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History of Ecological Sciences, Part 58B: Marine Ecology, mid‐1920s to about 1990: Carson, Riley, Cousteau, and Clark

2017; Ecological Society of America; Volume: 98; Issue: 2 Linguagem: Inglês

10.1002/bes2.1314

2327-6096

Frank N. Egerton,

Isotope Analysis in Ecology

Part 58A has a general bibliographic introduction for parts A and B (Egerton 2016b). I am sure that my own style and thought were deeply influenced, in certain critical years, by Henry Williamson, whose Tarka the Otter and Salar the Salmon are, I'm convinced, nature writing of the highest order. However, Carson only discovered Tarka (1927) by Englishman Henry Williamson (1895–1977) when she was in graduate school at Johns Hopkins University and Salar (1935) after graduation (Brooks 1972:5–6, Souder 2012:59–63). Rachel went to local schools until her last 2 years in high school, when she took a trolley to a school in Parnassus. She was always an exceptional student, and she received a scholarship to attend the Pennsylvania College for Women (later Chatham College) in Pittsburgh. There, she became closely attached to two faculty members, Grace Croff, who taught English, and Mary Skinker, who taught biology. She was conflicted about which subject in which to major, but biology won. After Carson's third year, both teachers left the college, and a year later Carson followed in Skinker's footsteps to both the Marine Biology Laboratory at Woods Hole, Massachusetts, and Johns Hopkins University in Baltimore for a master's degree in biology. Three of the six biologists on Johns Hopkins faculty were members of the Ecological Society of America and published articles in Ecology (Dritschilo 2006:357). At Woods Hole, Carson went out on the U.S. Bureau of Fisheries ship Albatross II for deep sea collecting of specimens. Skinker became a parasitologist at the United States Department of Agriculture, and she urged Carson to take civil service exams to qualify for a government position as a zoologist (Lear 1997:79). Carson took the exams in 1935, and when she applied for a job at the Bureau of Fisheries in Washington, Elmer Higgins, head of the Division of Scientific Inquiry, hired her part time to write the scripts for 52 radio programs (7 minutes each), entitled "Romance Under the Waters" which he had agreed to produce (Lear 1997:78, Souder 2012:52–53). He liked what she wrote. She also wrote feature articles for the Baltimore Sun Sunday magazine, based partly on her research for radio scripts—on Chesapeake Bay fisheries and related subjects. In April 1936, Higgins asked her to write an introduction for a brochure on fisheries, and when he read "The World of Waters," he commented that this essay should be sent to the Atlantic Monthly, and she could write something simpler for the brochure (Lear 1997:81). She did as advised, and it was published as "Undersea" (Carson 1937). Later she commented: From that essay, "everything else followed" (quoted by Lear in Carson 1998:3). Higgins hired her full time in August 1936. One of her newspaper essays was "Chesapeake Eels Seek the Sargasso Sea" (1938). She had first encountered eels at Woods Hole in 1929, and she had experimented at Johns Hopkins on the effects of changes in salinity on eel behavior (Lear in Carson 1998:15, Souder 2012:49–54). The mystery of eel migrations from Europe and America to reproduce in the Sargasso Sea had been solved by Danish biologist Johannes Schmidt (1877–1933) in the first two decades of the 1900s (Schmidt 1912, 1922, Jordan and Evermann 1923:76–80 + plate, Regan 1933, Spjeldnaes 1975). Carson was popularizing recent history of science. Wikipedia. Many of the fishes, as well as the bottom-dwelling mollusks and worms and starfish, begin life as temporary members of this roving company, for the ocean cradles their young in its surface waters. The sea is not a solicitous foster mother. The delicate eggs and fragile larvae are buffeted by storms raging across the open ocean and preyed upon by diminutive monsters, the hungry glassworms and comb jellies of the plankton. This essay "established her unique voice, scientifically accurate and clear, yet poetic and imaginative, capturing the wonder of nature's cycles and relationships" (Lear 1997:88). An editor at Simon and Schuster wrote, asking if she was writing a book. She had not thought of that, but liked the idea. She met with him and one of his authors, ocean traveler Hendrik van Loon, and they worked out a plan which included selling chapters to Atlantic or similar magazines when written. It was at this point that Williamson's Salar the Salmon became influential. Under the Sea-Wind (1941) was based upon her own investigations and library research (Souder 2012:89–95). Sea-Wind had no bibliography, but she listed in her "Foreword" (1941:xviii–xix) the main works and authorities on which she drew. She decided to tell three life histories, in three "books": sanderlings, mackerel, and eels (subjects of her radio scripts and newspaper articles). Sea-Wind was a complex "novel," with main characters and plots, but also other characters and subplots. Sanderlings nest near freshwater lakes in the Arctic and later migrate to southern South American beaches, stopping along Atlantic beaches going both ways. Foxes and Jaegers menace nests with eggs and later the young chicks, which parents avoid by carrying away egg shells after chicks hatch and diverting foxes with broken-wing ploys to lure them away (Carson 1941:65). Parent sanderlings migrated south before the young, and the latter migrated instinctively, without guidance. Subplots tell the stories of eagles, plovers, and more. The sanderling story ends in chapter 3, and chapters 4 and 5 of Book I are mainly on mullet. The common mullet (Mugil cephalus) inhabits the coasts of southern Europe and northern Africa as well as American coasts from Cape Cod to Brazil (Jordan and Evermann 1923:252–253). Mullets spend summers in North Carolina sounds and estuaries, living singly, but in the fall they enter the ocean, gather into schools, and migrate southward to reproduce (Carson 1941:79–83). Fishermen await them with open nets. Ospreys also await them, but had a negligible impact on populations, compared to fishermen. Bald eagles robed ospreys when they saw opportunities. Not only do fishermen haul in many mullet, their nets scoop up small fish without commercial value which fishermen discarded on beaches (Carson 1941:100). Book II, on mackerel (Scomber scombrus), is misleadingly entitled "The Gull's Way." Mackerel, in deep water at the edge of the continental shelf, virtually hibernated during four cold months, living on fat stored during eight active months (Carson 1941:105–106). They lived on both sides of the Atlantic and ranged more northerly than mullet. In America, they lived from Straits of Belle Isle to Cape Hatteras, and in Europe from Norway to the Mediterranean and Adriatic Seas (Jordan and Evermann 1923:273–275). In spring, they rise toward the surface and swim toward shore, in such numbers that they brought to Carson's mind long gone flights of passenger pigeons (1941:111). She observed a migration of mackerel a mile wide and 20 miles long (Lytle 2007:45). Near shore, they release eggs and milt—a female sheds 40 to 50 thousand eggs—then all head for feeding grounds off New England. Their fertilized eggs become part of plankton, where vast numbers of them are needed for the species to survive, since so many are eaten by plankton consumers (Carson 1941:116–119). After about a week, eggs hatch, and when young gain independent motion, and are no longer part of plankton, they move down into deep waters during day and swim toward the surface at night. They then begin eating plankton. As they grow, they eat larger prey, but also become vulnerable to predators that feed on larger organisms than plankton, such as comb jellies (ctenophore). Growing mackerel cluster into schools, which provide some advantages, but also provide targets for predators, such as octopi (Carson 1941:146–148). The unusual life history of eels, which Carson had summarized in 1938, she elaborated in "Book III: River and Sea," in which she did not mention the Sargasso Sea (though mentioned on page xvi), where eels reproduce. There is some overlap between breeding grounds of the European (Anguilla anquilla) and American eels (Anguilla rostrata); but many American eels breed west and south of the Sargasso Sea (Zim and Shoemaker 1956:52). She emphasized the ignorance of zoologists about movements of adult eels from freshwater streams to breeding grounds (Carson 1941:231, 254–255), but the movement of the billions of larval eels to the rivers from which their parents had come was documented by Schmidt (Carson 1941:xviii). It takes a year for young to reach American rivers and 3 years for young to reach European rivers. Mature females swim up rivers and live 10 or 12 years before swimming downstream to meet males who had remained near the river mouth, in brackish water. Possibly, they all swim to the breeding grounds together, though the sexes might swim there separately. Eels are largely scavengers, but when they encounter live fish caught in a gill net, they gladly help themselves, often leaving little for the fisherman. Carson 1941: facing 144. Carson's Under the Sea-Wind (1941) was published on November 1 and received very positive reviews. However, Japanese forces snuffed out her limelight 36 days later by attacking Pearl Harbor, severely hurting sales and deeply disappointing Carson. A small consolation was Beebe's reprinting of her two chapters on eels as final selection in his The Book of Naturalists: An Anthology of the Best Natural History (1944), which began with a selection from Aristotle. According to an agreement with the State of North Carolina, two areas along the south shore of the lake have been set aside for managing hunting, where shooting is under the direct control of the State. Federal as well as State regulations are enforced. About 20 to 25 blinds are established, each in charge of a guide who is required to enforce hunting regulations. In Lear's edition of Carson's previously uncollected writings an abridged version of the Mattamuskeet pamphlet (lacking the above quotation). On this trip, Carson took along illustrator Kay Howe, whose seven illustrations add significant appeal to the pamphlet. Yet, Lear did not reprint any of them. (I worked at Mattamuskeet Wildlife Refuge in summer 1958, where I obtained a copy of Carson's Mattamuskeet [1947].) Her dozen pamphlets initiated a series of books on U.S. Wildlife Refuges (including Laycock 1965, Murphy 1968, Riley and Riley 1979, Grove 1984). Audubon 1840–1844. On a trip to Parker River Refuge, Massachusetts coast, Carson conferred with Harvard zoologist-oceanographer Dr. Henry B. Bigelow (1879–1967), whose Fishes of the Gulf of Maine (coauthor, W. Welsh, 1925) and Plankton of the…Gulf of Maine (1926) she had cited in Under the Sea-Wind. She mentioned her hope of writing another book on oceanography and he encouraged her to do so. She did extensive library research in order to explain recent progress in oceanography. She included a bibliography in The Sea Around Us (1951:217–220), which lists outstanding works. However, she decided that the public was uninformed about many advances made during and after World War II, and The Sea Around Us became a best seller, which none of the works she listed had achieved. Looking at her book today, that might seem surprising, because Oxford University Press produced a book that is physically unimpressive. The title page credited Katherine L. Howe for the drawings, which are only two, neither being especially noteworthy. Yet, in 1951 it was an impressive book, which won the National Book Award, and remained on the best-seller list 86 weeks. When Carson discussed plans for this book with Beebe, he had urged her to go to Bermuda and arrange to study marine life under water (Lear 1997:162–165). She was unable to do so; instead, she went to Miami Marine Laboratory and took a class under Dr. F. G. Walton-Smith. She once descended underwater wearing an 84-pound diving helmet, with poor visibility, in a current so strong she was afraid to let go the ladder; but she did see live coral and was grateful for the experience. She apparently never used Cousteau's aqualung. Carson feared that Oxford Press would produce a book that resembled an oceanography textbook, and although it did not, what she wrote was actually a popularized oceanography textbook, though readers might not realize it (Kroll 2008:95–123). Unlike Sea-Wind, The Sea Around Us is mostly about oceans, with marine life being a minor theme (though emphasized in this summary). Scientists thought that the earth was about 2.5 billion years old (Carson 1951:4)—around half the current estimate—but she cautioned that still older rocks than were then known might be found in the future. Life arose in the oceans, but no details were known in 1951. Soon, "some of the living things that floated in the sea must have developed the magic of chlorophyll" (Carson 1951:8). She explained that chlorophyll uses carbon dioxide from air and water from sea to "build the organic substances they needed for life." However, she failed to mention that plants then release oxygen into air. The theory of continental drift and plate tectonics was not widely accepted in 1951, and the current theory of mountain building was that as the earth cooled, its core shrank and its outer shell "fell into folds and wrinkles—the earth's first mountain ridges" (Carson 1951:9). One drawing Howe provided was apparently Carson's "Chart of the History of the Earth and Its Life," with representative fossils illustrated for each geological period. She explained what paleontological records revealed about the evolution of life. From the plankton the food chains lead on, to the schools of plankton-feeding fishes like the herring, menhaden, and mackerel; to the fish-eating fishes like the bluefish and tuna and sharks; to the pelagic squids that prey on fishes; to the great whales who, according to their species but not according to their size, may live on fishes, on shrimps, or on some of the smallest of the plankton creatures. Unmarked and trackless though it may seem to us, the surface of the ocean is divided into definite zones, and the pattern of the surface water controls the distribution of its life. Fishes and plankton, whales and squids, birds and turtles, all are linked by unbreakable ties to certain kinds of water—to warm water or cold water, to clear or turbid water, to water rich in phosphates or in silicates. For the animals higher in the food chains the ties are less direct, they are bound to water where their food is plentiful, and the food animals are there because the water conditions are right. Tropical botanist Ray Fosberg and geologist Harry Ladd advised her on formation of oceanic islands (Lear 1997:155), and that topic was subject of an outstanding chapter. Carson was mindful of Darwin's contributions to the subject and mentioned both his and Challenger naturalists' observations of life on apparently barren St. Paul's Rocks in the mid-Atlantic (Carson 1951:88). She was intrigued with ways in which plants and animals colonized distant islands. The volcanic island Krakatoa provided a natural opportunity to follow colonization, because its explosion in 1883 obliterated all life. It was within Dutch East Indies, and Dutch scientists followed its recolonization. For a quarter century, only a spider and a few blades of grass appeared, but by 1908 it was recolonized by birds, mammals, snakes, lizards, insects, mollusks, and earth worms. Ninety percent of arrivals were by air (Carson 1951:91–92). She told a sad story of how unique island species of plants and animals were quickly exterminated by invasive species which humans intentionally or unintentionally introduced from continents (Carson 1951:94–97). The Sea Around Us was a best seller, with numerous foreign editions and translations, and Oxford University Press republished Sea-Wind in 1952, which led to its translation in foreign countries (Brooks 1972:339–340). In a way, the paucity of illustrations in The Sea Around Us was compensated for in an extremely well-illustrated edition for young readers, having some of Carson's discussions simplified by Anne Terry White (Carson 1958). Even that edition found foreign translations (Brooks 1972:341). Royalties, honors, and fame enabled Carson to resign from the Fish and Wildlife Service and buy an ocean-side lot at Boothbay Harbor, Maine, where she built a summer cottage. She wrote another book, The Edge of the Sea (1955), unlike either previous book. It explained ecology of shore life in three environments along America's Atlantic coast: rocky shores of New England, beaches of middle states, and mangrove and coral coasts of Florida Keys. In 1948, she had hired Bob Hines as an illustrator for Fish and Wildlife Service, and they developed a good working relationship. He welcomed collaboration on her third book, and he went to shores with her and participated in collecting animals and observing them in their habitats. His numerous illustrations in Edge of the Sea were widely praised (Lytle 2007:107, Souder 2012:183–185). Brooks 1972: plate 5. This book celebrated marine invertebrate animals, either in tidal (littoral) or in neritic (sublittoral) zones. From these zones, organisms first crawled out of the sea onto land (Carson 1955:7). Tides vary significantly between Florida Keys and Maine, and effects of varying tides are most dramatic along New England's rocky coast. She explained that this region once had sandy beaches, but vast glaciers compressed the land and ocean flowed into rocky lands behind former beaches (Carson 1955:42–43). Rocky coast's first inhabitants were probably barnacles and mussels that strain plankton from seawater, and spores of green algae, that coated rocks. Grazers that scraped algae off rocks followed. Plankton strainers and grazers attracted predatory whelks, starfish, crabs, and worms (Carson 1955:45). Above the tide line lichens clung to rocks and slowly eroded rock surfaces. Interspersed among colorful lichens were bands of blue-green algae. Periwinkle snails grazed on lichens and algae; snails lived out of the water as much or more as in it. Other snails made the transition to become land snails and moved inland (Carson 1955:50). Barnacles also lived where they alternately are submerged in water and exposed to air at low tides. Barnacles were first classified as mollusks because their protective plates resemble a shell, but their swimming larvae, nauplius, resembled those of other crustacean, indicating their true relations. Carson speculated that nauplii settle on "rock surfaces that are rough and pitted better than smooth ones; probably they were repelled by a slimy film of microscopic plants, or even sometimes by the presence of hydroids or large algae" (1955:54). After settling down, their bodies metamorphose as drastically as caterpillars do when turning into butterflies. After 3–5 years when barnacles die, their shells become refuges for baby periwinkles, tide-pool insects, young anemones, tube-worms, or new barnacles. Dog whelks sometimes prey on mussels or periwinkles, but prefer barnacles. And so on, through various denizens of rocky shores, Carson describes their life cycles and how they interact with other residents. In 1971, this chapter became a separate book, including Hine's drawings, but additionally with striking photographs—some in color, some in black and white—by Charles Pratt, who earlier had shown them to her (Carson 1971). Carson 1955:31. But almost always the essence of the lives—the finding of food, the hiding from enemies, the capturing of prey, the producing of young, all that makes up the living and dying and perpetuating of this sand-beach fauna—is concealed from the eyes of those who merely glance at the surface of the sands and declare them barren. Carson described these surface sand inhabitants and their life styles, and Hines drew them to decorate her pages. On a Georgia beach, the most abundant inhabitants were ghost shrimps, inhabiting holes "considerably smaller than a led pencil" (Carson 1955:140–141). She explained numerous life histories of these, the original beach combers, then noted that shells of animals living beyond the tides also wash ashore on beaches. "One such mysterious link between the open sea and the shore is the ramshorn shell, Spirula" (Carson 1955:165). The creatures that built these shells were unknown until Johannes Schmidt systematically towed plankton nets across the Atlantic at various depths in search of migrating eels. His nets captured far more than eels, including ten-armed squid-like Spirulas which lived at depths of 900–1,500 feet. Another commonly washed ashore shell was of female eight-armed paper nautiluses (males lack shells). Carson bought sea shells at a Nags Head shell shop (1955:168), possibly the same shop I patronized during summer 1962, when I was a seasonal naturalist at Cape Hatteras National Seashore. Hurricanes deposit unusual shells on nearby beaches which year-around collectors scoop up. Carson 1955:141. Florida Keys are fringed by mangrove trees close by and further out by coral reefs, for nearly 200 miles, and some islands are remnants of reefs (Carson 1955:191–194). Corals live where the water temperature seldom drops below 70°F and to depths of about ten fathoms, since calcareous algae which contribute to reefs cannot photosynthesize at greater depths. Each species of coral has a characteristic shape. The coral is white but can show the color of internal algal cells that secrete oxygen absorbed by coral; algae, in turn, absorb carbon dioxide secreted by coral (Carson 1955:200–201). Coral reefs provided a basis for complex and very productive ecological communities. Carson described their diverse species, and Hines drew them, mostly individually, adding charm to her pages. He did not provide a panorama of a coral reef comparable to his low tide panorama of the rocky shore (fig. 7, above), perhaps because without an aqualung he could not see such a panorama. However, three his illustrations provide glimpses of complexity: (1) loggerhead sponge, spiny lobster, and black sea urchins; (2) gorgonian sea whip, basket starfish, sea fan, and young angelfish; and (3) horse conch, octopus, pipefish, sea horses, sea hare, giant starfish, cowfish, and sea grass (Carson 1955:217, 226–227, 234–235). Edge of the Sea did not have a bibliography, but ends with classification of invertebrate animals—from protozoa to sea squirts—reminiscent of a glossary that ended Sea-Wind. Hine's illustrations had a more three-dimensional appearance than Frech's 1941 outline drawings (Carson 1955:251–270). Carson 1955:201. Two other books published in 1955 had excellent illustrations in color. Herbert Zim and Lester Ingle's Seashores was a very good guide book to species. Lincoln Barnett and Editors of Life Magazine collected a very impressive The World We Live In, from 13 issues of Life, 1952–1954, with chapter 7 on "Creatures of the Sea," and chapter 8 on "The Coral Reef." It included two spectacular fold-out illustrations, one almost entirely on fish and the other on invertebrates inhabiting the ocean in the sublittoral (neritic) zone, pages 130–135. The climax of Carson's publishing and of her influence came with Silent Spring (1962), to be discussed in part 63, on history of pollution ecology. Evelyn and I were particularly interested in the work at the Plymouth Laboratory in England where a bright and innovative group of investigators had developed a variety of quantitative methods and had applied them to good effect in ecological analyses. Fortuitously, Professor Albert Parr, Director of Bingham Oceanographic Laboratory at Yale, invited Riley, who had taken his course in oceanography, to participate in a brief cruise of Atlantis (Wüst 1964:10, Schlee 1978) off the mouth of the Mississippi in March 1937, to do pigment analyses similar to what he had done in his thesis using freshwater plankton. That was Riley's hands-on introduction to oceanography, and Parr was pleased with his work, which led to a paper by Riley on phosphate enrichment and chlorophyll. My only interest was in dynamic aspects of ecological relations. The descriptive work of the older generation, including Bigelow, seemed dull and trivial. Not until some years later did I achieve sufficient historical perspective to realize that Bigelow and his contemporaries were doing work that was innovative in their time and was a prerequisite for everything that followed. Martin was a native of New Orleans, of Spanish-Jewish descent. He shared Yngve [Olsen]'s dislike of the sea, but the resemblance ended there. Yngve was an easy-going plodder. Martin was brilliant, erratic, and somewhat paranoid. He was sophisticated and could be a suave and charming companion when he chose to be, which was not by any means all the time. Although he had been kicked out of Tulane, short of getting his bachelor's degree, he had managed, on his own, to become a respected shrimp systematist. …my relations with Martin in the Bingham Lab had their ups and downs. I liked him a lot despite a good many qualities I didn't like, and we could have been casual good friends had we not been thrown together so closely. As I remarked earlier, he could be a lively, entertaining companion or a nasty son of a bitch, depending on his mood. His moods also were reflected in his work. Sometimes he worked night and day for considerable periods, but he was also subject to black depressions when he didn't come into the lab for some days. In his personal life, he was a Don Juan, charming and attractive enough to make easy conquests during the generally brief periods when he was intensely interested in a woman, but I never knew him to achieve a stable, emotional relationship with anyone. Riley contrasted his extracurricular life with Burkenroad's: "My sex life at the time was nil." Martin "fixed him up" on a double date, but the girl did not appeal to Riley. Later, another friend tried, and that time he met his future wife, Lucy, a nurse at the New Haven Hospital. (They had three sons.) The Wikipedia summary on Burkenroad seems consistent with Riley's characterizations; however, Burkenroad married in 1945 (a few years after he and Riley had gone their separate ways). World War II motivated Riley to conduct research for the U.S. Navy, 1942–1947, stationed at WHOI. Simultaneously, he continued thinking about his own researches. His three papers dated 1941 were obviously pre-war submissions, but he published three papers—1942, 1943, 1944—that indicated his oceanographic thinking during the war. Manuscripts and Archives, Yale University Library. George Clarke was a budding young scientist at the time when he helped bring the Atlantis over [from Copenhagen in 1931], a member of the new generation that believed in the importance of quantitative ecological methods. In addition to the plankton sampler, he put meters on the larger nets; he built and used a submarine photometer; he and his associates made experimental studies grazing rates of zooplankton. Over the years, he accumulated a lot of data. However, he was essentially a follower rather than a leader. He was doing things that the Plymouth people had already done, with only slight modification, and his interpretations tended to be pedestrian. I liked George as an acquaintance, but we didn't become close friends. There was no real warmth of the man, and in some respects, he wasn't a very nice person. He was lazy, always trying to grab a major share of the credit for things in which others—Bump, students, research associates—had done most of the work. He was a negligent thesis supervisor, often leaving students to flounder or seek help elsewhere. He thought it was fun to invite a seminar speaker to dinner before the evening lecture and get him drunk, and he gave secretaries and female technicians a hard time. Bump's private nickname for him was Georgie Porgie. Nevertheless, none of that touched me directly, and I was grateful to him for inviting me to participate in a program that looked promising and proved to be so. …work on Georges Bank was largely done by George Clarke, a zoologist, Gordon Riley, a marine physiologist from the Bingham Oceanographic Foundation, and two laboratory technicians from the Oceanographic—Dean Bumpus for plankton and Dayton Carritt for chemical work. Clarke, with his knowledge of plankton and light penetration, outlines a foresighted program of quantitative measurements which, if made on Georges Bank for 3 years in a row, and at all seasons, might point to the factors that enabled the area to support such a rich population of fish. Between September 1939 and June 1941, when the project was prematurely ended by the war, Atlantis made 11 trips to Georges Bank… Clarke's textbook Elements of Ecology (1954) had its merits—translated into three foreign languages and reprinted until 1980—but it had to compete with Eugene Odum's very popular Fundamentals of Ecology (1953). Riley is most remembered for his pioneering mathematical models in marine biology. Eric Mills (1982) placed Riley's models in historical context, and later wrote a long discussion of his career from this perspective (1989:266–313). Recently, Thomas Anderson and Wendy Gentleman decided that there was more to say on the subject (2012:1–25). Mills critiqued their paper before publication; they discussed the mathematics of Riley's 1946 model in much more detail than was practicable in Mill's book surveying 90 years of biological oceanography. Since Anderson and Gentleman find Riley's mathematics has more than historical interest, others should also have a second look at his reasoning. Two biologists at the Plymouth Laboratory of the Marine Biological Association (Egerton 2014:356–357) who most directly influenced Riley were H[ildebrand] W[olfe] Harvey (1887–1970) and Richard H. Fleming (1909–). Riley and Harvey corresponded "extensively" (Mills 1989:305), and Riley