Eugen Warming
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Johannes Eugenius Bülow Warming (3 November 1841 – 2 April 1924), known as Eugen Warming, was a Danish botanist and a main founding figure of the scientific discipline of
ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
. Warming wrote the first textbook (1895) on
plant ecology Plant ecology is a subdiscipline of ecology that studies the distribution and abundance (ecology), abundance of plants, the effects of environmental factors upon the abundance of plants, and the interactions among plants and between plants and ...
, taught the first university course in ecology and gave the concept its meaning and content. Scholar R. J. Goodland wrote in 1975: “''If one individual can be singled out to be honoured as the founder of ecology, Warming should gain precedence''”.Goodland, R.J. (1975) The tropical origin of ecology: Eugen Warming’s jubilee. ''Oikos'', 26, 240-245.
/ref> 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''.


Early life and family life

Warming was born on the small
Wadden Sea The Wadden Sea ( ; ; or ; ; ; ) is an intertidal zone in the southeastern part of the North Sea. It lies between the coast of northwestern continental Europe and the range of low-lying Frisian Islands, forming a shallow body of water with tida ...
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 Vejle () is a city in Denmark, in the southeast of the Jutland Peninsula at the head of Vejle Fjord, where the Vejle River and Grejs River and their valleys converge. It is the site of the councils of Vejle Municipality (''Municipalities of De ...
in eastern
Jutland Jutland (; , ''Jyske Halvø'' or ''Cimbriske Halvø''; , ''Kimbrische Halbinsel'' or ''Jütische Halbinsel'') is a peninsula of Northern Europe that forms the continental portion of Denmark and part of northern Germany (Schleswig-Holstein). It ...
. He married Johanne Margrethe Jespersen (known as Hanne Warming; 1850–1922) on 10 November 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 University The Royal Veterinary and Agricultural University (, abbr. KVL, , abbr. RVA) was a veterinary and agricultural science university in Denmark. It was founded in 1856 and operated until 2007, when it became a part of the University of Copenhagen. It ...
, 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).Prytz, S. (1984) ''Warming – botaniker og rejsende''. Lynge, Bogan. 197 pp. A personal account by Warming’s granddaughter, based on family-owned letters. External link: Ancestors and descendents


Education and career

He attended high school at Ribe Katedralskole and commenced 1859 studies of natural history at the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
, but left university for three and a half years (1863–1866) to act as secretary for the Danish
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Peter Wilhelm Lund Peter Wilhelm Lund (14 June 1801 – 25 May 1880) was a Danish Brazilian paleontologist, zoologist, and archeology, archeologist. He spent most of his life working and living in Brazil. He is considered the father of Brazilian paleontology as wel ...
, who lived and worked in Lagoa Santa,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. After his return to
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, he studied for a year under K.F.P. Martius,
Carl Nägeli Carl Wilhelm von Nägeli (26 or 27 March 1817 – 10 May 1891) was a Swiss botanist. He studied cell division and pollination but became known as the man who discouraged Gregor Mendel from further work on genetics. He rejected natural selecti ...
and Ludwig Radlkofer in
Munich Munich is the capital and most populous city of Bavaria, Germany. As of 30 November 2024, its population was 1,604,384, making it the third-largest city in Germany after Berlin and Hamburg. Munich is the largest city in Germany that is no ...
and, in 1871, under J.L. von Hanstein in
Bonn Bonn () is a federal city in the German state of North Rhine-Westphalia, located on the banks of the Rhine. With a population exceeding 300,000, it lies about south-southeast of Cologne, in the southernmost part of the Rhine-Ruhr region. This ...
. Later in the same year (1871), he defended his
Doctor of Philosophy A Doctor of Philosophy (PhD, DPhil; or ) is a terminal degree that usually denotes the highest level of academic achievement in a given discipline and is awarded following a course of Postgraduate education, graduate study and original resear ...
thesis at
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
. The professorship in botany at the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
became vacant with the death of A.S. Ørsted and Warming was the obvious candidate for a successor. However, he was passed over and the chair given to the older, but much less productive and original Ferdinand Didrichsen. Warming then became docent of
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
at the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
, the polytechnic ( Polyteknisk Læreanstalt) and the Pharmaceutical College 1873–1882. He became professor in botany at ''Stockholms högskola'' (later
Stockholm University Stockholm University (SU) () is a public university, public research university in Stockholm, Sweden, founded as a college in 1878, with university status since 1960. With over 33,000 students at four different faculties: law, humanities, social ...
) 1882–1885. As the eldest professor, he was elected
rector magnificus A rector (Latin for 'ruler') is a senior official in an educational institution, and can refer to an official in either a university or a secondary school. Outside the English-speaking world, the rector is often the most senior official in a u ...
. In 1885, he became professor in botany at the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
and director of the Copenhagen Botanical Garden and held these positions until his retirement on 31 December 1910. He was
rector magnificus A rector (Latin for 'ruler') is a senior official in an educational institution, and can refer to an official in either a university or a secondary school. Outside the English-speaking world, the rector is often the most senior official in a u ...
of the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
1907–1908. He was a member of the
Royal Danish Academy of Sciences and Letters The Royal Danish Academy of Sciences and Letters ({{Langx, da, Det Kongelige Danske Videnskabernes Selskab or ''Videnskabernes Selskab'') is a Danish academy of science. The Royal Danish Academy was established on 13 November 1742, and was create ...
from 1878 to his death. As such, he served on the board of directors of the
Carlsberg Foundation Carlsberg Foundation () is a not-for-profit organization that was founded by J. C. Jacobsen in 1876, by allocating some of his shares in the Carlsberg Brewery to fund and operate the Carlsberg Laboratory and the Museum of National History at ...
1889-1921 and, because a biologist, on the board of the
Carlsberg Laboratory The Carlsberg Research Laboratory is a private scientific research center in Copenhagen, Denmark under the Carlsberg Foundation. It was founded in 1875 by J. C. Jacobsen, the founder of the Carlsberg brewery, with the purpose of advancing bioche ...
. He also served on the board of the Geological Survey of Denmark 1895–1917. Eugen Warming was a frequent visitor to foreign universities, e.g. a travel to
Strasbourg Strasbourg ( , ; ; ) is the Prefectures in France, prefecture and largest city of the Grand Est Regions of France, region of Geography of France, eastern France, in the historic region of Alsace. It is the prefecture of the Bas-Rhin Departmen ...
and
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, largest city of France. With an estimated population of 2,048,472 residents in January 2025 in an area of more than , Paris is the List of ci ...
in 1876 and another to
Göttingen Göttingen (, ; ; ) is a college town, university city in Lower Saxony, central Germany, the Capital (political), capital of Göttingen (district), the eponymous district. The River Leine runs through it. According to the 2022 German census, t ...
,
Jena Jena (; ) is a List of cities and towns in Germany, city in Germany and the second largest city in Thuringia. Together with the nearby cities of Erfurt and Weimar, it forms the central metropolitan area of Thuringia with approximately 500,000 in ...
,
Bonn Bonn () is a federal city in the German state of North Rhine-Westphalia, located on the banks of the Rhine. With a population exceeding 300,000, it lies about south-southeast of Cologne, in the southernmost part of the Rhine-Ruhr region. This ...
,
Strasbourg Strasbourg ( , ; ; ) is the Prefectures in France, prefecture and largest city of the Grand Est Regions of France, region of Geography of France, eastern France, in the historic region of Alsace. It is the prefecture of the Bas-Rhin Departmen ...
and
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, largest city of France. With an estimated population of 2,048,472 residents in January 2025 in an area of more than , Paris is the List of ci ...
in 1880. He participated in several Scandinavian Scientist Conferences between 1868 and 1916 and in the similar
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
meeting in Breslau in 1874. He joined the
International Botanical Congress International Botanical Congress (IBC) is an international meeting of Botany, botanists in all scientific fields, authorized by the International Association of Botanical and Mycological Societies (IABMS) and held every six years, with the locatio ...
es in
Amsterdam Amsterdam ( , ; ; ) is the capital of the Netherlands, capital and Municipalities of the Netherlands, largest city of the Kingdom of the Netherlands. It has a population of 933,680 in June 2024 within the city proper, 1,457,018 in the City Re ...
1877, in
Vienna Vienna ( ; ; ) is the capital city, capital, List of largest cities in Austria, most populous city, and one of Federal states of Austria, nine federal states of Austria. It is Austria's primate city, with just over two million inhabitants. ...
1905 and in
Brussels Brussels, officially the Brussels-Capital Region, (All text and all but one graphic show the English name as Brussels-Capital Region.) is a Communities, regions and language areas of Belgium#Regions, region of Belgium comprising #Municipalit ...
1910 and was president of the ‘Association internationale des botanistes‘ (1913). He attended the
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
celebration in
Uppsala Uppsala ( ; ; archaically spelled ''Upsala'') is the capital of Uppsala County and the List of urban areas in Sweden by population, fourth-largest city in Sweden, after Stockholm, Gothenburg, and Malmö. It had 177,074 inhabitants in 2019. Loc ...
1907 and the Darwin celebration in
London London is the Capital city, capital and List of urban areas in the United Kingdom, largest city of both England and the United Kingdom, with a population of in . London metropolitan area, Its wider metropolitan area is the largest in Wester ...
1908. He was honorary fellow of the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
in London, was elected member of the
Royal Swedish Academy of Sciences The Royal Swedish Academy of Sciences () is one of the Swedish Royal Academies, royal academies of Sweden. Founded on 2 June 1739, it is an independent, non-governmental scientific organization that takes special responsibility for promoting nat ...
in 1885 and honorary member of the Danish Botanical Society. He was a corresponding member of the botanical section of the
French Academy of Sciences The French Academy of Sciences (, ) is a learned society, founded in 1666 by Louis XIV at the suggestion of Jean-Baptiste Colbert, to encourage and protect the spirit of French Scientific method, scientific research. It was at the forefron ...
. He was made Commander 1st Degree of the
Order of the Dannebrog The Order of the Dannebrog () is a Denmark, Danish order of chivalry instituted in 1671 by Christian V of Denmark, Christian V. Until 1808, membership in the Order was limited to fifty members of noble or royal rank, who formed a single cla ...
, Commander of the
Royal Victorian Order The Royal Victorian Order () is a dynastic order of knighthood established in 1896 by Queen Victoria. It recognises distinguished personal service to the monarch, members of the royal family, or to any viceroy or senior representative of the m ...
and the Brazilian Imperial Order of the Rose. He is buried in Assistants Cemetery in
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
.


Expeditions

* 1863-1866:
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
, Lagoa Santa * 1884
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
( Fylla expedition - view images on arktiskebilleder.dk * 1885:
Norway Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
,
Finnmark Finnmark (; ; ; ; ) is a counties of Norway, county in northern Norway. By land, it borders Troms county to the west, Finland's Lapland (Finland), Lapland region to the south, and Russia's Murmansk Oblast to the east, and by water, the Norweg ...
* 1887: Norway,
Dovre Dovre is a List of municipalities of Norway, municipality in Innlandet county, Norway. It is located in the Traditional districts of Norway, traditional district of Gudbrandsdal. The administrative centre of the municipality is the Dovre (villag ...
* 1891-1892:
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
,
Trinidad Trinidad is the larger, more populous island of the Republic of Trinidad and Tobago, the country. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is the southernmost island in ...
and the
Danish West Indies The Danish West Indies () or Danish Virgin Islands () or Danish Antilles were a Danish colony in the Caribbean, consisting of the islands of Saint Thomas with , Saint John () with , Saint Croix with , and Water Island. The islands of St ...
* 1895:
Faroe Islands The Faroe Islands ( ) (alt. the Faroes) are an archipelago in the North Atlantic Ocean and an autonomous territory of the Danish Realm, Kingdom of Denmark. Located between Iceland, Norway, and the United Kingdom, the islands have a populat ...
In addition, shorter visits to the
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
and other proximate destinations.


''Plantesamfund'' ('Oecology of plants')

The book '' Plantesamfund'' was based on Warming's lectures on plant geography at the
University of Copenhagen The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
. 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 origin) in different regions of the world. This was a remarkably modern view—completely different from the merely descriptive floristic plant geography prevailing during his time. * Warming, E. (1895) ''Plantesamfund - Grundtræk af den økologiske Plantegeografi''. P.G. Philipsens Forlag, Kjøbenhavn. 335 pp. The subtitle alludes to the title of the book ''Grundtræk af den almindelige Plantegeografi'', published in 1822 (German edn 1823: ''Grundzüge einer allgemeinen Pflanzengeographie'') by J. F. Schouw, co-founder of the scientific
phytogeography Phytogeography (from Greek φυτόν, ''phytón'' = "plant" and γεωγραφία, ''geographía'' = "geography" meaning also distribution) or botanical geography is the branch of biogeography that is concerned with the geographic distribution ...
. ''Plantesamfund'' was translated to
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
in 1896 as * ''Lehrbuch der ökologischen Pflanzengeographie—Eine Einführung in die Kenntnis der Pflanzenvereine'' by Emil Knoblauch. Berlin, Gebrüder Borntraeger, 1896. 412 pp. This edition, which was approved by Warming, rapidly ran out of print. A second, unauthorized, edition was issued during 1902 by Paul Graebner, who put his own name after Warming's on the book's frontispiece, despite no changes to the contents. * ''Lehrbuch der ökologischen Pflanzengeographie - Eine Einführung in die Kenntnis der Pflanzenvereine];'' "Nach der neuesten Litteratur Vervollständigt bei Paul Graebner"; Berlin, Gebrüder Borntraeger. This edition was expanded in third and fourth editions: * Warming, E. & Graebner, P. (1918) ''Eug. Warming's Lehrbuch der ökologischen Pflanzengeographie'', 3 ed. Berlin, Gebrüder Borntrager. Fourth edn (1933) - 1158 pp. A Polish language, Polish translation of ’Plantesamfund’ (from Knoblauch's German translation) appeared in 1900: * Warming, E. (1900) ''Zbiorowiska Roślinne zarys ekologicznej geografii roślin'' by Edward Strumpf and Jósef Trzebiński. Warszawa, 1900. 451 pp. Two independent
Russian Russian(s) may refer to: *Russians (), an ethnic group of the East Slavic peoples, primarily living in Russia and neighboring countries *A citizen of Russia *Russian language, the most widely spoken of the Slavic languages *''The Russians'', a b ...
(
Moscow Moscow is the Capital city, capital and List of cities and towns in Russia by population, largest city of Russia, standing on the Moskva (river), Moskva River in Central Russia. It has a population estimated at over 13 million residents with ...
and
St. Petersburg Saint Petersburg, formerly known as Petrograd and later Leningrad, is the second-largest city in Russia after Moscow. It is situated on the River Neva, at the head of the Gulf of Finland on the Baltic Sea. The city had a population of 5,601, ...
) editions appeared in 1901 and 1903 * Вармингъ, Е. (1901) Ойкологическая географія растеній – Введеніе въ изученіе растительныхъ сообществъ by M. Golenkin and W. Arnol'di. Moskva, 542 pp. * 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. An extended and translated edition in English first appeared in 1909: * Warming, E. with M. Vahl (1909) ''Oecology of Plants—an introduction to the study of plant-communities'' by P. Groom and I. B. Balfour. Clarendon Press, Oxford. 422 pp. (2nd edn 1925). 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 15 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.


Warming as a teacher

Warming was a skillful and dedicated pedagogue, whose presentation of the subject was useful far beyond his lecture theatre in
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
. He wrote a number of
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
textbooks for the university level, as well as school books.


Handbook of systematic botany

Warming's textbook on systematics for his lectures of
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
in
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
appeared in several editions and was translated to
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
,
Russian Russian(s) may refer to: *Russians (), an ethnic group of the East Slavic peoples, primarily living in Russia and neighboring countries *A citizen of Russia *Russian language, the most widely spoken of the Slavic languages *''The Russians'', a b ...
and English and used in foreign universities. *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 issued as * Warming Eug. (1912) ''Frøplanterne (Spermatofyter)'' ranslated title: Seed Plants Kjøbenhavn, Gyldendalske Boghandel/Nordisk Forlag. 467 pp. (2nd edn 1933). The sections on spore plants were updated and published separately as * Rosenvinge L. Kolderup (1913) Sporeplanterne (Kryptogamerne). Kjöbenhavn, Gyldendalske Boghandel/Nordisk Forlag. 388 pp.


Handbook of general botany

Warming's textbook on plant
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
,
anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
and
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
was translated to Swedish and
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
: Warming Eug. ''Den almindelige Botanik: En Lærebog, nærmest til Brug for Studerende og Lærere'' ranslated title: ''General Botany'' Kjøbenhavn, 1880. (2nd edn 1886; 3rd edn by Warming and
Wilhelm Johannsen Wilhelm Johannsen (3 February 1857 – 11 November 1927) was a Danish pharmacist, botanist, plant physiologist, and geneticist. He is best known for coining the terms gene, phenotype and genotype, and for his 1903 "pure line" experiments in ...
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. Also, Warming's schoolbook on
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
was used abroad: Warming Eug. (1900) ''Plantelivet: Lærebog i Botanik for Skoler og Seminarier'' ranslated title: 'Plant Life' København. (2nd edn 1902; 3rd edn 1905; 4th and 5th edns by C. Raunkiær and Warming 1908 and 1914, respectively; 6th edn (1920) by E. Warming and Johs. Boye Petersen). 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 Russian(s) may refer to: *Russians (), an ethnic group of the East Slavic peoples, primarily living in Russia and neighboring countries *A citizen of Russia *Russian language, the most widely spoken of the Slavic languages *''The Russians'', a b ...
edn 1904: Растение и его жизнь (Началный учебник ботаники). (from the 2nd edn by L.M. Krečotovič and M. Golenkin). Moskva. Dutch edns 1905, 1912 and 1919 ''Kern der plantkunde'' (by Dr. A.J.M. Garjeanne).


Excursions

Warming felt a strong need to take students of
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
out of the lecture theatre. He used the
botanic garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens. is ...
to demonstrate live plants, but to teach plant
ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
he needed students to get out in nature. The action radius from
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
offered by trips by foot was far too small, however. He applied to the
government A government is the system or group of people governing an organized community, generally a State (polity), state. In the case of its broad associative definition, government normally consists of legislature, executive (government), execu ...
and obtained a grant to take students on longer excursions every year from 1893; every third year these went to western
Jutland Jutland (; , ''Jyske Halvø'' or ''Cimbriske Halvø''; , ''Kimbrische Halbinsel'' or ''Jütische Halbinsel'') is a peninsula of Northern Europe that forms the continental portion of Denmark and part of northern Germany (Schleswig-Holstein). It ...
, once to
Bornholm Bornholm () is a List of islands of Denmark, Danish island in the Baltic Sea, to the east of the rest of Denmark, south of Sweden, northeast of Germany and north of Poland. Strategically located, Bornholm has been fought over for centuries. I ...
, otherwise to
Zealand Zealand ( ) is the largest and most populous islands of Denmark, island in Denmark proper (thus excluding Greenland and Disko Island, which are larger in size) at 7,031 km2 (2715 sq. mi.). Zealand had a population of 2,319,705 on 1 Januar ...
. His excursion notes were published and are instructive introductions to the environment and plant
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
in
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
s,
salt marsh A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. I ...
es and other
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s: *Warming, E. (1890) Botaniske Exkursioner 1. Fra Vesterhavskystens Marskegne. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 1890. *Warming, E. (1891) Botaniske Exkursioner 2. De psammophile Formationer i Danmark. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 1891: 153–202. *Warming, E. (1891) Botaniske Exkursioner 3. Skarridsø. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 1891.


Further scientific works of E. Warming


Plant systematics

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 science, 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, mainly in Europe. * 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. E.g. Particula XXXIX
Enumeratio Myrtacearum Brasiliensium
by Hjalmar Kiærskou. They were all published as volumes in the series ’''Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn''’. In addition, Warming treated the families
Vochysiaceae Vochysiaceae is a plant family belonging to the order Myrtales. Description Trees or shrubs with opposite leaves; flowers are zygomorph 1-(3)-5 merous; ovary inferior or superior; one fertile stamen; fruits samara or capsules. Biogeography ...
and Trigoniaceae for the Flora Brasiliensis: *
Vochysiaceae Vochysiaceae is a plant family belonging to the order Myrtales. Description Trees or shrubs with opposite leaves; flowers are zygomorph 1-(3)-5 merous; ovary inferior or superior; one fertile stamen; fruits samara or capsules. Biogeography ...
et Trigoniaceae. Flora Brasiliensis, Eichler, A.G. ed.: Vol. XIII, Part II, Fasc. 67, Column 16–116. Monachii, 1875.


His favourite plant family: Etudes sur la famille des Podostemacees

Warming held a special interest in the family
Podostemaceae Podostemaceae (riverweed family), a family in the order Malpighiales, comprise about 50 genera and species of more or less thalloid aquatic herbs. Distribution and habitat They are found mostly in tropical and subtropical areas worldwide ...
, with which he had become acquainted during his stay in
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. The plant species of this family are extremely modified by the harsh environment in which they live - they are
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...
that resemble
liverworts Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
. *Warming, E. (1881–1899) ''Familien Podostemaceae - Etudes sur la famille des Podostemacees''. Part I-V. All published in Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 6. Rk.


Lagoa Santa

Having finished the taxonomical work, Warming finally published his ecological study of plant communities in the Lagoa Santa area, with
cerrado The Cerrado () is a vast ecoregion of Tropics, tropical savanna in central Brazil, being present in the states of Goiás, Mato Grosso do Sul, Mato Grosso, Tocantins, Maranhão, Piauí, Bahia, Minas Gerais, São Paulo (state), São Paulo, Paraná ...
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 The Cerrado () is a vast ecoregion of Tropics, tropical savanna in central Brazil, being present in the states of Goiás, Mato Grosso do Sul, Mato Grosso, Tocantins, Maranhão, Piauí, Bahia, Minas Gerais, São Paulo (state), São Paulo, Paraná ...
system and reissued as: Warming, E. & Ferri, M.G. (1973) ''Lagoa Santa – a vegetação de cerrados brasileiros''. University of São Paulo.


Organogenetic studies

Early on in Warming's scientific career, the morphological-organogenetic point of was the leading principle in
botanical Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
research, and he soon became one of the most prominent workers in this branch of
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
. His main works from the early period are his thesis on floral development in
Euphorbia ''Euphorbia'' is a large and diverse genus of flowering plants, commonly called spurge, in the family (biology), family Euphorbiaceae. Euphorbias range from tiny annual plants to large and long-lived trees, with perhaps the tallest being ''Eu ...
and on
seed plant A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. la ...
ovules. Warming's doctoral thesis (in Danish) dealt with ontogeny of the cyathia of
Euphorbia ''Euphorbia'' is a large and diverse genus of flowering plants, commonly called spurge, in the family (biology), family Euphorbiaceae. Euphorbias range from tiny annual plants to large and long-lived trees, with perhaps the tallest being ''Eu ...
(
Euphorbiaceae Euphorbiaceae (), the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of Euphorbia, the type genus of the family. Most spurges, such as ''Euphorbia paralias'', ar ...
). * Warming, J. Eug. B. 1871. ''Koppen hos Vortemælken en Blomst eller en Blomsterstand? En organogenetisk morfologisk Undersøgelse''. French summary: Le cyathium de l"Euphorbia est-il une fleure ou une inflorescence? Kjøbenhavn, G.E.C. Gad. Part of the work was published in
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
the year before the thesis: * Warming, E. 1870. ''Über die Entwicklung des Blütenstandes von Euphorbia''. Flora 53: 385–397. His studies of
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
and
anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
formation i
Angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...
and on the inflorescence of
Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...
were published in von Hanstein's ''Botanische Abhandlungen'': *Warming, E. (1873) ''Untersuchungen über pollenbildende Phyllome und Kaulome''. Botanische Abhandlungen aus dem Gebiet der Morphologie und Physiologie, 2 (2): 1-90. *Warming, E. (1876) ''Die Blüte der Kompositen''. Botanische Abhandlungen aus dem Gebiete der Morphologie und Physiologie, 3 (2): 1–167. His studies on
seed plant A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. la ...
ovules were published in French as * Warming, E. 1878. ''De l’Ovule''. Annales des Sciences Naturelles - Botanique et Biologie Vegetale sér. 6: 177–266. Through the 1870s, Warming became much-influenced by
Darwinism ''Darwinism'' is a term used to describe a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others. The theory states that all species of organisms arise and develop through the natural sel ...
. The scope of his research changed. First towards understanding
ontogenesis Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
in the light of a common descent as seen in ''De l’Ovule'', later towards plant
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
to environmental conditions. Again, his unparalleled ability to observe plants paired with his tropical experiencewas decisive to the route he chose.


Plant life-form

Although Warming did not coin the term ''life-form'' until 1895 (in Plantesamfund Ch. 2), he commenced work on
plant life-form Plant life-form schemes constitute a way of classifying plants alternatively to the ordinary species-genus-family scientific classification. In colloquial speech, plants may be classified as trees, shrubs, herbs (forbs and graminoids), etc. The sci ...
already during his
Stockholm Stockholm (; ) is the Capital city, capital and List of urban areas in Sweden by population, most populous city of Sweden, as well as the List of urban areas in the Nordic countries, largest urban area in the Nordic countries. Approximately ...
years. In the paper * Warming, E. (1884) ''Om Skudbygning, Overvintring og Foryngelse'' ranslated title: ''On shoot architecture, perennation and rejuvenation'' Naturhistorisk Forenings Festskrift: 1–105. Line drawings, he presented a classification based on longevity of the plant, power of
vegetative propagation Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is a form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or specializ ...
, duration of tillers,
hypogeous Hypogeal, hypogean, hypogeic and hypogeous (; ) are biological terms describing an organism's activity below the soil surface. In botany, a seed is described as showing hypogeal germination when the cotyledons of the germinating seed remain no ...
or
epigeous Epigeal, epigean, epigeic and epigeous are biological terms describing an organism's activity above the soil surface. In botany, a seed is described as showing epigeal germination when the cotyledons of the germinating seed expand, throw off the ...
type of shoots, mode of wintering, and degree and mode of branching of
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s. The observation were made while raising wild plants from seed under garden conditions. In the late 1880s, after Warming's return to
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
, he swopped research topic with his student Christen Raunkiær, who had traveled along the
North Sea The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium, and France. A sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Se ...
coast from
Jutland Jutland (; , ''Jyske Halvø'' or ''Cimbriske Halvø''; , ''Kimbrische Halbinsel'' or ''Jütische Halbinsel'') is a peninsula of Northern Europe that forms the continental portion of Denmark and part of northern Germany (Schleswig-Holstein). It ...
to the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
and published on the
phytogeography Phytogeography (from Greek φυτόν, ''phytón'' = "plant" and γεωγραφία, ''geographía'' = "geography" meaning also distribution) or botanical geography is the branch of biogeography that is concerned with the geographic distribution ...
of coastal
vegetation Vegetation is an assemblage of plants and the ground cover they provide. It is a general term, without specific reference to particular Taxon, taxa, life forms, structure, Spatial ecology, spatial extent, or any other specific Botany, botanic ...
. Warming now worked on plant
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s in
dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
and
salt marsh A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. I ...
es, while Raunkiær studied the morphology of Danish plants, eventually leading him to his plant life-form scheme. Nevertheless, after Raunkiær had published his life-form scheme, Warming return to this topic in the work *Warming, E. (1908) ''Om planterigets livsformer'' ranslated title: ''On the life forms in the vegetable kingdom'' G.E.C. Gad, København. Warming's new scheme was less simple than Raunkiær's, taking other environmental factors than wintering into account, especially water/drought stress. Warming did not approve of what he saw as over-simplification in the Raunkiær scheme. Warmings last published work was a renewed attempt to put all plant (including bacteria and algae) life forms into a system. * Warming, E. (1923) ''Økologiens Grundformer – Udkast til en systematisk Ordning'' ranslated title: ''Fundamental ecological forms - draft for a system'' Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 8. Rk., vol. 4: 120–187.


Greenland, Iceland and Faroe Islands

Warmings published a number of treatises based on his expedition to Southwest
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
in 1884. One of the most important ones is his observations of the
vegetation Vegetation is an assemblage of plants and the ground cover they provide. It is a general term, without specific reference to particular Taxon, taxa, life forms, structure, Spatial ecology, spatial extent, or any other specific Botany, botanic ...
of
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
and the history of the flora: *Warming, E. (1887) Om Grønlands Vegetation ranslated title: On the vegetation of Greenland Meddelelser om Grønland 12: 1–223. A summary was published as: *Warming, E. (1888) Über Grönlands Vegetation. Englers Botanische Jahrbücher, 10. Following the publication of this paper, Warming entered a dispute with A.G. Nathorst over the history of the
flora of Greenland The flora of Greenland consists of a total of 583 species or 614 taxa (species and subspecies) of vascular plants, of which 13 are endemic, and 87 taxa introduced by humans, most of which are naturalized. Apiaceae *''Angelica archangelica'' – n ...
. Warming's collections of leaves, stems and flowers, made during the brief expedition, were examined in detail and the
anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
of a number of species described in a series of papers in Danish. Later, Warming distributed the material family-wise, now ameliorated with collectections made later expeditions and elsewhere in the
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
, to students, who made further investigations and published the results in English: Warming, E. ed. (1908–1921) '' The structure and biology of Arctic flowering plants''. Meddelelser om Grønland vol. 36: 1-481 and 37: 1–507. *Warming, E. ed. (1901–1908) '' Botany of the Færöes - based upon Danish investigations, vol. I-III. Copenhagen and London. * Rosenvinge, L. Kolderup & Warning, E. (eds) (1912–1932) '' The Botany of Iceland'', vol. 1–3. Copenhagen, J. Frimodt. Continued in vols 4-5 edited by Johannes Grøntved, Ove Paulsen and Thorvald Sørensen. Full text of Vol. 1
part 1
an
part 2
) and Vol. 2
part 1
.


Vegetation of Denmark

* Warming, E. 1904. Bidrag until Vadernes, Sandenes og Marskens Naturhistorie (with contributions of C. Wesenberg-Lund, E. Østrup &c). Kongelige Danske Videnskabernes Selskabs Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 7. Rk., 2: 1-56. * Warming, E. 1906. Dansk Plantevækst. 1. Strandvegetationen. - Gyldendalske Boghandel Nordisk Forlag. each vegetation* Warming, E. 1909. Dansk Plantevækst. 2. Klitterne. - Gyldendalske Boghandel Nordisk Forlag. unes* Warming, E. 1917. Dansk Plantevækst. 3. Skovene. - Gyldendalske Boghandel Nordisk Forlag. orests


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, Kurt (2001) History of Ecology. In: Encyclopedia of Life Sciences. John Wiley & Sons.
/ref>
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 Sir Arthur George Tansley FLS, FRS (15 August 1871 – 25 November 1955) was an English botanist and a pioneer in the science of ecology. Educated at Highgate School, University College London and Trinity College, Cambridge, Tansley taught ...
was extremely influenced by reading ’Plantesamfund’ (or rather the 1896 German edition). Reading the book made him jump from anatomy to ecology. Tansley used the book as textbook in a university course as early as 1899.Tansley, A.G. (1947) The early history of modern plant ecology in Britain. ''Journal of Ecology'', 35, 130-137.
/ref> Similarly, Warming's book was decisive in forming the careers of North American naturalists like
Henry Chandler Cowles Henry Chandler Cowles (February 27, 1869 – September 12, 1939) was an American botanist and ecological pioneer. A professor at the University of Chicago, he studied ecological succession in the Indiana Dunes of Northwest Indiana.Smith, S. & ...
.Coleman, W. (1986) Evolution into ecology? The strategy of Warming’s ecological plant geography. ''Journal of the History of Biology'', 19(2), 181-196. Cowles' now classic studies of
Lake Michigan Lake Michigan ( ) is one of the five Great Lakes of North America. It is the second-largest of the Great Lakes by volume () and depth () after Lake Superior and the third-largest by surface area (), after Lake Superior and Lake Huron. To the ...
sand dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
plant communities were directly inspired by Warming's studies of Danish dunes. Also
Frederic Clements Frederic Edward Clements (September 16, 1874 – July 26, 1945) was an American plant ecologist and pioneer in the study of both plant ecology and vegetation succession. Biography Born in Lincoln, Nebraska, he studied botany at the University o ...
was much inspired by Warming when starting to working with
succession Succession is the act or process of following in order or sequence. Governance and politics *Order of succession, in politics, the ascension to power by one ruler, official, or monarch after the death, resignation, or removal from office of ...
, but more by Oscar Drude in formulating his concept of vegetational climax in his 1916 book. A more unexpected avenue of influence went through the American sociologist Robert E. Park, who read Warming's '' Oecology of Plants'' and used the ideas of
ecological succession Ecological succession is the process of how species compositions change in an Community (ecology), ecological community over time. The two main categories of ecological succession are primary succession and secondary succession. Primary successi ...
as inspiration for a notion of ''succession'' in human communities – a
human ecology Human ecology is an interdisciplinary and transdisciplinary study of the relationship between humans and their natural, social, and built environments. The philosophy and study of human ecology has a diffuse history with advancements in ecolo ...
. 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 The University of Copenhagen (, KU) is a public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in Scandinavia, after Uppsala University. ...
. In addition, he had a direct influence on Danish research, scientific and other, for a couple of decades. After his appointment to the professorship in
Copenhagen Copenhagen ( ) is the capital and most populous city of Denmark, with a population of 1.4 million in the Urban area of Copenhagen, urban area. The city is situated on the islands of Zealand and Amager, separated from Malmö, Sweden, by the ...
, he gradually took over Japetus Steenstrups power base, most notably as one of three members of the board of the
Carlsberg Foundation Carlsberg Foundation () is a not-for-profit organization that was founded by J. C. Jacobsen in 1876, by allocating some of his shares in the Carlsberg Brewery to fund and operate the Carlsberg Laboratory and the Museum of National History at ...
for 32 years. Thus, Warming had the upper hand in who should be granted money and who should not.


Warming and evolution

Warming was a firm believer in adaptation. However, he was a declared Lamarckist. In his popularizing book ''Nedstamningslæren'' (The theory of descendence; 1915),Warming, E. (1915) Nedstamningslæren. Copenhagen, Udvalget til Folkeoplysnings Fremme, G.E.C. Gad. he reviewed the direct and indirect evidence for common descent of living organisms and for
Darwinian ''Darwinism'' is a term used to describe a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others. The theory states that all species of organisms arise and develop through the natural sele ...
natural selection as a process involved in
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. His keen observations of how differently the same plant is grown under different circumstances (now known as
phenotypic plasticity Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
) led him to question the change of species by infinitesimally small steps as advocated by his contemporary Darwinists of the Biometry school, e.g.
Karl Pearson Karl Pearson (; born Carl Pearson; 27 March 1857 – 27 April 1936) was an English biostatistician and mathematician. He has been credited with establishing the discipline of mathematical statistics. He founded the world's first university ...
. Warming summarized his view on the ways in which new species could may arise: #By inheritance of acquired characters; #By hybridization; #By
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
, with the latter mechanism being the least important.


Warming, religion and politics

Warming was raised in a Christian
Protestant Protestantism is a branch of Christianity that emphasizes Justification (theology), justification of sinners Sola fide, through faith alone, the teaching that Salvation in Christianity, salvation comes by unmerited Grace in Christianity, divin ...
home and he continued to be religious throughout his life. He accepted the evolution by descent of living beings, but believed that laws governing planets’ orbits and other laws governing organic evolution were God-given. In his popular book ''Nedstamningslæren'' (translated title: Evolution by descent), he concludes the section on hypotheses about the origin of life writing that, no matter what hypothesis is considered, it just “defers the grand question: how did life first come into existence, »in the beginning«? ... as if we human beings thereby obtained understanding and explanation for anything at all, or circumvented the almighty power that, incomprehensibly to our mind, must have created matter, force, time and infinite space. Science has not disproven the
Bible The Bible is a collection of religious texts that are central to Christianity and Judaism, and esteemed in other Abrahamic religions such as Islam. The Bible is an anthology (a compilation of texts of a variety of forms) originally writt ...
that says: » In the beginning God created …«!”. Warming shared this view with many prominent contemporary naturalists, e.g.
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was an English naturalist, explorer, geographer, anthropologist, biologist and illustrator. He independently conceived the theory of evolution through natural selection; his 1858 pap ...
. Politically, Warming was national-conservative, Scandinavist and anti-
Prussia Prussia (; ; Old Prussian: ''Prūsija'') was a Germans, German state centred on the North European Plain that originated from the 1525 secularization of the Prussia (region), Prussian part of the State of the Teutonic Order. For centuries, ...
n. Warming was able to visit his birthplace only a few times in his life because
Schleswig The Duchy of Schleswig (; ; ; ; ; ) was a duchy in Southern Jutland () covering the area between about 60 km (35 miles) north and 70 km (45 mi) south of the current border between Germany and Denmark. The territory has been di ...
was conquered by
Prussia Prussia (; ; Old Prussian: ''Prūsija'') was a Germans, German state centred on the North European Plain that originated from the 1525 secularization of the Prussia (region), Prussian part of the State of the Teutonic Order. For centuries, ...
and
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
in 1864 and (
Northern Schleswig Northern may refer to the following: Geography * North, a point in direction * Northern Europe, the northern part or region of Europe * Northern Highland, a region of Wisconsin, United States * Northern Province, Sri Lanka * Northern Range, a ra ...
) returned to
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
in 1920. Warming expressed support, in letters, for France in the 1870
Franco-Prussian War The Franco-Prussian War or Franco-German War, often referred to in France as the War of 1870, was a conflict between the Second French Empire and the North German Confederation led by the Kingdom of Prussia. Lasting from 19 July 1870 to 28 Janua ...
. He made financial contributions to a secret fund that should support Danish-minded
Schleswig The Duchy of Schleswig (; ; ; ; ; ) was a duchy in Southern Jutland () covering the area between about 60 km (35 miles) north and 70 km (45 mi) south of the current border between Germany and Denmark. The territory has been di ...
ian farmers in buying farms and prevent
Germanization Germanisation, or Germanization, is the spread of the German language, German people, people, and German culture, culture. It was a central idea of German conservative thought in the 19th and the 20th centuries, when conservatism and ethnic nati ...
of
Northern Schleswig Northern may refer to the following: Geography * North, a point in direction * Northern Europe, the northern part or region of Europe * Northern Highland, a region of Wisconsin, United States * Northern Province, Sri Lanka * Northern Range, a ra ...
. In a letter of 1898 to his son Jens, he regrets that the Højre – the conservative party – would lose an upcoming election and expresses concern that ''anarchy and socialism'' will eventually rule.


Miscellaneous

The Orchid genus '' Warmingia'' Rchb.f. and dozens of vascular plant species (IPN]) has been named to his honour. The same is the case for a number of fungi, e.g. the Ustilaginales, smut fungus ''Microbotryum warmingii'' ( Rostr.) Vánky and the gall fungus ''Arcticomyces warmingii'' ( Rostr.) Savile. Warming Land - a
peninsula A peninsula is a landform that extends from a mainland and is only connected to land on one side. Peninsulas exist on each continent. The largest peninsula in the world is the Arabian Peninsula. Etymology The word ''peninsula'' derives , . T ...
in northernmost
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
is named for him.
Universidade Federal de Minas Gerais The Federal University of Minas Gerais (, UFMG) is a federalIn the Brazilian Higher Education context, ''Federal'' does not mean ''collegiate'' (even though most Federal Universities in Brazil enjoy a similarly collegiate system), but it means ...
has organized a series of 'Eugen Warming lectures in Evolutionary Ecology' since 1994.


See also

* History of ecology


Biographies and obituaries

* Raunkiær, C. (1904) Biography i
Dansk Biografisk Lexikon, vol. XVIII
* Urban, Ignatius (1906
Vitae for Warming in Flora Brasiliensis, enumeratio plantarum in Brasilia hactenus detectarum :quas suis aliorumque botanicorum studiis descriptas et methodo naturali digestas partim icone illustratas /ediderunt Carolus Fridericus Philippus de Martius et Augustus Guilielmus Eichler ; iisque defunctis successor Ignatius Urban; Fasc. CXXX (ultimus) - VITAE ITINERAQUE COLLECTORUM BOTANICORUM Etc.
* Obituary in ''
Nature Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
'', 113, 683-684 (1924) by William G. Smith * Obituaries in '' Botanisk Tidsskrift'', 39 (1924): ** L. K. Rosenvinge: ''Eug. Warming og Dansk Botanisk Forening'', pp. 1–6 ** Carl Christensen: ''Eug. Warming, en Levnedsbeskrivelse'', pp. 7–30 ** C. H. Ostenfeld: ''Warmings almindelige botaniske Virksomhed'', pp. 31–38 ** A. Mentz: ''Warming som plantegeografisk Forsker'', pp. 39–44 ** Charles Flahault, O. Juel, C. Schröter and A.G. Tansley: ''Eug. Warming in memoriam'', pp. 45–56. * Christensen, C. (1924–26) ''Den danske botaniks historie, med tilhørende bibliografi. I. Den danske botaniks historie fra de ældste tider til 1912. II. Bibliografi''. * Christensen, C. (1932) ''Eugenius Warming'', pp. 156–160 in: Meisen, V. Prominent Danish Scientists through the Ages. University Library of Copenhagen 450th Anniversary. Levin & Munksgaard, Copenhagen. * Müller, D. (1980) ''Warming, Johannes Eugenius Bülow''. In: Gillespie, C.G. (ed.)
Dictionary of Scientific Biography The ''Dictionary of Scientific Biography'' is a scholarly reference work that was published from 1970 through 1980 by publisher Charles Scribner's Sons, with main editor the science historian Charles Coulston Gillispie, Charles Gillispie, from Pri ...
, vol. 16. New York, NY: Charles Scribner and Sons. * Klein, Aldo Luiz (2000)
Warming e o cerrado brasileiro um século depois
'. São Paulo, UNESP. 156 pp. 


References


Bibliography

* * *


External links

* * Wikimedia Commons: "Om Skudbygning, Overvintring og Foryngelse"
Encyclopædia Britannica Online: Warming
{{DEFAULTSORT:Warming, Eugenius 19th-century Danish botanists Danish ecologists Danish phycologists Danish mycologists Danish microbiologists Botanists with author abbreviations Botanists active in South America Botanists active in the Arctic Danish scientists Academic staff of the University of Copenhagen University of Copenhagen alumni Members of the French Academy of Sciences Members of the Royal Swedish Academy of Sciences 1841 births 1924 deaths Honorary Fellows of the Royal Society 20th-century Danish botanists Members of the Royal Society of Sciences in Uppsala