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

David Wake, an evolutionary biologist and a pre-eminent authority on salamanders who raised an alarm in the 1980s about the loss of amphibians to climate change and other causes, died on April 29 at his home in Oakland, Calif. He was 84.

His wife, Marvalee, also an evolutionary biologist, said the cause was organ failure.

A longtime professor of integrative biology at the University of California, Berkeley, Dr. Wake used salamanders as vehicles to study how animals diversify and develop new species over millions of years. The focus of his studies was a species of lungless salamander called plethodontids.

“He was emotionally excited about salamander biology,” said James Hanken, director of the Museum of Comparative Zoology at Harvard and a former graduate student of Dr. Wake’s. “If you sat down to dinner with him, he would talk about and describe salamanders and you’d want to drop everything to go with him.”

Dr. Wake named 144 salamander species, some of which he discovered, on his own or with colleagues. One, Nototriton wakei, was named for him.

Two lizards and a frog also bear his name.

Jonathan Losos, a biology professor at Washington University in St. Louis, said in an email that Dr. Wake “promoted the view that to understand how species evolve, you have to understand how their genes guide the process by which a fertilized egg develops ultimately into an adult organism.” He added, “Evolution, at its heart, works by changing the way organisms develop, and Wake was at the forefront in demonstrating that the developmental process must be understood to figure out how evolution works.”

Dr. Wake’s amphibian expeditions took him to the Appalachians, California, the Pacific Northwest, Costa Rica and Guatemala. But in the 1980s, he realized that a species of salamander that he had found to be plentiful in Mexico a decade earlier had become difficult to find. Other herpetologists had by then had similar experiences of amphibian scarcity.

In response, Dr. Wake organized the first World Herpetology Congress in Canterbury, England, in 1989, where scientists shared information about dying amphibians. A year later, he chaired a meeting in Irvine, Calif., of 20 scientists from the United States and abroad, in which participants assessed the loss of amphibians from ponds, rivers, mountains and rain forests around the world.

“I wish there were a death star to explain it,” Dr. Wake told The New York Times Magazine in 1992. “I don’t see a single toxin, a single virus. My theory is that it’s general environmental degradation. That’s the worst thing.”

He added: “Frogs are telling us about the environment’s overall health. They are the medium and the message.”

In 1998, a chytrid fungus was found to have caused a lot of the deaths, especially of frogs, in the rain forests of Central America and Australia. But Dr. Wake and others pointed to other factors as well, including climate change, pollution and habitat loss.

Dr. Wake’s Berkeley seminar on declining amphibian populations led him in 2000 to help start AmphibiaWeb, an online compendium of information about the conservation status of thousands of species of amphibians as well as their biology, natural history and distribution.

“He considered AmphibiaWeb part of his legacy,” said his wife, who studies the limbless amphibians called caecilians and collaborated on a few papers with him. “He also thought the naming of so many species would be a lasting contribution.”

David Burton Wake was born on June 8, 1936, in Webster, S.D. and grew up in nearby Pierpont. His mother, Ina (Solem) Wake, was a schoolteacher, and his father, Thomas, sold hardware and farm implements. His family moved to Tacoma in 1953.

His maternal grandfather, a self-taught botanist, exerted a strong early influence on him.

“He had ‘Gray’s Manual of Botany,’ and we keyed up plants together,” Dr. Wake said in an interview with the UC Berkeley Emeriti Association in 2019. “So I early on developed an interest in the natural world.”

When he attended what is now Pacific Lutheran University, in Tacoma, his focus shifted from botany to zoology; his path narrowed when he took a course in entomology. While in the field collecting beetles, he recalled, “I would stumble onto salamanders and I was charmed by them.”

After graduating in 1958 with a bachelor’s degree in biology, Dr. Wake earned a master’s and a Ph.D. in biology at the University of Southern California, where he met Marvalee Hendricks. They married in 1962. He taught anatomy and biology at the University of Chicago until he was hired as an associate professor of zoology by the University of California in 1969.

Two years after his arrival, he was named director of the campus’s Museum of Vertebrate Zoology, a facility dedicated to research on the biology of amphibians, reptiles, birds and mammals. Dr. Wake, who remained the director through 1998, started its first collection of frozen tissues to augment its holdings of preserved specimens. He also established its molecular laboratory.

“He never saw the museum as a place where taxonomists figured out what’s species A or species B,” Michael Nachman, the museum’s current director, said by phone. “He used the collection to determine how evolution works and how biodiversity develops over time.”

Dr. Wake also taught a course on evolution that had a major impact on the career choice of Nancy Staub, a professor of biology and an expert on salamanders at Gonzaga University in Spokane, Wash. She was one of many students who followed in his path.

“I started in an ecology lab, but it didn’t capture me,” Dr. Staub said by phone. “I was still floundering and took Dave’s evolution course in my second year and everything clicked. I thought, ‘This is fascinating; this is the field I want to be in.’”

In addition to his wife, Dr. Wake is survived by his son, Thomas; a granddaughter; a sister, Marcia Wake Sherry; and a brother, Thomas.

Dr. Wake’s enthusiasm for salamanders never wavered. In 2003, an American high school teacher named Stephen Karsen who was working in South Korea found a lungless salamander, which was not known to have existed in Asia, in a rocky crevice in a wooded area. Mr. Karsen first showed it to salamander experts; it was subsequently sent to Dr. Wake, who determined that it was a plethodontid and named it Karsenia Koreana.

“For me, this is the most stunning discovery in the field of herpetology in my lifetime,” Dr. Wake was quoted as saying in a 2005 article on the Berkeley website. He added: “People have gone on expeditions looking for terrestrial salamanders in places like Kazakhstan and other Central Asian republics. They didn’t bother with northern China or Korea or Japan because we thought we knew everything that was there.”

The discovery seemed to remind Dr. Wake of how he himself became focused on salamanders: “Some guy who’s a high school teacher from Illinois goes out with his class and says, ‘Let’s look for salamanders, let’s see what we can find when we turn over rocks and logs.’”

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