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Christen Christensen Raunkiær (March 29, 1860 –
March 11, 1938) was a Danish botanist, who was a pioneer of plant ecology. He
is mainly remembered for his scheme of plant strategies to survive
an unfavourable season ("life forms") and his demonstration that
the relative abundance of strategies in floras largely corresponded
to the Earth's climatic zones. This scheme, the Raunkiær system, is still
widely used today and may be seen as a precursor of modern plant
strategy schemes, e.g. J. Philip Grime's CSR system.
Life
He was born on a small heathland farm, named Raunkiær, in Lyhne
parish in western Jutland,
Denmark. He later took his
surname after it. He succeeded Eugen Warming as professor in botany
at the University of Copenhagen and
the director of the Copenhagen
Botanical Garden, a position he held from 1912 to 1923. He was
married to the author and artist Ingeborg Raunkiær (1863-1921), who
accompanied him on journeys to the West Indies and the
Mediterranean and made line drawings for his botanical works. They
divorced in 1915, the same year as their only son Barclay
Raunkiær died. Raunkiær later married the botanist Agnete
Seidelin (1874-1956). After his retirement, C. Raunkiær made
numerical studies of plants and flora in the literature ("The flora
and the heathland poets", "The dandelion in Danish poetry", "Plants
in the psalms"). In these studies, he applied strict quantitative
criteria, like in his ecological studies. For example, he defined a
poet as a person who has written 1,000 or more lines of verse.
Plant life
forms
Raunkiær devised a system for categorising plants by life-form as a
way of ecologically meaningful comparison of species and vegetation
in regions having different floras.
Life form
spectra
Raunkiær compared statistically local life form spectra
(relative abundance) with the world average, which he called "the
normal spectrum" (Raunkiær 1918 – see below). Thereby, he devised
the first null model in the history of ecology. Raunkiær was a keen
naturalist, who described the flora and funga of Denmark, the Virgin Islands, Tunisia, and other countries. In contrast to
many contemporary naturalist, however, he strongly promoted
quantitative and numerical approaches and experimentation. He
devised a method to quantify the abundance of plants in vegetation
as frequency in subplots and used it for quantitative studies of a
range of plant communities.
Raunkiær's life forms: 1. Phanerophyte.
2-3. Chamaephytes.
4. Hemicryptophyte.
5-6. Geophytes (cryptophytes).
7. Helophyte.
8-9. Hydrophytes.
Therophyte (annual, surviving as seed) not shown
Raunkiær's
law
When plotting the number of species in a plant community that
fell in each 20-percentile frequency class from very frequent, i.e.
numerically dominant, to very infrequent, Raunkiær discovered that
most species were either very common or very rare. This came to be
known as "Raunkiær's law" and is related to Fisher's logseries
distribution and to Frank Preston's lognormal distribution of the
number of individuals of each species in a community[1][2]. The
significance of his idea was, however, disputed already by some of
his contemporaries[3].
Raunkiær's research axiom was that everything countable in nature
should be subjected to numerical analysis, e.g. the number of male
and female catkins in monecious plants and the number of male and
female individuals in dioecious plants. Raunkiær also was an early
student of apomixis in
flowering plants and hybrid swarms. In addition, he studied the
effect of soil pH on plants
and plants on soil pH, a work his apprentice Carsten Olsen
continued.
Scientific works of C.
Raunkiær
- Raunkiær, C. (1887) Frøskallens Bygning og Udviklingshistorie
hos Geraniaceerne.
Botanisk
Tidsskrift 16, 152-167.
- Raunkiær, C. (1888-89) Myxomycetes Daniae
eller Danmarks Slimsvampe. Botanisk Tidsskrift 17, 20-105
Description in English of some new and of some unsatisfactorily
known species of Myxomycetes described in the preceding treatise.
Botanisk
Tidsskrift 17, 106-110.
- Raunkiær, C. (1889) Notes on the vegetation of the North-Frisian Islands and a
contribution to an eventual flora of these islands. Botanisk
Tidsskrift 17, 179-196.
- Raunkiær, C. (1893) Vegetationsorganernes Morphologi og Biologi
hos de danske Cyperaceer - Bestøvning og Frugtsaetning hos
de danske Cyperaceer.
Botanisk
Tidsskrift 18, 19-23.
- Raunkiær, C. (1895) De danske Blomsterplanters Naturhistorie.
l. Enkimbladede. Gyldendalske Boghandels Forlag, København.
- Raunkiær, C. (1901) Om Papildannelsen hos Aira caespitosa. Botanisk
Tidsskrift 24, 223-238.
- Raunkiær, C. (1902) Statistisk Undersøgelse over Forholdet
mellem Han- og Hunrakler hos vore Alnus-arter. Botanisk Tidsskrift 24,
289-296.
- Raunkiær, C. (1903) Anatomical Potamogeton-Studies and Potamogeton fluitans.
Botanisk
Tidsskrift 25, 253-280.
- Ostenfeld, C.H. & Raunkiær,
C. (1903) Kastreringsforsøg med Hieracium og andre Cichorieae [Eng. summ.].
Botanisk
Tidsskrift 25, 409-413.
- Raunkiær, C. (1904) Comment les plantes géophytes à rhizomes
apprécient la profondeur où se trouvent placés leurs rhizomes.
Oversigt over Det Kongelige Danske Videnskabernes Selskabs
Forhandlinger, 1904, 330-349.
- Raunkiær, C. (1904) Om biologiske Typer, med Hensyn til
Planternes Tilpasninger til at overleve ugunstige Aarstider. Botanisk
Tidsskrift 26, XIV. Ch. 1 in Raunkiær (1934): Biological types
with reference to the adaption of plants to survive the
unfavourable season, p. 1.
- Raunkiær, C. (1905) Types biologiques pour la géographie
botanique. Oversigt over Det Kongelige Danske Videnskabernes Selskabs
Forhandlinger, 1905, 347-438.
- Raunkiær, C. (1905) Om Talforholdene mellem Kønnene hos tvebo
Planter og om Talforholdet mellem hanlige og hunlige individer i
Afkommet af Hunplanter og tvekønnede Planter hos Gynodiøcister. Botanisk
Tidsskrift 26, 86-88.
- Raunkiær, C. (1906) Edderkopper og blomsterbesøgende Insekter.
Botanisk
Tidsskrift 27, 313-316.
- Raunkiær, C. (1906) Sur la transmission par hérédité dans les
espèces hétéromorphes. Oversigt over Det Kongelige Danske Videnskabernes Selskabs
Forhandlinger, 1906, 31-39.
- Raunkiær, C. (1907) Planterigets Livsformer og deres Betydning
for Geografien. Gyldendalske Boghandel - Nordisk Forlag, København
and Kristiania. 132 pp. Ch. 2 in Raunkiær (1934): The life-forms of
plants and their bearings on geography, p. 2-104.
- Raunkiær, C. (1907) Om Livsformen hos Tussilago
farfarus. Et lille Bidrag til Følfodens Naturhistorie. Botanisk
Tidsskrift 28, 210. Ch. 3 in Raunkiær (1934): The life-form of
Tussilago
farfarus, p. 105-110.
- Raunkiær, C. (1908) Livsformernes Statistik som Grundlag for
biologisk Plantegeografi. Botanisk Tidsskrift 29, 42-43.
German translation: Statistik der Lebensformen als Grundlage für
die biologische Pflanzengeographie. Beiheft zum Bot. Centralbl., 27
(2), 171-206 (1910). Ch. 4 in Raunkiær (1924): The statistics of
life-forms as a basis for biological plant geography, p.
111-147.
- Raunkiær, C. (1908) Fungi from the Danish West
Indies collected 1905-1906. Botanisk Tidsskrift 29, 1-3.
- Raunkiær, C. (1909) Livsformen hos Planter paa ny Jord. Kongelige Danske Videnskabernes Selskabs
Skrifter - Naturvidenskabelig og Mathematisk Afdeling, 7.Rk., 8,
1-70. Ch. 5 in Raunkiær (1934): The life-forms of plants on new
soil, p. 148-200.
- Raunkiær, C. (1909a) Formationsundersøgelse og
Formationsstatistik. Botanisk Tidsskrift 30, 20-132. Ch.
6 in Raunkiær (1934): Investigations and statistics of plant
formations, p. 201-282.
- Raunkiær, C. (1911) Det arktiske og antarktiske
Chamaefyteklima. In: Biologiske Arbejder tilegnede Eug. Warming
paa hans 70 Aars Fødselsdag den 3. Nov. 1911. Kjøbenhavn. Ch. 7 in
Raunkiær (1934): The Arctic and Antarctic chamaephyte climate, p.
283-302.
- Raunkiær, C. (1912) Measuring-apparatus for statistical
investigations of plant-formations. Botanisk Tidsskrift 33, 45-48.
- Raunkiær, C. (1913) Formationsstatistiske Undersøgelser paa Skagens Odde. Botanisk
Tidsskrift 33, 197-243. Ch. 8 in Raunkiær (1934): Statistical
investigations of the plant formations of Skagens Odde (The Skaw), p. 303-342.
- Raunkiær, C. (1914) Sur la végétation des alluvions
méditerranéennes françaises. Mindeskrift i Anledning af
Hundredeaaret for Japetus Steenstrups Fødsel (eds H.
F. E. Jungersen & E. Warming), pp. 1-33. København. Ch.
9 in Raunkiær (1934): On the vegetation of the French mediterranean
alluvia, p. 343-367.
- Raunkiær, C. (1916) Om Bladstørrelsens Anvendelse i den
biologiske Plantegeografi. Botanisk Tidsskrift 34, 1-13. Ch.
10 in Raunkiær (1934): The use of leaf size in biological plant
geography, p. 368-378
- Raunkiær, C. (1916) Om Valensmetoden. Botanisk
Tidsskrift 34, 289-311.
- Raunkiær, C. (1918) Om Løvspringstiden hos Afkommet af Bøge med
forskellig Løvspringstid [Eng. summ.: On leaftime in the
descendants from beeches with
different leaftimes]. Botanisk Tidsskrift 36,
197-203.
- Raunkiær, C. (1918) Recherches statistiques sur les formations
végétales. Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 1
(3), 1-80. Ch 11 in Raunkiær (1934): Statistical researches on
plant formations, p. 379-424.
- Raunkiær, C. (1918) Über das biologische Normalspektrum.
Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 1
(4), 1-17. Ch. 12 in Raunkiær (1934): On the biological normal
spectrum, p. 425-434.
- Raunkiær, C. (1918) Über die verhältnismässige Anzahl
männlicher und weiblicher Individuen bei Rumex
thyrsiflorus Fingerh. Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 1
(7), 1-17.
- Børgesen, F. & Raunkiær, C.
(1918) Mosses and Lichens collected
in the former Danish West Indies. Dansk
Botanisk Arkiv 2 (9): 1-18.
- Raunkiær, C. (1919) Über Homodromie und Antidromie insbesondere
bei Gramineen. Biologiske
Meddelelser / Kongelige Danske Videnskabernes Selskab, 1
(12), 1-22.
- Raunkiær, C. (1920) Egern, Mus, og Grankogler. En
naturhistorisk Studie. Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 2
(4), 1-90.
- Raunkiær, C. (1920) Om Kryptogamernes Betydning for
Karakteriseringen af Planteklimaterne. Botanisk
Tidsskrift 37, 151-158. Ch. 13 in Raunkiær (1934): On the
significance of Cryptogams for characterizing Plant climates, p.
435-442.
- Raunkiær, C. (1922) Forskellige Vegetationstypers forskellige
Indflydelse paa Jordbundens Surhedsgrad (Brintionkoncentration).
Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 3
(10), 1-74. Ch. 14 in Raunkiær (1934): The different influence
exercised by various types of vegetation on the degree of acidity
(Hydrogen-ion concentration), p. 443-487.
- Raunkiær, C. (1926) Nitratindholdet hos Anemone
nemorosa paa forskellige Standpladser. Biologiske Meddelelser /
Kongelige Danske Videnskabernes Selskab, 5
(5), 1-47. Ch. 15 in Raunkiær (1934): The Nitrate content of Anemone
nemorosa growing in various localities, p. 488-516.
- Raunkiær, C. (1926) Om danske Agropyrum-Arter. (Isoreagent-Studier II). Botanisk
Tidsskrift 39, 329-347.
- Raunkiær, C. (1926) Vegetationen paa Maglehøj (vest for Arresø)
og lidt om vore Kaempehøjes Flora i det hele taget. Botanisk
Tidsskrift 39, 348-356.
- Raunkiær, C. (1928) Dominansareal, Artstaethed og
Formationsdominanter. Biologiske Meddelelser / Kongelige Danske Videnskabernes Selskab, 7
(1), 1-47. Ch 16 in Raunkiær (1934): The area of dominance, species
density, and formation dominants, p. 517-546.
- Raunkiær, C. (1928) Myxomycetes from the West Indian
Islands St. Croix, St. Thomas and St. Jan. Dansk
Botanisk Arkiv 5 (16), 1-9.
- Raunkiær, C. (1930) Variation hos Tussilago farfarus L. Botanisk
Tidsskrift 41, 257-258.
- Raunkiær, C. (1934) Om de danske Arter af Stellaria
media-Gruppen. Botaniske Studier, 1. haefte (ed C. Raunkiær),
pp. 3-30. J.H. Schultz Forlag, København.
- Raunkiær, C. (1934) The Life Forms of Plants and
Statistical Plant Geography. Introduction by A.G. Tansley.
Oxford University Press, Oxford. 632 pp. Collection of 16 of
Raunkiær's publications plus one new.
- Raunkiær, C. (1934) Botanical studies in the Mediterranean region. Ch. 17 in The Life
Forms of Plants and Statistical Plant Geography, pp. 547-620.
- Raunkiær, C. (1935) The vegetation of the sand dunes north of
Sousse. Botaniske Studier, 3.
haefte (ed C. Raunkiær), pp. 244-248. J.H. Schultz Forlag,
København.
- Raunkiær, C. (1936) The life-form spectrum of some Atlantic
islands. Botaniske Studier, 4. haefte (ed C. Raunkiær), pp.
249-328. J.H. Schultz Forlag, København.
- Raunkiær, C. (1937) Artstal, Artstaethed og Praedominanter i
danske Plantesamfund. Botaniske Studier, 5. haefte (ed C.
Raunkiær), pp. 357-382. J.H. Schultz Forlag, København.
- Raunkiær, C. (1937) Hudcellepapiller hos Carex arenaria.
Botaniske Studier, 5. haefte (ed C. Raunkiær), pp. 329-336. J.H.
Schultz Forlag, København.
- Raunkiær, C. (1937) Life-form, genus area, and number of
species. Botaniske Studier, 5. haefte (ed C. Raunkiær), pp.
343-356. J.H. Schultz Forlag, København.
- Raunkiær, C. (1937) Plantago intermedia og Plantago major.
Botaniske Studier, 5. haefte (ed C. Raunkiær), pp. 337-342. J.H.
Schultz Forlag, København.
Works
on C. Raunkiær
- Biography by Carl Christensen in: Dansk Biografisk
Leksikon, 3rd edn 1979-1984 (ed. Svend Cedergreen Bech).
References
External
links