Jay M. Savage: Tropical Herpetology Hero and Magnificent Mentor

Tropical biology lost one of its champions and strongest advocates when Jay M. Savage passed away on 3 November 2025 in San Diego, California. He is survived by his wife Rebecca Papendick, his son Charles (Joanne), his daughter Nancy, grandchildren Sean, Tori, and Tara, and many of his “academic children.” Jay was an evolutionary biologist with broad interests. He largely focused on questions related to Costa Rican amphibians and reptiles and was one of the founding fathers of the Organization for Tropical Studies (OTS). Jay was born on 26 August 1928 in Santa Monica, California, and passed away at 97 years of age after a sudden downturn in health. His herpetological tendencies were on full display at the age of eight in Zimbabwe when his family visited the gravesite of Cecil Rhodes during their year in southern Africa. Jay was chasing lizards on the memorial and the guards were not happy with the young boy scampering on the grave of their former leader. Although he was initially interested in mammalogy, his herpetological trajectory was fixed during his sophomore year in high school. That year, the Natural History Museum of Los Angeles County inaugurated a program called “Museum Juniors,” and Jay learned the thrill of the herpetological chase with his museum friends. Savage's love affair with museums and collections also bloomed during this transformative period. The authors were born in the Los Angeles area years after Jay and his companions collected across the canyons and creeks of southern California and often wish we could have experienced the Los Angeles of Jay's youth. In addition to collecting animals in the field, Savage started collecting scientific reprints before entering college, and today his massive paper reprint collection is housed at the University of Texas at Arlington in the Amphibian and Reptile Diversity Research Center. Jay was educated at Stanford University (A.B. in 1950, M.Sc. in 1954, Ph.D. in 1955), and studied with George Myers, an ichthyologist with an interest in amphibians and reptiles. Jay became a herpetologist with an interest in fish, and he successfully mentored herpetologists and ichthyologists. One evening during a graduate seminar at USC, Jay traced his and our lineage all the way back to Alexander von Humbolt, and he was very proud of that heritage. We are also proud of our heritage and grateful to Jay for cultivating collegiality among his students and colleagues. He helped us understand the importance of a historical lens in all aspects of our studies. After spending 2 years teaching at Pomona College following graduation from Stanford, Jay had two successful careers at doctoral-granting institutions: he spent 27 years at the University of Southern California (USC) in Los Angeles, California and 17 years at the University of Miami (UM) in Coral Gables, Florida. He often expressed his awe at getting paid for something he loved doing. Savage was also associated with the Museo de Zoología at the Universidad de Costa Rica (UCR) where he interacted with several students who studied with the late Douglas Robinson including his student, the recently retired Federico Bolaños. In retirement after moving from Florida to California, Savage immediately secured an adjunct position at San Diego State University because association with the academy was something he valued. In San Diego, Savage attended seminars and “herp group meetings,” and continued publishing scientific papers and books. His curiosity and desire to make sense of the world kept him active long past his retirement from UM in 1999. Jay emulated his mentor's approach to graduate education over the course of his career. Like George, Jay encouraged experienced students to help novice students adjust to academic life while studying in and outside the laboratory. Jay was a varsity boxer at Stanford, and he continued his love of sports at USC and UM by concluding his work day with an hour of racquetball followed by beer consumption. At USC, the daily happy hour occurred in the graduate student office with a refrigerator (MAD occupied that office at USC). In Miami, the graduate student office with the mini refrigerator was too small to accommodate more than the two resident graduate students (MAD occupied that office at UM), so we sat outside the office with beverages on salvaged chairs and a couch after sporting activities. At one point, we had a small blackboard and chalk attached to the wall next to the men's restroom. In Miami, it seemed fitting to name the venue “The Verandah” and hours were spent there during Jay's tenure in Coral Gables. In addition to his own graduate students, Savage mentored several students in Coral Gables during his decade of service as the Graduate Program Director for Biology at UM. Jay and his wife Rebecca regularly hosted parties open to faculty and graduate students. Cinco de Mayo, the winter holidays, and summertime cookouts were festive occasions with delicious food and ample drink. Taking care of students was a well-developed sense in Jay. He was deeply committed to graduate education and did all that he could to support the academic efforts of students. The family-like atmosphere Jay created fostered several collaborations with his students (his academic children), among his students, and Jay also collaborated with some of his students' students. During an interview for his Historical Perspective, he admitted that his proudest achievements were his students (Donnelly 2013). Many of Savage's students have tried to emulate him as they built their labs because we wanted to try and achieve some fraction of his success. Jay's retired life was full of scientific inquiry and working with colleagues on two books and additional projects focused on anole systematics and taxonomy, snake taxonomy, and zoological nomenclature. His final publication (Savage et al. 2025) was completed a few months before he passed away. Jay's love affair with Costa Rica was initiated when he and two colleagues from USC secured National Science Foundation (NSF) funds to study amphibians and reptiles in Costa Rica in 1959. He loved that his legal name in Costa Rica was Jay Savage Bird (Bird was his mother's maiden name), and his affection for the country is often captured in his writings like this: “The first two are now regarded as members of our fauna.” (Savage 2002, 41). Savage made over 60 visits to Costa Rica from 1960 to 2000 and much of that time in the 1990s was spent with his Costa Rican colleagues in the Museo de Zoología at UCR looking at specimens. Jay has many fond memories of Costa Rican fieldwork including his participation with the sampling of the Zona Protectora transect (1985) and a workshop hosted by Dan Janzen in the Guanacaste Conservation Area (1995). Three of Jay's students (Arnold Kluge, Arden Brame, Jr., and Robert Lavenberg) did the bulk of the 1959 field work associated with Savage's first grant for research in Costa Rica. The trio of Savage students met important, influential biologists at UCR (Rafael Lucas Rodríguez: Ph.D. from University of California Berkeley and John de Abate: Ph.D. from Tulane University) who became key in collaborations that led to the formation of the Organization for Tropical Studies. Savage, another faculty member (Thomas Pray), and John Mohr, Chair of Biological Sciences at USC traveled to Costa Rica in 1960 to discuss education and research with Rafael Lucas Rodriguez (Director of the School of Biology) and John de Abate (Vice-Dean of the College of Science and Letters) at UCR. These men developed a collaborative educational program designed to introduce faculty from the United States to the tropics by bringing them to Costa Rica for a six-week summer course. The UCR-USC team wanted to get faculty excited about tropical biology so that they could excite their graduate students about tropical biology and develop this fascinating area of scientific inquiry. The NSF funded course ran in 1961, 1962, and 1963. The Fundamentals of Tropical Biology course was a combination of classroom lectures and a few overnight field trips using small hotels or hostels because there were no field stations in the early 1960s. The road system of modern Costa Rica did not exist and transit among locations presented complicated logistical challenges. While Jay was developing his connections in Costa Rica at UCR, other tropical biologists in the United States were trying to figure out how and where to conduct research in tropical landscapes. A meeting was called in Coral Gables to discuss the formation of a consortium because these academic leaders understood they would be stronger together than any institution would be on its own (Stone 1988). The wisdom of their decision was recently validated by a publication that shows field stations are important for conservation efforts (Eppley et al. 2024). Jay was one of the founders of OTS representing USC. All of the hard work of the early founders and participants in OTS was rewarded when the organization won the Tyler Prize for Environmental Achievement in 2001 (with Thomas Lovejoy and Jared Diamond). While Savage and others worked to establish OTS, the Association for Tropical Biology (now the Association for Tropical Biology and Conservation ATB/ATBC) formed in 1962. The ATBC and OTS have fostered tropical research, education, and conservation for more than 60 years along with those who manage the Revista de Biología Tropical, a key outlet for research in Costa Rica (established at UCR in 1953). Jay was active in OTS over the course of his career and served as the President during one of the turbulent financial times that have afflicted the Organization during its history. He encouraged all his students to take an OTS course, and many of Savage's students taught the OTS course that followed the design developed by Daniel H. Janzen and Norman J. Scott, Jr. in the mid 1960s. Dan and Norm changed the course from a classroom-centered course with occasional field trips to the immersion course OTS is famous for today. One measure of OTS success is the great number of OTS-like courses being taught globally. Alumni have been successful in sharing the OTS immersion model with new generations of students, and the field of tropical biology has blossomed over the last 60 years. Jay served on the OTS Board of Visitors once he retired from his academic position at UM to help guide fund-raising efforts. Jay did all that he could to promote and facilitate tropical education and research during his tenures at USC and UM. Jay's academic interests in evolution and biogeography were explored in Costa Rica and elsewhere in the Caribbean and his academic achievements built nicely on the work of those who preceded him in Costa Rica (E.D. Cope, E. R. Dunn, and E. H. Taylor). Jay was supported by a Guggenheim Fellowship in 1963 and 1964 to promote his work in Costa Rica and that support helped guide OTS in the early days of the organization. Savage's life work on the herpetofauna of Costa Rica (Savage 2002) has stimulated others to advance our knowledge of the herpetofauna of Costa Rica (Guyer and Donnelly 2005; Kubicki 2007; Leenders 2016, 2019; Sasa et al. 2025; Solórzano 2004), and we know his foundational research will continue to inspire others in the future. In addition to Jay's herpetological work in the Caribbean, he served on eight different working groups related to development in the humid tropics and oversaw publication of some of the findings and reports of those groups (Savage 1982). Jay's friends and colleagues have many “Savage stories,” and we share two of our stories for this obituary. One of us (MAD) spent considerable time in the field in Costa Rica and Mexico with Jay, but my most memorable experience was visiting David Bickford in the highlands of Papua New Guinea in 1998 with Jay and my husband Steven Oberbauer. We relied on helicopters to transport us from Goroka to the Wara Sera field station (Bickford 2005) where we spent two weeks “camping in the house” while learning about the fiercely undulating landscape and the frogs David encountered as he conducted his field studies. Jay enjoyed watching his last doctoral student manage a large project in a remote field location, and several times he expressed how proud he was of David and gave him a compliment for what he was accomplishing in the wilds of Papua New Guinea. Savage motivated all of us with his stories of field adventures, and he was captivated by the frogs David was working with and their amazing parental care behaviors. Brian Crother and Jay Savage shared adventures in the United States and abroad, in the pub and on the playing field, and of course while doing field work. This is Brian's story: One of my favorite stories took place in the states before I joined his lab as a graduate student. In 1982, I was an undergraduate work study student with Jay at USC. The ASIH meetings that year were in DeKalb, IL at Northern Illinois University. Two graduate students wanted to drive up and fly back, so Jay asked me if I wanted to take a van and drive to Illinois to attend the meeting, and then drive the van back to Los Angeles. I agreed and Jay let me bring a buddy along as company for the long drive back. The trip to DeKalb was uneventful; the meetings were unlike anything I had experienced before, but the drive back topped it all. As far as I knew, I had no time limit to return the van, so my buddy and I took about 2–3 weeks to get back because we visited just about every National Park possible. Upon my return, I learned a lot about Jay and why I wanted to work with him in graduate school. Even though I missed the deadline for return by at least 2 weeks, Jay was even keeled and understanding and defended me to a couple of furious faculty members. Wow was I impressed! But the next time I “borrowed” the van that is another story! Several of Jay's students (his academic children) continued along his path by conducting research in Costa Rica, and they had students (his academic grandchildren) who work or worked in Costa Rica, and some of their students (his academic great grandchildren) continue working in his beloved Central American Republic located between two continents and between two seas. Jay infected all of us with his enthusiasm and “seize the day” spirit. We and countless others are eternally grateful to Jay M. Savage for helping us search for our “golden frogs” as we try to follow in his footsteps. We thank E.L. Webb for inviting us to write the obituary and we thank R.E. Papendick for critical comments on the document. The ASIH granted us permission to use Figure 1 that was originally published in Copeia by Donnelly (2013, 764, fig. 7) and used here with permission from the American Society of Ichthyologists and Herpetologists (ASIH). The authors declare no conflicts of interest. Data sharing not applicable to this article as no datasets were generated or analyzed during the current study.