Vol. 66, No. 2 | Fall 2021

In Memoriam

Barry Sinervo (November 24, 1960 – March 15, 2021)

Written By: Bruce E. Lyon and Alexis S. Chaine

Barry Sinervo with lizard (Photo from Internet – UCSC). Barry Sinervo watches while students count male vocalization rates in western chorus frogs as part the Behavioral Ecology Field Course taught by Sinervo and Lyon for two decades. Natural Bridges State Park in Santa Cruz, California, March 2001. (Photo B. Lyon).

Barry Sinervo, pioneering animal behaviorist and evolutionary biologist, died on March 15, 2021 at the age of 60. Barry had been fighting cancer for six years but remained very active in research and teaching until the very end.

Barry was born in Port Arthur, Ontario, Canada, on November 24, 1960. His father was a lumberjack and put his sons to work as lumberjacks in their early teens. Barry's brother Ken recalls the work as grueling and very dangerous but they made up for it by having a great deal of fun together. Ken suspects the early experience in the forest was "probably a reason that we all excelled at school so that we never had to do that again, and our dad would always joke, there was always one of his skidders waiting for us if we didn’t want to go to university."

Barry received a BSc at Dalhousie University in Nova Scotia, majoring in both Biology and Mathematics. He studied under the influential mathematical biologist E.C. Pielou. He told us often the huge impact she had on his development as a biologist, particularly his realizing the power that modeling brings to questions in the natural sciences.

Barry's PhD, with Ray Huey at the University of Washington in Seattle, focused on life history evolution in fence lizards. Huey recalls the day Barry came into his office with an idea to experimentally manipulate lizard offspring size to investigate optimal offspring size. Barry had successfully manipulated offspring size in urchins (with Larry McEdward) by removing yolk from eggs and wanted to try this with lizards. Huey told Barry that lizard eggs are far too sensitive for this to work and not to waste his time. Weeks later Barry returned to Huey's office with a small box containing several fence lizard eggs. Some of the eggs were tiny but when held up to the light Huey could see that they contained developing embryos. Looking at Barry he "saw one of the biggest and brightest grins" he had ever seen—Barry had ignored his advice and pulled off a remarkable experimental manipulation. Pulling off seemingly impossible studies became a hallmark of Barry's career. The Science paper that resulted from these experiments, Allometric Engineering: An Experimental Test of the Causes of Interpopulational Differences in Performance, brought Barry to the attention of evolutionary biologists around the globe.

Barry continued working on offspring size with side-blotched lizards (Uta stansburiana) as a Miller Postdoc at University of California, Berkeley. He discovered that he could also experimentally increase offspring size by surgically removing yolk completely from some follicles in females. Based on experiments that both increased and decreased lizard offspring size, Barry and colleagues tested Lack and Williams' ideas about optimal offspring size. The resulting Science paper, “Allometric Engineering: A Causal Analysis of Natural Selection on Offspring Size”, is considered to be among the finest experimental life history evolution studies ever done.

Barry was an assistant professor at Indiana University from 1992 to 1997. Work with the Uta lizard system continued, now requiring an annual trek between Bloomington and his study site near Los Banos, California, lovingly dubbed 'Lizardland'. Five years of work on male color morphs yielded a blockbuster study that would go on to define much of Barry's career. With Curt Lively, Barry published the first example of a mating system with rock-paper-scissor dynamics. Lively recalls the epiphany: "We were both assistant professors at Indiana University and we talked science all the time. We were especially interested in the coexistence of discrete types. We had coffee once or twice a week when we focused on our respective research projects. Barry had discovered a male color polymorphism in his lizards (orange, blue, and yellow) and he knew that male color was highly heritable. He also had very convincing data showing a complete cycle in the frequencies of the three morphs over a five-year period. We talked a lot about this intriguing oscillation. One day we were talking about John Maynard Smith's book on evolutionary game theory. Maynard Smith had shown that two morphs would come into a stable equilibrium under negative frequency-dependent selection, but three morphs could oscillate over time, especially if they had a non-transitive relationship, like in the rock-paper-scissors game. As soon as this came up, Barry jumped up and very excitedly exclaimed: ‘Dude, it's rock-paper-scissors!’ Barry invited me to be a coauthor on the paper which was very generous of him. He was a very giving collaborator."

Lively also notes that part of Barry's greatness lay in having interests outside of research. Barry was an avid cyclist (along with his wife Jeanie Vogelzang) and got Lively interested in cycling—a pattern he continued after moving to Santa Cruz and even inspiring one of his graduate students to become a professional cyclist and US cycling champion. One of us (ASC) commuted 45 min by bike to campus for over two years at Santa Cruz with Barry—rain or shine—with excellent scientific discussions along the way that lead to several collaborations. Barry was also an excellent chef and artist. Barry's family was extremely important to him and, when he shifted work to broad geographic patterns, Jeanie and their son Ari often accompanied Barry on his lizard expeditions to exotic sites around the world.

The three male color morphs of side-blotched lizards (Uta stansburiana): orange, blue and yellow. The morphs differ in mating strategies, their fitness is frequency-dependent and non-transitive, and the dynamics fit a rock-paper-scissors mating game as first shown by Sinervo and Lively in a now classic Nature paper in 1996. Photo B. Sinervo.

In 1997 Barry moved to University of California in Santa Cruz, joining what would soon become the Department of Ecology and Evolutionary Biology, and where we first crossed paths with him. At Santa Cruz the work on Uta ramped up considerably and, with the help of many talented graduate students and post docs, the scope of the program expanded dramatically. With the morphs as the underlying conceptual linchpin, the research addressed a broad diversity of important questions and topics in evolutionary biology and animal behavior: the role of adaptive maternal effects to deal with changes in population densities, density-dependent selection and selection on immune function, cryptic and context-dependent female mate choice, correlational selection favoring the evolution of morphs, cooperation within morphs via 'greenbeard' recognition, game theory, the Chitty hypothesis that population cycles can favor the evolution of a mixture of r and K female life history strategies in a population, among others. Barry’s interests and reading were very broad and he had an unusual gift for understanding diverse domains and making creative links across them—a characteristic that made our joint lab meetings exciting and dynamic. Accordingly, his research was unique in the breadth of disciplines he investigated—evolution, ecology, genetics, behavior and physiology—essentially the entire breath of the EEB department, and he made landmark contributions to each of these fields.

During the lizard field season the Sinervo lab was a hub of activity, and a large number of undergraduate students got their first taste of science helping with the research and lizard husbandry. Barry maintained annual surveys of morph frequencies on his core sites through 2020, brought gravid females to the lab to build pedigrees from wild lizards, and simultaneously conducted experiments in the lab or in the field. Some of the experiments were audacious—one study, with his PhD student Ryan Calsbeek, involved moving rocks from territories of some males to those of others (Barry dubbed this a true “Sisyphean effort”) to decouple male quality from habitat quality, whereas another moved lizards around the landscape to create entire experimental populations with altered frequencies of the morphs. One of us (BEL) got to see Barry in action at one of these populations—Yellow World—and was shocked by his extraordinary talent at catching lizards. The lizards are wary and hard to catch without spooking, but with the standard lizard catching kit—a pole with fishing line with a slip noose on the end—Barry effortlessly snatched lizards at double the rate of even the most experienced grad students and postdocs. Barry liked to joke that to study lizards one had think like a lizard—if his lizard catching skills are any indication, Barry was part lizard!

Morphs are common in many lizard groups and Barry expanded his focus to the broader macroevolutionary patterns. With Mexican colleagues Barry began working with the Sceloporus spiny lizard radiation in Mexico and Central and South America, and the common lizard in Europe. These two projects brought the impact of climate change to Barry's attention, specifically the discovery that some of the populations of lizards he was studying were disappearing. Don Miles, one of Barry’s longest-term collaborators, was checking field sites in Mexico with one of Barry's students to sample Sceloporus lizard morphs. They visited promising sites based on previous census data from their Mexican collaborators, but they could not find the lizards. Their absence was so dramatic that Miles initially wondered if they had mixed up locations, but instead the lizards had simply vanished. Similar patterns were emerging with the European study, and prompted a massive analysis showing an alarming global erosion of lizard diversity—evidence suggested that climate change impacts on lizard thermal niches was responsible. The focus on climate change led Barry to help establish in 2014, and then direct, the Institute for the Study of Ecological and Evolutionary Climate Impacts (ISEECI), a University of California wide institute that sought to leverage UC’s fabulous Natural Reserve System to study the effects of climate change on California ecosystems. Through this collaborative work, Barry forged deep friendships with colleagues around the globe—many of whom say that his creativity, enthusiasm, and especially his joie de vivre will be missed most.

Barry loved teaching and his endless enthusiasm was infectious for the students. He taught the material at a high level and the students loved it. Barry was technologically savvy and used these skills to enhance his teaching, including the development of a rock-paper-scissors video game for students to learn about the dynamics of alternative strategies. For years he had been developing his own online textbook for his signature Behavioral Ecology class and the book, Behavioral Genetics to Evolution, was just published by Top Hat Marketplace. He also co-taught a Behavioral Ecology Field Class with one of us (BEL) and hunting for Ensatina salamanders with Barry made everybody feel like a kid on Christmas morning.

Barry made a diverse array of important contributions but his name will forever be associated with rock-paper-scissor (RPS). Barry sometimes got carried away thinking that RPS explained everything in the universe but he was often surprisingly right with his instincts. For example, Barry gave a talk at Simon Frazer University in British Columbia where our mutual friend Dov Lank had a captive breeding colony of ruffs, a lekking shorebird famous for its two color morphs—darker males that defend small territories on the lek and paler satellites. Dov had recently worked out the genetics of the morphs. Sitting in Dov's office discussing the genetics and natural history of ruff matings, Barry suddenly proclaimed that there must be a third morph. Dov expressed his doubts, given that the species had been studied by ethologists and behavioral ecologists (including himself) for 40 years. However, a few years later a Dutch group in fact discovered a third male morph with a large scale banding project—it had been missed by behaviorists because it is rare and is a cryptic female mimic. When Barry and Dov spoke again after the discovery, Barry suggested that the low frequency meant that it was a 'degenerate RPS' ... but Dov remained annoyed —''Sinervo was right!'

Numbers have always been mythically important to people. In the Hitchhiker's Guide to the Galaxy, Douglas Adams playfully suggested that the meaning of life is 42. For Barry Sinervo, however, the meaning of life was always three—rock, paper, scissors.


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