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Volume 102, Issue 2 e01846
Resolutions of Respect
Open Access

Resolution of Respect Joseph Hurd Connell (1923–2020)

Wayne P. Sousa

Corresponding Author

Wayne P. Sousa

Department of Integrative Biology, University of California, Berkeley, 3040 Valley Life Sciences Building #3140, Berkeley, California, 94720-3200 USA

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Kyle E. Harms

Kyle E. Harms

Department of Biological Sciences, Louisiana State University, 202 Life Sciences Building, Baton Rouge, Louisiana, 70803 USA

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Claudia M. Tyler

Claudia M. Tyler

College of Creative Studies, University of California, Santa Barbara, Santa Barbara, California, 93106-6110 USA

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Margaret D. Lowman

Margaret D. Lowman

TREE Foundation, P.O. Box 48839, Sarasota, Florida, 34230-5839 USA

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Michael J. Keough

Michael J. Keough

School of BioSciences, University of Melbourne, Parkville, Victoria, 3010 Australia

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Peter T. Green

Peter T. Green

Department of Ecology, Environment and Evolution, La Trobe University, Bundoora, Victoria, 3086 Australia

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Allan Stewart-Oaten

Allan Stewart-Oaten

Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, 4311 Life Sciences Building, Santa Barbara, California, 93106-9620 USA

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First published: 09 February 2021

On 1 September 2020, we lost a legend in the science of ecology: Dr. Joseph (Joe) Hurd Connell, who died aged 96 (Fig. 1). Joe’s research and conceptual writings have shaped the field since the publication of his highly novel PhD study about factors controlling the abundance and vertical distribution of two barnacle species on the intertidal seashore of Scotland. His pioneering field experiments and unmatched long-term monitoring studies transformed the field of community ecology. His example moved the discipline from a predominantly descriptive endeavor of cataloging and interpreting spatial and dynamic patterns in nature to an experimental, hypothesis-driven effort aimed at understanding mechanisms responsible for these observed patterns. He also wrote several highly synthetic review papers that refocused conceptual perspectives of the discipline, constructively challenged status quo paradigms, and identified important questions for future researchers.

Details are in the caption following the image
Joe Connell at Davies Creek, Queensland, Australia, in 1988. Photo credit: M. Keough.

An insatiably curious, highly creative, warmly gregarious, and wickedly funny human being, Joe enriched and forever changed the lives of hundreds of friends and scientific colleagues, not to mention literally thousands of professional ecologists and their students who studied his publications or learned about his research from the pages of every ecology textbook and many introductory biology texts. In this essay, we share and celebrate the rich history of his life and his contributions to science and its practice. Some of the personal details we include come from the invited autobiographical accounts he wrote for Current Contents in recognition of five papers that had been designated “Citation Classics” (Connell 1981, 1987, 1988, 1989, 1992) and a personal profile he wrote for Peter Stiling’s textbook, “Ecology: Theories and Applications” (Stiling 2002, pp. 118–119). These essays reveal a lot about his motivations and personal circumstances. Margaret Connell, Joe’s spouse, generously provided additional details and helped us complete the timelines.

His Early Life

Joe’s path to an extraordinary career in ecology was far from linear and included a rich assortment of life experiences. He was born the fifth of October 1923 in Gary, Indiana, USA, and attended a Catholic elementary school in Fort Wayne, Indiana, USA. Later, his family moved to Ellwood City, outside Pittsburg, Pennsylvania, USA, where his father was employed as an engineer in the steel manufacturing industry. Joe attended high school there. In fall 1941, he enrolled at Carnegie Institute of Technology, planning on becoming an engineer like his father. However, his life changed dramatically after the attack on Pearl Harbor and the United States’ entry into WWII in December 1941. In the following year, he joined the war effort and enlisted in the US Army Air Corps in December 1942. The Air Corps had a critical need for weather forecasters, so Joe was enrolled in a specialized training program in Meteorology at the University of Chicago (1943–1944). He was then stationed in the Azores as a commissioned officer (1944–1946) with the 1st Weather Reconnaissance Squadron, serving as meteorologist on weather surveillance flights across the North Atlantic in a modified B-25D Mitchell Bomber. He gathered data essential to the safe passage of American convoys crossing the Atlantic in support of the European Theater of the war.

From an early age, Joe had enjoyed watching birds and identifying trees, but growing up in a small industrial town, where most professionals he knew were medical doctors, lawyers, or engineers, he did not recognize field biology as a viable career option. However, conversations with other army enlistees, who had pursued civilian careers in biology and wildlife management before joining the war effort, convinced him that a career as a field biologist was feasible. While in the Azores, Joe hiked around the islands observing birds and other wildlife. This was where his love of natural history became an enduring part of his makeup.

After the war ended, Joe returned to the University of Chicago and completed his BSc in Meteorology in 1946, but now he was determined to pursue a career in field biology.

The Curvy Path to an Unparalleled Career in Ecology

The temperate zone years

Supported by funds from the GI Bill, Joe enrolled in a master’s program in Zoology at UC Berkeley in 1947 and earned his MSc degree in 1953 under the supervision of wildlife biologist Dr. Aldo Starker Leopold. For his master’s thesis, he set out to document the movements and home range of the brush rabbit, Sylvilagus bachmani, in a chaparral-grassland-dominated canyon just east of the Berkeley campus. Leopold recommended this study organism because he thought the rabbit was abundant and a potential game animal that had been little studied. As it turned out, this proved to be dubious advice. The rabbits were very difficult to capture; over the course of his 14-month study (March 1948 to May 1949), he sampled for 2251 trap nights, but caught only 40 rabbits, and several of these became “trap-happy,” returning 6-14 times to the same trap (Connell 1954). Although his findings were an important contribution to our knowledge of brush rabbit natural history at that time, Joe found the project “frustrating” and the results “pretty dull” (Connell 1981, Stiling 2002, p. 118). This discouraging experience may explain why, after collecting his rabbit trapping data in 1948–1949, it was not until June 1953 that Joe filed his master’s thesis. (By then, he had already started, in 1952, to collect data for his doctoral dissertation!) The serendipitous consequence of this tedious rabbit project was that Joe “vowed then to adopt a simple rule of thumb, namely, never again to study anything bigger than my thumb” (Connell 1981). This pledge and a fortuitous introduction to the little-known field experimental studies of the French marine ecologist, Harry Hatton (Hatton 1938) motivated Joe’s famous study of competitive interactions between two species of barnacles on the shores of Scotland, described below. Ironically, he later immersed himself in long-term studies of corals and rainforest trees, which spend only a small fraction of their lives at a size smaller than a human thumb. Unlike brush rabbits, however, they stay put and it is relatively easy to collect data from many of them!

Taking a needed break from research after his master’s work, Joe tried his hand at secondary school teaching and taught biology for two years (1949–1951) at C. K. McClatchy Senior High School in Sacramento, California, USA. Teaching was hard work, but rewarding, and he might have continued in that profession had he not received notice that he had one year left of his GI Bill funds to use immediately or lose (Stiling 2002, p. 118). Two experiences he had while in graduate school at Berkeley were pivotal in determining his next career move (Connell 1992, Stiling 2002, p. 118–119). The first was a graduate seminar he had taken as a beginning student, in which he reviewed what he described as a “wonderful” paper published in 1947 by Edward Smith Deevey (Stiling 2002, p. 118). Deevey’s paper summarized and compared the limited number of life table datasets that had been collected from natural animal populations by that time. The most complete dataset included in the review came from Hatton’s study of settlement and survival rates in multiple populations of the intertidal barnacle, Balanus (= Semibalanus) balanoides, which lived on sheltered to exposed shores adjacent to St. Malo on France’s Brittany coast (Hatton 1938). Deevey was very impressed with Hatton’s results and specifically pointed out (p. 312) that B. balanoides “is a very favorable object for population research.” Joe took note that barnacles offered many advantages over brush rabbits for quantitative experimental studies of factors controlling the distribution and abundance of natural populations (and were smaller than his thumb!). The second event that shaped the trajectory of his career was his introduction to Dr. Charles Maurice Yonge, a sabbatical visitor to Berkeley from the University of Glasgow and a renowned specialist in the physiology and morphology of marine invertebrates, especially corals, mollusks, and crustaceans. With one year of GI Bill funding in his pocket and a clear vision of the kind of research he wanted to pursue, Joe moved back across the Atlantic to pursue a PhD in Zoology with Professor Yonge.

For his dissertation research (1952–1955), Joe studied barnacle populations on the shores of the Isle of Cumbrae in the Firth of Clyde, Scotland, based at the Marine Station at Millport. Because his Scottish landlady, Mrs. Plant, charged him very modest room and board (only £9 per week), he was able to stretch his one year of GI Bill funding to three (Connell 1981). This is also where, in 1952, he met Margaret Harvey, a visiting graduate student researcher from Oxford University who was studying ctenophore biology. They married in 1954 in Exeter, England, Margaret’s hometown (Fig. 2).

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Joe and Margaret Connell at the celebration of Joe’s 90th birthday in 2013. Photo credit: S. Holbrook.

Wanting to better understand the details of Hatton’s (1938) study, Joe painstakingly translated from French to English the entirety of Hatton’s 107-page paper, discovering that not only had Hatton gathered detailed observational data on barnacle demographics, but had used controlled field experiments to examine the factors that shaped patterns of post-settlement survival. This was a highly novel approach, perhaps the first time that such experiments had been conducted in the field under natural conditions. Prior to that time, ecological experiments had largely been relegated to the laboratory environment. Hatton scraped clean patches of the rock surface and monitored larval recruitment and subsequent survival of B. balanoides. He similarly monitored co-occurring populations of the barnacle, Chthamalus stellatus, which lives higher on the shore than B. balanoides. Hatton primarily studied the effects of density and physical factors on survival, employing controlled transplant experiments to measure the effects on survival of tidal elevation, rock surface aspect with respect to sun exposure, and an individual’s age/size. He also performed surface wetting and shading manipulations to assess the effects of heat and desiccation.

Deevey’s review paper and Hatton’s research greatly inspired Joe, who always gave credit where credit was due: “my career was shifted into a new direction by Hatton and Deevey, unbeknownst to them” (Connell 1992). Since Hatton had studied the effects of physical factors in controlling barnacle distributions across the tidal gradient, Joe decided to focus on the effects of biotic interactions on the same barnacle species, initially planning to investigate predation and intraspecific competition. Professor Yonge thought this an appropriate scope for Joe’s PhD dissertation, and cautioned him not to take on too much (Connell 1981). Joe, however, had taken a field ecology course at Oxford University, taught by Charles Elton, which convinced him that competition between species was an important biotic interaction structuring natural communities. Disregarding his advisor’s counsel, Joe surreptitiously added an experimental study of interspecific competition for space between B. balanoides and C. stellatus, species that were differentially distributed along the gradient of tidal height (Connell 1961a). Amazingly, this “side study” was not included in his dissertation, but turned into probably the most widely cited and influential study that Joe conducted. Joe surmised (Connell 1981) that this study received so much attention because there was a growing body of theory about interspecific competition, but little in the way of direct experimental tests demonstrating its influence on a natural animal population. Joe felt that the central study of his dissertation (Connell 1961b), while cited less frequently, was a more substantial and better paper. The two studies are highly complementary and employed controlled manipulations of barnacle densities, transplant experiments, and predator exclusion treatments to demonstrate the impacts of competition for space and predation on density and size structure.

Adult C. stellatus are most abundant on the upper shore and rare below, even though their larvae recruit over a range of lower tidal levels. Conversely, B. balanoides adults are densest at mid-shore to low shore levels and rare at upper levels, despite their larvae settling over much of the tidal range. Joe’s experiments demonstrated that, while predation by the snail, Thais (=Nucella) lapillus, reduced the density of large B. balanoides, their preferred prey, the mortality they caused was not sufficient to prevent the faster-growing B. balanoides from competitively excluding C. stellatus from the mid-shore to low shore. When B. balanoides were manually removed from the mid-shore plots, C. stellatus survived and grew well, while suffering high mortality due to competition for space with B. balanoides in unmanipulated control plots. In fact, C. stellatus grew better at mid-low shore levels, where they were more frequently submerged and filter feeding, than on the upper shore, where they were exposed to the air for longer periods. B. balanoides cannot survive the desiccating conditions characteristic of the upper shore, so the more desiccation-tolerant C. stellatus occupies this spatial refuge, free of competitive pressure from B. balanoides, thereby ensuring coexistence of the two species on the same shore.

Application of these innovative yet simple experimental techniques clearly revealed the mechanisms underlying the differential distribution of species along an environmental gradient. Joe’s approach inspired the use of controlled experimentation in hundreds, if not thousands, of similar studies across a wide variety of marine, freshwater, and terrestrial habitats. In a word, Joe’s dissertation study revolutionized the science of community ecology. It is no wonder his study’s results are featured in so many introductory ecology texts. In recognition of the special significance of the work, Joe was awarded the 1963 Ecological Society of America’s Mercer Award, given for “an outstanding ecological research paper published by a younger researcher.”

Joe received his PhD in Zoology from the University of Glasgow in 1956. Following a postdoctoral study at Woods Hole Oceanographic Institute (1955–1956), where he studied the spatial distribution of two species of clams, he accepted a two-year instructorship in Zoology (1956−1958) at the University of California, Santa Barbara (UCSB), and was subsequently hired as an Assistant Professor of Zoology in 1958. He spent the rest of his career at UCSB, earning tenure to Associate Professor of Zoology in 1961, and a promotion to Full Professor of Zoology in 1966. He retired in 1991 as Professor Emeritus of Zoology and held the appointment of Research Professor of Biology from 1996 to the time of his death.

When he returned to the west coast of the United States, Joe sought to test the generality of the patterns he had documented in Scotland. Soon after being appointed to the faculty at UCSB, Joe headed north to the Friday Harbor Marine Laboratories on San Juan Island off the northwest coast of Washington State, where a comparable assemblage of barnacles and their snail predators occupy the intertidal zone. Working at two sites on the shore of San Juan Island, Joe conducted a series of experiments, similar to those he had used in Scotland, to study the impact of competition for space among three species of barnacles. The interactions proved quite different in this site, where three species of predatory snails (Thais spp.) preyed on the barnacles, as opposed to the single species of predatory Thais in Scotland. Their combined impact strongly reduced the density of the most common barnacle species, Balanus glandula, so that competition for space among barnacle species on the low to mid-shore was insignificant (Connell 1970), a very different outcome than Joe observed in his dissertation research. This study was among the first to demonstrate that predation can mediate the intensity of interspecific competition and thereby promote the coexistence of potentially competing prey species. The site-dependent nature of the interactions that Joe documented in Scotland versus Washington State became a major conceptual theme (i.e., context-dependence) in community ecology 40 years later.

The tropical years

After being awarded tenure in 1961, Joe decided he “would strike out in a new research direction—try something new or outrageous—without worrying about whether it might cause a hiatus in my publication record” (Connell 1987). After studying relatively low-diversity temperate marine intertidal ecosystems for more than a decade, Joe became increasingly curious about the mechanisms that maintain high levels of species diversity in tropical ecosystems. At that time, Joe subscribed to the prevailing theory that, over evolutionary time, the high productivity and purportedly stable climatic conditions of the tropics had selected for narrow, specialized niches and high levels of species packing along resource gradients. He recognized, however, that the assumptions and predictions of this model needed to be challenged with data. By collecting long-term, spatially explicit data on demographic dynamics, including patterns of recruitment, growth, and mortality, Joe aimed to evaluate the stability of these communities and discover the mechanisms that structure them. He chose to go “all-in,” successfully applying for a Guggenheim Fellowship and moving with his family to the tropics of Australia in 1962 to study the ecological processes structuring coral assemblages on the Great Barrier Reef. During that first year in Australia, Joe met John Geoffrey Tracey and Leonard Webb of the CSIRO Rainforest Ecology Unit, the only two Australian ecologists working in rainforest at that time (Connell 1987). Their discussions gave birth to the idea of a collaborative study of mechanisms maintaining species diversity in this iconic tropical habitat, and in 1963, Joe added rainforest ecology to his portfolio.

These more species-rich communities do not lend themselves as readily to the small-scale experiments that Joe had so successfully used on rocky seashores to identify the mechanisms structuring those communities. Nevertheless, he and colleagues were able to conduct manipulations of avian and mammalian seed predators and litter disturbers at his forest site in North Queensland to assess their effects on seedling assemblages (Theimer et al. 2011). The major results of his tropical work, however, come from the analysis of long-term census records collected from permanent plots established on the Great Barrier Reef and in two rainforest sites in the state of Queensland. In 1962, during his first visit to Australia, Joe established replicate permanent 1-m2 plots, line-intercept transects, and belt transects in several different sub-habitats across the reef on Heron Island near the southern end of the Great Barrier Reef. Changes in the 1-m2 plots, including the recruitment of new colonies, were quantified from 35-mm color slides taken with an SLR camera positioned vertically over each plot during 36 visits in 26 of the 38 years between 1962 and 2000 (Fig. 3; Connell et al. 1997a, 2004, Tanner et al. 2009). A year later in 1963, Joe, working with Tracey and Webb, laid out two large permanent rainforest plots, one (1.7 ha) at Davies Creek in tropical North Queensland southwest of Cairns, and the other (1.9 ha) in subtropical South Queensland near O’Reilly’s Rainforest Resort within Lamington National Park just south of Brisbane (Connell et al. 1984, Connell and Green 2000, Green et al. 2014). Seedlings and small saplings were identified, measured, and mapped within several belt transects at each location, and larger trees were individually marked and censused throughout both plots. These plots have been re-censused by teams of field assistants every few years; surviving trees are remeasured, new recruits tagged, and deaths recorded (Figs. 4-6). Through the years, literally hundreds of graduate students and postdocs "groveled" on the rainforest floor with Joe, measuring seedlings and getting leech bites as part of a ritual for aspiring field biologists. In celebration of the 50th anniversary of the establishment of these plots, Harms and Green (2014) published a description of the history and contributions of what is now known as the Connell Plots Rainforest Network. These records of community structure and dynamics may be the longest ever collected from these habitat types.

Details are in the caption following the image
Joe Connell preparing to photograph a 1-m2 plot on the coral reef flat at Heron Island, Great Barrier Reef, Australia, in 1967. The wooden frame around the quadrat reduces water surface rippling, improving the clarity of the image. Photo credit: D. Potts.
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Joe Connell carefully mapping a treefall gap in the Davies Creek rainforest plot in 1999. Photo credit: K. Harms.
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Joe Connell (right) and Geoff Tracey (left) censusing seedlings along a transect at the Davies Creek rainforest site in 1977. Photo credit: W. Sousa.
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Davies Creek rainforest plot census crew in 1977. Photo credit: W. Sousa.

Joe immediately recognized that, to make meaningful headway in studying the mechanisms maintaining diversity in these hyper-diverse communities, he needed the expertise of skilled taxonomists to accurately distinguish the many morphologically similar species co-occurring in each habitat. In addition to being accomplished ecologists, Tracey and Webb were experts at identifying rainforest plants, including their juvenile stages. Drawing on Tracey’s extensive field observations and his help compiling large sets of forest survey data, Webb had published the first physiognomic-structural classification of Australian rainforest vegetation in the Journal of Ecology in 1959. On the coral side, Dr. Carden Wallace, of James Cook University and the Museum of Tropical Queensland, provided essential help identifying corals in his plots and belt transects.

Several events raised questions in Joe’s mind about the generalization that tropical communities should be considered equilibrial or “balanced” systems, tightly regulated by density or frequency-dependent biotic interactions such as interspecific competition for limited resources, predation, and disease. The first was a teatime conversation Joe had with the renowned Australian population biologist, Charles Birch, during which Joe presented his working hypothesis of stable, tightly regulated tropical communities. Birch replied: “Fine, but also keep your eyes open for tropical weather up in Queensland—some of those cyclones might shake things up a bit” (Connell 1987). The second event that challenged his assumption was in fact the impact of a severe cyclone that passed over both Heron Island and one of his rainforest plots in 1967, causing considerable damage (Connell et al. 1997a, 2004). At that time, many researchers considered these episodes rare, annoying events that “wrecked my study,” rather than drivers of community dynamics whose effects might not be recognized without long-term monitoring studies. Joe subsequently broadened his view of the possible mechanisms that could maintain diversity in these systems to include disturbance.

Joe’s detailed coral and rainforest studies yielded novel insights into the processes that maintain the extraordinary levels of species diversity in these two tropical ecosystems. The studies challenged the long-standing view that these were equilibrial assemblages of tightly co-evolved species. From his 38-year record of changes in the coral system (Connell et al. 1997a, 2004, Tanner et al. 2009), Joe documented the damage caused by multiple cyclones and subsequent patterns of recovery. Large storm waves associated with these storms battered the reef, breaking and displacing colonies, while shifting sediments harmfully abraded them. Partial or complete death and displacement of colonies opened space for new recruits; competitively dominant species were knocked back and recolonization by competitively inferior species enriched the coral assemblage. This episodic reset of coral assemblages to a younger, more diverse state by disturbances of intermediate frequency and intensity as a general mechanism by which diversity is maintained is what Joe called the “Intermediate Disturbance Hypothesis” (Connell 1978).

In the Queensland rainforests, a different story emerged. When storms and other natural disturbances damaged or killed trees (Connell et al. 1997b), they opened light gaps that enhanced local recruitment and growth of juvenile trees. But other diversifying processes were also at work. Joe’s good sense to study all rooted life-cycle stages of his rainforest trees arose from his earlier research, which convinced him that both the supply side and what might be called the “sorting side” (i.e., the competition, predation, disease, and habitat filtering that can non-randomly sort among individuals) were important in community ecology. The youngest seedlings on his transects clearly recruited in spatial clumps, most often near the parent tree, and recruitment rates varied year to year (e.g., Connell and Green 2000). Even so, relatively quickly the pattern diversity of cohorts of seedlings increased as they matured into later-stage saplings. These observations suggested to Joe that a combination of limited seed dispersal and the frequency-dependent influence of natural enemies—especially those that are relatively host specific—could be important diversifying mechanisms for trees (and other sessile organisms). Joe published his idea in a book chapter (Connell 1971) soon after tropical biologist Dan Janzen (1970) independently published similar thoughts. Their ideas have been linked ever since as the “Janzen-Connell Hypothesis” (Hubbell 1980). The concept has had a resounding influence on tropical forest ecology and remains one of the key elements in our collective working model for tropical rainforest diversity and dynamics (Wright 2002).

Impact on the Discipline of Ecology

Joe’s empirical findings and conceptual writings forever changed the science of ecology. With a well-deserved nod to his little-known predecessor Harry Hatton, Joe deserves credit for introducing and popularizing the use of controlled, replicated experiments to investigate ecological hypotheses in nature. One measure of the impact of his work on the discipline is how heavily his papers have been referenced by other scientists: 11 papers or book chapters have been cited over 1,000 times each. To date, these 11 papers combined have been cited 32,136 times!

Joe also made major contributions to the growth of ecological theory and concepts by writing highly synthetic and critical reviews that evaluated published research results pertinent to key ecological phenomena. Especially impactful reviews addressed the role of natural enemies in preventing competitive exclusion (Connell 1971), community interactions on rocky intertidal shores (Connell 1972), shifts in species interactions along environmental gradients (Connell 1975), mechanisms of ecological succession (Connell and Slatyer 1977), hypotheses explaining the maintenance of high diversity in tropical communities (Connell 1978), the prevalence and relative importance of interspecific competition (Connell 1983), and the evidence needed to judge ecological stability and persistence (Connell and Sousa 1983). Each of these reviews has been cited more than 1,000 times to date, with the succession paper receiving more than 5,200 citations and the tropical diversity paper receiving more than 11,600!

Ironically, while Joe provided perhaps the most famous demonstration that interspecific competition can shape community structure and species distributions, he also actively challenged the notion that it was the overriding and ubiquitous factor structuring communities. Several of his reviews questioned this prevailing perspective (Connell 1980, 1983) and provoked those invested in the “competition” paradigm.

In the arena of applied ecology, Joe served as inaugural Chair of the Marine Review Committee (1974–1990), which was charged by the California Coastal Commission with assessment of the environmental impacts of the San Onofre Nuclear Power Plant on the coast of southern California.

Joe received many honors and awards, among them two Guggenheim Fellowships, the Ecological Society’s Mercer and Eminent Ecologist awards, Fellow of The American Academy of Arts and Sciences, and corresponding member of the Australian Academy of Sciences.

Curiosity and Objectivity Drove Joe’s Science

Joe was insatiably curious about the world around him (natural and human), and he wanted to know about everything. Few, if any, ecologists have studied as many different organisms and natural communities as Joe did, including small mammals in chaparral, rocky intertidal invertebrates, soft-sediment clams, rainforest trees, corals, and desert shrubs. He thoroughly enjoyed opportunities to immerse himself in new natural history experiences and loved engaging with students over their research ideas and results, regardless of the study organism or habitat.

He was unfailingly objective about his own ideas and data, adhering to the Popperian scientific method more closely than many scientists. He did not cling to pet hypotheses; if evidence falsified one of them, he gladly moved on to consider another. Ego did not drive his scientific judgments. Joe was also highly egalitarian in his interactions with students and colleagues, especially when it came to seeking critical comments on his manuscripts. He would relentlessly pepper colleagues and students with multiple drafts of every paper he wrote. He carefully considered every comment, no matter whether it came from an undergrad or a full professor. If something was unclear, it was his fault, not the reader’s. He worked hard to clearly communicate his results and conceptual ideas in his writing,

Joe as Mentor, Colleague, and Friend

Joe gave his graduate students a lot of freedom in their choice of study organisms and field sites. He mostly cared that the questions and hypotheses were clearly stated and testable and that the findings would make a novel and valuable contribution to our understanding of the natural world. Of his 18 PhD and 2 MA thesis students, 12 worked in marine rocky intertidal habitats, two in marine soft-sediment environments, two in subtidal kelp forests, one in chaparral shrubland, one studied salmon in aquatic habitats, one studied terrestrial snails, and one studied stream flatworms.

Fifteen postdoctoral researchers worked with Joe. Most collaborated on data collection, data analysis, or manuscript writing associated with either his coral reef or rainforest study. Several were involved with assessment of the potential impacts of the San Onofre Nuclear Power Plant. Others pursued their own research projects. Joe considered them peers and their collaboration greatly enhanced and expedited the analysis and publication of some of his most important research results.

Joe gave detailed feedback on his students’ work. When he finished working over their papers, the bath of red ink could be visually and psychologically jarring, but the feedback always resulted in a clearer exposition of ideas. Some of his most frequent corrections were marked “No NUAs” (nouns used as adjectives) or “EUW” (eliminate unnecessary words), mixed with a few good-natured curse words. Learning to accept constructive criticism was a key part of his regimen of scientific training.

He was so committed to the value of scientific data that he actually wrote up and published the dissertation chapters of two of his PhD students who were unable to do so, assigning himself second authorship. Even though he spent long hours advising on research questions and study design, he never asked to be included as an author on his students’ dissertation publications, believing that his mentoring efforts were part of his job description and that sole authorship was in the best interest of his students’ careers. It was not only the professional success of his students and colleagues that he cared about; he was also attentive to their general well-being by offering thoughtful support and kindness during hard times and, on occasion, providing practical material support for fieldwork. For example, when one of his graduate students had no means of transport to off-campus field sites, Joe donated an old, but fully functional, family car to the cause.

Joe was a very social person, whether it be a campfire “billy up” at lunchtime on the side of the dirt road to the Davies Creek site (Fig. 7; https://www.nma.gov.au/exhibitions/symbols-australia/billy), or a gathering at his home to celebrate a visitor or special event. He loved hearing what people had to say, and the conversations often moved to non-scientific topics like favorite movies and books. He loved Japanese samurai films, classical music, and Marx Brothers movies, often imitating Groucho’s witty mannerisms and clever remarks. And Joe was a huge fan of the LA Dodgers. He probably cheered them to a World Series victory shortly after his own final inning in 2020.

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Lunchtime break for the 1977 census crew at Davies Creek. Note the classic Australian billycan in left foreground. Photo credit: W. Sousa.

Joe is survived by his wife Margaret, their four children, Andrew, Jane, Tim, and Kate, and his niece Mary Rollins Jones, along with 11 grandchildren and seven great grandchildren. We are deeply appreciative to Joe’s family for sharing him with us; it is such a precious gift.

A graduate research fund has been established in Joe’s honor at UCSB: The Joseph H. Connell Field Ecology Research Fund. Donations may be made in two ways. Those who wish to donate online can use this link: https://giving.ucsb.edu/Funds/Give?id=78 (please add "for JH Connell Fund" in the notes.). For donations by check, please make it payable to the UCSB Foundation (with “for JH Connell Fund” in the memo line) and mail to: Office of Development #2013, Attn: Bethany Innocenti, UCSB, Santa Barbara, CA 93106-2013. Questions may be directed to [email protected].

Acknowledgments

We thank Margaret Connell for providing important details about Joe’s life. We are also grateful to Betsy Mitchell for comments that improved the manuscript.