Portal de Periódicos da CAPES

History of Ecological Sciences, Part 48: Formalizing Plant Ecology, about 1870 to mid‐1920s

2013; Ecological Society of America; Volume: 94; Issue: 4 Linguagem: Inglês

10.1890/0012-9623-94.4.341

2327-6096

Frank N. Egerton,

Bryophyte Studies and Records

Click here for all previous articles in the History of the Ecological Sciences series by F. N. Egerton A German zoologist, Ernst Haeckel, coined and defined a new branch of biology, "Oekologie," in 1866 (Egerton 2013). In the later 1800s, four ecological sciences emerged: (terrestrial) plant ecology, (terrestrial) animal ecology, limnology, and marine biology. Plant ecology evolved out of phytogeography and physiology (Billings 1985:5–6), to which was added during the 1890s investigations of plant communities and vegetation succession. Physiology was important, but played a relatively passive role: its significance often depending on how well phytogeographers understood it. European botanists and publications led the way. Phytopathology during the 1800s is discussed in part 44 (Egerton 2012) and omitted here. For phytopathology during the early 1900s, see G. C. Ainsworth, Introduction to the History of Plant Pathology (1981) and C. L. Campbell et al., The Formative Years of Plant Pathology in the United States (1999:225–339). The history of aquatic plant ecology is part of limnology and marine ecology and will be discussed in parts 50 and 51. The ecology of lichens is deferred to part 52 on the history of symbiosis studies. After these exclusions, the remaining plant ecology, about 1870 to mid-1920s, is still vast in scope and detail, and surveys by Becking (1957), Whittaker (1962), Shimwell (1972), Tobey (1981), Nicolson (1989, 1996), and Cittadino (1990) provide introductions to additional developments and literature. Phytogeography along ecological lines laid down by Humboldt (Egerton 2009b) and Augustin-Pyramus de Candolle (Egerton 2010:26–29) continued throughout the 1800s (Pritzel 1871–1877:458–462, Nicolson 1996) and can be seen in such treatises as Joakim F. Schouw's Grundtraek til en almindelig Plantgeographie (1822, German, 1823), Franz Meyen's Grundriss der Pflanzengeographie (1836, English 1846, English 1977), Arthur Henfrey's Vegetation of Europe, Its Conditions and Causes (1852, 1977), and Alphonse de Candolle's Géographie botanique raisonée (1855) (Reed 1942:128–129). German botanists led the way, with Russian botanists following a parallel, rather independent, course (Becking 1957:417–419). An Austrian, Anton Kerner von Marilaün (1831–1898), studied medicine at the University of Vienna, but soon gave up his practice to become a botanist (Kronfeld 1908, Stafleu and Cowan 1976–1988, II:525–530, Nicolson 1996:297–298), who was strongly influenced by Humboldt (Cittadino 1990:119). He became a professor at Budapest, then Innsbruck, and finally in Vienna. While in Hungary he had collected plants in botanically little-known eastern Hungary and Transylvania and published Das Pflanzenleben der Danaulaender (1863). Its American translator claimed that this book "is the immediate and direct parent of all later works on Plant Ecology" (Conard 1951:vii). Is this translator merely hyping a work he has translated or is there some validity to his claim? R. J. Goodland (1975:241) reserved that "parental honor" for Warming (1995). Dwight Billings (1985:5) and Malcolm Nicolson (1996:304–305) agreed with Conard. Clearly, Kerner's Plant Life of the Danube Basin is a good landmark with which to begin this history. Whereas Humboldt considers the north German heather-covered coastal plains as steppes, and Koch applies that name to a formation composed of high sod-forming grasses, and Willkomm restricts the term to formations on saline soils of both highlands and lowlands, Grisebach—whose usage of the word steppe we shall follow—conceives of a steppe as a region in which, because of extreme heat, sterility and drought of summer, no tree can survive, and in which the most diverse other formations composed of grasses, herbs, etc., may develop. Kerner argued that others had failed to choose as examples definite genera, but had portrayed physiognomic differences on the broadest lines of resemblance. Furthermore, others had used vernacular names as scientific terms. He developed a new terminology, but without resorting to Latin, as Linnaeus had, because the German language was quite adequate for the purpose (Kerner 1951:7). He did not consider that this might be a problem for people who knew no Germanic language. Kerner did not become the Linnaeus of plant ecology, but Patrick Matagne (1998:428) thinks that Kerner's effort "laid the theoretical basis for a transition from phytogeography to phytosociology." In bitter struggle with the unyielding elements the first settlers gain possession of the lifeless earth. Step by step they press forward over the waste talus of the mountains or over the shifting sands of coastal plains and clothe them with a sparse plant cover. Years must pass before a second generation can unfold with greater strength and variety on the prepared soil. But ceaselessly plant life works, and builds its green structure greater and greater. On the remains of fallen races the germs of other younger, newer plant forms take root, and so it goes on with tireless change, until at last the shady crowns of a high forest whisper above the deep mull soil. The challenge was to unravel these changes and their causes within each formation. As he crossed the Hungarian plains, he could see where a village had stood 200 years previously, because there were nettles there foreign to the prairie flora, which accompany human settlers, and also locusts, oaks, and apparently willows that villagers had planted (Kerner 1951:16). He was particularly interested in explaining why trees could invade the prairie only along rivers or in lowlands covered with fens, and he compared the treeless prairie with the treeless mountains above treeline. In both cases, the season for tree growth was only about two months, limited on the prairie by drought and on mountains by frost (Kerner 1951:26–27). On the other hand, treeless alpine regions had scant prostrate perennial vegetation, but the treeless prairie had abundant vertical annuals. He often could explain the causes of vegetative dynamics, and where he could not, he gave good descriptions of vegetation. Many trees obviously had their seeds distributed by the wind. He was puzzled by oaks. Those growing along streams had their acorns moved by water, but what about those away from streams? He postulated that in past times streams had gone in different directions (Kerner 1951:39–40). Thoreau had published his insight about squirrels burying more acorns than they ever recover during winter, in the Middlesex Agricultural Society's Transactions (1860), and republished in the New York Weekly Tribune (Egerton 2011:269); neither American publication was likely to reach a botanist in the Austrian Empire. Puzzling acorn dispersal was not a typical example of Kerner's reasoning; his book is indeed an impressive achievement. Probably no one followed his example by writing a similar book on another European region because no one could—until 1904–1908, when Carl Schröter published his Pflanzenleben der Alpen (see below). (a) Anton Kerner. Kronfeld 1908: frontispiece. (b) Alphonse de Candolle. (c) August H. R. Grisebach. Magin-Gonze 2004:165, 202. Instead, conventional phytogeography and physiology progressed in leading European countries. Matagne sees four phytogeographers as most important in the later 1800s—Kerner, Alphonse de Candolle, Grisebach, and Flahault—extracts from whose works Matagne reprinted following his introductory essay (in Acot 1998:439–519). Swiss Alphonse de Candolle (1806–1893) continued the great work of his father, Augustin-Pyramus de Candolle (Pilet 1971a, b, Stafleu and Cowan 1976–1988, I:433–452, Dajoz 1984:39–46, Drouin 1993:60–62, Isley 1994:178–180, Matagne 1998:432–433, Magnin-Gonze 2004:165–167, Egerton 2010:26–29), which culminated in his Géographie botanique raisonée (two volumes, 1855). He corresponded with Charles Darwin and Hewett Watson, among others. Candolle judged heat and moisture to be the dominant factors affecting plant distribution. Matagne (1998:433) thinks Candolle's development of a classification based on plants' mode of life contributed to the emergence of a new ecological approach, centering on plant–environmental interactions (published before Kerner's classification). Nicolson (1996:300) viewed Alphonse de Candolle as more of a floristic phytogeographer than as a follower of Humboldt. (a) Charles-Henri-Marie Flahault. Virville 1954:247. (b) Ragnar Hult. Collander 1965: plate 4. German August Heinrich Rudolf Grisebach (1814–1879) was a professor of botany at the University of Göttingen, influenced by Humboldt, who studied taxonomy as a basis for his phytogeography (Reed 1942:131, Wagenitz 1972, Stafleu and Cowan 1976–1988, I:1007–11, Morton 1981:433, Cittadino 1990:113–114, 119–122, Matagne 1998:429–431, Magnin-Gonze 2004:202). He traveled widely in Europe, but not elsewhere. His numerous publications (five early ones are reprinted in Egerton 1977b), culminated in his authoritative Die Vegetation der Erde und ihrer klimatschen Anordnung (two volumes, 1872, French, 1877). French botanist Charles-Henri-Marie Flahault (1852–1935) received his doctorate at Paris in 1878, then traveled with botanist Gaston Bonnier (1853–1922) to Sweden to study the relationship of plants and climate (Emberger 1936, Jovet 1954:256–257, Stafleu and Cowan 1976–1988, I:843, Matagne 1998:434–437, 1999:49–50). Bonnier and Flahault's "Observations sur les modifications des végétaux suivant les conditions physiques du milieu" (1878) compared the similarities and differences between alpine and polar climates and plants, somewhat reminiscent of Kerner's comparisons (1863) of treeless alpine and prairie regions and plants. Flahault returned to Sweden on four occasions and became friends with Warming. Flahault lived through the period of transition in which plant ecology evolved from earlier disciplines, and he participated in the progress, while remaining a phytogeographer. He accepted the chair in botany at Montpellier in 1881 and occupied it for 47 years. In 1891 he started a school of Mediterranean phytogeography (Matagne 1999:169–171), and in 1897 he sent to the Académie des Sciences six maps on the phytogeography of the French coastal region, which were not published until 1937. Another persistent subject of investigation was plant succession (Clements 1916:8–32, Egerton 2009). Norwegian Professor of Botany Alex Blytt, at Kristiana (Oslo), in 1876 and later, continued investigations of paleo-vegetational history, pioneered by Japetus Steenstrup (1842, Fries 1950:70, Egerton 2009:49–52), after documentation of the waxing and waning of ice ages provided an opportunity to understand causes of vegetational changes; his conclusions were controversial at the time (Clements 1916:21–22, 382–384). Finnish botanist Ragnar Hult (1857–1899) studied at Uppsala University before receiving his doctorate in 1881 from Helsingfors University (Fries 1950:70, Collander 1965:74–77, Stafleu and Cowan 1976–1988, I:361). His dissertation was on the physiognomy of northern Ostrobothnia and Kemi-Lappmark, in which he distinguished seven vegetation layers. His stratification system was later modified by Sernander and is now known as the Hult-Sernander scale of cover (Becking 1957:415, 419–420). Hult's most important work analyzed "Blekinges Vegetation" (1885) in southern Sweden. This study on the developmental history of Blekinge's plant communities "first fully recognize[d] the fundamental importance of development in vegetation," and made "a systematic study of a region upon this basis" (Clements 1916:22). Hult "traced the succession of each intermediate formation through its various stages to the supposed climax." Clements provided an English summary of Hult's Swedish-language study, and Nicolson (1996:303) quotes passages translated by Sigridur Oladottir. Becking (1957:415) stated that "Hult (1881) published, for the first time, a complete species list with numerical evaluation of cover degree." In summer 1886, Hult taught a field course in plant geography, taken by Swedish student Rutger Sernander, who "adopted Hult's ideas with great enthusiasm" (Fries 1950:70–71, Collander 1965:77). Sernander is discussed below. In the 1890s, three botanists guided the transition from phytogeography to plant ecology: Germans Drude and Schimper and a Dane, Warming (Reed 1942:132–134, Magnin-Gonze 2004:203). (Carl Georg) Oscar Drude (1852–1933) studied under Grisebach and received his doctorate from Göttingen in 1873 (Stafleu and Cowan 1976–1988, I:682–683, Matagne 2009: see index). Drude apparently did not know Haeckel's term "oecologie" when he published Handbuch der Pflanzengeographie (1890, 1977), and his Handbuch was little known in America until after he published Deutschlands Pflanzengeographie (1896), which received an eager audience at the University of Nebraska, mainly from Roscoe Pound and Frederic Clements, who were conducting a botanical survey of Nebraska (Tobey 1981:63–64, McIntosh 1985:28, Nicolson 1989:153–154). Drude provided them with a set of definitions, including "habitat" and "formation," and "a somewhat mechanical theory relating physical and biological factors" that could account for geographical distribution of species. When they obtained Drude's Handbuch, however, they found his vegetational boundaries of the Great Plains inaccurate (Kendeigh 1954:158). "Schouw (1823) proposed a qualitative arrangement of species into groups of 'plantae sociales, gregariae, copiosae, sparsae and solitarine', in which Drude (1890) followed him" (Becking 1957:415). Drude was active in Germany during its flourishing empire, when maps were very important, and he was able to exploit that interest for plant geography maps (Gütler 2011). (a) C. G. Oscar Drude. Web site. (b) Karl F. Schimper. Schimper 1903: frontispiece. (c) J. Eugenius B. Warming. Web site. The first botanist known to have used "oecologie" in print was H. Reiter, Die Consolidation der Physiognomick als Versuch einer Oekologie der Gewaechse (1885). In 1893, the term "ecology" was introduced to two English-speaking audiences: by animal physiologist and professor of medicine John Burdon-Sanderson (1828–1905) in his presidential address (1893:465) to the British Association for the Advancement of Science, and by a Committee on Terminology of Physiology at the Madison Botanical Congress (McIntosh 1985:29). However, not until publication of Warming's Plantesamfund: Grundträk af den okologiske Plantesgeografi (1895, German 1896; extracts from both editions reprinted in Acot 1998, II:827–901). Warming substantially revised his book for the English edition (1909, 1977), utilizing literature that had appeared since 1895, including Schimper's Pflanzengeographie (1898). (Johannes) Eugenius (Bülow) Warming (1841–1924) was from a Frisian island in the North Sea (Christensen 1924–1926, I:617–665, 776–806, Rosenvinge et al. 1927, Müller 1976, Isley 1994:227–229, Matagne 2009: see index). While a student at the University of Copenhagen, he spent three years, 1863–1866, as secretary to a Danish zoologist in Brazil, where Warming studied the flora (Goodland 1975). He received a Ph.D. from the University of Copenhagen in 1871, and he taught in Stockholm, 1882–1885. He became professor of botany and director of the botanic garden at the University of Copenhagen, 1886–1911. Müller (1976) called Warming "founder of plant ecology," and Pascal Acot (1998:675) states: "Historians of biology concur that [Plantesamfund] marked the birth of scientific ecology." (Billings 1985:5 and Nicolson 1996:304–305 cited instead Kerner 1863.) Andreas Schimper (1856–1901) was the son of botanist Wilhelm Schimper, director of the Strasbourg Natural History Museum and, after the Franco-Prussian War, Professor of Natural History at the University of Strasbourg. Andreas received his doctorate in natural philosophy from that university in 1878, then studied under Julius Sachs at Würzburg (Sanders 1975, Cittadino 1990:97–115, Isley 1994:271–273, Matagne 2009: see index). He traveled in North and South America, and in Brazil he studied the symbiotic relationship between ants and Cecropia trees (Schimper 1888). He died in Basel, Switzerland, age 45, from diabetes and from malaria contracted earlier along the African coast. He was a prolific author. For ecology, his most important work was Pflanzengeographie auf physiologischer Grundlage (1898, English, 1903, 870 pages, 502 illustrations, and 4 color maps)—one of the largest ecology monographs written by one author (surpassed much later by G. E. Hutchinson's four-volume Treatise on Limnology). It illustrates the synthesis of phytogeography and physiology that became the foundation of plant ecology. In his preface, Schimper stated (1903:vi) that "The connexion between the forms of plants and the external conditions at different points on the earth's surface forms the subject-matter of oecological plant-geography," and that Grisebach's point of view had become obsolete! Schimper identified a schism in biogeography that persisted well into the 1900s (Worster 1977:194–195, Hagen 1986). Schimper credited Haeckel for naming this ecological science "of biological adaptations." Cowles (1909:356) judged Schimper "a prophet as well as an ecologist of the first magnitude." However, Schimper borrowed heavily from Warming, with scant acknowledgment (Goodland 1975:243). Warming and Schimper's books were more complementary than duplicative (Dajoz 1984:108–111). Schimper's was encyclopedic and well illustrated with many photographs and drawings. Warming's book is more like a textbook, with no illustrations, and half the size of Schimper's. Both books were worldwide in scope, differing in that respect from Kerner's. Warming's chapter 3 (1909:12–13), only 1.5 pages, might be the earliest general discussion of plant communities. It is brief because it is meant to orient one's thinking to the following chapters on different kinds of plant communities. Equally brief is chapter 24 (Warming 1909:83–84), "Symbiosis of Plants with Animals," which merely identifies different kinds of symbiotic relationships, without describing or explaining them. Chapter 25 (Warming 1909:84–91), "symbiosis of plants with one another, mutualism" is more detailed, discussing parasitism, helotism (fungi), mycorhiza and endophytes, epiphytes, saprophytes, and lianes. Chapter 26 (Warming 1909:91–95), "Commensalism, plant-communities," discusses plants that have an aggregate existence but no interactions except a common struggle for life requirements. A historian of symbiosis, Jan Sapp (1994:30) faults Warming for only devoting nine pages to the subject, but if one counts all the pages in Section II, "Communal Life of Organisms," (chapters 22–26), it is 14 pages, and Warming mentioned many biologists who had published on the subject. Sapp also faults him for too strong an emphasis on struggle for existence and skepticism about reciprocity between species. Chapter 34 (Warming 1909:131–136), "Oecological classification," divides formations into 13 groups, each of which are discussed in following chapters: hydrophytes, helophytes, oxylophytes, psychrophytes, halophytes, lithophytes, psammophytes, chersopohytes, eremophytes, psilophytes, sclerophyllous, coniferous, and mesophytes. If some of these terms are unfamiliar to modern readers, it shows that not all of Warming's suggested terms were adopted by other plant ecologists. Chapter 35 (Warming 1909:137–148), "Physiognomy of vegetation, formations, associations, varieties of associations," insists, with no substantial evidence, that physiognomy (landscape appearance) is of scientific interest and is determined by seven characteristics: dominant growth forms, density, height, and color of vegetation, seasonal relationships, duration of species life, and number of species. The term "formation" had been introduced by Grisebach (1838), but had come to mean for Warming (1909:140), "a community of species, all belonging to definite growth-forms which have become associated together by definite external (edaphic or climatic) characters of the habitat…" If that seems a bit vague, his six examples of formations are not: microphyte, moss, herb, dwarf-shrub and undershrub, bush-wood or shrub-wood, and forest. But then he discusses simple formations, compound formations, secondary formations, and sub-formations. Humboldt had introduced the term "plant association" in 1807. For Warming (1909:145), "An association is a community of definite floristic composition within a formation; it is, so to speak, a floristic species of a formation which is an oecological genus." An association may occur as patches in a formation or exhibit zonal arrangement. But there are also varieties of association, such as edaphic or geographical varieties. By proposing straightaway in his Projet in 1900 that nomenclature must apply to geographical and topographical as well as to biological units, Flahault situated the problem clearly within the framework of the ecological approach to the relationship between living beings and the environment. However, Flahault's system was not definitive either, and the Second International Botany Congress (1905) charged him and German-born Swiss botanist Carl (Joseph) Schröter (1855–1939) with heading a committee to establish terminology "that integrated the floristic and physiognomic components of botanical geography" (Matagne 1998:436) for the Third International Botany Congress (1910). Schröter had studied botany at the University of Zurich under Professor Oswald Heer (1809–1883), who also taught entomology. Schröter succeeded his mentor and was Professor of Botany at the University of Zurich from 1879 to 1925; he coined the terms "autecologie" (1896) and "synecologie" (1902), according to Chapman (1931:5–6). Furthermore, Schröter's Planzenleben der Alpen (1904–1908) "was the start of an understanding of relationships between the growth of high mountain plants and their patterned and severe environments" (Billings 1985:6). His book "contains most of the knowledge of the time on the influence of elevation, soil, climate, and other environmental factors on alpine plants and plant communities" (Gigon et al. 1981:314). At last, Europe had a worthy companion to Kerner's Das Pflanzenleben der Danaulaender (1863). However, Schröter's monograph has 1288 pages, and no one has translated it into English. Although the 1910 Congress did not fully accept Schröter and Flahault's proposals, in 1913 their proposal provided the basis for the Zurich-Montpellier school of phytosociology (Becking 1957:426–463, Shimwell 1971:45–47, Lévêque 2003:18–19). Yet their methods "were not applicable everywhere, especially in Scandinavia, with its limited flora" (Becking 1957:419–426, Matagne 1998:437). Therefore, other regional schools formed, and a uniformity of concepts and methods became an elusive dream. Josias Braun-Blanquet (1884–1980), Swiss, was a student of both Flahault and Schröter, and he became Flahault's successor at Montpellier (Nicholson 1989:142–143, 146–148, 181). His important influence came from publication of his Pflanzensociology (1928), which is beyond the scope of this part 48 survey, but he had begun publishing his version of phytosociology in 1913. (a) Carl J. Schröter. Web site. (b) Leonard Cockayne. Thomson 1983. (c) Józef Paczoski. Web site. Three modern plant ecologists responded to this awkward nomenclature-classification situation by explaining the different systems and seeking some common understanding. A Dutch plant ecologist working in the United States, Rudy Becking (1957), explained the Zurich-Montpellier system to Americans, but found he also needed to explain competing systems. Robert H. Whittaker, at Brooklyn College, took a similar approach, with more details, in his "Classification of Natural Communities" (1966, 1977). English phytogeographer David W. Shimwell thought it would help to go into even more detail than either Becking or Whittaker in his Description and Classification of Vegetation (1971). Phytogeography did not fade away with the growth of plant ecology. German systematic botanist H. G. Adolf Engler (1844–1930) collaborated with Drude in editing Die Vegetation der Erde (15 volumes, 1896–1923), to which Engler contributed volume 9: Die Pflanzenwelt Afrikas (1910–1915). Engler recruited other botanists to write volumes on parts of the world matching their expertise (Stafleu and Cowan 1976–1988, I:757–797, Stafleu 1978). An example of this recruitment was Leonard Cockayne (1855–1934), whom Engler recruited to write the volume on New Zealand (Hill 1935, Moore 1967, Thomson 1983). Cockayne was from Derbyshire, England, and he had attended Owen's College, Manchester, 1872–1874, then immigrated to Australia in 1876 and taught school there until 1880, when he went to New Zealand, where he continued teaching for four or five years. Afterwards, he bought land for farming and gardening. In 1877, he obtained a book on New Zealand ferns by G. M. Thomson, which turned his attention to native plants. He decided to devote his life to gardening and New Zealand botany. "The Cockayne period in New Zealand botany may be said to have begun in 1896 when Diels of Berlin published his 100-page account of Vegetations-Biologie von Neu-Seeland" (Moore 1967:3). Diels had not been to New Zealand (!) and depended upon accounts which Cockayne sent him. Diel's publication aroused the interest of Prof. Karl Ritter von Gobel, Munich University, who visited New Zealand in 1898, and Cockayne took him around the country, showing him the plants and their environments. This expedition of several weeks was mutually beneficial, and in 1903 Gobel had his university confer upon Cockayne an honorary Ph.D. For his part, Gobel conveyed to Cockayne the ecological perspective that was developing in Europe, and Cockayne read Warming's text, in German (Star 2006:198). In 1890 there was an extensive fire in a subalpine beech forest near Arthur's Pass (930 m). In 1898 Cockayne published "On the burning and reproduction of subalpine scrub and its associated plants with special reference to Arthur's Pass District." That study initiated "the longest recorded and best documented example of plant succession in [New Zealand's] ecological literature" (Mark 1981:424). He also published on an ecological experiment in 1898, and in 1900 he published "A sketch of the plant geography of the Waimakariri River Basin, considered chiefly from an Oecological Point of View." He published a series of ecological surveys, then synthesized them into New Zealand Plants and Their Story (1910, edition 3, 1927). However, his earlier papers and Gobel's conferring of an honorary degree on him attracted Engler's attention, and he asked Cockayne to write the New Zealand volume of Die Vegetation der Erde. Cockayne was thrilled and reported to a friend in May, 1904: "Of course it is a most high honour for a colonial botanist to be invited to contribute towards such a series, where every work is supposed to be of the highest excellence…" (quoted in Moore 1967:5). He sent in his manuscript in 1914 to Leipzig, and then worried throughout World War I whether it would ever be published. After the dust and ashes of war had settled, The Vegetation of New Zealand appeared (Cockayne 1921, 364 pages, edition 2, 1928, 456 pages). Cockayne used his ecological knowledge to persuade New Zealand's government to establish or enlarge national parks (Star 2006:198). Besides his books, Cockayne published 184 articles, 40 being on ecology and 25 on plant distributions (Thomson 1982,1983:15–17). In 1932, he and J. D. Calder published "The Present Vegetation of Arthur's Pass (New Zealand) as Compared with that of Thirty-Four Years Ago." Another example of Engler's recruitment was of John Harshberger to write the North American volume (see below). The 15 volumes that Engler and Drude edited was a monumental climax for a century of phytogeography, which began with Humboldt. Phytosociology arose in the 1890s from the work of Jozef Konradovich Paczoski (or Pachoskii 1864–1942), born in Bialograd, southwestern Ukraine to Polish parents (Maycock 1967:1031, Kajak and Pieczyska 1981:287, Kozlowski and Sevedrzyuski 2009). He attended the School of Agriculture and Horticulture in Human, then the University of Kiev, 1888–1894. He was assistant curator of the Botanical Garden, Leningrad, 1894–1895, assistant in the School of Agriculture in Dublany, 1895–1897, director of the Natural History Museum in Cherson, 1897–1920, director of Steppe Reserve at Ascania Nova, 1922–1923, director of the National Park of Bialowieza, Poland, 1923–1928, and professor of plant systematics, sociology and geography at University of Poznan, 1925–1931, where he founded the first Institute of Plant Sociology. …which includes a definition of plant sociability, exclusion due to competition, simple and complex communities, the mutual exclusion principle, plant succession at the autecological level, root layering and competition, seasonal competition and aspection in steppe communities, dominance and subdominance, characteristics of community boundaries, community complexity and variability, influence of vegetation on soils, soil moisture and microclimate, shade tolerance, tolerance ratings of forest trees, tree reproduction and forest succession as related to tolerance, protophilous and sciophilous species, mutual intolerance, fire succession, the organismic concept, the plant formation and lower phytosocial groupings, actual and climatic limits of a species, restriction of ecological amplitude due to competition, etc. In 1921, Paczoski synthesized his studies in The Principles of Phytosociology (346 pages, in Russian), and applied these principles in his "Phytosociology and Forestry" (1925, in Polish). A controversial aspect of his concept was drawing an analogy between plan