Dr Graham L Banes

About Me

Since Fall 2017, I am an Assistant Professor in the School of Veterinary Medicine at the University of Wisconsin - Madison. In collaboration with the nearby Henry Vilas Zoo, my Molecular Ecology and Evolution Laboratory is based in the Zoo's Animal Health Center, integrating research, teaching and outreach with the University's wider community. Our research group includes post-doctoral scientists, graduate students and numerous undergraduates, principally focused on studying orang-utans and other endangered wildlife. I concurrently maintain an honorary appointment at the Chinese Academy of Sciences and Max Planck Society Partner Institute for Computational Biology in Shanghai, People's Republic of China. I return to Borneo annually, where my studies of wild and ex-captive orang-utans are now in their twelfth year.

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 CAS-MPG Partner Institute for Computational Biology in Shanghai, where I served three-and-a-half years as a postdoctoral fellow of the Chinese Academy of Sciences.

My work to date has been generously funded with more than 1,000,000 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, Henry Vilas Zoo, 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, The Orang-utan Conservation Genetics Trust, Orangutan Outreach, Rock of Apes II and III, the Morris Animal Foundation, and by the generosity of private donors. I am delighted to extend my research into 2018 with new grants from the Association of Zoos and Aquariums, with the renewed support of the Ronna Noel Charitable Trust, and with my third grant from the Arcus Foundation.

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 serve as Director and Trustee of The Orang-utan Conservation Genetics Trust, UK-registered charity number 1155284, and as Chair of the Scientific Advisory Committee of the Orang Utan Republik Foundation, a California-registered non-profit co-founded by Dr Gary and Inggriani Shapiro. In 2016, I joined the Steering Committee of the Biomaterials Banking Advisory Group of the Association of Zoos and Aquariums. 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 served 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?"), and have strict rules on kiss-squeaking and nest-building in the laboratory.


My studies of wild and ex-captive orang-utans on Borneo are now in their twelfth year, and are conducted in grateful ongoing collaboration with the government of the Republic of Indonesia. Since 2010, I have concurrently focused on ex-situ conservation of orang-utans, having collected more than 3,000 DNA samples from orang-utans in 10+ countries and across four continents: comprising among the largest biomaterials collections in the United States derived from any critically endangered mammal. My ongoing analysis of these samples falls under the broad umbrella of The Orang-utan Conservation Genetics Project, a programme approved and branded by the World Association of Zoos and Aquariums, which I founded with support from the Max Planck Society and the Chinese Academy of Sciences. I am now involved in a wide variety of research projects, and frequently 'branch out' to smaller studies of other taxa.

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 1960s and 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 February 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.

Effects of inter-breeding orang-utans of genetically distant origins on health and reproductive success

Kawan, an orang-utan at Henry Vilas Zoo Having discovered that orang-utans of multiple subspecies were unwittingly reintroduced in the wild and have inter-bred (left), I am now working to determine if outbreeding depression occurs — in short, if such hybridizations adversely affect viability. If that is the case, future reintroductions — however well-intentioned — could harm the long-term fitness and survival of wild populations. Zoo orang-utans have been indiscrimnately inter-bred for decades, however, and so present an ideal model population for studying potential outbreeding depression. With funding from the Association of Zoos and Aquariums (2017), therefore, I am combining necropsy, health and reproductive histories with full-genome genetic data and pedigree records dating back to 1926, in collaboration with Megan Elder, the Orangutan International Studbook Keeper, from Como Park Zoo and Conservatory. We are concurrently developing new breeding recommendations that pair the most genetically appropriate individuals.

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. The 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 2018, in the interests of statistically supporting correlations.

Genetic correlates of Chronic Respiratory Disease (air sacculitis) in great apes

An infant female orang-utan In collaboration with Jennifer Taylor-Cousar of National Jewish Health, and with Garry Cutting and Neeraj Sharma of Johns Hopkins University, I am working to study potential genetic correlates of Chronic Respiratory Disease (CRD, formerly known as air sacculitis) in great apes. Orang-utans have especially high incidence of CRD, which causes chronic respiratory problems that — if not fatal — can severely affect the individual's quality of life. Symptoms are similar to cystic fibrosis in humans, and thus may have a similar underlying genetic cause. Our preliminary data are now in preparation for publication (2018), and will ideally serve as the precursor for a proposed larger and longer-term research programme.

How many wild Bornean orang-utans remain in Tanjung Puting National Park?

A Pertamina oil can I 2008, I led a Park-wide survey of orang-utan population densities, updating the overall population estimate for Tanjung Puting National Park, Indonesia. An initial analysis of our findings was presented at the European Federation of Primatology meeting in Zürich, Switzerland in 2009. The underlying data have since been incorporated into three peer-reviewed manuscripts (two published, one in prep), and were also analyzed as part of the 2016 range-wide orang-utan Population and Habitat Viability Assessment. I have since drafted a fine-scale, longitudinal density re-assessment of Tanjung Puting's orang-utan population, currently in review, which asserts the Park as home to the largest contiguous population of wild orang-utans on Borneo.

A genomics-based regional breeding programme for orang-utans in Asian zoos

An adolescent female orang-utan Informed genetic management of species in zoos is essential to maintaining — or even increasing — their genetic diversity, ensuring viable and healthy populations long term. I am currently collecting data from Asian zoos for inclusion in the International Orang-utan Studbook, and thus work closely with Megan Elder, the International Studbook Keeper from Como Park Zoo and Conservatory. By genetically determining the species of orang-utans of unknown origin — either Bornean or Sumatran — we hope to facilitate breeding recommendations, domestic exchanges between zoos, and potentially international transfers. At present, I am occupied with genetic samples from all orang-utans in the Philippines, Singapore and Taiwan, plus most from mainland China, with multiple in-country collaborators. An invited review of our efforts and findings to date was published in International Zoo Yearbook (2018), written in collaboration with Wendy Chua and Jim Kao of the South East Asian Zoos Association.

Chinese-language Orang-utan Husbandry Manual

Sign from an orang-utan exhibit in China Since 2013, I have worked with more than 150 zoos across mainland China (where I am known as 林伊: Lín Yī) to assist in welfare, husbandry and management of all great apes. I proposed and now serve as Editor in Chief of the forthcoming Chinese-language Orang-utan Husbandry Manual, to which chapters have been contributed by zoo professionals across the United States and abroad. The Manual will shortly be published by the Chinese Academy of Sciences, the Chinese Association of Zoological Gardens and Nanjing Hongshan Forest Zoo. A similar manual for chimpanzees is now also in the early stages of preparation. In January 2017, I led a delegation of keepers from US zoos — including Janine Steele (Sacramento Zoo, CA), Megan Fox (Los Angeles Zoo and Botanical Gardens, CA) and Linda Jacobs (Jungle Island, FL) — to facilitate a week-long workshop in chimpanzee and orang-utan care. Their visit served as a successful precursor to China's first National Orangutan Husbandry Workshop, scheduled for October 2018.

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. Two other manuscripts on female orang-utan reproduction are in preparation or review as of 2018.

Screening symptomatic chimpanzees for chondrodysplasias

An adult chimpanzee I was approached by Melanie Bond, formerly of Smithsonian's National Zoo, to consider the size of several unusually short-statured chimpanzees in North American zoos. We are now investigating whether or not these chimps might suffer from chondrodysplasias (e.g. 'dwarfism'), which occur in humans in approximately 1 in every 25,000 births. Genetic testing is presently underway to determine if these chimpanzees exhibit similar genetic mutations, or if their condition might be linked to other conditions that could affect bone development, such as rickets.

Publications and Talks

Please click on each category to view selected research outputs. Peer-reviewed publications are displayed by default. A detailed curriculum vitae is available upon request.
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The Kingfisher Encyclopedia of Life (2012, 2014 paperback)

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." After five years of research, my second book — a popular-science guide to animal sex — is 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."


My research group, laboratory and primary office are all located at Henry Vilas Zoo, a short walk from the School of Veterinary Medicine at the University of Wisconsin-Madison. As I continue to work frequently in China and Indonesia, I am best reached by e-mail (banes [at] wisc [dot] edu), or through the form below.

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