Eugenius Warming

Johannes Eugenius Bülow Warming (November 3, 1841 - April 2, 1924), known as Eugen Warming, was a Danish botanist and a main founding figure of the scientific discipline of ecology. Warming wrote the first textbook (1895) on plant ecology, taught the first university course in ecology and gave the concept its meaning and content. “If one individual can be singled out to be honoured as the founder of ecology, Warming should gain precedence” (Goodland 1975).

Warming wrote a number of textbooks on botany, plant geography and ecology, which were translated to several languages and were immensely influential at their time and later. Most important were Plantesamfund and Haandbog i den systematiske Botanik.

Life
Warming was born on the small Wadden Sea island of Mandø as the only child of Jens Warming (1797-1844), parish minister, and Anna Marie von Bülow af Plüskow (1801-1863). After the early death of his father, he moved with his mother to her brother in Vejle in eastern Jutland. He attended highschool at Ribe Katedralskole and commenced 1859 studies of natural history at the University of Copenhagen, but left university for three-and-a-half year (1863-1866) to act as secretary for the Danish palaeontologist Peter Wilhelm Lund, who lived and worked in Lagoa Santa, Brasil. He finished his Dr.Phil. in 1871. He was passed over when the professorship in botany became vacant with the death of A.S. Ørsted. He then was a docent of botany at the University of Copenhagen, Polyteknisk Læreanstalt and the Pharmaceutical College 1873-1882. He became professor in botany at Stockholms högskola (later Stockholm University) 1882-1885. As the eldest professor, he was elected vice chancellor. In 1885, he became professor in botany at the University of Copenhagen and director of the Copenhagen Botanical Garden and held these positions until his retirement in 1911. As such, rector magnificus of University of Copenhagen 1907-1908. He was a member of the Royal Danish Academy of Sciences and Letters from 1878 to his death. As such, he served as a board member of the Carlsberg Foundation 1889-1921 and, because a biologist, on the board of the Carlsberg Laboratory. He also served on the board of the Geological Survey of Denmark 1895-1917. Eugen Warming was president for the ‘Association internationale des botanistes‘ (1913), honorary fellow of the Royal Society in London and honorary member of the Danish Botanical Society. He was made Commander 1st Degree of the Order of the Dannebrog, Commander of the Royal Victorian Order and the Brazilian Imperial Order of the Rose.

Expeditions: Brasil, Lagoa Santa: 1863-1866 Greenland: 1884 Norway, Finnmark: 1885 Norway, Dovre: 1887 Venezuela, Trinidad and the Danish West Indies: 1891-1892 Faroe Islands: 1895

He married Johanne Margrethe Jespersen (known as Hanne Warming; 1850-1922) on November 10, 1871. They had eight children: Marie (1872-1947) married C.V. Prytz, Jens Warming (1873-1939), who became a professor in economy and statistics at the Royal Veterinary and Agricultural College, Fro (1875-1880), Povl (1877-1878), Svend Warming (1879-1982), engineer at Burmeister & Wain shipyard, Inge (1879-1893), Johannes (1882-1970), farmer, and Louise (1884-1964).

External link: Ancestors and descendents

'Plantesamfund' or 'Oecology of Plants'
The book 'Plantesamfund' was based on Warming’s lectures on plant geography at the University of Copenhagen. It gives an introduction to all major biomes of the world. Warming’s aim, and his major lasting impact on the development of ecology, was to explain how nature solved similar problems (drought, flooding, cold, salt, herbivory etc.) in similar way, despite using very different ‘raw material’ (species of different decent) in different regions of the world. This was a remarkably modern view – completely different from the merely descriptive floristic plant geography prevailing at his time. The book was translated to German in 1896 as This edition, which was approved by Warming, rapidly ran out of print. A second, unauthorized, edition was issued in 1902 by Paul Graebner, who put his own name after Warming’s on the book’s frontispiece, despite no changes to the contents (Goodland 1975). This edition was expanded in third and fourth editions: A Polish translation of ’Plantesamfund’ (from Knoblauch’s German translation) appeared in 1900: Two independent Russian (Moscow and St. Petersburg) editions appeared in 1901 and 1903 An extended and translated edition in English first apperead in 1909: The German ecologist A. F. W. Schimper published ”Pflanzengeographie auf physiologisher Grundlage” in 1898. "'This work not only covered much of the same ground as Warming did in 1895 and 1896 but in fact also leaned heavily on Warming’s research. Schimper (1898) quoted extensively from more than fifteen of Warming’s works and even reproduced Warming’s figures. Yet nowhere did Schimper acknowledge his profound debt to Warming, neither in the list of picture credits, nor in the acknowledgements section of the Vorwort, nor in his list of major sources, and not even in a footnote! ... Although replete with Warming’s data, it contains few ideas and did not advance ecology beyond what Warming had done earlier.” (Goodland 1975)"
 * Warming, E. (1895) Plantesamfund - Grundtræk af den økologiske Plantegeografi. P.G. Philipsens Forlag, Kjøbenhavn. 335 pp.
 * Lehrbuch der ökologischen Pflanzengeographie - Eine Einführung in die Kenntnis der Pflanzenverenie by Emil Knoblauch. Berlin, Gebrüder Borntraeger, 1896. 412 pp.
 * Warming, E. & Graebner, P. (1918) Eug. Warming's Lehrbuch der ökologischen Pflanzengeographie, 3 ed. Berlin, Gebrüder Borntrager. Fourth edn (1933) - 1158 pp.
 * Warming, E. (1900) Zbiorowiska Roślinne zarys ekologicznej geografii roślin by Edward Strumpf and Jósef Trzebiński. Warszawa, 1900. 451 pp.
 * Вармингъ, Е. (1901) Ойкологическая географія растеній – Введеніе въ изученіе растительныхъ сообществъ by M. Golenkin and W. Arnol'di. Moskva, 542 pp. Full text link
 * Вармингъ, Е. (1903) Распредъленіе растений въ зависимости отъ внъшнихъ условій - Экологическая географія растеній by A. G. Henkel' and with a treatise of the vegetation of Russia by G. I. Tanfil'ev. St. Petersburg, 474 pp.
 * Warming, E. with M. Vahl (1909) |Oecology of Plants - an introduction to the study of plant-communitiesby P. Groom and I. B. Balfour. Clarendon Press, Oxford. 422 pp. (2nd edn 1925).

Haandbog i den systematiske Botanik
Warming, E. (1878) Haandbog i den systematiske Botanik (nærmest til Brug for Universitets-Studerende og Lærere). København. (2nd edn 1884; 3rd ed with Algae by N. Wille and fungi by E. Rostrup 1891). German edn 1890: Handbuch der systematischen Botanik by E. Knoblauch (2nd edn 1902, 3rd edn 1911, 4th edn 1929 all by M. Möbius). Russian edn 1893: Систематика растеній (from the 3rd Danish edn by S. Rostovzev and M. Golenkin; 2nd edn 1898). English edn 1895: A handbook of systematic botany (by M.C. Potter; several editions, latest 1932). The section on seed plants was later expanded and issue as
 * Warming Eug. (1912) Frøplanterne (Spermatofyter). Kjøbenhavn, Gyldendalske Boghandel/Nordisk Forlag. 467 pp. (2nd edn 1933).

Warming Eug. Den almindelige Botanik: En Lærebog, nærmest til Brug for Studerende og Lærere. Kjøbenhavn, 1880. (2rd edn 1886; 3rd edn by Warming and Wilhelm Johannsen 1895; 4th edn by Warming and Johannsen 1900-01). Swedish edn 1882: Lärobok i allmän botanik (by Axel N. Lundström). German edn 1907-09: Lehrbuch der allgemeinen Botanik (from the 4th edn, by E. P. Meinecke). Berlin, Borntraeger. 667 pp.

Warming Eug. (1900) Plantelivet: Lærebog i Botanik for Skoler og Seminarier. København. (2nd edn 1902; 3rd edn 1905; 4th edn by C. Raunkiær and Warming 1908). English edn 1911: Plant Life - A Text-book of Botany for Schools and Colleges (from the 4th edn by M.M. Rehling and E.M. Thomas). London. Russian edn 1904: Растение и его жизнь (Началный учебник ботаники). (from the 2nd edn by L.M. Krečotovič and M. Golenkin). Moskva.

These works show Warming as a skillful and dedicated pedagog, whose presentation of the subject was useful far beyond his lecture theatre in Copenhagen.

Floral development of Euphorbia
Warming's doctoral thesis (in Danish) dealt with ontogeny of the cyathia of Euphorbia (Euphorbiaceae). Part of the work was published in German the year before: Both these works a still cited in scientific papers by scholars of botany every now and then.
 * Warming, J. Eug. B. 1871. Koppen hos Vortemælken en Blomst eller en Blomsterstand? En organogenetisk morfologisk Undersøgelse. Copenhagen, G.E.C. Gad.
 * Warming, E. 1870. Über die Entwicklung des Blütenstandes von Euphorbia. Flora 53: 385-397.

Lagoa Santa
His early experience with vegetation in a tropical region was decisive for his future work. His collections from Lagoa Santa, 2600 plant species, of which some 370 turned out to be new to scinece (Goodland 1975), were treated in a monumental 40-volume and 1400-page work, Symbolæ ad Floram Brasiliæ centralis cognoscendam. For this work, Warming farmed out plant families to more than fifty plant taxonomists around Europe. They were all published as volumes in the series ’Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn’.
 * Symbolæ ad Floram Brasiliæ centralis cognoscendam, particulæ 1-10, 1873
 * Symbolæ 11-20, 1875
 * Symbolæ 21-30, 1886
 * Symbolæ 31-40, 1893
 * Symbolæ 31-40, 1893.

Having finished the taxonomical work, Warming finally published his ecological study of plant communities in the Lagoa Santa area, with cerrado as the main vegetation type.

Warming, E. (1892) Lagoa Santa: Et Bidrag til den biologiske Plantegeografi med en Fortegnelse over Lagoa Santas Hvirveldyr. Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 6. Rk. vol. 6 (3): 153-488.

Warming issued a lengthy summary in French (1893): Lagoa Santa – Étude de Geographie Botanique. Revue Générale de Botanique 5: 145-158, 209-233.

Portuguese translation: Warming, Eugenio Lagoa Santa: Contribuição para a geographia phytobiologica, by Alberto Löfgren Belo Horizonte, 1908. This edition was augmented by the Brazilian ecologist M.G. Ferri with more recent research on the cerrado system and reissued as: Warming, E. & Ferri, M.G. (1973) Lagoa Santa – a vegetação de cerrados brasileiros. University of São Paulo.

Life forms
1-105. These works may be seen as precursors of the life-form scheme of Warming’s pupil C. Raunkiær. However, Warming’s scheme was more complicated, taking other environmental factors than wintering into account, especially water/drought stress. Warming did not approve of what he saw as over-simplification in Raunkiær’s scheme. Warmings last published work was a renewed attempt to put all plant (including bacteria and algae) life forms into a system.
 * Warming, E. (1884) Om Skudbygning, Overvintring og Foryngelse (On shoot architecture, perennation and rejuvenation). Naturhistorisk Forenings Festskrift:
 * Warming, E. (1908) Om planterigets livsformer (On the life forms in the vegetable kingdom). G.E.C. Gad, København.
 * Warming, E. (1923) Økologiens Grundformer – Udkast til en systematisk Ordning (Fundamental ecological forms - draft for a system). Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 8. Rk., vol. 4: 120-187.

Structure and biology of arctic flowering plants
Warming, E. ed. (1908-1921) The structure and biology of Arctic flowering plants.
 * Meddelelser om Grønland, 36,
 * Warming, E. (1908) 1. Ericinæ (Ericaceae, Pirolaceae). 1. Morphology and biology, p. 1-71.
 * Petersen, H. E. (1908) 1. Ericinæ (Ericaceae, Pirolaceae). 2. The biological anatomy of them leaves and stems, p. 73-138.
 * Petersen, H.E. (1908) 2. Diapensiaceae. Diapensia lapponica, p. 139-154.
 * Mentz, A. (1909) 3. Empetraceae. Empetrum nigrum, p. 155-167.
 * Warming, E. (1909) 4. Saxifragaceae. 1. Morphology and biology, p. 169-236.
 * Galløe, O. (1909) 4. Saxifragaceae. 2. The biolocical leaf-anatomy of the Arctic species of Saxifraga, p. 237-294.
 * Seidelin, A. (1909) 5. Hippuridaceae, Halorrhagidaceae and Callitrichaceae, p. 295-332.
 * Jessen, K. (1909) 6. Ranunculaceae, p. 334-440.
 * Heide, F. (1909) 7. Lentibulariaceae, 441-481.
 * Meddelelser om Grønland, 37,
 * Jessen, K. (1913) 8. Rosaceæ, p. 1-126.
 * Olsen, C. (1914) 9. Cornaceae, p. 127-150.
 * Hagerup, O. (1915) 10. Caprifoliaceae. Linnaea borealis L., p. 151-164.
 * Mathiesen, F.J. (1916) 11. Primulaceae, p. 165-220.
 * Warming, E. (1920) 13. Caryophyllaceæ, p. 229-342.
 * Porsild, M.P. (1920) 14. Liliales, p. 343-358.
 * Mathiesen, F.J. (1921) 15. Scrophulariaceae, 359-507.

Iceland and Faroe Islands
Rosenvinge, L. Kolderup &  Warning, E. (eds) (1912-1945) The Botany of Iceland, vol. 1-3. Copenhagen, J. Frimodt. 675, 607 and 547 pp. Warming, E. (1901) Botany of the Faeroes (based upon Danish investigations), vol. I-III. London.

His favourite plant family - Podostemaceae
Warming, E	(1881-1899) Familien Podostemaceae - Etudes sur la famille des Podostemacees.
 * I. 1881 Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 6. Rk., vol. II (1)
 * II. 1882. Kong. Dan. Vid. Selsk. Skr. – Naturv. Math. Afd., 6. Rk., vol. II. (3)
 * III. 1888 Kong. Dan. Vid. Selsk. Skr. – Naturv. Math. Afd., 6. Rk., vol. IV (8)
 * IV. 1891. Kong. Dan. Vid. Selsk. Skr. – Naturv. Math. Afd., 6. Rk., vol. VII (4)
 * V. 1899. Kong. Dan. Vid. Selsk. Skr. – Naturv. Math. Afd., 6. Rk., vol. IX (2)
 * VI. 1901. Kong. Dan. Vid. Selsk. Skr. – Naturv. Math. Afd., 6. Rk., vol. XI (1)

Vegetation of Denmark

 * Warming, E. 1906. Dansk Plantevækst. 1. Strandvegetationen. - Gyldendalske Boghandel Nordisk Forlag. [beach vegetation]
 * Warming, E. 1909. Dansk Plantevækst. 2. Klitterne. - Gyldendalske Boghandel Nordisk Forlag. [dunes]
 * Warming, E. 1917. Dansk Plantevækst. 3. Skovene. - Gyldendalske Boghandel Nordisk Forlag. [forests]

Warming’s influence
"It was Eugenius Warming's Lehrbuch der ökologischen Pflanzengeographie that must be considered as the starting point of self-conscious ecology. This book was the first to use physiological relations between plants and their environment, and in addition biotic interactions to explain the moulding of the assemblages that plant geographers had described and classified, and it would set up a research agenda for decades to come. Jax (2001)"

Despite the language barrier, Warming’s influence on the development of ecology is remarkable, not the least in Britain and the USA. The British ecologist Arthur Tansley was extremely influenced by reading ’Plantesamfund’ (or rather the 1896 German edition). Reading the book made him jump from anatomy to ecology (Goodland 1975).

"The German translation was widely read in England and America and played an important part in stimulating fieldwork in both, countries. It certainly did in my own case: I well remember working through it with enthusiasm in 1898 and going out into the field to see how far one could match the plant communities Warming had described for Denmark in the English countryside; and I also made the book the basis of a course of University Extension lectures at Toynbee Hall in 1899. (Tansley 1947)"

Similarly, Warming's book impregnated North American naturalists like Henry Chandler Cowles and Frederic Clements (Coleman 1986). Cowles appear to have been completely taken: "Charles J. Chamberlain, who attended Coulter’s lectures as a student and later joined the University of Chicago faculty, recalled in a memoir that ‘none of us could read Danish except a Danish student, who would translate a couple of chapters, and the next day Coulter would give a wonderful lecture on Ecology ... Cowles, with his superior knowledge of taxonomy and geology, understood more than the rest of us, and became so interested that he studied Danish and, long before any translation appeared, could read the book in the original ... The treatment of such sand dunes as Warming knew, started Henry on his study of the comparatively immense moving dunes south of the University (Cassidy 2007)."

Warming’s influence on later Scandinavian ecology was immense. Especially significant was his inspiration to Christen Raunkiær – his pupil and successor on the chair of botany at the University of Copenhagen.

Miscellaneous
The standard botanical author abbreviation Warm. is applied to species he described.

The Orchid genus Warmingia and dozens of species has been named to his honour.

Universidade Federal de Minas Gerais has organized a series of 'Eugen Warming lectures in Evolutionary Ecology' since 1994.