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David Burton Wake
In Memoriam

David Burton Wake

Professor of Integrative Biology, Emeritus, and Professor in the Graduate School
Director of the Museum of Vertebrate Zoology, Emeritus

UC Berkeley

David Burton Wake died of renocardiac syndrome on 29 April 2021 at his Oakland home, with his immediate family by his side. He was less than two months shy of his 85th birthday. One of the foremost evolutionary biologists of recent generations, he pursued his science with vigor until the time of his death. Personal testimonials from around the world, and in publications that range from the New York Times to prestigious academic journals have poured in, all emphasizing how deep his loss has been, especially within the communities of evolutionary biology and herpetology, the two cornerstones of his academic life.

Born on 8 June 1936 in Webster, South Dakota, David was raised in nearby Pierpont, a small farming community (1940 census population of 326 people) of mostly Norwegian immigrants, and into a culture that valued a set of principles that became the ethos upon which David lived his life: a strong work ethic, high moral and ethical standards, and compassion for those whom he encountered. At an early age, his maternal grandfather, a Lutheran pastor, immersed David in local natural history, especially of plants, introduced him to the concept of evolution, and encouraged him to read the Washington Post that was mailed to his grandfather’s home every week, a habit David continued throughout his life. Education in David’s world was highly prized. His mother, herself a college graduate, taught high school biology; three of his class of 11 earned a PhD and became college professors. Before his final year in high school, his family relocated to Tacoma, Washington, where David completed high school and then attended Pacific Lutheran College. There, while collecting insects for an entomology course, he encountered salamanders for the first time, the group of amphibians that would occupy his scientific world for the remainder of his life.

After graduating summa cum laude in 1958, David entered graduate school at the University of Southern California, where he obtained both masters (1960) and doctoral (1964) degrees under Jay Savage, a tropical ecologist and biogeographer, who encouraged David’s interest in salamanders, especially the lungless Plethodontidae that had their centers of diversity in Central America and the US east and west coasts. There he met his future wife, Marvalee Hendricks, who also pursued graduate work under Savage; they would have celebrated their 59th anniversary in June 2021. He left USC to assume a faculty position at the University of Chicago in anatomy and biology in 1964 before being recruited to UC Berkeley in 1968, arriving on campus in the summer of 1969 as a tenured associate professor in the Department of Zoology and associate curator of herpetology in the Museum of Vertebrate Zoology (MVZ). He was appointed MVZ Director in 1971 and promoted to professor and curator two years later. He held the directorship for 27 years, a span second only to that of founding director Joseph Grinnell (1908-1939). David held the John L. and Margaret B. Gompertz Chair from 1991 through 1997.

Two recent retrospectives by former PhD and postdoctoral students, published in the Proceedings of the National Academy of Sciences (J. Hanken, in press) and Science (K. Zamudio, 372:1399) emphasized David’s legacy as one of the foremost organismal biologists of his generation, one who pursued integrative research that had substantive impact on multiple disciplines within the broad sweep of biology. He sought to explain the evolution of diversity by understanding underlying processes, such as development, that had been largely neglected in the modern synthesis of evolutionary biology. His 1979 collaborative paper in Paleobiology (5:296-317) built a conceptual framework for the evolution of size and shape that became part of the foundation upon which a new field, evolutionary development (or “evo-devo”) was built. David argued that achieving a satisfactory understanding of the evolution of any group required a holistic view, one that must incorporate all relevant information. Plethodontid salamanders became one of the most comprehensively investigated and fully documented cases of an adaptive radiation in biology. Today, we have a richer understanding of the many mechanisms that may mediate lineage diversification. Yes, he was a synthetic biologist, but he was also a systematist who immersed himself in the description of new species and genera, most of which he had first encountered during his many field expeditions. He named over 140 salamander species and nine new genera, nearly all plethodontids. He authored 471 papers, reviews, and opinion pieces in scholarly journals and books. More remained in the pipeline at the time of his death.

His focus on plethodontids took him to the areas where they lived, particularly the Neotropical countries of Mexico, Guatemala, and Costa Rica. One unanticipated result of decades in the field was the realization that salamanders were becoming increasingly difficult to find, and not just because of habitat loss. In 1989, David was among the few who called the world’s attention to a precipitous decline in many amphibian populations (D. B. Wake, 1991, Science 253:860). At his initiation, the National Research Council convened a 1990 meeting to highlight the problem and recommend mitigation steps. This meeting rightfully garnered global interest through public media exposure. He then became Founding Chair (from 1991 to 1994) of the Task Force on Declining Amphibian Populations for the World Conservation Union (IUCN). Amphibian declines remained a central focus of David’s activities for the remainder of his life. As an outgrowth, in 2000 he created AmphibiaWeb (, the highly successful on-line repository of “everything amphibian,” a compendium of information on all known species, their natural history, conservation, distribution, and taxonomy, that has allowed both scientists and the lay public to learn where species occur, and the dangers they face. As of 1 July 2021, the website contained accounts for 8,350 species.

It comes as no surprise that someone of David’s stature garnered significant honors. Three of the most prestigious are his 1996 election to the American Philosophical Society (founded by Benjamin Franklin in 1743), 1997 election to the American Academy of Arts and Sciences, and 1998 election to the National Academy of Sciences, where he worked diligently to expand representation in underrepresented fields, such as ecology and evolution. Honors that gave him the greatest personal pleasure were receipt of the Fellows Medal of the California Academy of Sciences (2012), the Joseph Leidy Medal of the Academy of Natural Sciences, Philadelphia (2006), the Joseph Grinnell Medal in Scientific Natural History (1998), awarded at the time of his retirement as MVZ director, and two from his peers in the herpetological community: the Henry S. Fitch Award from the American Society of Ichthyology and Herpetology (1999) and Honorary Membership from the Herpetologists’ League (2012). The campus fittingly recognized his numerous contributions by presenting him with the Berkeley Citation in 2005, following his formal retirement from teaching in 2003.

The legacy David left to the campus is profound, both materially and conceptually. He played a primary role in the reorganization of the biological sciences in the 1980s, including service as chair of the Program Planning Committee overseeing the reconstruction of the venerable Valley Life Science Building. Through his 27-year directorship of the Museum of Vertebrate Zoology, his conceptual leadership led to the development of facilities that brought state-of-the-art laboratory technological opportunities to what, before then, had been largely field-based programs. Faculty, staff, and students from multiple campus units now use this shared facility. He oversaw the development of the first archived collection of tissues for use in biomolecular studies for any museum in the country, expanded the physical specimens to nearly three quarters of a million, and led the effort to make the MVZ collections one of the first to be digitized in the world. He viewed the collections as the intellectual heart of the museum; believed that the MVZ and other biological collection units were central to the teaching and research mission of the campus; and positioned the MVZ as the institution of choice for generations of graduate and post-doctoral students from around the world. His vision centered on providing both physical and intellectual space to optimize collaboration between individuals with different research questions or agendas. He made MVZ’s Wednesday noon colloquium series, “Museum Lunch,” a campus focal point in evolution and ecology. He took great interest in the work of his colleagues within and well outside zoology and was supportive of their efforts, including scientific and educational initiatives in the other Berkeley natural history museums.

David’s service to his profession was equally impactful. He was president of the Society for the Study of Evolution (1983), the American Society of Naturalists (1989), and the American Society of Zoologists (1992) that, under his guidance, expanded its mission conceptually and became renamed the Society for Integrative and Comparative Biology. He served as a member of the National Board of the National Museum of Natural History (1990-1997), the Board of Biology, National Academy of Sciences/National Research Council (1986-1992), and the National Research Council Committee on the Value of Biodiversity (1995-1998), and as editor or member of the editorial board of several national and international journals. Outside of academia, he was elected to the Council of the Save the Redwoods League, the century-old organization responsible for protecting the remaining old-growth redwoods and giant sequoias.

Two things dominate any description of David’s teaching paradigm: the first was to build a questioning mind, not just an encyclopedia of facts. He taught students to focus on ideas, and to understand both the history of those ideas and how evidence was marshaled in support. The second was to care for every student, to recognize each as the individuals they were. If he identified any lacunae in their knowledge , he would work to help fill those holes. As one colleague commented, David’s hallmark combined soft-spoken intellectualism with kindness. His primary course, Evolution (IB 160), was taught every fall semester to an overfilled auditorium. It was legendary for pushing students well beyond what each considered their own limits when the class began, and it changed the lives of many by the time the final exam was over. He always told this writer that the legacy he most valued would be the students he was fortunate enough to know, and mentor. He trained 42 PhD and 30 postdoctoral students during his long career.

A decent, caring man, David valued integrity above all else and gave himself to all whom he encountered. He loved to be “out in nature”, and would be unhappy if a week passed when he (most often with Marvalee) was unable to visit even well-known local places to see the change of seasons. Being in a new place, anywhere in the world, was always a stimulating adventure, both for him and anyone who was fortunate enough to accompany him. His breadth of interests was huge, supported by voracious reading habits, not just of science but history, politico-economics, and beyond – he was a true Renaissance scholar.

David Wake’s loss has been impactful to the campus, his department, and the MVZ, but more so to those of us who were fortunate enough to call him a friend and to have had him as a close colleague or teacher. He is survived by his wife, Marvalee H. Wake (Professor Emerita of Integrative Biology and former Chair of both the Departments of Zoology and Integrative Biology), his son Thomas A. Wake (Director of the Zooarchaeology Lab, UCLA), granddaughter Summer Wake, daughter-in-law Chrissy Campbell (Professor of Anthropology, CSU-Northridge), brother Thomas (of Bow, Washington), and sister Marcia (of Tacoma, Washington).

James L. Patton