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History of Ecological Sciences, Part 50: Formalizing Limnology, 1870s to 1920s

2014; Ecological Society of America; Volume: 95; Issue: 2 Linguagem: Inglês

10.1890/0012-9623-95.2.33

2327-6096

Frank N. Egerton,

Aquatic Ecosystems and Phytoplankton Dynamics

Click here for all previous articles in the History of the Ecological Sciences series by F. N. Egerton Limnology is unique among sciences in having one scientist name, define, and write the first monograph, first survey, and first textbook of that science (Egerton 1962). Before Forel formally founded limnology, naturalist–philosopher Henry Thoreau had conducted a limnological study at Walden Pond (Egerton 2011:257–259), and others conducted studies that now could be called "limnological" (Beeton and Chandler 1963:542–544, Carlander et al. 1963:324–325, Eddy 1963:303–304, Fry and Legendre 1963:498–506, Pennak 1963:349, Yount 1963:269–270, Elster 1974:8–10, Steleanu 1989:26–99, 111–123), but isolated studies do not establish an organized science. Limnology is fortunate in having a general history of limnology (Steleanu 1989), and although it is heavily focused upon Europe, there is also a magnificent collaborative history of North American limnology (Frey 1963). I have not been able to call my study limnography because the word limnograph is already appropriated for some instruments used in lake measurements; therefore I have coined limnology. Limnology, then is the oceanography of lakes. That poor worm, a Mermis aquatilis, was a revelation for me. If one organism could live in that clay, in depths of 40 m, a perpetually cold region, far from littoral vegetation, others must be able to do so. If the ooze is inhabited down to the greatest depths, the profundal region is not a desert; there is an abyssal society. …our lakes are isolated from the sea and are connected to each other only by surface drainage and that they have been filled relatively recently by glaciers from the last glacial epoch means the origin of the species of the profundal fauna and their differentiation goes back no further than this period in the history of the earth. Forel proceeded to answer his own questions. In 1872 he outlined a decade-long research program, with costs. He wanted to determine the bottom topography of the lake, in both deep and shallow waters, profundal fauna and flora, lake temperatures, and study the deltas of the Rhone and Dranse Rivers. His subsequent profundal fauna studies included, besides Lac Léman, lakes Constance, Neuchâtel, and Zurich. His profundal studies lasted longer than a decade, culminating in La faune profonde des lacs Suisse (Forel 1884) and extended in "L'origine de la faune des poissons du Léman" (Forel 1901a). Forel's great monograph, Le Léman, three volumes, devoted two volumes to physical aspects of the lake. Volume 1 (Forel 1892) included geography, hydrography, geology, climate, and hydrology (sources and changes in water); volume 2 (1895) included hydraulics (seiches), temperature, optics, acoustics, and chemistry—an aquatic Humboldtian project. Volume 3 (1904) devoted over 400 pages to fauna, followed by human history, navigation, and fishing. In contrast to that encyclopedic work, Forel also published a concise Handbuch der Seenkunde: Allgemein Limnologie (Forel 1901, 1977), which summarized three decades of his research, placed in the larger context of contemporary investigations by others. An even larger-scale survey of limnology than Forel's Le Léman was the Bathymetrical Survey of the Scottish Fresh-water Lochs (six volumes, 1910). This survey of 562 lochs, which followed a survey of saltwater lochs (1884–1891), was sponsored by the Royal Societies of London and Edinburgh and the British Association for the Advancement of Science, and supervised by John Murray (1841–1914), who is famous for participating in the voyage of the Challenger (1873–1876) and editing the 50-volume Report on the Scientific Results (Burstyn 1974, Price 2004), and conducted during 1897–1909. The general results are all in volume one, the other volumes being devoted to descriptions and maps of the lochs. Four of the articles in volume one, 163 pages, emphasized bio-limnology: James Murray, "Biology of the Scottish lochs," John Hewitt, "Some distinctive characters in the fresh-water plankton from various islands of the North and West Coasts of Scotland," William A. Cunnington, "On the nature and origin of fresh-water organisms," and C. Wesenberg-Lund, "Summary of our knowledge regarding various limnological problems," with the last article not being limited to biology. James Murray's survey was limited to invertebrates and microscopic algae, as the Scottish vertebrates were already well studied. John Murray contributed "Characteristics of lakes in general, and their distribution over the surface of the globe" (45 pages). Additionally, James Chumley compiled an impressive 95-page bibliography of limnology. (a) François Alphonse Forel in his study–laboratory, Morges. By Ernest Biéler. (b) Water mite Campognatha (now Hygrobates) foreli from Lake Geneva benthos. Forel 1879. From Vincent and Bertola 2012:52, 54. In Germany in 1891, E. Otto Zacharias (1846–1916) founded a Biologische Station on the Grossen Plöner See, at Plön, which opened 1 April 1892 (Thienemann 1917, Wetzel 2004). Also in 1891, he published a text, Die Tier–und Pflanzenwelt des Süsswassers (Leipzig), with contributions from other limnologists, including Forel. He was "an independent scholar and former teacher who supported the station from soft money grants and his earnings as a popular writer" (Nyhart 2009:321). In 1893 he began publishing an annual Forschungsberichte aus der Biologischen Station zu Plön (Chumley 1910:750–751), which evolved into Archive für Hydrobiologie (Elster 1974:9). Zacharias also published two more books on plankton in 1907 and 1909. Also in 1909, Bruno Eyferth and Alfred Kalberlah published their rather encyclopedic work on European fresh-water plankton. In 1917, after Zacharias's death, the Biologische Station he had founded became affiliated with the Kaiser Wilhelm Gesellschaft, under the excellent direction of August Thienemann (1882–1960), who had studied at Heidelberg under Robert Lauterborn (1869–1952), "Germany's leading river biologist" (Nyhart 2009:321). Thienemann joined the faculty of the University of Kiel, and eventually became Europe's leading limnologist (Thienemann 1959, Ohle 1961, Steleanu 1989:300–308, 385–404, Egerton 2008). He remained director at Plön for forty years and edited Archiv für Hydrobiologie for more than forty years. In Sweden, Finnish immigrant fisheries manager Oscar Nordqvist (1858–1925) organized a Södra Sveriges fiskeriförening (South Sweden Fishery Association) in 1907, with laboratory at Aneboda (Söderqvist 1986:80). In summer 1910, Nordqvist hired two assistants at Aneboda, students Gunnar Alm (1891–1962), Uppsala University, who studied Ostracoid Crustacea and received his Ph.D. in 1915, and Einar Naumann (1891–1934), Lund University, who studied lake plankton and bottom fauna and received his Ph.D. in 1917 (Thienemann 1937, Söderqvist 1986:80–81, Steleanu 1989:382–385). Alm pursued a career in fisheries management and studied the biota and environment of lakes in relation to fish populations. Naumann became a leading European limnologist. In 1921 Thienemann traveled to Sweden and met Naumann. They developed a close working relationship (Stelleanu 1989:381–404), and Naumann suggested that they organize an international association. Thienemann agreed, and in January 1922 they sent a prospectus to 100 colleagues and got a very favorable response (Rodhe 1974, 1975, Steleanu 1989:405–411). A foundation meeting was held in August, and the Internationale Vereinigung für Limnologie (later, Societas Internationalis Limnologiae) was formed with 188 members in 23 countries, and at Thienemann's suggestion it published an annual Verhandlungen. Naumann was active in this society during the remaining 12 years of his life. In 1907, C. A. Weber introduced the German terms for eutrophic (nährstoffreichere), mesotrophic (mittelreich) and oligotrophic (zuletzt nährstoffarme) regarding the chemical nature of the soil solution in German bogs (Hutchinson 1969:17, Rodhe 1969:50). Naumann became a leading limnologist who introduced Weber's concept, but with different terminology ("eutrophic" and "oligotrophic") into limnology to indicate poor or rich phytoplankton communities (Naumann 1919, Hutchinson 1969:17–18, Rohde 1969:50–51, Elster 1974:10). In 1921, Thienemann accepted Naumann's terms but applied them to clear lakes rather than phytoplankton and added "dystrophic" for lakes with brown, humic waters with little lime. Frenchman Emile Gadeceau (1845–1928) of Nantes at age 16 participated in a field trip of the Société botanique de France at Nantes, which led to his becoming a disciple of English botanist James Lloyd (1810–1896), who had settled in Nantes (Desmond 1977:390, Perrein 1994:181). Lloyd had published Flore Loire-inférieure (1844) and Flore de l'ouest de la France (1854). In 1893 Gadeceau met Université de Montpellier plant geographer Charles Flahault (who is discussed in Egerton 2013:344–345) at a meeting of the Société botanique de France, held at Montpellier (Perrein 1994:187–188). They began to correspond, and Flahault became an important influence on Gadeceau. Flauhault (Flauhault 1900:444–445) introduced the term "écologiques" into French botanical discussion, and he wrote a preface for Gadeceau's Le Lac de Grand-Lieu (1909), which monograph won a prize from the Société Botanique de France. Le Lac de Grand-Lieu is 13 kilometers south of Nantes, and Nantes is on the lower Loire River in western France. According to the scale on Gadeceau's foldout map of the lake, it is roughly 6.5 km wide by 8 km long, but has a very irregular shape. Two small rivers, la Boulogne and l'Ognon, flow into the lake from the southwest and northwest respectively, and l'Acheneau River flows out at the northeast, to the Loire (Gadeceau 1909:17). In his introduction, he stated that he had access to James Lloyd's unpublished journal of observations, 1837–1889, and his own observations, 1879–1908, from all seasons, but mostly from summers. He also thanked a number of botanists and others who had assisted his research. His title page indicates that his book is a "monographie phytogeographique," but it does devote 11 pages to an inventory of the lake's birds, fish, and insects. The book's 120 pages of description and discussion are divided into three parts: geography, 30 pages (including 10 pages on animals); aquatic plants, 55 pages (7 pages on ecological influences, 48 pages of annotated plant list); and biological ecology (in three chapters: physiographic classification, ecological groups, and phytogeographic history). Aside from Flahault (whose physiographic nomenclature, 1901, flora and vegetation of France, 1901 were cited), who influenced Gadeceau's thinking, observations, and organization of his monograph? His general bibliography (as opposed to his local sources) provides clues: Forel's Le Léman was obviously important, as also was Schröter and Kirchner's Die Vegetation des Bodensees, both parts, and two works by Ant. Magnin: "Les Lacs du Jura et notes additionnelles sur la limnologie juras-sienne" (1893) and La vegetation des Lacs du Jura (1904). The only British title he cited was the French translation of Darwin's Origin of Species (1887). He cited plant ecology works by Oscar Drude (in French translation, 1897), A. F. W. Schimper (1903, in English), and Eugene Warming (in German translation). He cited two articles by Henry Cowles in the Botanical Gazette (1899, 1901) and Conway MacMillan's "Observations on the distribution of plants along shore at Lake of the Woods" (1897). Gadeceau achieved a synthesis of his own and Lloyd's observations with the leading limnological–ecological literature of the day. Cambridge University zoologist J. T. Saunders apparently taught the first course in hydrobiology in Britain, 1924–1925, and with W. Pearsall and algologist F. E. Finch, organized The Freshwater Biological Association, which established a station at Lake Windermere in 1926 (Jack 1945:43, Hiatt 1963:194, Slack 2010:see index). In 1901, Eustace Gurney had built a station on Sutton Broad, near Stalham (Juday 1910:1260, Kofoid 1910:166–168). Jack (Jack 1945:43–44) did not include either station in his survey of world stations around 1940. Hiatt (Hiatt 1963:194) cited the date of Windermere Laboratory founding as 1931—perhaps the date its current building was erected. In July 1928, Thienemann, Austrian limnologist Franz Ruttner (1882–1961), director of the Biological Station at Lunz, and German zoologist H. J. Feuerborn embarked on an expedition to the Sunda Islands (Java, Sumatra, and Bali) for 10 months to collect plants, animals, water samples, and environmental data, believing that "only knowledge of the primordial, tropical conditions can give the scientist a deeper understanding of how life proceeds at our latitudes" (Thienemann 1959:129, translated in Rodhe 1974b:540). They studied lakes and running waters. Later, collaborating with over 100 European specialists, they described 1100 new species. They published their findings in 11 supplementary volumes to Archiv für Hydrobiologie, 1931–1958, entitled Tropische Binnengewässer, in 7920 text pages and 348 plates (Thienemann 1959:263). Ruttner provided hydrographic, physical, and chemical data for 15 lakes that could be correlated with their flora and fauna. That breadth of knowledge of temperate and tropical limnology enabled Ruttner to write an introductory textbook (1940) that was worth translating into English from the second and third German editions (Ruttner 1963), the textbook Art Hasler used to teach me limnology in spring 1963. Despite detailed investigations, however, Thienemann and Ruttner evaluated productivity in tropical lakes somewhat differently, leaving it to later researchers to resolve the differences (Rodhe 1974b:545). (a) August (Friedrich) Thienemann. Web page. (b) Food web in a lake. Thienemann 1925:57. [Egerton 2007:60.] The more variable the biotope (environmental) conditions, the greater the number of species in the biocenosis (biotic community); The more biotope conditions deviate from optimal for most species, the smaller the number of species, but the greater the number of individuals of species represented; The longer a locality retains the same conditions, the richer and more stable is its biocenosis. Thienemann in 1926 first used the concepts of producers, consumers, and reducers to organize his biological data, and included a diagram that would influence Raymond L. Lindeman in his epochal "Trophic–Dynamic Aspect of Ecology" (1942) that cited this (1926) and two other Thienemann papers (Egerton 2007:60–61). The earliest biological research stations were marine (Jack 1945:9–11). In 1910, two American aquatic biologists, Chancey Juday and Charles Kofoid (on both, see below), published—independently of each other—illustrated surveys of European biological stations, freshwater and saltwater. Juday's was an article, Kofoid's a book. The earliest founding date Juday listed of 11 freshwater stations is 1888 for the Bohemian Portable Laboratory under Dr. Karl Fritsch, first located near several Bohemian lakes and later moved to the Elbe River (Juday 1910:1274). Zacharias' station at Plön may have been the earliest permanent freshwater station. In The Netherlands, several pleas before World War I for government to establish a hydrobiological laboratory (like other countries) were unsuccessful, and the Netherland's first notable advance was Professor H. J. Jordan's 1918 text on animal life in freshwater (Cramer 1987:68–70). Limnology in America began, like plant and animal ecology, in the upper Midwest. Stephen Forbes' "The Lake as a Microcosm" (1887) provided "the first concept of the limnological ecosystem" (Elster 1974:10; Hagen 1992:7–10, Golley 1993:36–37). In one respect, establishment of field stations, limnology was ahead of plant ecology and animal ecology (Kofoid 1898). In 1893, Jacob Reighard (1861–1942), with support from the Michigan Fish Commission, led a team in a biological survey of Lake St. Clair, located between Lakes Huron and Erie (Chandler 1963:98–99, Bocking 1987:81–91, 1990:478–495, Burgess 1996:91). In 1894 the survey moved to Traverse Bay, Lake Michigan. The two reports from these collaborations were substantial contributions (Reighard 1894, Ward 1896) which dealt with aquatic fauna, biota, and fisheries, but there were no funds for a third year. However, in 1898–1901, the U.S. Fish Commission funded Reighard and his team in a biological survey of Lake Erie, operating from the Fish Commission's Hatchery at Put-in-Bay, Ohio, and using the Hatchery's boats (Stuckey 1988, Bocking 1990:485–497). That survey produced a series of valuable reports. Ohio also had a fish hatchery at Put-in-Bay, to which Ohio State University provided funds to Prof. David S. Kellicott (1842–98) in 1896 to build a second floor for a Lake Erie biological station, which operated for two summers before Kellicott's death (Langlois 1949:1). Kellicott's successor at Ohio State was Herbert Osborn, who operated the station in that location, 1899–2 July 1903, in its own building until 1918, when it returned to the State Fish Hatchery (Langlois 1949:3–6). Now, it has separate facilities and is named Franz Theodore Stone Laboratory (Kofoid 1898:394–395, Gerking 1963:239). Prof. Stephen Forbes, whom we met in part 45 (Egerton 2013a:72–73), established a University of Illinois Biological Research Station on the Illinois River at Havana (originally three rooms and a boat) in 1894 (Kofoid 1898:398–406, Bennett 1958:165–166, Gunning 1963:167–168, Bocking 1987:101–117, 1990:472–478). It was America's first research station on a river. By 1900, the University of Minnesota had a temporary station on a houseboat (Kohler 2002:68). Charles A. Kofoid (1865–1947), an Illinois native with a Harvard Ph.D., became head of the Havana station until 1900, when he joined the Department of Zoology at the University of California, Berkeley (Mullen 1973, Burgess 1996:62), and the Illinois station continues to conduct valuable research. Kofoid's specialty was plankton, and he was one of the very few who studied plankton in rivers. His publications on Illinois river plankton extended from 1897 to 1908, with his most substantial and general results appearing in 1903. By 1900, his studies extended throughout the Illinois River watershed In 1895, German immigrant Carl Eigenmann (1863–1927), who had studied ichthyology under David Starr Jordan, established an Indiana University field station on Wawasee (earlier called Turkey) Lake, which was moved four years later to Winona Lake (Kofoid 1898:395–398, Frey 1955, Gerking 1963:239, 263, Shor 1971). The U.S. Fish Commission sponsored a physical and biological survey of Lake Maxinkuckee, in northern Indiana, 1899–1908, which was said to be "one of the most complete ecological surveys ever conducted" (Gerking 1963:263), though publication of the two-volume report was delayed (Evermann and Clark 1920). Botanist Conway Macmillan had a University of Minnesota station at Gull Lake (Kohler 2002:51–52), but before 1900 another one opened at Lake Itasca in a state park (Hiatt 1954:181) and the Gull Lake one probably ended then. Edward A. Birge (1851–1950), from Troy, New York, earned A.B. (1873) and A.M. (1875) degrees from Williams College in Massachusetts, then entered Harvard to work on a doctorate. Agassiz set him to work on the Museum of Comparative Zoology collection of sea urchins. Three months later, Agassiz died and Birge was saved from becoming a specialist on echinoderms. One of his professors at Williams was John Bascom, who soon became president of the University of Wisconsin. He invited Birge to become instructor in natural history and curator of the university cabinet (museum). Birge accepted and began teaching in January 1876. Later, he received time off to complete his doctorate at Harvard (1878), and he was promoted to professor at Madison in 1879 (Sellery 1956:4–14, Noland 1970, Beckel 1987:1–2, Egerton 1987:86–94, 1999a, Burgess 1996:16). During July, only the upper 12 m. of Lake Mendota are tenanted by crustacea, and over 90 per cent are in the upper 9 m. Nearly 50 per cent are in the upper 3 m.; 30 per cent between 3 and 6 m. and over 15 per cent between 6 and 9 m. There is, therefore, apparently an "abyssal" region with little crustacean life. This region is only temporarily unoccupied, being peopled by the crustacea later in the year, as the temperature of the lake falls. I employ two new words in this paper, which seem convenient in writing of the temperature and other phenomena of lakes. These terms are epilimnion, for the warm layer of water which develops in the lake in summer, and hypolimnion, for the lower colder water. These two parts of the lake differ widely in their temperature changes, as well as in their chemical and biological phenomena. It seems advisable, therefore, to assign definite names to them. The word thermocline, first used by me in 1897, is the equivalent of Richter's term Sprungschicht, or the discontinuity layer of Wedderburn. It lies at the top of the hypolimnion. Birge's bibliography (Birge 1910b:1014–15) cited three papers by E. M. Wedderburn, 1907–1908, but one must consult Birge's previous paper in the same volume (1910a:1000) for a citation to Eduard Richter's paper of 1897. Birge's career resembled Stephen Forbes' career, with an accumulation of academic and government positions and responsibilities, but with little or no diminishing of research and publishing. He was Chairman of the Zoology Department (1875–1906), Dean of the College of Letters and Science (1891–1918), Acting University President (1900–03), President (1918–25), Director of the Geological and Natural History Survey (1879–1919), and Commissioner of Fisheries (1895–1919). He needed an assistant, and in 1900 hired Chancey Juday (1871–1944), from Indiana, as biologist with the Survey (Frey 1963:4–6, Noland 1973, Beckel 1987:4–5, Burgess 1996:59–60, Egerton 1999). Juday had B.A. and M.A. degrees from Indiana University, where he had studied under Eigenmann. In 1908 Juday also became instructor in zoology at the University of Wisconsin, and in 1931, when he became professor of limnology, he resigned from the survey. Birge and Juday's first substantial collaboration was on dissolved gases in Wisconsin lakes (Birge and Juday 1911, 1977), which "is the single most outstanding single contribution of the Wisconsin school" (Mortimer 1956:188). The central question raised in this bulletin was why lakes differed so substantially in their ability to support plankton? Birge and Juday devoted the rest of their limnological careers to providing answers to such environmental questions. A major part of the answer was in their bulletin on the quantity and chemical composition of plankton (Birge and Juday 1922, 1977), followed by studies on penetration of sunlight into lake waters (listed in Juday and Hasler 1946). Birge and Juday. Beckel 1987:cover. Birge's mature point of view is expressed in the concept of the heat budget, which, though derived from ideas of Forel and others, represented a highly original and important contribution because it first called attention in the lake as a natural system with an input and output. This point of view has tended to underlie most of what has been done in lake chemistry and in the study of primary productivity during the past three or four decades. Wisconsin's south-east quarter. Lake Mendota is at the left side of map. Birge and Juday 1911:3. Since the University of Wisconsin-Madison is located on the shore of Lake Mendota (largest of five interconnected lakes), Birge and his collaborators first studied the limnology of Lake Mendota without establishing a biological station, making it (eventually) the most studied lake in the world. In 1925, they established a station at Trout Lake, in far northern Wisconsin, to facilitate comparative studies (Frey 1963:5). My work as an assistant professor is increasingly demanding with extra hours of lab for research as a result of "fishing" for Birge. He has developed a fine net, 1/20th of millimeter gauge, which is used to gather crustacean from varying depths in the lakes. She had obviously taken the position in the Zoology Department fairly recently, as her cousin Nora on 28 April 1901 lamented: "we thought you should remain at Downer as head of the science dept" (Hartridge 1997:4). Why Merrill in 1902 went to South America, and alone, is unclear: "at age 39, Merrill seemed to experience a mid-life self-appraisal that unexpectedly prompted her to demonstrate an independent attitude in a man's world" (Hartridge 1997:xv). Her professional curiosity was possibly an excuse for the trip, 1902–1903, for her collections and notes during that trip were rather modest. However, that trip prepared her for her more serious trip to South America in 1907–1909, when her scientific journals ran to 14 notebooks, compared to one notebook in 1902–1903 (Hartridge 1997:185–190). She collected Cladocera and other invertebrates, and her samples and notebooks are preserved in the Milwaukee Public Museum. It might be more accurate to say her work was zoological than limnological, but it was done within a limnological context. She corresponded with Birge during her trips. One unknown species she collected was named for her. She was working on a Ph.D. at the University of Chicago in 1915, when she died, age 52. Juday in February 1910 visited lakes in Guatemala and El Salvador, making "one of the first studies in tropical limnology" (Juday 1915; Frey 1963:6). Perhaps Merrill's successful collecting trip to Brazil encouraged Juday to take this trip. Victor Shelford (1877–1968), whom we met in part 49 (Egerton 2014:69–71), was an animal ecologist whose researches included aquatic as well as terrestrial species. His Animal Communities in Temperate America, as Illustrated in the Chicago Region (1913) had more pages devoted to aquatic than to land animals. The chapter titles of his aquatic chapters were: (4) Conditions of Existence of Aquatic Animals, (5) Animal Communities of Large Lakes (Lake Michigan), (6) Animal Communities of Streams, (7) Animal Communities of Small Lakes, and (8) Animal Communities of Ponds. In addition, he also wrote two chapters on transition zones—(10) Animal Communities of the Tension Lines between Land and Water, (11) Animal Communities of Swamp and Flood-Plain Forest, that discussed amphibians and insect species that spend part of their life cycle in water and another part on land, as well as discussing terrestrial feeders that eat aquatic foods. James G. Needham (1868–1957) participated in a limnological survey of Walnut Lake, Michigan in summer 1907 (Chandler 1963:99, Mallis 1971:174–178, Burgess 1996:80), then returned to Cornell University as Assistant Professor of Limnology, and since he studied freshwater insects, he joined the Department of Entomology (Berg 1963:209, Needham 1946.). In 1908 he began teaching a course in limnology. In 1914 when Professor Comstock retired as chairman of the Department, Needham succeeded him. The Life of Inland Waters (Needham and Lloyd 1916) was a well-illustrated, widely used introductory textbook of limnology, with a bibliography of English and German titles. Needham also served as president of the Entomological Society of America in 1919 and vice-president of ESA in 1936. However, it was Birge and Juday (Birge and Juday 1914, 1921) who initiated limnological research on the New York Finger Lakes, as that was beyond the scope of Cornell's limnologists. Two years after Needham's textbook appeared, he and Birge were among 25 limnologists and zoologists who contributed chapters to Fresh-Water Biology (1918), edited by Henry Baldwin Ward (1865–1945) and George Chandler Whipple (1866–1924). Ward was Professor of Zoology at the University of Illinois (Burgess 1996:109), and Whipple was retired Professor of Sanitary Engineering at both Harvard and M.I.T. The encyclopedic Fresh-Water Biology (1121 pages) is ostensibly limited in scope to the 48 contiguous United States, though it would have been of some use to Canadians and Mexicans. Its coverage includes bacteria, algae, vascular plants, and aquatic invertebrates, and is well illustrated, though not all species are illustrated. It does not cover aquatic vertebrates. (a) Victor E. Shelford, 1917. Croker 1991. (b) Food web for ponds in northern Illinois and Indiana. Shelford 1913:70. [Egerton 2007:54] Conochilus unicornis Rousselet. A. colony. x40. B. Single animal. x150. (After Weber.) Ward and Whipple 1918:616. University of Michigan Biological Station, Douglas Lake. Chandler 1963:101. Another equally remarkable limnological collaboration was Limnology in North America (Frey 1963), with 26 chapters by 32 authors. This volume encompasses Canada, Alaska, Mexico, and the Caribbean Islands, as well as the contiguous 48 U.S. states. All chapters have ample bibliographies. Some chapters were used to write the above account on North America, but chapters not already used deserve notice here if they discussed the period 1870s–1920s. These chapters are discussed in their sequence in the volume. David Chandler's chapter, "Michigan," discussed more than Reighard's group surveys (see above). Reighard's own researches continued beyond those turn-of-the-century surveys, and he also helped plan a survey in 1908 that Thomas Hankinson performed at Walnut Lake to learn about whitefish reproduction, as guidance to a fish-stocking program. Reighard was chairman of the University of Michigan's Department of Zoology, and he also encouraged courses in ecology and limnology and establishment of a Biological Station at Douglas Lake, established in 1909, which encouraged limnological research and teaching (Chandler 1963:99–100). John Brooks and Edward Deevey (1963) focused their 47-page chapter on the limnological feat