anthropology (2013), also at the University of Cambridge, for which I performed genetic analyses of orang-utan social structure, mate choice and reproductive success, primarily determining the extent to which Kusasi — the dominant male at Camp Leakey for more than a decade — fathered offspring over socially subordinate, non-cheek-padded males. To date, I am one of only three Western scientists to earn their PhD wholly from studies of orang-utans at Camp Leakey, after its co-founder, Dr Biruté Galdikas (1978), and Dr Gary Shapiro (1985).

During my doctorate, I completed an 18-month residency in the Department of Primatology at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, before moving to the Department of Evolutionary Genetics as a post-doctoral scientist. I then joined the Chinese Academy of Sciences and Max Planck Society Partner Institute for Computational Biology in Shanghai, China, where I served three-and-a-half years as a post-doctoral fellow. I joined the University of Wisconsin - Madison in Fall 2017.

My work to date has been generously funded with more than 1,250,000 USD in research support, from the University of Aberdeen (Expedition, Alumni Annual and Small Grants Funds), the Royal Geographical Society with Institute of British Geographers, the Royal Scottish Geographical Society, the Gilchrist Educational Trust, the Orangutan Foundation UK, the North of England Zoological Society at Chester Zoo, the John Reid Trust, the Primate Society of Great Britain (through a Charles A Lockwood Memorial Grant), the Arcus Foundation, Darwin College, the Leverhulme Centre for Human Evolutionary Studies at the University of Cambridge, the Division of Biological Anthropology at the University of Cambridge, the Cambridge Ridgeway-Venn studentship, the Miss Millie Foundation, the Max Planck Society, the American Association of Zoo Keepers Los Angeles Chapter, the Orang Utan Republik Foundation (as recipient of the 2012 LP Jenkins Memorial Fellowship), the German Academic Exchange Service, Hendrix College, the Chinese Academy of Sciences, the National Natural Science Foundation of China (Grant Nos. 31450110070 and 91331203), the Sacramento Zoo Conservation Fund, Como Park Zoo and Conservatory, Orangutan Outreach, The Ronna Noel Charitable Trust, the Morris Animal Foundation, the Association of Zoos and Aquariums and by the generosity of private donors. I am delighted to extend my research into 2020 with the renewed support of the Ronna Noel Charitable Trust, and with grants from the Eppley Foundation for Research and the Shanghai Municipal Forestry Department; the latter in collaboration with Shanghai Zoo.

My work has been especially well received by the media, particularly following publication of our study on male orang-utan bimaturism in September 2015, which was featured by The Washington Post, The Daily Mail, IFL Science and Serious Science, among more than 200 other outlets, plus by the German-language Leipziger Volkszeitung. Our 2016 manuscript on orang-utan subspecies hybridization was highlighted in the March print edition of Science magazine, covered separately online by ScienceShots, and featured by The Daily Mail and Mongabay, among others. My career as a scientist has been featured in Poland's national daily newspaper, Gazeta Wyborcza, in the Hebrew-language magazine, Mishpacha, and by Psychology Today magazine. In response to my collaborations with Chinese zoos, I was profiled in Chinese by The New York Times.

I currently Chair the Scientific Advisory Committee of the Orang Utan Republik Foundation, a California-registered 501(c)(3) non-profit (since 2013), and am a Member of the Association of Zoos and Aquariums' (AZA) Biomaterials Banking Advisory Group (since 2016) and the AZA's Molecular Data for Population Management Scientific Advisory Group (since 2018), among other advisory appointments. I also serve as Executive Director of The Orang-utan Conservation Genetics Project, a Wisconsin-registered 501(c)(3) non-profit. From 2009-2015, I taught on the faculty of 'The Cambridge Tradition' summer school with Oxbridge Academic Programs, teaching Zoology every July at Jesus College, Cambridge, and on the subsequent French programme, 'The Paris Connection'. To my knowledge, I am the only Cambridge doctoral graduate with a thesis signed by Dolly Parton ("Well are you going to hold me, or that old book?").


I have long transitioned to addressing questions that are useful, not simply interesting. Thus, all of my current research projects have real-world sustainability outcomes, with opportunities for students to make broader impacts far beyond the confines of our campus. In the face of declining wild populations, I am increasingly interested in what happens when we interfere with natural populations — in particular, through translocations, reintroductions and the creation of ex-situ populations. Much of my ongoing work is conducted in the context of my long-term research programme, The Orang-utan Conservation Genetics Project, and capitalizes on ~3,200 DNA samples from wild, ex-captive and zoo-housed orang-utans that I have collected globally. Together, these comprise the largest biomaterials collection in the United States from a critically endangered mammal.

Genetic viability of orang-utans in zoos: is the 'backup' population sustainable?

Kawan, a zoo orang-utan Concern is mounting that zoo orang-utans are becoming increasingly inbred, having derived from a small number of wild-caught founders in the 1920s. Through whole genome and exome sequencing, ddRAD, and targeted baits assays, we have learned that zoo orang-utans are predominantly outbred, however, and are highly admixed between distinct species and subpopulations that diverged up to ~3.4 million years ago in the wild. Four resulting manuscripts are currently in revision at peer-reviewed journals; these add to an earlier invited review for International Zoo Yearbook (2018). This long-term project has been supported with ~500,000 USD in extramural funds, principally from the National Natural Sciences Foundation of China (2014, 2015) and the Arcus Foundation (2016, 2017-2018), and with a grant from the Association of Zoos and Aquariums (2017-2018). This work is performed in collaboration with Megan Elder, the Orangutan International Studbook Keeper, from Como Park Zoo and Conservatory.

Genetic causes of chronic respiratory and cardiovascular diseases in great apes

An infant female orang-utan Chronic respiratory and cardiovascular diseases are the primary causes of death in zoo-housed orang-utans, accounting for 29% and up to 40% of all mortalities, respectively. The former is also increasingly prevalent in rehabilitation centres in-situ, and is hampering the reintroduction of displaced orang-utans, hindering efforts in range countries to bolster declining wild populations. In collaboration with colleagues at the Wisconsin National Primate Research Center; California State University, Fullerton; and The Great Ape Heart Project (GAHP) at Zoo Atlanta, I am working to determine the etiology and pathogenesis of these conditions through a comparative genomic approach. Preliminary data are now presented in a manuscript currently under review, and will ideally serve as the precursor for a proposed larger and longer-term research programme. Since 2018, I am also working with The Great Ape Heart Project to analyze whole genomes from other great ape species with cardiovascular disease. Preliminary data from a familial group of bonobos suggests a Mendelian inheritance pattern.

Wild orang-utan population and conservation status: how many remain on Borneo and Sumatra?

A Pertamina oil canIn collaboration with a consortium of orang-utan researchers, I performed line-transect surveys across Tanjung Puting National Park as part of a Borneo-wide census of the species. Pooled data from across the island led to publication of the 2016 Population and Habitat Viability Assessment, which guided the new 'Strategy and Orangutan Action Plan' (SRAK) of the government of the Republic of Indonesia. We found that only ~80,000 orang-utans are expected to remain on Borneo; fewer than ~15,000 are thought to exist on Sumatra. We later performed the first integrative trend analysis of a great ape species on Borneo, concluding that orang-utan populations have declined at a rate of 25% over the last 10 years (Scientific Reports, 2017). We have since determined that 100,000 orang-utans were potentially killed in the wild in just 17 years, from 1999-2015 (Current Biology, 2018). These findings led to a change in Bornean orang-utan conservation status, from 'Endangered' to 'Critically Endangered' in 2016. My fieldwork was funded with ~60,000 USD in small grants.

Subspecies and geographic origins of orang-utans reintroduced into wild populations

Percy, an orang-utan in Tanjung Puting National Park Bornean orang-utans comprise three geographically and reproductively isolated subspecies, yet nobody knew this in the 1970s at the dawn of orang-utan reintroduction programmes. In collaboration with Biruté Mary Galdikas of Simon Fraser University and Linda Vigilant of the Max Planck Institute for Evolutionary Anthropology, I investigated subspecies composition at Camp Leakey in Tanjung Puting National Park, Indonesia, finding that multiple subspecies were unwittingly reintroduced and have since had hybrid offspring that have further introgressed. The results were published in 2016 in the journal, Scientific Reports, in the context that reintroductions of displaced mammals may serve as de facto genetic translocations. Given the magnitude of potential ill effects, our findings were featured in the print edition of Science. I am staunchly opposed to such inter-breeding within wild populations, as the effects on health and reproductive success are not yet remotely understood.

Why do male orang-utans have cheek pads? Male bimaturism and reproductive success

Tom, an orang-utan in Tanjung Puting National Park Despite being one of the most prominent ornaments in the animal kingdom, the purpose of male orang-utan cheek pads is poorly understood. The theory that cheek pads make dominant males more attractive to females, and thus result in them fathering more offspring, is complicated in that subordinate, non-cheek-padded males are also able to achieve paternities. In collaboration with Biruté Mary Galdikas of Simon Fraser University and Linda Vigilant of the Max Planck Institute for Evolutionary Anthropology, I investigated male reproductive success in orang-utans at Camp Leakey in Tanjung Puting National Park, Indonesia, in the largest longitudinal orang-utan paternity study conducted to date, in-situ, in a single population. We concluded that male bimaturism is likely an evolutionarily stable reproductive strategy, in which unflanged males simply bide their time until periods of rank instability. Our findings were published in September 2015 in the journal, Behavioral Ecology and Sociobiology.

A search for chemical signals in orang-utans: how and why do cheek pads (not) develop?

Collecting samples at a zoo Having better elucidated the role of male orang-utan cheek pads (left), I am now keen to understand how and why they develop in the first place. In particular, I hope to determine the underlying psychoendoneurocrinological mechanism that might facilitate developmental suppression of subordinate males: it has long been theorised that dominant, cheek-padded male orang-utans are somehow able to inhibit cheek-pad development in younger rivals. In collaboration with Tom Goodwin of Hendrix College and Melanie Bond, formerly of Smithsonian's National Zoo, we are analysing cheek pad, sweat and urine samples using GC-MS to determine potential compounds that might be useful in chemical signalling and olfactory communication. This research was profiled in 2014 on PBS (US) and Channel 4 (UK) in the documentary series, Sex in the Wild: Orang-utans. Sample collection and analysis is ongoing in 2020, to expand on promising pilot data.

Reproductive parameters of ex-captive and translocated female orang-utans

A female orang-utan carrying her infant on her head The success of orang-utan reintroduction programmes can be assessed on the basis of reproductive success — that is, if individuals survive and go on to have their own healthy and viable offspring. As part of a larger collaboration headed by Dr Noko Kuze, I contributed data from Camp Leakey in Tanjung Puting National Park, Republic of Indonesia, which I collected through fieldwork and via a meta-analysis that spanned multiple decades of reintroductions. The resulting paper, in the journal, Primates, was published in November 2012.

...and now for something completely different

Two-toed sloth Under growing pressure from students in my research group, I have recently reluctantly acquiesced to study other (less orange) taxa. Over the last year, with support from the Linnean Society and Systematics Association, post-doc Emily Fountain has been studying taxonomic diversity in two- and three-toed sloths. Meanwhile, DVM student Jenny Mathe has developed a new molecular method to test for sex and species in two-toed sloths, and is now testing the entire US captive population at the request of the Species Survival Plan (SSP). Other students are currently working on gorillas, bonobos, chimpanzees and South China tigers.


Selected publications and conference presentations are detailed below, including those by mentored students. As of 2020, six of my students have co-authored manuscripts in preparation or submission; one has a first-authored paper in review; two have presented at national conferences; two have presented posters (one winning 'best poster'); and one has won an extramural award for outstanding research with a faculty mentor. My detailed curriculum vitae is available upon request.
Selected peer-reviewed publications:
Selected conference presentations:
** indicates sole keynote speaker; * indicates invited speaker:
  • * Banes, G. L. (2019). Outbreeding depression in orang-utans: is it real, and does it kill? June 10. Association of Zoos and Aquariums Molecular Data for Population Management Workshop, San Diego, CA, USA: San Diego Zoo.
  • * Banes, G. L. (2018). Genomic analysis of orang-utans in zoos reveals unexpected barriers to species conservation. September 26. Association of Zoos and Aquariums 2018 Annual Conference, Seattle, WA, USA: Washington State Convention Center.
  • Banes, G. L. (2017). Towards a sustainable future for great apes in zoos. October 3. Eleventh Annual Orangutan Species Survival Plan Husbandry Workshop, New Orleans, LA, USA: Audubon Nature Institute.
  • * Banes, G. L. (2016). A national strategy for the population management of critically endangered orang-utans in zoos. August 24. Chinese Association of Zoological Gardens Annual Meeting, Shijiazhuang, P. R. China.
  • ** Banes, G. L. (2016). Ten years with 'tans: celebrating a decade of care and conservation. August 17. Tenth Annual Orangutan Species Survival Plan Husbandry Workshop, Madison, WI, USA: Henry Vilas Zoo.
  • * Banes, G. L. (2015). Orang-utans in South-East Asian zoos: where they’re from, and where we’re at. November 27. Regional Species Management Program of Primates Conference. Taipei, Taiwan: Taipei Zoo.
  • Banes, G. L. (2015). “We don’t EAT the orange chimpanzees; we ENJOY them”: in search of ‘lost’ orang-utans in Chinese zoos. October 12. Ninth Annual Orangutan Species Survival Plan Husbandry Workshop, Wichita, KS, USA: Sedgwick County Zoo.
  • Banes, G. L. (2014). The Orang-utan Conservation Genetics Project: a leap from Mean Kinship to DNA for managing orang-utan populations in zoos. October 26. Twenty-second Annual Conference of the South-East Asian Zoological Association, Taipei, Taiwan: Taipei Zoo.
  • Jang-Liaw, N. H., Tsao, E. H. S., Huang, H. M. and Banes, G. L. (2014). Where are they from? The next step for ex-situ orang-utan conservation: genetic assignment of captive orang-utans in Taiwan. October 26. Twenty-second Annual Conference of the South-East Asian Zoological Association, Taipei, Taiwan: Taipei Zoo.
  • * Banes, G. L. (2013). The Orang-utan International Studbook Genetics Project: ex-situ conservation of Asia's great ape. November 2. Regional Species Management Programs of Primates Conference. Taipei, Taiwan: Taipei Zoo.
  • Banes, G. L. (2013). Does my face look big in these? The role of cheek pads in male orang-utan reproductive success -- and other fun findings from faeces. October 15. Seventh Annual Orangutan Species Survival Plan Husbandry Workshop. Los Angeles, CA, USA: Los Angeles Zoo.
  • Banes, G. L. (2011). You want WHAT from our orang-utans? October 19. Fifth Annual Orangutan Species Survival Plan Husbandry Workshop. Waco, TX, USA: Cameron Park Zoo.
  • Banes, G. L., Piertney, S. B., Galdikas, B. M. F. and Knapp, L. A. (2010). Orang-utan reintroduction: are we getting it right? September 30. 4th Annual Orangutan Species Survival Plan Husbandry Workshop. Denver, CO, USA: Denver Zoo.
  • Banes, G. L., Piertney, S. B., Galdikas, B. M. F. and Knapp, L. A. (2010). Mitochondrial DNA diversity and reproductive success in the orang-utans at Camp Leakey, Tanjung Puting National Park. September 16. International Primatological Society 23rd Congress. Kyoto, Japan: Kyoto University.
  • Banes, G. L. (2009). The effects of forest disturbance on the Bornean orang-utan in Tanjung Puting National Park, Central Kalimantan, Republic of Indonesia. August 15. European Federation of Primatology Third Congress. Zürich, Switzerland: University of Zürich.
Conference presentations by mentored students:
  • Karklus, A., Rayment, K., Dedolph, T., Fountain, E. and Banes, G. L. (2018). Assessing admixture between historically isolated orang-utan lineages in North American zoos. August 20. 12th Annual Orangutan Species Survival Plan Husbandry Workshop, Kansas City, MO, USA: Kansas City Zoo.
  • Prince, L. and Banes, G. L. (2018). Towards a best-practice framework for orang-utan facility design. August 20. 12th Annual Orangutan Species Survival Plan Husbandry Workshop, Kansas City, MO, USA: Kansas City Zoo.
Conference posters by mentored students:
  • Mathe, J., Fountain, E. and Banes, G. L. (2018). Determining cryptic speciation in two- and three-toed sloths. August 2-4. National Veterinary Scholars Symposium. College Station, TX, USA: Texas A&M University.
  • Karklus, A. and Banes, G. L. (2017). Admixture between historically isolated lineages of orang-utans in North American zoos. August 3-6. National Veterinary Scholars Symposium. Bethesda, MD, USA: National Institutes of Health.
  • Ravikumar, P., Umuhire-Juru, A., Bradley, S. B., Banes, G. L. and Goodwin, T. E. (2017). Searching for chemical signals in orang-utans (Pongo spp.). April 2-6. 253rd American Chemical Society National Meeting and Exposition. San Francisco, CA, USA: Moscone Convention Center.
  • Umuhire-Juru, A., Bond, M., Jacobs, L., Banes, G. L. and Goodwin, T. E. (2014). Searching for Chemical Signals in Orang-utans (Pongo spp.). March 16-20. 247th American Chemical Society National Meeting and Exposition. Dallas, TX, USA: Dallas Convention Center.


Outreach is a defining pillar of my service work to date, particularly in underserved communities. I have now visited more than 200 schools, community centres and zoos to give presentations, and have repeatedly opened my lab to hold science outreach days and inquiry-based activities. I am especially active in working with students from backgrounds traditionally underrepresented in science, particularly at 'last chance' high schools, with a view to cultivating their interest and pursuit of STEM careers.

Further afield, I have trained more than 40 students and local people in Indonesia and Malaysia since 2008, building capacity for scientific research in orang-utan range countries. Since 2013, I have worked with 187 zoos across mainland China (where I am known as 林伊: Lín Yī) to assist in welfare, husbandry and management of all great apes. Most recently, I proposed and am Editor-in-Chief (with co-editors Carol Sodaro, Megan Fox and Bai Yali) of the Chinese-language Orang-utan Husbandry Manual (left), to which chapters were contributed by zoo professionals across the US and abroad. The Manual was published by the governmental Chinese Association of Zoological Gardens in October 2018, and is the first ever official Chinese-language guide for the care of a non-endemic species. In the same month, I co-organized and hosted the China National Orang-utan Workshop at Nanjing Hongshan Forest Zoo; the first ever national workshop for a non-endemic species. The Workshop was attended by 70+ staff from 37 Chinese zoos, plus a delegation of seven experts from zoos across the US.

I am also passionate about popular science, and making otherwise 'boring' subjects more relatable to the public. My popular-science writing has been published by the World Association of Zoos and Aquariums, the Orangutan Foundation UK, the Orang Utan Republik Foundation, Serious Science, and Orangutan Outreach. Meanwhile, my video — 'Waiting for the Centrifuge' (right) — offers a light-hearted introduction to The Orang-utan Conservation Genetics Project, and won 'Most Comical' in DNA Genotek's 'Express Your Groove Gene' contest.


The Kingfisher Encyclopedia of Life (2012, 2014 paperback; co-editions through 2019)

My first book, 'The Kingfisher Encyclopedia of Life', was first published in the United Kingdom on 5 July 2012, by Macmillan. It was released in North America on 31 October 2012, and has since been translated and published in 13 languages in a further 16 territories: Russia, Slovakia, Colombia, Germany, Spain, Brazil, the Czech Republic, Poland, Portugal, Georgia, Turkey, Lebanon, Israel, Romania, Taiwan, and mainland China. The German-language edition, 'Leben!' ('Life!') was one of nine national finalists in the 'Perspectives' category for the 2013 Knowledge Book of the Year (Wissensbuch des Jahres) award in Germany, given by the magazine, Bild der Wissenschaft. The book was also nationally shortlisted in Austria in the junior-knowledge category for the 2013 Science Book of the Year award, given by the country's Federal Ministry for Science and Research. Booklist described the book as a "powerful interdisciplinary resource, encompassing lessons from biology, geography, natural history, and the social sciences." School Library Journal starred the book and concluded, "This work is a spectacular success." I have ghost-written other children's non-fiction, and most recently, was a contributing author to Kingfisher's My First Encyclopedia (2019). After six years of research, my second book under my own name — a popular-science guide to animal sex — is now in preparation for publication.

"This book presents a jaw-dropping look at 30,000 years of biological evolution. Packed with the latest science, it is designed to engage readers emotionally - from a sense of astonishment that the mayfly can enjoy a full life in less time than it takes to read the book, to the awe that is inspired by trees over a thousand years old, to a sense of humble self-reflection when encountering colonies of simple life forms which have existed since before recorded time. At the heart of the book is the human story, examining the many and varied factors which influence our longevity. At the end, readers of all ages will emerge with a new sense of wonder at the natural world's amazing diversity, and a new appreciation for their own place among it."


I am best reached by e-mail (banes [at] wisc [dot] edu) or through the form below.

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