Richard Bambach was a leading figure in the “paleobiology revolution” of the late 1960s and 1970s, keeping the movement grounded with his keen geological and ecological insights. With interests ranging from the functional biology of individual organisms to the largest macroevolutionary trends in the history of life, he was especially adept at linking paleoecological and macroevolutionary patterns across spatiotemporal scales. He authored seminal publications during five different decades and was recognized with both the Moore Medal from the Society for Sedimentary Geology and the Paleontological Society Medal.

Figure 1. Richard Bambach on his first day in his office at the National Museum of Natural History in 2005.

Richard spent his early years in Cincinnati but professed to have no interest as a child in the classic Ordovician fossils of that region. Instead, his childhood interest in geology was kindled in the meteorite exhibit at the Smithsonian Institution after his family moved to the Washington, D.C., area. A staff member was cleaning the meteorites when a small piece fell off one of the larger specimens; the kindly staff member gave it to Richard and ended up taking him on a behind-the-scenes tour. Richard took the fragment home and carefully labeled it “1”—the first specimen in his collection. His first exposure to paleontology came about through his childhood crush on the daughter of Gus Cooper, who happened to be the curator of fossil brachiopods at the Smithsonian.

Richard earned his bachelor’s degree in biological sciences from the Johns Hopkins University in 1957. He was a member of the lacrosse team, and he recalled playing against Jim Brown before Jim became famous in the NFL. Richard was not the most diligent student at Hopkins, and he loved to recount how a frustrated dean warned him that he was in danger of not graduating. Thankfully, he did, although it took two senior years. During his college years, he worked for a couple summers as a fire lookout in Glacier National Park, and he gained valuable experience after college working in the Geology Department at the Smithsonian and serving in the U.S. Navy in signal intelligence. Richard then moved north to Yale University for graduate school, studying Paleozoic bivalves under Lee McAlester. He thrived during his time in New Haven; major influences included geologist (and fellow jazz lover) John Rodgers, ecologist G. Evelyn Hutchinson, and an amazing group of fellow graduate students, many of whom also went on to become influential paleobiologists. He completed his Ph.D. dissertation on the Silurian bivalves of Arisaig, Nova Scotia, in 1969.

Richard briefly taught at Smith College before moving in 1970 to Virginia Polytechnic Institute and State University (Virginia Tech), where he remained on the faculty until he retired in 2000. During this time, he spent a sabbatical at the University of Chicago in 1978–79 and a semester at Harvard in 1995. He spent the first five years of his retirement at Harvard, teaching occasionally, helping to mentor graduate students, and continuing to do research, mostly in collaboration with Andy Knoll. In 2005, he returned to the Smithsonian as a research associate.

Richard was both a talented geologist and an excellent biologist, a combination that proved a major secret to his success. His multidisciplinary perspective is evident in his early work on community paleoecology, as the degree to which ecological information is preserved in a fossil assemblage is largely a sedimentological question. He coauthored an influential paper on Silurian communities with Fred Ziegler and Robin Cocks that appeared on page 1 of the first issue of Lethaia, and another with Jeff Levinton in the first issue of Paleobiology (Ziegler et al. Reference Ziegler, Cocks and Bambach1968; Levinton and Bambach Reference Levinton and Bambach1975). In a 1971 GSA abstract that has been cited hundreds of times (Walker and Bambach Reference Walker and Bambach1971), he and Ken Walker coined the term “time-averaging” to describe the mixing of many generations of organisms into a single stratum—a foundational concept in our understanding of the fossil record that has inspired numerous research programs in the years since. He often emphasized the benefits of a time-averaged record that filtered out annual to decadal fluctuations in species abundances. His interest in distinguishing biological and geological signals is also evident in an early paper on the retrodeformation of bivalve fossils that stemmed from his dissertation research (Bambach Reference Bambach1973).

Even as his interests expanded over the course of his career, Richard continued to think about the preservation of fossil assemblages and the controls on paleocommunity change, particularly working with graduate students, including Ron Kreisa (M.S. 1972, Ph.D. 1980), Dale Springer (Ph.D. 1982), Arnie Miller (M.S. 1981), Bret Bennington (Ph.D. 1995), and Gwen Daley (Ph.D. 1999). He also coauthored an important paper in Reference Sepkoski, Bambach, Droser, Einsele, Ricker and Seilacher1991 with Jack Sepkoski and Mary Droser on secular changes in bedding fabrics related to increasing bioturbation through time (Sepkoski et al. Reference Sepkoski, Bambach, Droser, Einsele, Ricker and Seilacher1991), as well as a review paper in 2003 with Michał Kowalewski on temporal resolution in the fossil record (Kowalewski and Bambach Reference Kowalewski, Bambach and Harries2003).

Richard was also interested in the functional morphology and ecology of individual organisms, as seen in an early publication on the Silurian bivalve Grammysia obliqua (Bambach Reference Bambach1971), as well as in his paleocommunity studies. In a 1974 paper that anticipated much of his later work, he and Ken Walker devised a scheme for classifying benthic invertebrates based on their feeding behavior (Walker and Bambach Reference Walker and Bambach1974). This initial paper focused only on the interpretation of local paleocommunities and ecosystems, but it laid the foundation for later analyses of larger-scale paleobiological questions.

As Richard was exploring an ecological view of the fossil record that was firmly grounded in fieldwork and observations of specimens, other founders of the paleobiology movement were taking a broader-scale view of the fossil record. Jim Valentine proposed that the global species richness of marine animals might have increased 10-fold over the course of the Phanerozoic, driven by continental movements explained by a new theory called plate tectonics. Dave Raup, worried about secular biases in fossil preservation, suggested that diversity might have been similar in the Paleozoic and Cenozoic. Jack Sepkoski began collecting data for his famous compilation of marine animal diversity.

These developments set the stage for one of Richard’s most important and enduring contributions: a Reference Bambach1977 paper in Paleobiology titled “Species Richness in Marine Benthic Habitats through the Phanerozoic” (Bambach Reference Bambach1977). Richard realized that his interest in paleocommunities provided an elegant test of the competing proposals of Valentine and Raup—all else being equal, an increase in species richness at the level of local communities would drive an increase at the global level, and an analysis of local diversity would avoid the biases that most concerned Raup. After compiling data on hundreds of fossil assemblages, he found that species diversity increased about two- to threefold from the Paleozoic to the late Cenozoic, splitting the difference between the models of Valentine and Raup. Because it appeared to transcend the secular-bias conundrum, Richard’s analysis of alpha diversity was a key pillar in 1981’s famous “Consensus Paper,” authored with Sepkoski, Raup, and Valentine (Sepkoski et al. Reference Sepkoski, Bambach, Raup and Valentine1981). Improvements in databases and analytical methods in subsequent decades have continued to support his general conclusions (e.g., Bush and Bambach Reference Bush and Bambach2004, Reference Bush and Bambach2015).

Richard’s 1983 paper “Ecospace Utilization and Guilds in Marine Communities through the Phanerozoic” integrated many of his prior interests and insights (Bambach Reference Bambach, Tevesz and McCall1983a). In fact, the paper could be considered the epitome of Bambachian scholarship: drawing on his deep knowledge of marine ecology and paleontology, he constructed a coherent framework for understanding the history of marine animal life that elegantly spanned a breathtaking range of scales, from individuals to communities to global diversity trends over half a billion years of deep time. Specifically, he argued that the alpha-diversity increase he documented in 1977 was driven by an increase in the number of ecological guilds (or functional groups) within these assemblages, and that, perhaps, the global increase in marine animal diversity over Phanerozoic time could likewise be ascribed to ecological and functional diversification. Richard continued to explore these themes throughout his career, focusing at times on traits like predation, motility, and physiology (e.g., Bambach Reference Bambach and Valentine1985, Reference Bambach, Kowalewski, Kelley and Dodson2002; Bambach et al. Reference Bambach, Knoll and Sepkoski2002, Reference Bambach, Bush and Erwin2007; Bush et al. Reference Bush, Bambach and Daley2007; Bush and Bambach Reference Bush and Bambach2011).

But what drove changes through time in the dominant clades and functional groups, as well as in species richness? In his classic 1993 paper “Seafood through Time: Changes in Biomass, Energetics and Productivity in the Marine Ecosystem” (Bambach Reference Bambach1993), Richard argued that an increase through time in food availability to marine consumers had driven increases in biomass and activity levels, contributing to the diversification of predators in the Mesozoic and Cenozoic that had been documented by Gary Vermeij, Steve Stanley, and others—an argument that seems more prescient with each passing year. Richard continued to explore these themes in several additional papers, and in his last full-length journal article in 2016, he, Andrew Bush, and Gene Hunt expanded on these ideas by tying the diversification of active, motile animals to reproductive biology (Bush et al. Reference Bush, Hunt and Bambach2016). The ideas in “Seafood through Time” also inspired many other paleobiologists; for example, lunchtime conversations with Richard about biomass and energetics helped launch major research projects by Jon Payne and Seth Finnegan.

In 1995, Richard was hired by Harvard as a sabbatical replacement for Steve Gould, and he spent his time in Cambridge working with Andy Knoll and John Grotzinger to develop a novel hypothesis for the end-Permian mass extinction (Knoll et al. Reference Knoll, Bambach, Canfield and Grotzinger1996, Reference Knoll, Bambach, Payne, Pruss and Fischer2007). Geological and geochemical records at the Permian/Triassic boundary suggested elevated CO₂ levels in the shallow oceans, so Richard and Andy dove into the literature to evaluate the effects of this stressor on the marine fauna. In their 1996 publication, they explained the selective nature of the extinction in terms of differential physiological sensitivity to high CO₂, which eventually led to the recognition of ocean acidification as an extinction kill mechanism, a timely finding that dovetailed with growing concerns about its effects on modern marine organisms. Richard followed up with several additional papers on mass extinctions (e.g., Bambach et al. Reference Bambach, Knoll and Wang2004), including a well-cited review paper (Bambach Reference Bambach2006).

In several other projects during his career, he teamed up with colleagues for large-scale studies of diversity history, including several papers in the 1980s with his Virginia Tech colleague Norm Gilinsky on the interplay of origination, extinction, and diversification dynamics during the Phanerozoic (Gilinsky and Bambach Reference Gilinsky and Bambach1986, Reference Gilinsky and Bambach1987), as well as a series in the 2010s with Adrian Melott on periodicity in marine diversity (Melott and Bambach Reference Melott and Bambach2010, Reference Melott and Bambach2011a,Reference Melott and Bambachb, Reference Melott and Bambach2013, Reference Melott and Bambach2014; Melott et al. Reference Melott, Bambach, Petersen and McArthur2012). He was also intimately involved in the establishment of the Paleobiology Database and was an important contributor to discussions in August 1998 at the initial meeting of the group that went on to develop the project. As always, Richard helped keep these efforts grounded with his detailed knowledge of the fossil record, the paleontological literature, and the geological timescale.

Paleoecology and diversity history were the main themes of Richard’s career, but he made important contributions to other topics as well. Most notably, in the late 1970s and early 1980s, he collaborated extensively with Fred Ziegler, Chris Scotese, and others on influential reconstructions of Paleozoic paleogeography (Ziegler et al. Reference Ziegler, Scotese, McKerrow, Johnson, Bambach and West1977, Reference Ziegler, Scotese, McKerrow, Johnson and Bambach1979, Reference Ziegler, Bambach, Parrish, Barrett, Gierlowski, Parker, Raymond, Sepkoski and Niklas1981; Bambach and Scotese Reference Bambach and Scotese1979; Scotese et al. Reference Scotese, Bambach, Barton, vanDerVoo and Ziegler1979; Bambach et al. Reference Bambach, Scotese and Ziegler1980). In 1990, he connected his interest in paleogeography back to Phanerozoic diversity change, testing Valentine’s hypothesis that increased provinciality was a major driver of diversification in marine animals (Bambach Reference Bambach, McKerrow and Scotese1990). Over the years, he also participated in a number of other geological studies, many focused on southwest Virginia and surrounding areas (e.g., Kreisa and Bambach Reference Kreisa and Bambach1973; Bambach Reference Bambach and Shumaker1983b, Reference Bambach and Shumaker1987; Sundberg et al. Reference Sundberg, Bennington, Wizevich and Bambach1990; Dennison et al. Reference Dennison, Bambach, Dorobek, Filer, Shell, Dennison and Stewart1992; Dorsch et al. Reference Dorsch, Bambach and Driese1994; Bennington and Bambach Reference Bennington, Bambach, Schultz and Henika1994; Pope et al. Reference Pope, Read, Bambach and Hofmann1997). He also took an interest in evolution, as seen in student projects by Fred Sundberg (Ph.D. 1990), Dave Jacobs (Ph.D. 1990), Delpfine Welch (Ph.D. 1991), Susan Phillips (M.S. 1993), and Andrew Bush (M.S. 1999).

Richard will be remembered as a key figure in the growth and success of analytical paleobiology. Although far from an expert in statistical methods (as he was always the first to acknowledge), he had an intuitive, almost uncanny grasp of paleontological data—what was signal, what was noise, and what the data meant. Thus, it was not a coincidence that he frequently collaborated with some of the world’s most talented analytical paleontologists, who recognized that Richard brought a sophisticated tool kit to the table: his understanding of geology, biology, and paleontology was broad and deep, and he frequently saw connections among disparate phenomena that others did not. We hope his publications will continue to illustrate how sophisticated analyses of the fossil record are enhanced by a rich understanding of natural history in all its messy details.

In particular, Richard deserves considerable credit for incorporating ecological and biological information into large-scale, quantitative analyses of the fossil record—he operationalized biological properties of organisms, finding a level of detail that was informative while being straightforward enough to apply to long-dead organisms and large databases. In effect, he bridged the gap between more traditional, idiographic approaches to paleontology (treating all events and species as unique) and the extreme versions of the newer, nomothetic approach, which sought general laws of history but treated all species as identical. His approach was ultimately built into the structure of the Paleobiology Database.

Richard also deserves credit for relentlessly encouraging his students and colleagues to present their statistical analyses clearly. As an example (according to Richard), Jack Sepkoski originally numbered his famous three evolutionary faunas in reverse: (3) Cambrian Fauna, (2) Paleozoic Fauna, and (1) Modern Fauna. These were the numbers spit out by the factor analysis: the Modern Fauna was the largest factor, so by default it was number 1. Richard convinced Jack to renumber the faunas in chronological order—a simple, thoughtful change that helped make Sepkoski’s analysis a classic.

Richard was generous in supporting students, colleagues, and the broader community. To pick a few examples from a long list, he was the president of the Virginia Tech Faculty Senate, he served on the editorial board of Paleobiology, he co-organized a short course for the Paleontological Society along with Patricia Kelley, and he coedited the Critical Moments and Perspectives in Earth History and Paleobiology book series for Columbia University Press along with Dave Bottjer. He also helped mentor a wide variety of students at Virginia Tech, Chicago, Harvard, the Smithsonian, and elsewhere; after Steve Gould’s death in 2002, he unofficially took over the advising of Steve’s remaining graduate students. Richard’s generosity with his time and ideas helped launch and support the careers of generations of students and colleagues. He continued to offer guidance and encouragement into his final years, as he became the last remaining “wise elder” to a generation of paleontologists who were themselves aging into the elder ranks of the discipline.

Beyond science, Richard was kind and supportive, adding joy to our personal lives even as he enhanced our professional endeavors—he was as eager to discuss Duke Ellington as Phanerozoic diversity history. A couple of us fondly remember him joining us to watch the opening-night premiere of Star Wars: The Phantom Menace in 1999 … we didn’t realize it as we crowded into the theater a few minutes before midnight, but it was his 65^th birthday—and he was happily staying up to the wee hours of the morning to watch a movie. And he loved baseball, insisting that at least some of his visits to Chicago coincide with Cubs homestands so he could take in a game at Wrigley Field. He liked to sit in upper-deck seats that were directly aligned with the first-base line and, because Richard was Richard, he charted every single pitch in his scorecard, with a filled-in circle if the pitch was a ball and an open circle if it was a strike.

As Richard liked to say when teaching biostratigraphy, every species divides time into three segments: the time before it existed, the time during its existence, and the time after it has passed on. So it is with people, and so we mark the passage of time. We’re deeply saddened that paleontology has entered its post-Bambach era, but immensely thankful that we were able to spend so many years with him. We’ll miss his belly laughs, his bolo ties, his goofy humor, his innumerable stories, his avuncular advice, his imagination, his kindness, his optimism, and all the other facets of his friendship. When you get a chance, join us in raising a can of Dr. Pepper in his memory.