David Wake, Expert on Salamanders and Evolution, Dies at 84

David Wake, who passed away on April 29, 2021, was largely responsible for making modern evolutionary biology what it is today. It was estimated that he was 84 years old at the time.

Who is David?

David, a leading biologist of his time, conducted research that tried to explain the origins of species diversity by combining disparate lines of inquiry into a single, coherent whole.

He kept his attention squarely on processes like organismal growth that the contemporary synthesis accomplished in the middle of the 20th century had downplayed, if not ignored totally.

David spent the last half of his career raising awareness about the worldwide collapse of amphibian populations, particularly those of the salamander family Plethodontidae, the members of which he held dear (and it was love).

 Early Life and Education of David

David’s birthdate is June 8th, 1936 and he was born in Webster, South Dakota. He grew up in the nearby rural village of Pierpont, which in the 1940 census counted only 326 residents, the vast majority of whom were Norwegian immigrants .

The value placed on education was evident in the fact that three of the eleven classmates in his high school class went on to get their Ph.D.s and take faculty positions at universities.

David and his two siblings moved to Tacoma, Washington, before his senior year so that all three of them could attend Pacific Lutheran College (now Pacific Lutheran University) on full scholarships while still living at home.

David found plethodontid salamanders, but especially Ensatina, a rare species confined to the Pacific Coast ranges and the Sierra Nevada, and best known to evolutionary biologists as a typical example of Rassenkreis, or ring species, while collecting insects for an entomology class.

David quickly lost interest in insects and plants in favour of salamanders, which he studied for the following 60 years and wrote over a dozen publications on, exploring the significance of spatial variation in Ensatina for our knowledge of speciation.

David continued his education with Jay Savage to the master’s (1960) and doctoral (1964) levels after his graduation from high school in 1958.

David’s interest in plethodontids was stoked by Savage, a tropical ecologist and biogeographer who was just getting started with a large-scale study of the amphibians and reptiles of Central and South America.

David’s early education in taxonomy and systematics benefited from Savage’s proximity to the Los Angeles County Museum of Natural History, where he had access to study collections in herpetology.

And it was at USC that David would meet Marvalee Hendricks, a premed student who opted to undertake her doctoral work with Savage on caecilians rather than the more popular frogs and toads. As of 2020, David and Marvalee have been married for 58 years.

Career of David

In academia, David had a very comfortable life. When he was nearly done with graduate school, he sent letters to a wide variety of universities asking whether they had a job for which he would qualify. This was done at a period when most faculty searches weren’t posted and on-site job interviews were uncommon.

Seven offers came his way rapidly, and he ultimately settled on one from the University of Chicago. Five years later, he got a call asking him to give a seminar in Berkeley, and that was the beginning of his relocation there.

His host mentioned the opening at the Museum of Vertebrate Zoology (MVZ) in passing the morning he came, and by the time his visit was over, the Zoology Department had voted to grant him a position as assistant professor of zoology and assistant curator at MVZ.

Some days later, back in Chicago, David politely declined the offer, citing his four years of service toward tenure at Chicago and Marvalee’s academic appointment at the University of Illinois at Chicago.

After two weeks, his seminar host called back to tell him that David had been offered a tenured position at Berkeley.

In less than a year, David and Marvalee moved from the South Side to the Berkeley Hills, and shortly after, David was named director of MVZ, a post he would occupy for the next 27 years. In time, Marvalee would become a full-fledged member of the zoology faculty and even head that division.


David Wake’s lengthy study of plethodontid salamanders has given us a model-taxon approach and a deeper knowledge of the mechanisms that drive diversification of lineages. Now that he is gone, the rest of us have to do what we can to protect Earth’s precious biological diversity.