Glen Everett Woolfenden, Fellow, Patron, and Past President of the AOU, died on 19 June 2007 at the age of 77, following surgery for a severe hiatal hernia. Glen was best known for his pioneering long-term study of the cooperatively breeding Florida Scrub-Jay, but he also was a towering figure in Florida ornithology. Among regular attendees of ornithological meetings, he is remembered as the consummate raconteur, a witty and gregarious promoter of the informal social gathering.
Glen was born in 1930 in Elizabeth, New Jersey, and fell in love with birds as an 11-year-old after his parents (Lester and Ethyl Woolfenden) moved to Westfield, New Jersey. As a youngbirdwatcher, Glen accompanied (and frequently impressed) members of the venerable Urner Ornithological Club through the woods, swamps, and shorelines of northern New Jersey. Too young to enlist during World War II, Glen sometimes birded along the coast of New Jersey and imagined himself crucial to national security while watching for enemy submarines. He never lost his deep fascination with the history and details of that era. Glen attended Westfield High School through junior year and then spent his senior year at Peddie School in Hightstown, New Jersey. By this point he had attained a robust size (6′2″ and 220+ pounds) and distinctive build (enormous shoulders and chest, narrow hips, strong but slender legs). As a lineman on the varsity football team, he incurred several knee injuries that would progressively hobble him later in life. (While getting in and out of cars, or as an airplane passenger, the elderly Glen's locked right leg frequently caused him to snarl “damn football knee.”)
From 1949 to 1953, Glen attended Cornell University, where he enrolled in R.O.T.C. and was an active member of the Sigma Alpha Epsilon fraternity. It was there that Glen explored in earnest his “work hard, play hard” philosophy. By his own admission, he worked hard and often struggled with academic fields outside of biology. However, he never struggled with being near the center of social activities from Friday afternoon through Sunday evenings. At Cornell, Glen was influenced by Arthur A. “Doc” Allen, as well as the many other student-ornithologists around Fernow Hall, including Richard Fisher, Brina Kessel, Kenneth C. Parkes, William Dilger, Walter J. Bock, and Robert W. Dickerman. Glen also spent a summer as a volunteer at the American Museum of Natural History. He admits that he felt intimidated by the likes of Robert Cushman Murphy, Roger Chapin, Thomas E. Gilliard, and Ernst Mayr, but the friendly and unassuming young Dean Amadon (fresh from his Cornell Ph.D.) became one of Glen's lifelong friends and role models.
Glen's ornithological training continued at Kansas University (KU) as one of the first graduate students of Harrison B. (Bud) Tordoff. From 1953 to 1956, Glen spent countless hours in the Dyke Museum, skinning birds under the guidance and wit of Robert M. Mengel. Besides the influence of Tordoff and Mengel, Glen credited E. Raymond Hall, who forced him to parse his words carefully and succinctly. Later, every one of Glen's students would experience this influence, occasionally to exasperating degrees. While at KU he participated in numerous collecting expeditions to the prairies and Rocky Mountains, including a memorable one to Colorado on which his father was invited along to be the cook. For his Master's thesis, Glen returned to the shorelines of New Jersey, where—at the urging of Amadon—he studied the comparative breeding biology of Seaside and Sharp-tailed sparrows.
In 1956, Glen entered the doctoral program at the University of Florida. Under Pierce Brodkorb's tutelage, Glen became a master of avian skeletal anatomy and could name every bone in a bird's body until his last living day. His Ph.D. dissertation was the first exhaustive comparison of the postcranial skeletal anatomy of the waterfowl. Glen's knowledge of the birds of the world and his obsession with taut writing were both enhanced by Oliver L. Austin, Jr., longtime editor of The Auk, whose research for his Birds of the World exactly coincided with Glen's period at Gainesville. By the time Glen received his Ph.D. in 1960, he had prepared more than 2,000 bird specimens.
In 1960, Woolfenden became a charter faculty member of the newly established University of South Florida (USF), in Tampa, Florida, where he would remain until his retirement in 1999. A lively lecturer, Glen taught ornithology, comparative anatomy, biogeography, and evolution. His size and outgoing nature made him a formidable colleague. For his robustly expressed, strong opinions, he was admired by some peers and feared by others. He once embarrassed a dean attempting to railroad an unpopular decision in a faculty meeting by storming to his feet and asking, “Did you, in fact, ever count the votes?” Glen won both teaching and research awards at USF, eventually being named Distinguished Research Professor. He used specimens regularly in all his classes, and routinely held court about ornithology and birds of the world with undergraduate andgraduate students in a well-worn lab-turned-museum at USE His lasting educational legacies are the many undergraduates whom he motivated to become professionals, and more than 30 graduate students, many of whom became successful scientists, educators, or conservationists. Among all his students, Ralph W. Schreiber (“pelican man” as Glen called him) deserves special mention. Despite the occasional pitched battle with Glen during Ralph's graduate career, the two became close friends, professional colleagues, and worldwide travel companions before Schreiber's untimely death.
Until his last few years at USF, Glen was active at volleyball, handball, and racquetball, each giving way to the next as a consequence of steadily worsening knees. Throughout his career, he was active in Florida ornithology and birding. He participated in countless Christmas Bird Counts, often proudly leading the charge into chilly, thigh- or waste-deep marshes just past dawn to flush bitterns, rails, and marsh-wrens. He was the meticulous compiler of counts in St. Petersburg (1962–1984) and Lake Placid (1993–2003). He helped establish the Florida Ornithological Society (FOS) and its quarterly journal, The Florida Field Naturalist, purposely insisting that the name would encourage articles on subjects other than birds. Glen served the FOS in numerous capacities as an officer, committee member, and President (1991–1992). In 1994, the FOS named Glen its sixth Honorary Member. Preceding him in this honor was William B. Robertson, chief biologist for Everglades National Park. During the 1970s and 1980s, Glen helped Robertson, a close friend and colleague, marshal the field crews for an intensive banding study of Sooty and Noddy terns on the Dry Tortugas. Over two decades, numerous established and prospective ornithologists assisted in those annual spring and summer Tortugas trips— which, Glen helped ensure, included concentrated hard work by day and convivial insobriety by evening. During a legendary FOS “roast” of Robertson in the late 1980s, Glen served as master of ceremonies dressed only in a makeshift toga crafted from a bed sheet. In 1992, Robertson and Glen jointly published Florida Bird Species: An Annotated List, the definitive catalogue of species documented from this biologically rich state.
During the 1960s, Glen spent considerable field time monitoring and trapping breeding birds for encephalitis surveillance along the Gulf coast of Florida. Glen recognized that Florida's bird communities were poorly described, so he conducted and published standardized breeding-bird censuses in numerous habitats even during graduate school. At Archbold Biological Station in the distinctive, sandy “highlands” of central Florida, Glen located several Florida Scrub-Jay nests and noticed that more than two adult jays appeared to be attending these nests. Aware of the phenomenon of “helpers” (cooperative breeding), and mindful of the superb opportunities for detailed study afforded by an excellent ecological research station, Glen began color-banding Florida Scrub-Jays in February 1969. That year, five of the nine nests he monitored had helpers, and a historic long-term study had begun. For 20 years, he regularly commuted the two hours between Tampa and Archbold to band, census, and observe the jays.
Among his strengths as a field biologist, Glen was compulsively organized, a meticulous record-keeper, and driven to completeness. These traits served him perfectly for designing one of the most thoroughly detailed field studies of a bird population ever undertaken. His cleverly simple color-banding scheme has been used by researchers worldwide. His compulsion to “censusevery month, find every nest, band every jay” defines the ongoing study to this day (to which we eventually added “map every territory, plot every breeding attempt, blood-sample every young, locate every disperser” and other such “Glenisms”). Woolfenden also recognized very early the privilege and scientific advantages of working with wild organisms that do not fear humans. By providing tiny bits of peanut each time he visited a jay family, Glen established a population of hundreds of birds that were easy to locate on monthly censuses, and—more importantly—extraordinarily easy to observe for collecting scientific data and behavioral insights. Because corvids learn quickly and empathically, especially from their parents, this revolutionary field technique led to the self-perpetuating tameness for which generations of Florida Scrub-Jays at Archbold have become famous.
In 1972, Glen hired me (a college sophomore) as a summer intern, which began a close friendship and professional collaboration that we both cherished for the rest of his life. Our attention during those early years was devoted to documenting the demographic details of the cooperative breeding system and interpreting these in light of conflicting theories about the origins of cooperation. Ever since first reading articles by Hamilton, Trivers, Alexander, and others, Glen remained suspicious that gains to inclusive fitness were not necessary to explain the advantages of group living, shared territorial defense, and even the helping at the nest exhibited by Florida Scrub-Jays. Reflecting especially our discovery of “territorial budding” and other direct benefits of group living and helping, Glen's view of the controversy was best summarized in an article entitled “Selfish behavior of Florida Scrub-Jay altruists.”
Woolfenden was not just a meticulous field biologist; he was also a devoted, sometimes single-minded, natural historian. He reveled in asking questions, both little and big, about nature. He also read enough, and forced his students to read enough, to keep engaged in timely questions about behavior, ecology, evolution, and sociobiology.
Although he continued to be active in Florida ornithology, Glen's attention grew increasingly focused on the Florida ScrubJays at Archbold. Through scores of technical and popular articles, Glen and his coworkers documented the intricate biology of Florida Scrub-Jay family groups, in which offspring remain in the natal group as a survival strategy while seeking breeding vacancies in a confined, overcrowded ecological setting. Strictly dependent on stunted, fire-maintained oak scrub unique to Florida, the species became seriously threatened as a result of agricultural, residential, and commercial development. Work by the Woolfenden group established the biological basis for the Florida Scrub-Jay joining the federal Endangered Species List in 1987. The attendant publicity greatly elevated public awareness of the need to protect the last, remnant tracts of oak scrub throughout peninsular Florida.
A steady series of articles about Florida Scrub-Jays made Glen an internationally recognized ornithologist. He was regularly invited to lecture around the United States and abroad, invariably scheduling extra days on such trips to see the local habitats and birds. He was named Corresponding Fellow of the Deutsche Ornithologen-Gesellschaft, member of the International Ornithological Committee, Distinguished Animal Behaviorist by the Animal Behavior Society, and recipient of the Margaret Morse Nice Award from the Wilson Ornithological Society. In 1985, our joint papers and book, The Florida Scrub-Jay: Demography of a Cooperativebreeding Bird, were recognized with the William Brewster Memorial Award from the American Ornithologists' Union. Glen always revered this award as his highest professional achievement.
Glen married his second wife, Jan, in 1981, and in 1989 they moved to live permanently near his jays at Archbold Biological Station. At the “Jay House,” Jan and Glen hosted dozens of distinguished visitors from around the world, plus regular, lively gatherings of local students, colleagues, and friends. Besides serving his simple joie de vivre, Glen used such occasions to link his past friends and colleagues with current ones, and to mentor his students socially as well as scientifically.
Glen was devoted to several ornithological societies, and was an early advocate of the occasional joint meetings that eventually spawned today's North American Ornithological Congresses. He attended annual AOU meetings faithfully from the 1940s through 2006. He was an Associate Editor for periodical literature (1965– 1971), was twice elected Vice-President (1974–1975,1977–1978), was twice an Elective Councilor (1972–1974,1976–1979), served as Assistant to the Treasurer (1974–1983), as President (1990–1992), and on the advisory board for the Birds of North America project, and was an AOU delegate to the North American Bird Banding Council. As a past president, Glen rarely missed even a single session of the AOU Council, where he steadfastly defended high scholarly standards, as well as legacy and tradition. During the 1984 AOU meeting at his alma mater KU, Glen surprised the assembled masses by appearing in white tie, tails, and top hat to serve as auctioneer.
Glen's lifelong unbridled enthusiasm and charisma attracted countless students and friends to the study of birds and nature. His mentorship of students was all-encompassing, encouraging attendance at meetings, freely introducing professional colleagues without airs or status, socializing together, and hosting many a sleepover on his living room couches before bird counts. Without question, he was a demanding mentor who expected his students to share his devotion to their chosen field. More than once, his students heard him declare that “scrub-jays don't take weekends off, why should we?” To students unprepared for this exuberance, and especially as he aged, Glen often came across as impatient and intolerant. Indeed, even to his closest friends, his views of the world could seem unrealistically absolute. But he never lost his enjoyment of nature, birding, and a good debate, especially with friends who recognized and loved the twinkle in his eye. His devotion to meticulous, long-term field study helped establish the importance of such projects in ecology, both at the Archbold Biological Station and elsewhere around the world. His study continues to this day, and his influence will continue indefinitely.
Glen is survived by his wife, Jan; a brother, Dan; three children (with his first wife, Gwen), Kim Woolfenden-Kaan, Scot Woolfenden, and Lisa Coker; grandchildren Michael Woolfenden, Grant Bucker, and Faith Coker; and a healthy, well-managed population of hundreds of Florida Scrub-Jays at Archbold.
R. Bowman and F. E. Lohrer, two of Glen's students, contributed to the preparation of this memorial.