Marine Ecology, mid‐ - Wiley Online Library

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History of Ecological Sciences, Part 58A: Marine Ecology, mid-1920s to about 1990, Featuring Beebe, Bigelow, Ricketts Frank N. Egerton Department of History, University of Wisconsin-Parkside, Kenosha, Wisconsin 53141 USA E-mail: [email protected] Part 51 discussed the origins of marine ecology and oceanography, 1870s to the 1920s (Egerton 2014). Vaughan et al.’s (1937) International Aspects of Oceanography included marine ecology. Jack (1945) wrote a briefer survey, limited to biology: “Biological Field Stations of the World”. Hiatt et al. (1954) wrote a larger, but more restricted Directory of Hydrobiological Laboratories and Personnel in North America. Later, Hiatt (1963) edited World Directory of Hydrobiological and Fisheries Institutions. Schlee’s (1973) The Edge of an Unfamiliar World: a History of Oceanography has some discussion of marine ecology and much background information concerning oceanography. C(harles) P. Idyll’s (1969a, b) Exploring the Ocean World and Ward’s (1974) Into the Ocean World are popular histories of oceanography, with Ward’s emphasizing marine ecology. An important collection of historical essays is Sears and Merriman, Oceanography, the Past (1980). The title of Mills’ (1989) Biological Oceanography: an Early History, 1870–1960, indicated coverage of marine ecology for the period now discussed. However, he concentrated on plankton studies at Kiel, Germany; Plymouth, England; and Woods Hole, Massachusetts. “Mac” McIntosh (1985:110–120) briefly surveyed the history of marine ecology for the period of this part 58, including numerous references, in The Background of Ecology. The American Institute of Biological Sciences (AIBS, 2010) reprinted from BioScience a booklet (ii + 72 pages) of articles—one from 2002, six from 2009)—of historical relevance, entitled Topics in biological field stations. Tydecks et al. (2016) collected worldwide data on biological field stations which they summarized and cited a Web site for details. Part 58 supplements Schlee’s (1973), Ward’s (1974), and Mills’ (1989) discussions. Ward has a c hapter on Ricketts, both Schlee and Mills discussed Henry Bigelow, Mills discussed Gordon Riley, yet neither Schlee nor Mills discussed popularizing scientists William Beebe, Edward Ricketts, Rachel Carson, or Jacques Cousteau. Bigelow, Riley, and Clark were university professors; Beebe, Ricketts, and Cousteau were independent ecologists; and Carson was a government biologist. Popularizers are 372 Bulletin of the Ecological Society of America, 97(4)

included because they published original observations and inspired youths to become marine ecologists. Just as Philip Gosse’s popularization of marine biology in the mid-1800s had advanced that science (Egerton 2010:190–194), so Beebe, Carson, and Cousteau advanced science with their popularizations of marine ecology a century later. Evidence? A Cousteau biographer reported (Matsen 2009:xv) the following: A marine biologist from Olympia, Washington, said he had watched every episode of The ndersea World, decided to devote his life to studying the sea and its creatures, and never wavered U in his passion, He gave me a program he had saved from a Cousteau Society environmental rally held in Seattle in 1977. William Beebe (Charles) William Beebe (1877–1962) was the best-known and most widely read American naturalist during the first half of the 1900s (Welker 1975:75, Sterling 1981, 1997, 1999). He is the subject of two biographies and an attractive children’s book (Sheldon 2009). The biographies are quite different: Welker’s (1975) Natural Man: The Life of William Beebe focused heavily on Beebe’s publications (24 books and 821 articles; Berra 1977), finding much to praise and to criticize; Gould (2004) had access to unpublished sources not previously available when writing The Remarkable Life of William Beebe: Explorer and Naturalist, and she found much to praise. Additionally, Matsen (2005) has written Descent: The Heroic Discovery of the Abyss, a superb study of Beebe’s oceanic adventures and investigations. Beebe was the only (surviving) child of a successful businessman and a nurturing mother, born in Brooklyn, but when he was ready for school, they moved to East Orange, New Jersey, motivated largely by his fondness for nature. He was a slightly above-average student in classwork, but was diligently self-educated in natural history and zoology. He was also an energetic, happy extrovert who excelled at sports and played musical instruments. In summers, he and his parents vacationed on the Bay of Fundy, where he enjoyed collecting and studying invertebrate animals. After graduating from high school, he enrolled in classes at Columbia University in September 1896, under the guidance of Professor Henry Fairfield Osborn, chairman of the Biology Department. Although Beebe never graduated, he acquired his biological outlook at Columbia, mainly from Osborn. His theoretical perspective was evolutionary, but evidences he collected were ecological. After his three years at Columbia, Osborn thought Beebe qualified for a position at the new Bronx Zoological Park. Osborn took him there and he became an assistant curator of birds in 1899 (Bridges 1974:63, Welker 1975:10, Gould 2004:54–55). Beebe published numerous articles on birds, beginning in 1906 (Berra 1977:27–41), and William T. Hornaday, Director of the Bronx Zoo, decided that Beebe needed a proper zoological journal in which to publish and founded Zoologica (Bridges 1974:290–292). Its opening article was Beebe’s (1918–1924) “Geographical Variation in Birds with Especial Reference to the Effects of Humidity.” He also published six books, 1905–1918, mostly on his second love, jungles, followed by a definitive, well-illustrated Monograph of the pheasants (4 volumes, 1918–1924) (Welker 1975:215, Berra 1977:109–113). In 1915, Beebe and two assistants travelled to Pará, Brazil, to collect specimens for the Zoo and New York’s Museum of Natural History. That brief trip convinced him that the tropics were too complex to really understand in brief visits. Beebe was perhaps the first biologist to establish a land-focused

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Fig. 1. William Beebe in British Guiana in 1917. Wikipedia.

research facility in the American tropics to study jungle life. In 1916, he arranged for his research team to use Kalacoon House, a “fine large house overlooking the confluence of the Mazaruni, Essequibo, and Cuyuni rivers, which [George] Withers had offered rent free,” near Georgetown, British Guiana (Welker 1975:67–69, Gould 2004:190). In 1923, he led an expedition to the Galapagos Islands. Beebe’s fascination for marine life, awakened as a youth at the Bay of Fundy, became a prominent research topic on this expedition to the Galapagos Islands in 1923. Land animals had somber coloring and often moved slowly. In contrast, marine animals were often brightly colored and moved quickly. Also, “three-fifths of the birds of this island [Indefatigable] were wholly dependent on water life for food, while a number of others included fish and crustaceans as a part of their diet” (Beebe 1924:127). Half dozen expedition members collected specimens by fishing, trolling, scooping up, dredging, towing plankton nets, and seining in different marine environments. Although scientists now use the Spanish names (Johnson and Johnson 1959:688), Beebe and his biographers used English names (as Darwin had), and for convenience, I do also. He was curious about Galapagos marine iguanas, which graze on sea weed—the world’s only marine lizards. There were also red crabs along rocky shores. One day while he was observing the placid marine iguana, he a crab crawled onto one of them and plucked off three ticks Amblyomma darwini (Beebe 1924:121–122). He assumed the crab ate the ticks but he did not see it do so. The iguana was undisturbed by the crab, and after the crab left, Beebe found the iguana still had 16 ticks. 374 Bulletin of the Ecological Society of America, 97(4)

Fig. 2. Map of Galapagos Islands, English names. Darwin (1844).

In three weeks, they collected 60 of 156 known species of near-shore fishes (Beebe 1924:128–134, + plate 3 and figure 20). They discovered two new species (Berra 1977:141–142 lists new species Beebe described) and added twenty species not previously known from the Galapagos Islands. Beebe described fish and other animals from low-tide pools. In Galápagos: World’s End, Beebe listed the impressive number of specimens collected (and included paintings and pages of notes) and also 22 articles either already published in Zoologica or pending publication there (Beebe 1924:xii–xiii). Beebe only wrote the article on birds; the other twenty-one were written by specialists. Beebe’s interest in species extended beyond collecting for the Bronx Zoo. He was fascinated by their behavior, seen in this comment about blennies (Beebe 1924:129): “Two closely related species had diametrically opposite escape reactions; one was found on wet rocks or seaweed out of water, and when alarmed dived to the pool bottoms, the other perched on stones or fronds beneath the surface and instantly leaped clear of the water on being disturbed.” He and his team were as interested in invertebrates as in fish and in those living in deeper water as well as in tide pools. A plankton net pulled through apparently empty water revealed “a tremendously rich haul” of plankton which they could observe in a laboratory aquarium in the yacht. Transparent, hermaphroditic Sagitta worms were “very voracious and feed on copepods, diatoms, small shrimps and even larval fish” (Beebe 1924:136). His plate illustrating plankton is likely one of 46 drawings made by Gilbert Broking, Artist, and General Preparateur. His Galapagos expedition was successful, and after he returned, a member of the Board of Managers of the New York Zoological Society offered use of another ship, Arcturus, for another expedition

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Fig. 3. Tide-pool fish from Eden Island. Beebe (1924: plate III).

(Beebe 1926a:v). Beebe accepted and it lasted from 11 February to 30 July 1925. On his Galapagos expedition, marine life had been an afterthought, but it became the focus of the Arcturus expedition. In the preface to The Arcturus Adventure, he explained that this book, like Galápagos: World’s End, is a “scientifically accurate, popular presentation of the high lights and vivid experiences of the expedition” (Beebe 1926a:viii) and that technical accounts would appear in Zoologica, volume 8. Before departing, he set two objectives, neither of which he accomplished as anticipated: to investigate the Sargasso Sea and the Humboldt Current. He discovered that February was the wrong time to explore the Sargasso Sea because of frequent storms, and the Humboldt Current had disappeared from around the Galapagos Islands (Beebe 1926a:ix–x). Nevertheless, the Arcturus was well equipped to study oceanic life and waters, and the expedition managed to collect enough specimens to provide a basis for Beebe’s speculations. He concluded from his examination of sargassum that this kelp could reproduce vegetatively in the Sargasso Sea without depending upon fragments being “storm-torn from the rocks of shallow coastal waters and poured forth by the Gulf Stream” (Beebe 1926a:7). Beebe’s party crossed the Panama Canal on March 28, and he expected to hit the Humboldt Current on the morning of the third day from Panama. However (Beebe 1926a:44), 376 Bulletin of the Ecological Society of America, 97(4)

Fig. 4. Plankton species caught with a net in waters near Galapagos Islands. Beebe (1924: facing 136).

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Fig. 5. Oceanic investigative apparatus of the Arcturus in 1925. Beebe (1926a: facing 5).

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…I soon found that, most astonishingly, that Antarctic river had nothing at all to do with the gigantic Current Rip, which was caused by the coming together of two warm, westwardly flowing streams of water. When we first detected the rip we were in 2° 36′ N Latitude, and 85° W Longitude, which placed us about two hundred miles southeast of Cocos Island. Temperature measurements showed that this was not the Humboldt Current. John Kricher (2002:24) provides a diagram indicating the impact of El Niño on the Humboldt Current, and Gould (2004:249) thought this was probably the first study of what became known later as El Niño–Southern Oscillation (ENSO). Peruvian fishermen were aware of the disruption of the Humboldt Current every 10 to 20 years. There was one other scientist who observed and published observations on the 1925 El Niño, and he knew this name from the fishermen (Wooster 1980). Robert Cushman Murphy was an ornithologist at the American Museum of Natural History; he and Beebe were acquainted, although their simultaneous expeditions had no contact. Murphy (1926) was in Peru studying oceanic birds, and by the time he published an article on El Niño, he was fairly well informed. He also had a discussion on it in his Oceanic Birds of South America (Murphy 1936: I, 2 volumes, 101–108). An understanding of the larger phenomena required an understanding of how the weather affected the Humboldt Current. Sir Gilbert T. Walker, while stationed at India’s Meteorological Department, saw the importance of understanding monsoon rains and published many papers on his findings, 1910–1936. He is credited with explaining the Southern Oscillation some years before Beebe published Arcturus, but not until 1969 did J. Bjerknes connect El Niño and Southern Oscillation (Bjerknes 1969, Katz 2002:98). Beebe may have been one of the first two scientists to call attention to El Niño, but it was Murphy who knew about its periodicity and what caused it. Beebe (1926c) did report in January that another American ship was at the Galapagos Islands in September 1925 and reported that the Humboldt Current had reappeared. Glynn (1988) has described the ecological impact of the 1982–1983 ENSO. In 1923, Beebe had been fascinated at discovering the Galapagos water-strider Halobates (new species in established genus), the only known insect genus that spends its life at sea (Beebe 1924:83–86). He found thousands in a tight cluster. He had seen other members of the genus in other oceans. He dropped them into a basin of water, in which they sank and drowned, because they could not regain the surface film on which they lived. He wondered how they survived a heavy rain storm at sea. He had previously read that they lay eggs on floating bird feathers, and on the Arcturus voyage, he found ten floating feathers with their eggs. When he placed feathers with eggs in an aquarium, the next day the eggs began hatching and continued hatching for four more days (Beebe 1926a:60–63). He estimated that there were about 20,000 eggs on one feather and thought it was a collective contribution from many females. Small fish in two aquaria in which he had placed feathers gladly ate the Halobates hatchlings. The feathers all appeared to be from booby wings. (Today, Halobates often deposits eggs on floating plastic: Madren 2012). Before embarking on Arcturus, Beebe bought a copper diving helmet with hose, to study ocean life underwater. Diving helmets were made for underwater salvage work. “One other scientist had tried a helmet, but he had never published his work in any but the driest of scientific terms” (Gould 2004:252). Biographer Gould did not name the previous scientist nor indicate whether Beebe had known about him before his own underwater explorations. Beebe’s other biographer claimed that “Beebe was simply the first famous and highly trained natural scientist to take to helmet diving as a new form of field work” (Welker 1975:111–112). To call his helmet explorations “diving” is to use the term loosely, as he always entered the water feet first down a ladder on the side of a boat, but Beebe himself referred to his descents as diving. His first descents were at Darwin Bay on Tower Island in the Galapagos Islands. He

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Fig. 6. Galapagos Halobates species. Beebe (1924: plate 4a).

took along a slate on which to take notes (which he found cumbersome) and a harpoon for protection (Beebe 1926a:78–86). He used the harpoon on his first dive to discourage a large grouper fish, which attacked the harpoon. He had his boat team lower a dead crab as bait, and he was soon surrounded by about 500 fish of 20–25 species. On his second descent, he harpooned “my first Pomacentrus leucurus” (Beebe 1926a:87). Then, thinking of the lay audience for his Arcturus book, he commented, “Although I am a scientist and a hunting scientist, I hate to take life.” Was this statement sincere? A few pages later he stated that “The best sport was to be had with the brilliantly colored wrasse” (Beebe 1926a:90). Meaning? “Twice I struck and marked them, and day after day the same individuals would come about as bold as ever, flaunting their wounds in my face” (Beebe 1926a:91). On one descent, he decided to collect rocks from the bottom. He filled five pails with rocks which were hauled aboard his boat. The pails were left standing out in the sun, and at the end of the day (Beebe 1926a:93): “There had crept out an amazing array of interesting beings—beautiful sea-worms, starfishes, squillas, hermit crabs, and shrimps of every hue, a number of strange larval fish,” including moray eels exactly like adults, but only 1.5 inches long. He realized that these animals could not be obtained by dredging along the bottom and that he had therefore opened a new door to exploration. Beebe (1926a:x) judged that: …the scientific accomplishment which, scientifically, proved the most valuable of all, was the result of my decision to make a ten-day stay in one spot in mid-ocean, Station 74 (Chapters XIII and XIV), where continual dredging yielded very remarkable collections of fish and crustacean, equivalent to any two months of the less intensive work….crustacea taken at Station 74 equal 80% of all…which we took in the Pacific. 380 Bulletin of the Ecological Society of America, 97(4)

Station 74 was where they were after steaming south from Cocos Island from late afternoon, May 24 until dawn May 25. A sounding indicated the sea floor was 771 fathoms below, so no helmet descent. This idea of intense study at a single place had come to him when he was studied jungles (Beebe 1916); now, he applied it to marine organisms. Station 74 was 4.5° N by 87° W of Greenwich, sixty miles south of Cocos Island. In ten days, Beebe saw 74 birds of 13 species, a good number of which had landed on the Arcturus during rain storms. Five species—three petrels and two shearwaters—were pelagic, six were shore birds of Cocos, and two species were storm-driven strays (Beebe 1926a:327). He provided observations of many of them. At Station 74, Arcturus steamed in circles so investigators could tow large nets, during which circling Beebe (1926a:331–338) saw two green sea turtles, three great schools of dolphins, a school of small whales, and a school of 27 blackfish. Surface nets caught seeds and insects, which he itemized, and gave him a perspective on how the Galapagos Islands acquired its plants and animals. Surface plants and animals were 412 individuals of 34 species. Below the surface, he collected 136 species of fish and 50 species of crustaceans, and he gave longer descriptions of them, including depths, because unfamiliar. He was especially intrigued with luminous species, and his undersea experience in Galapagos waters helped him visualize what he found at greater depths. He concluded (Beebe 1926a:362) the following: …a generous majority of the deep sea fish are little more than living eating-machines, with every function subordinated to that of capture with the appalling rows of teeth, engulfment in the cavernous mouth, and finally reception and digestion in a stomach which is beyond belief elastic and distensible. In 1926, Beebe published two books, one on pheasants and the other, The Arcturus Adventure; he also published 11 articles (scientific and popular), eight book reviews, and eight brief notes (Berra 1977:63– 65). One article (Beebe 1926b) gave a more scientific account of the Arcturus expedition than did the book. Another article, “The Guianas,” lists Henry A. Gleason as co-author (Beebe and Gleason 1926). He may have contributed some of the discussion of plants and vegetation. This is an impressive article in the ESA’s Naturalist’s Guide to the Americas, which Victor Shelford, edited. On Thanksgiving Day, a brief article in the New York Times announced that “Beebe to Explore Ocean Bed in Tank” (Anonymous 1926). A reader was (Frederick) Otis Barton, Jr. (1899–1992), son of a wealthy family, who had an ambition similar to Beebe’s (Matsen 2005:3–12). He had grown up spending summers at Cotuit, on Vineyard Sound, Massachusetts, where the sea captured his imagination. At age 16, he saw in a catalog equipment for underwater salvage work: a metal helmet with hose and an underwater suit. He sketched a wooden helmet and took the sketch to a Boston cabinet maker who built it. With his brother manning the hose pump, he donned his helmet and climbed down a ladder at Cotuit pier. That contraption provided entertainment for the remaining summer. He had then studied engineering at Columbia and Harvard Universities. After graduating, he spent a year sailing around the world. In the Philippines, he met native divers who hunted fish and pearls, and he was dazzled by the colors of coral reefs and a ssociated animals. While at Columbia University Graduate School, he searched the library for information on undersea exploration, and he pondered the physics of constructing equipment for doing so. Having already explored the problem, he concluded that Beebe’s proposed steel cylinder was inadequate, and he decided they should collaborate. Beebe was so busy he never answered Barton’s letters, but finally, someone introduced them and Barton convinced him of the wisdom of collaboration. Barton explained that the only shape that could withstand

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Fig. 7. Luminous surface fish in daylight and darkness: eater of stars Astronestaahes in pursuit of lantern fish Myctophids. (A) Daylight; (B) darkness. Beebe (1926a: plate V facing 214).

the pressure of deepwater was a sphere, and he volunteered to arrange for its manufacture. The concept for what Beebe named “Bathysphere” seems simple, but the production of one that was usable and effective became very complex and took several years (Matsen 2005:65–76). It absorbed Barton’s thoughts and fortune, but Beebe carried on his frenetic career until the Bathysphere was ready. In 1927, he found another ship, the Lieutenant, to take him and his staff to Haiti, mid-January to mid-May. Why Haiti? Neither biographer knew, but Gould (2004:266) guessed that “the 382 Bulletin of the Ecological Society of America, 97(4)

Fig. 8. William Beebe and Otis Barton with the Bathysphere. Wikipedia.

colonial governor, Sir John Russell, was eager to add to the knowledge of the colony’s resources.” Haiti had been independent since its slaves expelled their French masters in 1794. An American diplomat rolled out the welcome mat for Beebe and staff. However, they lived in tents on the deck of their schooner. Beebe’s own casual clues about why Haiti were as follows: “There is not a single list of Haitian fresh-water or marine fish” and “Fish are the dominant reason for my choosing this part of the world for a winter of work” (Beebe 1928:11, 7). This was Beebe’s last expedition using his underwater helmet. He was better prepared than on the Arcturus expedition. He had obtained help in developing a waterproof case for a motion picture camera underwater, to show life on a coral reef, and Bell Telephone had developed (gratis) an underwater phone with which he could communicate with his handlers aboard ship. He was not first to develop a method of underwater photography, but there had not been any scientific documentation in previous efforts (Matsen 2005:164–166, 258). Beebe returned to a discussion of helmet descents in a later article (1932a) and in his Zaca Adventure (1938) to the Sea of Cortez. Very common in Port-au-Prince Bay were small thimble jellyfish Linuche unguicalata, some of which Beebe scooped up and put in an aquarium containing small fish (Beebe 1928:20–23). The jellyfish tentacles captured the fish and consumed the bodies but not the heads or tails. On another occasion (1928:29), when he was preparing to don his helmet, an immense jellyfish floated by and someone scooped it up and flipped it upside down into a pail. Hovering under its body were about 350 small fish (1.5–2 inches long) which used the jellyfish as a safe haven. The jellyfish was [supposedly] C hiropsalmus quadrumanus; the fish were Choroscombrus chrysurus. He included in Beneath Tropic Seas (1928: facing 4) photographs

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of this jellyfish from the side, showing its size, and from the bottom, showing the sheltered fish. Yet, he did not ponder the contradictions between these stories, told in the same chapter: why one (small) species of jellyfish ate the small fish and the other (large) species, instead, sheltered them. He was a close observer, but instead of analysis, he went on to the next adventure. Beebe liked to take things apart to see what he could learn. An example was a black sponge that grew at depths of 20–30 feet (no scientific name given in Beneath Tropical Seas). When he sliced it into chunks, he found living in its cavities many shrimp, small crabs, and slim gobies (Beebe 1928:124–133). When he squeezed the sponge, the shrimp clicked their claws, and some came out to attack him. He then became fascinated with the shrimp Alpheus, as its responses were more rapid and interesting than those of the sponge. Beebe and his closest assistant colleague, John Tee-Van, assembled a collection of Haitian fish, and a book appendix (Beebe 1928:225–227) gave family names, common names, and numbers of species in each family; there were 277 species, which are not named in the book, but are in their technical report, which included illustrations of each species (Beebe and Tee-Van 1928). Seven years later they published a list of additions to their list (Beebe and Tee-Van 1935). Beebe’s first wife had left him before he became a celebrity, and in 1927, after he was one, he married a fashionable novelist half his age, whose pen name was Elswyth Thane (Gould 2004:262–268). Thane disliked tropical heat, and in spring 1929, the Beebes and two assistants went to balmy Bermuda to see if it might be suitable for marine research. They were entertained by Governor Louis Bols, whose other guest at the time was Prince George from England. During a series of parties, Beebe captivated everyone with stories of underwater exploration, and Prince George had no trouble persuading Beebe to take him down to the sea floor for his own adventure, which was quite successful (Matsen 2005:52). The governor took everyone on a tour of a small island a short distance from Bermuda, Nonsuch. It was attractive, with a large vacant building which Bols offered to Beebe to use as a research station. Bols’s generosity extended to also providing a tugboat, Gladisfen, and some motorboats (Gould 2004:272). Beebe gratefully accepted and then returned to New York City to raise funds to equip Nonsuch as a marine laboratory (Matsen 2005:53–54). Many patrons contributed to the project. Beebe bought what he needed, recruited a staff, and returned to Nonsuch. Although other researches also occurred at Nonsuch, it became the launching site for the Bathysphere, which stole the show. The Bathysphere was sent down empty on test runs, and minor problems were fixed before Beebe and Barton descended. Their first descent was on 6 June 1930, and their deepest descent in 1930 was to 1,426 feet; they had sent it down empty to 2000 feet deep (Welker 1975:127, Gould 2004:290). Beebe made arrangement with the National Geographic Society to produce an article with his narrative and photographs and colored illustrations by Else Bostelmann. As that article (Beebe 1931) was a success, three more followed (Beebe 1932a, b, 1934a). In August 1932, he arranged with the new National Broadcasting Company to have a live hour broadcast of their descent in the Bathysphere, which was broadcast throughout the United States and England to about 14 million listeners. There was no article in 1933, due to the Depression. These articles are more detailed and better illustrated accounts than his later book, Half Mile Down (Beebe 1934b, Second edition, 1951), although the book is well illustrated with drawing and photographs and eight of Bostelmann’s paintings in color. 384 Bulletin of the Ecological Society of America, 97(4)

Was Beebe the first discoverer of symbiotic fish cleaning? It would have been difficult for anyone not observing from below the surface to detect it. He was the earliest source known to Feder (1966:329), “Cleaning Symbiosis in the Marine Environment”, who cited Beebe’s (1928) Beneath Tropic Seas, but without citing a page, and I did not find an account in that book. In 1931, Beebe (1931:676) called fish cleaning “an amazing exchange of courtesy”, and later he generalized (Beebe 1951:142–143) the following: …an interesting exchange of courtesy, one which I have observed many times when diving near shore. The giant caerulean parrotfish browse on hard coral as a horse tears off mouthfuls of grass. After an interval of feeding, when the teeth and jaws and scales of the head are covered with débris, the fish upends in mid-water and holds itself motionless while a school of passing wrasse, all tiny in comparison with the big fish, rush from all sides and begin a systematic cleaning of the large fish’s head. As in most relationships between different species of animals, this is founded on mutual benefit, the parrotfish getting a free cleaning, and the wrasse finding a supply of particles of food ready at hand (Plate III). In 1923, he had observed a crab remove three ticks from a marine iguana. That differed from fish cleaning in that the iguana did not solicit the crab’s cleaning, but merely tolerated it. Both were interesting symbiotic relationships worthy of comment, but Beebe seems not to have noticed the similarity. National Geographic articles brought Beebe’s adventures to a broader audience than he reached with his many books and articles. As the National Geographic articles were all successful, why did he go on to other projects after the fourth article and stop using the Bathysphere? It was a matter of diminishing returns. The Bathysphere enabled him to see animals alive that he otherwise knew only as specimens dredged from a ship. His most important ecological discovery was probably the symbiosis between parrotfish and wrasse, which at the time would not have been obvious otherwise. The marine adventures, especially with the Bathysphere, were his most famous achievements (Gold and Warsaw 1980). After 1934, however, he returned to the coastal adventures described in Galápagos (1924) and Arcturus (1926), with two voyages of the Zaca, in 1936 into the Gulf of California, described in Zaca Adventure (1938) and in 1937–1938 to the Pacific coast stretching from Guadalupe Island, off of Baja California, south to Panama and Gorgona Island, off of Columbia, described in Book of Bays (1942). An interesting appendix in Book of Bays lists scientific articles published after the 1936 voyage which provided technical support of the narrative in Zaca Adventure. All 17 of the articles appeared in the New York Zoological Society’s Zoologica. Barton (1953) published his own memoir, The World Beneath the Sea, in which he told his side of the story of their partnering. He also used the Bathysphere to make undersea motion pictures, and later, he developed a more advanced undersea vehicle than the Bathysphere, which he called a Benthoscope. His book has three photographs of it. He did not abandon undersea exploration when Beebe did. Henry Bigelow Henry Bryant Bigelow (1879–1967) was son of a Boston banker and amateur naturalist (Bigelow 1964, Redfield 1976, Shor 1988, Brosco 1989, Lewandowski 1999). Henry went to Milton Academy and then spent a year at the Boston Natural History Museum, studying under Alpheus Hyatt. He was

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an avid hunter and fisherman. He earned three degrees from Harvard University (B.A., 1901; M.A., 1904; and Ph.D. 1906). During his senior year, he heard that Alexander Agassiz was preparing to lead an expedition to Maldive Islands, in the Indian Ocean. He introduced himself to Agassiz and asked permission to go along. Permission granted (Bigelow 1964:9, Schlee 1973:254–255). They sailed to Italy, left Genoa on 26 November 1901, and when they reached Colombo, Ceylon (Sri Lanka), on 16 December, they took charge of the Amra, with crew, which Agassiz had reserved beforehand (Agassiz 1913:384–385). Agassiz assigned jellyfish to Bigelow, to preserve and describe, and Bigelow (1904) obliged with a substantial article. Thus, began Bigelow’s substantial work as a taxonomist. After the Maldives expedition, Bigelow returned to Harvard; yet, for six months in 1904–1905, he accompanied Agassiz in exploring the Galapagos Islands, Easter Island, and Mexico’s west coast. He worked under Alexander Agassiz, 1901–1910, and after Agassiz died, Bigelow (1964:23) was at “loose ends” until John Murray visited the MCZ about 1911 and suggested that Bigelow study the oceanography of the Gulf of Maine, which stretches 200 miles between Cape Cod and Newfoundland. Coincidentally, the U.S. Bureau of Fisheries learned in 1912 that commercial fisheries in the Gulf of Maine were declining and was receptive to Bigelow’s proposal for research funds to study the area (Brosco 1989:245). In 1912, he used a small vessel from the U. S. Bureau of Fisheries at Woods Hole, Grampus, and began his studies. He was a diligent reader of ICES publications and knew about area studies being conducted by European oceanographers and marine ecologists in the North Sea, with the same motivation to understand fish biology and revive sagging fisheries. He had the knowledge, skill, and determination for the task, and his efforts culminated in three large volumes, 1925–1927, on fishes, plankton, and physical oceanography. The “Fishes of the Gulf of Maine” was co-authored by Dr. William W. Welsh (1850–1924), from the U.S. Bureau of Fisheries. Bigelow indicated (page 7) that most of the fish volume was by Welsh, who died before their monograph was completed (Bigelow and Welsh 1925). Fishes of the Gulf of Maine is well illustrated, mostly with previously published drawings. Ecologically valuable was information on habits and food, which were well known for most commercial species, but less so for non-commercial species. The plankton volume is well illustrated with maps and microscopic photographs of plankton. Revelle (1980:12) characterized these three volumes as “the first genuinely oceanographic study to be completed in North America”. Bigelow’s volumes made the Gulf of Maine “the best-known body of water in the world, oceanographically speaking” (Schlee 1978:78). Bigelow’s (1928) article, “Exploration of the Waters of the Gulf of Maine”, summarized his findings on temperature, salinity, and currents, but did not include biology. In 1924, the USBF vessel, Albatross, needed repairs which USBF could not afford, and it had no other for Bigelow’s use. He reverted to museum work until another opportunity arose. In 1923, he had become a member and was mostly chairman, of the American Commission of Fisheries Investigations until 1932. Those investigations paralleled those of ICES, and he was invited to attend ICES meetings and did, 1931–1940 (Bigelow 1964:30, Lucas 1980:502–506). Bigelow was asked by the National Academy of Sciences’ Committee on Oceanography in 1927 to write a report on U.S. contributions to oceanography (Bigelow 1927). That NAS committee had been established at the suggestion of Frank Lillie, Director of the Woods Hole Marine Biological Laboratory. Bigelow spent a year investigating and writing a report. He emphasized (1928) that America needed a program of environmental and biological marine investigations on the eastern coast equaling the Scripps Institution on the western coast. NAS sent it to the Rockefeller Institute that gave $2.5 million to build an 386 Bulletin of the Ecological Society of America, 97(4)

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Fig. 9. (a) Henry Bryant Bigelow at the helm of Grampus. Bigelow (1964: facing 22) and at web site. (b) Bigelow as administrator. Courtesy Woods Hole Oceanographic Institution.

oceanographic institute and $50,000 a year for ten years to run it. Bigelow was chosen as first director of the Woods Hole Oceanographic Institution (WHOI). He found a site in Woods Hole, build an institute, ordered a ship which M. I. T. Professor George Owen designed, and staffed both WHOI and its ship. Bigelow had a steel sailing vessel built in Copenhagen, and he appointed a former student (and in 1939 his successor as director at WHOI) Columbus O. Iselin (1904–2012) as captain of the 142-foot Atlantis (Schlee 1978:16–18, 1980; Rozadowski 2011:169-170). WHOI opened in 1931. Bigelow’s Gulf of Maine studies and NAS report got him elected to the National Academy of Sciences in 1931. His report became the basis for his published summary, “A Developing View-Point in Oceanography” (1930) and his book, Oceanography: its scope, problems and economic importance (1931). The book met a general need for an overview of American oceanography. Much of it was either based on his oceanographic work or was a wish list of what needed to be done. The most interesting part was his discussion of sea fisheries, because there were data available and he was skilled at seeing and explaining their implications for the future (Schroeder 1960). After presenting data on the extent of American and Canadian fisheries, he commented that “An industry of this magnitude deserves the most intelligent management possible” (1931:190). At the time when WHOI was being established, it was reasonable to expect that rational management was likely to follow. Bigelow remained director of WHOI for a decade, afterward became president of its board of trustees for another decade, and then chairman of the board for a third decade. In 1929, when Walter Fisher, Director of the Hopkins Marine Station, invited him to teach a marine biology course there, he went, and while there studied the temperatures and salinities of Monterey Bay from surface down to 600 m. In 1946, the U.S. Navy’s Hydrographic Office asked him to write a textbook on waves for naval officers. He could easily have said that this subject was beyond his expertise, but instead, he said he would need a co-author

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Fig. 10. Zooplankton near Georges Bank. Bigelow (1926: fig. 26, facing 29).

to handle aspects beyond his expertise (Bigelow 1964:37). The resulting Wind Waves at Sea, Breakers and Surf (Bigelow and Edmondson 1947) was obviously quite competent: Soviets published a Russian translation in 1951. Bigelow, the esteemed administrator, was also a conventional scientist. Gordon Riley discovered this when he asked Bigelow for research time on Atlantis (1979:74, quoted from Mills 1989:272–273): Henry and I [became] good friends, but we did not see eye to eye. I was very much under the influence of the oceanographers at the Plymouth Laboratory—Atkins and Harvey and the others— who were trying to develop quantitative measurements of plankton and ecological variables and to analyze seasonal cycles and regional variations in a quantitative way. To Bigelow this was a profound and distasteful oversimplification. This exchange of opinions reminds us of the Hensen–Haeckel conflict (Egerton 2013:236), with Riley defending a “Hensen-like” perspective and Bigelow defending a “Haeckel-like” perspective. 388 Bulletin of the Ecological Society of America, 97(4)

Fig. 11. Map of the Gulf of Maine. From New England Coastal Wildlife Alliance (NOAA).

Bigelow advanced both at the MCZ and at Harvard University, becoming a professor of zoology in 1931, and made Alexander Agassiz Professor of Zoology in 1944 (Bigelow 1964:33). He thought he set a longevity record as an officer of Harvard University for 55 years, 1905–1960 (1964:41). He won many awards and made professional friends readily. Alfred Redfield considered him “shrewd, wise, well-informed, a pragmatic realist, widely traveled, and the best informed naturalist that he had ever met” (quoted in Fye 1980:2, see also Redfield 1976).

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He was first recipient of The Henry Bryant Bigelow Medal in Oceanography (Anonymous 1961), awarded about the time of publication of Papers in Marine Biology and Oceanography Dedicated to Henry Bryant Bigelow by His Former Students and Associates on the Occasion of the Twenty-fifth Anniversary of the Founding of the Woods Hole Oceanographic Institution, 1955 (Graham et al. 1955). This volume of xx + 498 pages is a bibliographic orphan: no editor listed. It is listed here under Graham, as he is the first of 56 authors represented. No article was about Bigelow, but it has his bibliography (pages xviii–xx; more extensively in Redfield 1976:70–80). An environmentalist’s recent evaluation of him sees his science and sport fishing as complementary sources of knowledge (Dobbs 2000:8): …the scientist who merged the perspectives of fishermen and scientists most successfully,[was] an avid angler, sailor, zoologist, and oceanographer named Henry Bryant Bigelow. The most accomplished oceanographer and fishery scientist of his generation, Bigelow pioneered the scientific exploration of the Gulf of Maine in the early twentieth century. He came to know the Gulf perhaps as well as anyone has ever known any large natural system, and he did so largely by blending the mental habits and knowledge of the scientist and fisherman. This integration made his work, particularly his comprehensive field guide Fishes of the Gulf of Maine, so informative and inspiring that even today he is admired unreservedly by both sides. Oceanus is a popular magazine which WHOI publishes, and volume 14, issue 2 (July 1968) was named H. B. Bigelow volume and included remembrances and photographs by 18 authors, plus (some of) his bibliography, pages 29–32. The WHOI became a leading world oceanographic center and remains one: See retrospective talks at its fiftieth anniversary (Burstyn 1980, Deacon 1980, Haedrich and Emery 1980, Revelle 1980, Schlee 1980). Edward Ricketts Edward Flanders Ricketts (1897–1948) was from Chicago and had an early interest in nature (Ward 1974:195–211, Tamm 2004:7–9, Lannoo 2010). He attended the University of Chicago and was strongly influenced by a course in animal ecology under Prof. Warder Clyde Allee, who studied and published on the ecology of social animals (Burgess 1996:11). Like Beebe, Ricketts only finished three university years and left without a degree. He married and in 1923 moved his wife and son to Monterey, near Stanford University’s Hopkins Marine Station. Ricketts established a supply business, Pacific Biological Laboratories, Inc., based on specimens he and others collected along shore. In 1958, Monterey named a street Cannery Row, to honor John Steinbeck’s novel, set at Ricketts’ laboratory and the canneries. Steinbeck (1951:xiii–xiv) described Ricketts’ laboratory, between two canneries that canned sardines. Ricketts was gregarious. One friend was Jack Calvin, a teacher attracted to the sea. Calvin and wife Sasha travelled for 53 summer days in 1931 the Inside Passage from Tacoma to Juneau in a canoe with sail, Nakwasina (Calvin 1933). In June–July of 1932, he took Ricketts, Sasha, Sasha’s sister Xenia, and Joseph Campbell to Alaska on Calvin’s boat, Grampus (Tamm 2004:181–187).


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Fig. 12. (a) “Cannery Row” in the 1930s, as envisioned by Bruce Ariss (1984). Ricketts’ Pacific Biological Laboratories is in the center, squeezed between two canneries, with Ricketts at the head of his stairs, waving. Courtesy Monterey Bay Aquarium. (b) Map of Cannery Row and Pacific Grove. Courtesy E. D. Stephan.

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Most prominent Ricketts friends were novelist John Steinbeck (1902–1968) and mythology scholar Joseph Campbell (1904–1987). Ricketts had a strong influence on their thinking; indeed, he was the strongest influence on Steinbeck (Steinbeck 1951:x, Benson 1984:183). Ricketts was junior author of The Sea of Cortez (Steinbeck and Ricketts 1941), with Steinbeck as senior author (Shillinglaw and Gilly 2007). Steinbeck’s (1945) novel, Cannery Row, featured Ricketts as a main character, “Doc.” Before those books appeared, Ricketts turned his natural history notes into Between Pacific Tides (Ricketts and Calvin 1939), assisted by co-author Calvin, who photographed many specimens. Other specimens were drawn by local artist Richie Lovejoy. Ricketts ranged more widely than just Monterey Bay, and so his book was a guide to species along much of coastal California, Oregon, and Washington. It explained environments of all animals studied, what each species ate, and other ecological aspects. He had been preceded in studies on stratification of species on beaches by English ecologist Coleman et al.’s (1933) “The Nature of Intertidal Zonation in Plants and Animals”, which Ricketts and Calvin (1939:304), but whether Coleman’s article inspired Ricketts or was discovered in a literature search after he had begun his studies is unclear. Their original manuscript was sent to Stanford University Press in 1930, but changes were made before publication in 1939. Between Pacific Tides was Ricketts’ masterpiece. Other books on sea and shore animals organized descriptions according to zoological classification. Ricketts realized that no one on shore sees animals that way, and they want to know some of the natural histories of species encountered. He organized his accounts according to microenvironments in which animals lived: rocky shores, subdivided into exposed rock, protected rock, and rockweed, pool; bays and estuaries, subdivided into rocky shores, sand flats, eelgrass, mud flats; and sandy beaches required no subdivisions. Illustrations followed the same organization and had an unusual feature—mostly, drawings are on a left plate and photographs on a right (a)

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Fig. 13. (a) Edward Flanders Ricketts, explorer and author. Ricketts, Calvin, and Hedgpeth (1968:viii). (b) Ricketts as biological specimen supplier. Courtesy Pacific Biological Laboratory.


plate. Drawings are of different subjects than photographs and are of animals more easily displayed in drawings than photographs. The First edition (1939) had 320 pages, the Second edition (1948), which Ricketts revised but did not live to publish, had 365 pages; it acknowledged Lovejoy’s drawings on title page and carried color photograph of chitons as frontispiece. Although a few of Ricketts’ comments made Buchsbaum (1940) uncomfortable, he gave it a friendly review in Ecology.

Fig. 14. Chitons. Scientific names were listed on an accompanying transparent sheet which did not detract from this frontispiece. Ricketts and Calvin (1948).

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Accounts of individual species were based on both Ricketts’ observations and other cited sources. The final organisms discussed were plankton—plant and animal—with discussion of fluctuations in abundance and environmental factors that seemed important (using 17 charts). That section was immediately followed by a plankton bibliography. The remainder of the bibliography is systematically organized according to classification, because that is how most of the literature was organized by the authors. Finally, a brief separate bibliography listed works not limited to specific species. An interesting feature of some bibliographic citations was an annotation explaining content more fully than did titles and information correspondents had sent him. Between Pacific Tides became the basis of a mutual admiration professional relationship between Ricketts and British seashore ecologist Alan Stephenson (Tamm 2004:246–247). Stephenson had participated in a British Great Barrier Reef Expedition, 1928–1929, and later, he had spent a decade conducting an ecological survey of South Africa’s coast. Eventually, Stephenson and his wife, Ann, published a similar book, Life Between Tidemarks on Rocky Shores (Stephenson and Stephenson 1972), with worldwide coverage. Ricketts had long admired Stephenson’s published papers, and when the Stephensons visited Ricketts in 1947, he told Ricketts that he so admired Between Pacific Tides that he had read “every word” (quoted by Tamm 2004:247). To the uninitiated, that would seem to be like reading an encyclopedia, but Stephenson was so familiar with Ricketts’ subject that every line was potentially new information or new interpretation. In return, Ricketts judged Stephenson to be “one of the world’s greatest zoologists, probably the greatest ecologist” (in letter to Steinbeck, 22 Sept. 47, quoted by Tamm 2004:247). Ricketts, of course, was expressing the judgment of a seashore specialist, who had no ability to judge Stephenson in relation to ecologists in other specialties. The Sea of Cortez (Steinbeck and Ricketts 1941) was a kind of southern extension of Between Pacific Tides into the Gulf of California, with Steinbeck writing a 271-page narrative of their expedition, based on Ricketts’ notes, with Ricketts writing natural histories of specimens they obtained and an annotated bibliography similar to that in Between Pacific Tides, pages 272–589, and an index covering only Steinbeck’s part, pages 590–598. The illustrations are similar to Between Pacific Tides, except that there are eight plates of color photographs of mollusk shells, and drawings are separate from photographs. The Sea of Cortez, excepting a few quibbles, received a warm review in Ecology by Schmidt (1942), at the Field Museum of Natural History. Ricketts and Steinbeck’s expedition was an isolated exploration of the Gulf of California, isolated not only in the sense of they being the only explorers there in 1940, but isolated also in not being preceded or succeeded by other biology surveys until 2004 (Sagarin et al. 2008:372). Dungan et al. (1982) reported on a “Catastrophic Decline of a Top Carnivore in the Gulf of California Rocky Intertidal Zone,” in which they cited Steinbeck and Ricketts’ (1941:378) account of the sun star Heliaster kubiniji: “The most common, obvious, and widely distributed shore starfish in the Gulf”. That prevalence had ended by 1978, which they attributed to disease and temperature increases in 1977–1978. Dungan’s group studied only one species. The 2004 Sea of Cortez Expedition and Education Project retraced the sampling sites that Ricketts had used in 1940, took comparable samples and compared the results from the two expeditions. The 2004 expedition found (Sagarin et al. 2008:372): Diversity and abundance of large gastropod snails and echinoderms have declined at many intertidal sites and large pelagic species of tuna, sharks, billfish, and turtles also appear to be 394 Bulletin of the Ecological Society of America, 97(4)

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Fig. 15. (a) Gulf of California showing collection sites, 1940 and 2004. Gray circles indicate Ricketts’ sites not certainly located. (b) Number of echinoderm species observed at 12 sites sampled, 1940 and 2004 (below zero line is 1940). Sagarin et al. (2008:374 and 375).

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Fig. 15. Continued.

much less abundant. However, vermetid gastropods (tube snails) appear to be more abundant and widespread, and jumbo squid, not documented at all by Steinbeck and Ricketts, are currently very common in the Sea of Cortez and constitute a major fishery. Ricketts’ Gulf survey achieved in 2004 the importance he had hope for. Steinbeck enjoyed Ricketts’ blend of science and philosophy and was an equal participant. Their discussion at times was convoluted, as in this example (Steinbeck and Ricketts 1941:257): And in a unified-field hypothesis, or in life, which is a unified field of reality, everything is an index of everything else. And the truth of mind and the way mind is must be an index of things, the way things are, however much one may stand against the other as an index of the second or irregular order, rather than as a harmonic or first-order index. These two types of indices may be compared to the two types of waves, for indices are symbols as primitive as waves. This discussion has not yet been incorporated into philosophy textbooks. After Ricketts died, Steinbeck (1951) wrote an essay, “About Ed Ricketts,” in which he stated, “We worked and thought together very closely for a number of years so that I grew to depend on his knowledge and on his patience in research” (Steinbeck and Wallsten 1975:315). Ricketts saw how many sardines were being harvested and canned and warned in a newspaper article that the high level could not be sustained (Ricketts 1947, Tamm 2004:81–82). He lacked the authority to be taken seriously, and the sardine fishery collapsed (with partial revival in 1986: Gore 1990:8, Palumbi and Sotka 2011:120–122). He was worrying about the collapse in 1948, around the time he died after a train accident (Tamm 2004:266–267, 279–280). Although he never achieved Beebe’s eminence, he was more theoretically oriented than Beebe, and his theoretical ideas were as much philosophy as science. He accumulated observations and notes on the natural history of animals he collected, his curiosity guided somewhat by Allee’s (1931) Animal Aggregations. 396 Bulletin of the Ecological Society of America, 97(4)

Ricketts also made three expeditions along Canada’s coast to write a third volume on Pacific shore animals of Canada’s and some of Alaska’s Pacific coast. His first, on Jack Calvin’s boat, Grampus, travelled the inner passage to southern Alaska (1932); his other two trips (1945, 1946), along the west coast of Vancouver Island, reached Queen Charlotte Island. He accumulated notes and specimens for a book, but died after his car collided with a train in 1948. When Steinbeck received word in New York that Ricketts was dying, he lamented: “The greatest man in the world is dying and there is nothing I can do” (Benson 1984:615). Steinbeck reached Cannery Row in time for the funeral. In his will, he left his notebooks to Hopkins Marine Station; Steinbeck purged their non-biological content before turning them over (Steinbeck 1951:lii). Ricketts’ notes and other writings for a third volume are now published (Hedgpeth 1978, Rodger 2002, 2006). Ricketts’s lifetime project of documenting Pacific coastal invertebrates was a first installment in a series of works by many authors which culminated in Pacific Coast (Ricketts et al. 1985), edited by Bayard and Evelyn McConnaughey. Their bibliography (McConnaughey and McConnaughey 1985:610) lists Ricketts and Calvin’s Between Pacific Tides, Fourth edition. Katharine Rodger began publishing Ricketts’ posthumous works in 2002. Since then, Michael Lannoo (2010) has taken a closer look at his philosophical writings. These writings are interesting to historians of ecology; are they of more than historical interest to ecologists? They are to marine ecologist Rafe Sagarin, who admires Ricketts’ ecological philosophy as well as his data (Sagarin and Crowder 2009, Sagarin and Pauchard 2012:199). Ricketts’ book organization became popular, as seen in Herbert Zinn and Lester Ingle, Sea Shores: A Guide to Animals and Plants along the Beaches (1955), which includes aquatic species living near shore. Pacific coast zoology, of course, occurred within a larger zoological context. Libbie Hyman (1888– 1969) wrote an astonishing encyclopedia, The Invertebrates (Hyman 1940–1967, 6 volumes), perhaps surpassing Hutchinson’s (1957–1993) 4-volume Treatise on Limnology as a solo achievement (she did not have a professor’s responsibilities, as he did). Richard and Gary Brusca wrote a one-volume encyclopedia, Invertebrates (Brusca and Brusca 1990). A later specimen collector, Jack Rudloe (b. 1943), located in Florida, has written autobiographical books, the first being The Sea Brings Forth (Rudloe 1968). An early mentor of his was John Steinbeck. Acknowledgments I thank Professor Emeritus Eric L. Mills, Dalhousie University, for his critique of an early version of this survey; Edward A. Johnson, for help in obtaining funds for a student; Alejandro Quiroz, in assisting me with permissions for reproducing illustrations; Laura Mason, for reproducing illustrations; Thomas Anderson, for sending me a copy of Anderson and Gentleman on Ripley (cited in 58B); and Jerry Hershberger, for computer assistance. Literature Cited Agassiz, G. R., editor. 1913. Letters and recollections of Alexander Agassiz, with a sketch of his life and work. Houghton Mifflin, Boston, Massachusetts, USA. Allee, W. C. 1931. Animal aggregations: a study in general sociology. University of Chicago Press, Chicago, Illinois, USA.

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Bigelow, H. B. 1927 [for 1924]. Physical oceanography of the Gulf of Maine. United States Bureau of Fisheries Bulletin 40(Part 2):511–1027. Bigelow, H. B. 1928. Exploration of the waters of the Gulf of Maine. Geographical Review 18:232– 260. Bigelow, H. B. 1930. A developing view-point in oceanography. Science 71:84–89. Bigelow, H. B. 1931. Oceanography: its scope, problems and economic importance. Houghton Mifflin, Boston, Massachusetts, USA. Bigelow, H. B. 1964. Memories of a long and active life. Cosmos Press, Cambridge, Massachusetts, USA. Bigelow, H. B., and W. T. Edmondson. 1947. Wind waves at sea, breakers and surf. U.S. Navy Hydrographical Office Publication 602. Bigelow, H. B., and W. W. Welsh. 1925 [for 1924]. Fishes of the Gulf of Maine. United States Bureau of Fisheries Bulletin 40(Part 1):1–567. Bjerknes, J. 1969. Atmospheric teleconnections from the equatorial Pacific. Monthly Weather Review 97:163–172. Bridges, W. 1974. Gathering of animals: an unconventional history of the New York Zoological Society. Harper and Row, New York, New York, USA. Brosco, J. P. 1989. Henry Bryant Bigelow, the U.S. Bureau of Fisheries, and intensive area study. Social Studies of Science 19:239–264. Brusca, R. C., and G. J. Brusca. 1990. Invertebrates. Sinauer Associates, Sunderland, Massachusetts, USA. Buchsbaum, R. 1940. Review: E. Ricketts and J. Calvin, Between Pacific Tides (1939). Ecology 21:93– 94. Burgess, R. L. 1996. American ecologists: a biographical bibliography. Huntia 10:5–116. Burstyn, H. L. 1980. Reviving American Oceanography: Frank Lillie, Wickliffe Rose, and the founding of the Woods Hole Oceanographic Institution. Pages 57–66 in M. Sears and D. Merriman, editors. Oceanography: the past. Springer-Verlag, New York, New York, USA. Calvin, J. July 1933. Nakwasina goes north: a man, a woman, and a pup cruise from Tacoma to Juneau in a 17-foot canoe. National Geographic 64:1–16. Coleman, J. 1933. The nature of the intertidal zonation of plants and animals. Marine Biological Association of the United Kingdom Journal 18:435–476. Cited from Ricketts et al. 1985:597. Darwin, C. R. 1844. Geological observations on the volcanic islands: being the second part of the geology of the voyage of the Beagle. Smith, Elder, London, UK. Deacon, G. E. R. 1980. The Woods Hole Oceanographic Institution: an expanding influence. Pages 25–31 in M. Sears and D. Merriman, editors. Oceanography: the past. Springer-Verlag, New York, New York, USA. Dobbs, D. 2000. The great gulf fishermen, scientists, and the struggle to revive the world’s greatest fishery. Island Press/Shearwater Books, Washington, D.C., USA. Dungan, M. L., T. E. Miller, and D. A. Thomson. 1982. Catastrophic decline of a top carnivore in the Gulf of California rocky intertidal zone. Science 216:989–991. Egerton, F. N. 2010. History of ecological sciences, part 35: the beginnings of British marine biology: Edward Forbes and Philip Gosse. ESA Bulletin 91:176–201. Egerton, F. N. 2013. History of ecological sciences, part 47: Ernst Haeckel’s ecology. ESA Bulletin 94:222–244. Egerton, F. N. 2014. History of ecological sciences, part 51: formalizing oceanography and marine ecology. ESA Bulletin 95:347–430.

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