RAUNKIAER'S

" LIFE-FORMS)" AND STATISTICAL

METHODS
BY WILLIAM G. SMITH
(With Three Tables)

C. Raunkiaer, who on January 1, 1912, succeeded E. Warming as Professorof

Botany in the University and Director of the Botanic Garden and Museum at
Copenhagen,is one of that distinguishedDanish school which has done so much
to promote the study of the vegetative organs of plants in relation to their
environment. His naine is primarilyassociated with a systemof so-called "biological types" or "life-forms"and its application in phytogeography. The idea
underlyingthis interestingand valuable system was firstexpressed at a meeting
of the Danish Botanical Society in December 1903, and a short summarywas
it appeared in French in 1905 (2),
publishedin 1904 (1). In a moreelaborateformi
and in a still more extended form in 1907 (3) and 1908 (4)-this last paper
was translated into German by Tobler and so became more widelyknown than in
the less accessible Danish periodicals. The followingaccount will, it is hoped,
convey to English readers a general impressionof Raunkiaer's system,besides
indicating the evolution of the concept on which it is based.
In the 1905 paper (2) Raunkiaer reviews the work of Humboldt, Grisebach,
Schouw and Engler as representingwhat inay be termed the floristicaspect of
and points out that
phytogeography,
and that of Warming' on "growth-forms,"
somethingis still lacking to expressthe correlationbetweenvegetationand climate.
He proposes to circumscribethe domain of geographical botany as "that geographical science which endeavours to characterisethe earth by its climate in so far
as this is manifested by the adaptation of plants to the various seasons." As
expressed later (3), he aims at findinga method of estimating the value of the
climate of the various regionls of the earth by some standard which has a bearing
on plant-life. The usual physicalmethodsfail to do this,because different
physical
factors may have the same effecton plant growth,and the influenceof various
conlbinationsof factors may result in essentially the same growth-form.Thus
"physiological drought" may arise fromwidely differentcombinationsof physical
factors-cold and drought. The plant itself must be the recorderof the biological
value of any climate. Any valuation used must have a uniformstandard; nlothing
bud protection
can be gained by using cytologicaladaptations in one growth-form,
in others.
in another,and leaf-structure
The factorselected by Raunkiaer is the adaptation of plants to the critical or
I

Warming,E., OecologyofPlants (Oxford,1909), Ch. II.

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W. G. SMITH

17

rigorousseason as expressed by the nature and the degree of protectionpossessed

by the dormantperennatingshoot-apices. The preliminaryto this is a classificatioll
of plants according to shoot-apexor bud protection,and the biological types or
life-forms
formthe basis of the earlier memoirs(1, 2, 3, 4).
Phanerophytes have their dormantbuds on branches whichlproject freely
into the air; theyare the trees and shrubs. Amongst these several modifications
are recognised. (a) According to degree of protectionone canl distinguishevergreens with naked or with covered buds, and deciduous species with coveredbuds
these are fully illustrated in Raunkiaer's second and third papers (2, 3).
(b) According to size, since this is determined-bythe relation between the plants
atid the humidityof-the environment,one can distinguish (1) megaphanerophytes with a stature over 30 metres,(2) mesophanerophytes
8 to 30 m.,
(3) microphanerophytes
2 to 8 m., and (4) nanophanerophytes
less than
2 m. high. Combiningthe groups (a) and (b), we have fifteensub-typesof phanerophytes.
Chamaephytes include plants with their buds or shoot-apicesperennatingon
the surface of the ground or just above it (not exceeding 25 cm.), so that in
countrieswith snow theywill be protectedin winter,while in othercountrieswith
a dry season some protectionwill be affordedby plant remains. The buds are
thus better protected than in phanerophytes. The chamaephyte types include
(1) active chamaephytes,with shoots diageotropicand persistentthroughouttheir
whole length-Empetrum nigrurn, Lysirnachia iVummularia, etc.; (2) passive
chamaephytes with weak stems which lie on the ground-Stellaria Holostea,
Cerastiumtrigynrqrn,etc.; (3) suffruticose
chamaephytes,in which the perennating
parts remain on the surface of the ground after the herbaceous parts have died
away oil the approach of the critical season-many Mediterranean species of
Lahiatae, Papilionaceae, etc.; (4) cushion plants. Obviously, it is sometimes
difficultto draw a sharp line between this life-form,the phanerophyteswhich
precede it, and the next type.
Hemicryptophytes have their dormant buds in the upper crust of the soil,
just below the surface; the aerial parts are herbaceousand die away in the critical
period, so that they form an additional protection to the earth-buds. The
perennatingparts may be long or short, laterally extended or formingcoumpact
root-stocks,hence the group includes a large numberof our native woodlaiid and
hedgerowspecies (Lamium album, Mercurialis perennis,etc.), and many rosette or
half-rosette
species(Caitha palustris,Bellis pereninis,
CChrysanthemrum
Leucanthemum,
Taraxacumn,etc.). This type may be siabdividedto a considerableextent.
Cryptophytes include plants with their dormantparts subterraneanin the
case of geophytes with bulbs, rhizomes,tubers on stem and root, and root-buds
(Rumex A cetosella, etc.). Ainother division is characterised by semi-aquatic
The helophytes or marshand hydrophytes.
dormant buds-helophytes
plants do not include all so-called marsh species, but only such cryptophytesas
have their buds at the bottom of the water or in the subjacent soil-Typha.
Sparganium,Acorus,Phragrnites,Sagittaria,etc. The hydrophytes
(e.g. Nymphaea,
have either perennatirigrhizomes,etc., or
Zostera, Hippuris, Elodea, Potamnogeton)
winter-buds.
2
Journ.ofEcologyI

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18

n " and Statistical Methods

Raunkiaer's " Life-fjrs

Therophytes, or plants of the favourableseason, live throughthe unfavourable season as seeds; hence theyare annual plants. They are speciallycharacteristic
of deserts and of regions under high cultivation. In temperate regions, two
divisions are recognised: (a) summer-floweringannuals, (b) winter-flowering
annuals-e.g. Viola tricolor-which pass through the winter in a vegetative
condition.
Raunkiaer's papers, as well as an earlier paper by Warming in 1891, contain
a wealth of observation and illustrationsof the vegetativeparts of iimany
species,
alnd form a distinguishing feature of the Danish school. Whilst the above
groupingis easily enough determinedin most cases, there is sometimesdifficulty
in
deciding on the type,and-as Raunkiaer points out-a species may belong to one
type in one countryand to another type elsewhere. Thus, Ostenfeld(Botany of
theFaroes, 1908) includes Montia lamnprosperma
both as an annual of the corn-fields
and as a perennial helophytein natural formations; also Jfatricariainodora var.
phaeocephala as an annual in the potato-fieldsand as a perennial hernicryptophyte
in the sand-strandvegetation.
In his 1908 paper (4), Raunkiaer bases his analyses on the following ten
life-forms:
1.

= Stem-succulents.

6.

Ch

2.

= Epiphytes.

7.

3.

MM-

8.

Geophytes.

9.

HH

Helophytes and Hydro-

4.

= Microphanerophytes.

5.

Megaphanerophytes and
Mesophanerophytes.
Nanophanerophytes.

- Chamaephytes.
iHemicryptophytes.

phytes.
10.

Th

Therophytes.

This list, startingfromthe phanerophytes,formsa series in which each type is

on the whole better adapted for the rigour of the critical period than the one
precedingit. The succulentsand epiphytesstand apart by themselves.
The utilisation of these life-formsin plant-geographyis only indicated in a
general way in the earlier papers, but in the 1908 one (4) this aspect is elaborated.
In Table I we give a few of Raunkiaer's analyses for various types of climate; the
locality is indicated, also the total species actually analysed, while in the later
the results are given as percentages according to the grouping given
colutmins
above. Such an analysis for any region is termeda biological or phyto-climatic
spectrumn,. The normal spectrumis the base-line,and the outstandingfeaturesof
the other spectra are deduced by comparison,not by the highest percentagein
their own curve, but by the atmiount
of variation fromthe normal spectrum. The
latter is, ideally, the phyto-climaticspectrumof the whole earth; actually, it is
obtained by computation,and at present is given olulyas approximate. It was
arrived at by firstselecting 1000 representativespecies, and then taking 400 of
these which were carefully analysed. This number 400 has been carefully
controlled in various ways. Thus, taking the stem-succulents,the number of
species known is about 1300, while the total species of flowering-plants
(Raunkiaer
excludes pteridophytes)is taken as 13,000: this gives the 1 per cent. of the normal
spectrum. Again, the Compositae have been estimated as one-tenthof the total

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W. G.

SMITH

19

flowering-plant
population of the earth, and of the actual 400 plants analysed, 45
were Compositae, hence the spectrum in this respect approaches to an average.
Other evidenicededuced fromhis study of spectra leads Raunkiaer to reg,ardthe
normal spectrumas a good workinghypothesis.
What do the numbers signify? If the numbers for Seychellesin Table I are
considered,it will be seen that every life-formis represented,hence this is a rich
and varied flora,differingmost fromthe normal spectrumin the high percentage
of MM and M, and the low percentage of H. If Seychelles and the Danish
TABLE

Examples of Biological Spectra

Total
No. of
Species
400

...
NormalSpectrum
Eastern N. America(B)
Baffin'sLand
...
Labradorcoast
...
Georgia
...
...
Danish West Indies ...
WesternN. America(C)
St Lawrence(Alaska)
Sitka ...
...
...
Death Valley ...
WesternEurope, etc. (C)
Francis JosephLand
..
Spitzbergen ... .
Iceland
...
...
Denmark
...
...
...
Stuttgart ...
Madeiralowlands ...
LibyanDesert (Egypt)
Aden.
.
...
...
Seychelles.

129

246

Percentageof Species belongingto each Life-form

S

717

(O1)
2

126
222
294

904

25
110
329
1084
862
213
194
176
258

MIM

-..-

(0-4)
1

1
1

20

Ch

27

HU

1
8
11
30

30
L7
4
12

51
5:2
55
9

13
9
4
3

3
5
6
1

2
6
8
14

23
7
7

61
60
18

11
10
2

4
7
5

1
5
42

32
22
13
3
3
7
:21
27
6

60
60
54
50
54
21
20
19
12

13
10
11
10
4
3
3

MI

1
7
23

3
2
-

17

1
3
- _
3
1O

(0 1)
-

3
3
1
3
7
23

5
21
1
2
3
3
14
9
26
24

Th

2
10
11
7
3
1
2

13

2
11
18
17
51
4:2
17
16

the same varied flora

West Indian islands (St Thomas and St John) be conmpared,
is revealed, but the optimum point is shifted to the right, N and Ch being
distinctlyincreased in the tropical but drier climate of the Danish West Indies.
This is furtheremphasizedif the spectrumforAden be compared: more extreme
droughtconditionshave raised Ch and H. The moistertropicsshow in this as in
other cases a marked preponderanceof phanerophytes--inother words,they are
distinguishedby a phanerophyticclimate. With increasingdroughtthe conditions
become moreantagonisticto plant-life,and the less protected life-fornms
give place
to the more protected.
Raunkiaer's statisticsreferas yet onlyto the NorthernHemisphere,and within
this he recognisesthree principal regional climate-zones:
(A) a tropical area with uniformn
and high temperatures,but a varying
humidity; passing northwardsfrom this we lhave

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20

Raunkiaer's "Life-forms" and Statistical Methods

(B) an area of decreasing warmth correlated with an increasing difference

between summerand winter,but with a precipitationsuitable at most times for
plant-life--such areas distinguish the eastern parts of the continental masses
(America and Europe-Asia);
(C) warmth decreasing from equator to pole as in B, but with decreasing
precipitation,at least in sutmmer(although furthernorth the conditions do not
fromB)-characteristic of the westernparts of the continents.
differmuuch
These clinmate-zones
are characterisedas A, B, and C climate-seriesrespectively,
and they are limited by biochoresor plant-climateboundaries as suggested by
Koppen. As yet Raunkiaer has not attemptedto lay down the biochoresexcept
in a tentative manner. On exaiiining these climate-zones,it is seen in eastern
North America (B series) that the southern spectra of the West Indies and
Georgia have a large number of species with all the life-formsrepresented,and
with a large proportionof phanerophytes. The series suggests three types of
climnate-phanerophytic,
hemicryptophytic,
and chamaephytic. Western North
America shows a C series,in which desert regions are stronglyrepresented. In
the south, Death Valley correspondsto Georgia, and the better protectedforms
predominate,especiallythe stem-succulents
and therophytes. Furthernorththere
is closer agreementwith the B series. In the Old World the climate-zoneC( alone
has been followed out. There, is here a distinct contrast between therophytic
(above 30 per cent.) and hemicryptophytic(above 40 per cent.) climates. The
therophytesincrease in deserts,and again in the more cultivated areas (Denmark
and Stuttgart). The application of the therophytetest is shown in Paulsen's
recent memoir'; the Transcaspian lowlands lie near the southernRussian steppes
(Yekaterinoslaw) and the Pamir highlands,but the spectrumrevealsclose similarity
with desert regions adjoining the Mediterranean (Libya, Samos, Cyrenaica), and
this with otherconsiderationsleads Paulsen to place Transcaspia with the deserts
ratherthan with the steppes.
These statistical methodsare also applicable to more limitedareas, and a fairly
exhaustivestudy of the Arctic regionhas been made (4). Here the chainaephytes
become importantguides, and significanceis attached to the biochores indicated
by 10, 20, and 30 per cent. of chamaephytes. Thus in Nova Zembla the chamaephytepercentageof the whole florais 19, but a more detailed analysis shows that
in the zone 70-7I' N. Lat., and in each degree-zonenorthwardsthe chamaephytes
exceed 20 per cent. (75-76?, 31 per cent., 76-77?, 50 per cent.). The course
of the 20 per cent. Ch biochore has been traced by Raunkiaer throughall the
circunipolarlands: it lies between Scandinavia and Spitzbergen and fringesthe
arctic coasts of Asia, crossing Bering Straits between St Lawrence Island and
the CommanderIslands and followingthe arctic coasts of Canada. The 30 per
cent. Ch biochore includes BaffinLand, Francis Joseph Land, and Jan Mayen,
but does not touch the arctic coasts of Asia or Alaska. The 10 per cent. Ch
biochore lies between the Faroes and the Shetlands, in Norway it divides the
lowlands fromthe highlandsand the extremenorth,while in Siberia it crossesthe
Yenisei about 67? N. Lat., and in America it lies between Sitka and Chilcat
1 Paulsen, 0. "Studies on the vegetationof the Transcaspianlowlands." Second Danish
PantirExpeditionReports,Copenhagen,1912.

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W. G. SMITH

21

(Alaska)and through
SouthLabrador. Thesebiochores
are regardedas significant,
as theyagree approximately
with recognisedisotherms:the 20 per cent. Ch
biochorecorresponds
closelywiththe4.40 C. Juneisotherm,
and the 10 percent.
one with the 10?C. June isotherm. In orderto comparedifferent
clirnates,
Raunkiaerdevised" hydrotherimic
figures,"
by representing
in a singlediagramthe
curveof themonthly
averagesoftemperature
and thatofthemonthly
averagesof
rainfall;then,by determining
the biologicaltypesor combinations
of typescorre"
spondingwiththehydrothermic
figures,
he obtainedthe " biologicalexpressions
of thevariousclimates. For instance,the tropicalclimate,constantly
warmand
humid,is characterised
by the predominance
of phanerophytes
belongingto the
leastprotected
sub-types;thesub-tropical
withwinterrains,by evergreen
climate,
phanerophytes
with unprotected
buds. Temperateregionsare distinguished
by
the essentiallyhemicryptophytic
characterof the vegetation; desertsby the
therophytic
type.
Summarisiing,
it thus appears that fromthe equatornorthwardsthereis a
and therophytes
definiteseriesof biologicalspectra. The phanerophytes
develop
withwarmth,
and northwards
decreaseand disappear. The cryptophytes,
although
widelydistributed,
also disappearin the north. The hemicryptophytes
reach a
maximum
in temperateclimates,wheretheyremainconstantat aboutdoublethe
and theirbiochores
increasenorthwards,
normalspectrum. The chamaephytes
zones: (1) cold-temperate
delimitcertainphyto-climatic
zone,the hemicryptophyte
zone south of the 10 per cent. Ch biochore;(2) boreal zone of H and Ch
betweenthe 10 andthe 20 percent.Ch biochores;(3) arcticzone,thechamaephyte
zone from20 to 30 per cent. Ch biochores;(4) arctic-nivaldomainwithover
30 per cent. Ch.
TABLE II
Puschlav(Switzerland)
AttitudinalDistributionof Life-forms,
Altitude
Above2850 metres
2550-2$50,,
2250-2550,,
1900-2250,,
1550-1900,,
1200-1550 ,,
850-1200,,
,,
Below 850

...
...
...
...

NormalSpectrum

...

...
...
...
...

Total
No. of ,
Species S
51
199
348
492
487
449
604
447
400

Percentageof Species belongingto each Life-form

N
A
_
G
mm
N
HH Th
W
E
H
Ch
61
2
35
2
4
4
25
67
64
1
3
7
6
1
18
68
8
1
1
3
13
6
4
62
10
1
3
11
8
4
2-5 2-5
60
9
1
7
14
55
2
5
5
9
2
3
19
(0,2)
55
8
5
1
21
3
4
3
(0 2)
3

17

20

27

13

The applicationof thesenmethods

to altitudinalzones will be seen fromthe
detailsof thePuschlavvalleyin Switzerland,
shownin Table II. At lowerlevels
thereis a completeseriesfromthe phanerophytes;
passingupwardsthereis a
markeddecreaseof all groupsexceptCh and H, and the 10, 20, and 30 percent.
are revealed. The same successionholdsgood forothermountains
Ch biochores
whichhavebeen investigated;the variationin H is slight,thatof Ch great,but

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22

Raunkiaer's " Life-forms" and Statistical Methods

the altitudesof the biochoresvary greatly. Thus in the Western Alps the 20 per
cent. Ch biochore is at 2440 metres; on Tatra in the Carpathians at 2200 min;
in Noiway between 1000 aind 1200 m.; on Clova mountainsin Scotland, between
700 and 800 m.; and in the Faroes somewhatbelow 500 m;
As Raunkiaer remarks,much remains to be done towards perfectingthese
phyto-climaticregions. The necessary spectra can only be constructedwherethe
Uora is completelyknown, and recorded inuch better than is usually done in
lands there is thus a gap as regardsthe flora
floristicworks. In the circumpol-ar
of northernCanada, and growth-forms
are too often very imperfectlydescribed,
even in the standard British floras for instance. From such defective material
Raunkiaer has worked out a -system which all must admit is of the greatest
interest and importance.
The system works much better in limited areas such as islands. Yet it is
clear that our methods of subdividing continentalareas with varying altitudes
are so defectivethat any methodsuch as Raunkiaer's meritscarefulconsideration.
In 1909 (5) Raunkiaer applied his inethodsto the study of strand vegetation
in the Danish West IIndian islands where the soil is new, for instance in lagoons
which are being formedand gradually filledup and covered with plants, and on
sandy shores. He compared in detail a small peninsula of alluvial soil in Santa
Cruz with a sandv peninsula on the coast of Denmark, and found that the
on the new soil in both localities depends upon the
percentage of growth-forms
climatic and not upon the edaphic factors. The West Indian peninsula showed
69 per cent. of phanerophytesand the Danish peninsula 47 per cent. of hemicryptophytes-that is, in -both localities the same type predominatesas in the
countries to which they belong. He gives other examples to illustrate the fact
that the geological age of a flora has no influencein determiningits biological
spectrum,which is determinedsolely by climate. He then considersthe question
whetherthe biological spectrumof a countrycan be changed by immigrationfrom
other countries,and points out that immigratedand naturalised species belong to
and in the same mutual proportionas the native plants of
the same growth-forms
the country.
or the vegetation
The sixth paper cited (6) is a study in " Forrnationslaeren,"
on a given soil or substratum,in contrast to "plant-climatology,"the term
suggestedto definethe groupingwithin the larger domains determinedby climatic
factors,as dealt with in our earlier part. Raunkiaer points out that hithertothe
treatmentof plant formationshas been largelysubjective,each observerestimating
the value of the species, so that differentobservers may estimate the same
formationdifferently.It is true that any of the exact methods,e.g. the quadrat
method of Clements,the "gridiron" of Oliver and Tansley', and the Rothamsted
method of analysing and weighing grass herbage,need much time, and probably
have revealed the difficultiesratherthan solved theimi. Raunkiaer's methoddoes
not attempta completeanalysis of any unit-area,but it is claimed that it gives a
correctexpressionof the mass-conditionsof the species in any plant community
(formiation)by means of mechanical enumeration,so that comparable results are
obtained. The method consists in recording the species on a unit-area; each
I New Phytologist,
8, 1904, p. 228.

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W. G.

23

SMITH

species counts as one, and no attemptis made to distinguishbetween commonor

rare species. After 50, 100, or more unit-areas of a given plant communityare
thus recorded,the occurrencesare added; the dominantspecies may occur in each
unit-area,the rare on veryfew. Table III, slightlyabridged by omittingspecies
hardly represented,shows-resultsfromthe analysis of a beech-woodundergrowth.
Anemonenemorosaoccurs in 5 out of every 5 areas examined,the other species
are given on the same scale. The lettersslhowthat there are 7 geophytes,10 heminemorosais to all the others
cryptophytes,
and 1 chamaephyte,and that Ane,mone
as 6: 94.
TABLE

III

Analysis of the Undergrowthof a Beech Forest,from Raqunkiaer(abridged)

of Unit-Areas
Number
examined
Size ofUnit-Areain sq. m.

...

nernorosa
An7emozon7e
.
Gagea litea
RanuncidusFicaria
Oxalis Acetosella
Melica uniflora.H
Miljion effisum......
Corydalisintermedia
4 otherspecies ...
6
...
1 ,,i

...

...
..
...
...
...
...
...

...
...
...
.

Proportionof Aitentone
ntentorosa

10

20

5
3
2
1-5
1P5
1-5
05
05
1P5

5
1-5
0 25
0 75
025
1
05
05
-

10
G
G
H
H
H
G
G
H
h
_JL
9

29

100

9
491

T5

400

01

0 01

5
0-85
0 05
0.8
0-25
0-2
02
0-2
005

5
0-26
0-06
0-25
0 01
0 05
013
-

A(?

Shoot-count
-

5
0 03
0 09
0-2
0.008
-

0 03
-

The number at the head of each of the firstfour columns gives the actual
number of unit-areas examined and the size of the unit-area; the numbers are
reduced to a scale with highest number5, which Raunkiaer prefersto a decimal
or percentage scale. The last column gives results by counting the number of
shoots (leafyand flowering)of Anemonenemorosaon 1 sq. me.tre; 1281 were thus
counted and their distributionis shown on a quadrat figure. The smaller unitareas are obtained by means of a wooden frame of the-requisite size which is
thrown down and the included plants counted.
In a later paper (7) an improved type of apparatus is recommended. This
consists of a ring to be fixed on a walking-stick;a piece of metal is fixedon the
ring,with a screw-threadinto which a metal rod can be screwed,and the length
of this rod is equal to the radius of a circle of 01 sq. metreor any requiredunitarea. The stick is thrust into the ground at intervals,and the species occurring
withinthe circle describedby the tip of the rod are recordedand tabulated.
The importanceof Raunkiaer's contributionlies in the careful examination
of results obtained on unit-areas of differentsizes'. That variable results are
1 Pethybridgeand Praeger ("Vegetation south of Dublin," Proc. B. Irish Acad. 25, 1905,
areas
p. 141) used a similarmethod. Their lists were preparedfroma numberof representative
by a mechanical process; the lists for each association were tabulatedby numbers,and from
this the generallist foreach plant association was preparedaccordingto an examplegiven; the
methodwas not followedout in detail.

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24

Raunkiaer's "L?fe-forms"and >StatisticalMethods

obtained will be seen fromthe table, and careful interpretationis necessary. It

was found that the smiialler
the unit-area the more closelydoes the percentage of
Anem)onecorrespondto the actual couInt of shoots. The amount of time needed
for a 10 sq. m. area is an objection; on the other hand, with simallareas great
care must be exercised to exclude (as is done) tips of shoots withinthe framebut
not rooted in the unit-area. These and othervariationsare fullydealt with in a
series of tables. As the result of experience,Raunkiaer regards01 sq. m. as the
InoSt useful area. The numberof readingsis variable, but theymust be carried on
till a relativelyfixed scale is reached so that furtherreadings do not materially
alter the averages from precedingreadings; in the case of the beech-woodundergrowthdealt with, 50 readingsof 01 sq. m. gave stability.
Referringto these variations Raunkiaer says (5, p. 38):
"The table (Table III of this paper) shows that by using 0 01 sq. m. as unitarea, and also with 01 sq. m., each species acquires the same degree of frequency
as in a shoot-countingmethod if the frequencybe expressed in whole numbers
fromI to 5; if one uses half-degreesor a decimal scale in determiningfrequency
then Cagea lutea and Oxalis Acetosellaare greaterit is true by half a degree; but
in consideringthe affairsof nature, one cannot and ought not to place any value
on a variation between two formationswhich merely consists in one or more
species of one having a frequency,e.g. 2, while in the other formationtheyhave
2-5 or 1-5."
It seems then that this method has its limitations and, as with all other
methods,a certain amount of subjective common sense must be used, and this
of the memoir (pp. 41-129) is conbecomes more evident when the remiiainder
sidered. This takes up the examiinationof a number of representativetypes of
vegetation in Denmark. Besides the statistical method just described, there is
introducedthe distributionof life-forms;thus, the Anemonenernorosafacies of the
beech-woodis geophyticsince the 01 sq. m. readings show geophytes82 per cent.,
18 per cent.
hemicryptophytes
The firstseries of studies is on beech and coniferwoods, scrub and cultivated
lands on glacial clays and overlyinghumus soils. A second series deals with
heaths, bogs, etc., on glacial sands, and a third with formationson salt soils. The
may be used to
types are too numerousto take up separately,but several examnples
the workingof the statistical methods.
examnine
A successful application is evident in a comparisonbetween (a) a grass-field
one year sown, (b) grass-field8 years old, (c) the field margins. It appears that of
110 species recorded 10 occur in all three situations, while in each case 5 or 6
species constituteover 66 per cent. of the whole vegetation. There is also a sharp
distinctionof the one-yeargrass by the preponderanceof therophytes,while the
8-year grass and the field margins are mainly H and Ch. The decrease of
Rumex Acetosella after the first year, and the increase of Ranunculus repens
between the firstand eighth years, are also revealed.
The examination of retrogressivescrub, with oak, hazel and blackthorn,
destroyedby disforestingand grazing,a conditionwell-knownin Britain, reveals
some weaknesses of the mechanical method. In order to arrive at a list of the
scrub vegetation, there are deducted "humus-planits"aind species presumed to

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W. G.

SMITH

25

have come fromadjoining cultivated fields; there remains a reputedflora of the

mineralsoil. Here the experienceof the observeris called on to discriminate,and
one can imaginethat the resultsof various observersmightdifferconsiderably.
The studies of the ground-vegetationof woods show the methodin one of its
best aspects-the detection of variations due to soil and illumination. Thus in
one table (No. 22) readings are shown fromunit-areasin differentparts of spruce
woods; Aira,flexuosa and Oxalis Acetosetlawere the two dominant plants, but
theyoccur in different
proportionsand give rise to the followingfacies (each based
on 50 readings of 0 1 sq. m.):
(a)
(b)
(c)
(d)
(e)

itira facies in open spruce wood--Aira 50, Oxalis 1, 6 other species

varying from 1-4.
Aira-Oxalis facies-Aira 50, Oxalis 50, 7 other species 1-6.
Oxalis facies in denser and gloomier places-Aira 0, Oxatis 50,
a other species 1--4.
No floweringplants in quite dense and dark places.
Oxalis facies in transition froinspruce to beech with AnemonefaciesOxalis 50, Aira 0, 9 other species 1-4.

The contrastbetween spruce hemicryptophytic

wood and beech geophyticwood
is also revealed. These and other studies show that the method has great value
in showing up differencesin the plant covering; it does not claim to explain the
variations. Under the author's experienced guidance the results obtained are
valuable guides and enable one to followthe distribution. But in unskilledhands
a mechanical method may be made, by formidablenumerical lists, to prove what
statistics usually prove-namely, anything the speaker wishes! It will still be
possible to obscure importantfacts in plant distributionquite as much as any
subjective method has done. Thus, in woods one frequentlyfindsa drier type of
vegetationinterruptedby moistspring-flushes
with an entirelydifferent
vegetation.
If 50 readings are made at random and the results averaged, the existence of a
flushvegetation mightbe masked,and another observeron the same ground but
selectingonly the drier vegetationwould get a differentset of statistics. In the
same way, variations in illuminationand topographicalfeatures mightbe slurred
over by untrainedobservers. If, however,as Raunkiaer has shown,the divergent
readings are kept apart, a considerable advance is possible in ecological studies.
The heterogeneousnature of migratoryformationson non-stabilisedsoils' is at
once revealed,as in the salt-marshvegetationdescribedforDenmark. The method
also works well in determiningthe compositionof close vegetation like grassland,
but it is not intendedfordetermninations
of yield per acre foragriculturalpurposes.
In the case of uniformvegetation on stable soils, e.g. a Callunetum,the meagre
floristiclist and the openness of the vegetation should renderelaborate statistical
methodsunnecessary.
Raunkiaer's work appears to have been hitherto somewhat neglected by all
except Scandinavian botanists; among recentpublicationsby the latter,containing
results of general interest arising fromthe application of Raunkiaer's systemin
1 See C. B. Crampton: "The geological relations of stable and
migratoryplant formations." ScottishBot. Revietv,1, 1912. (Reviewin this Journal,1, 1913, p. 47.)

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26

Raunkiaer's "Life-forms"and Statistical Methods

the study ofrvegetation, mention may be made (in addition to t-hememoirby

Paulsen already referredto) of a paper by Simmonson Swedish Lapland (reviewed
in this Journal, 1, p. 64). There can be little doubt that Raunkiaer has opened
up a promisingfield for investigation,and it seems no less certain that when
broughtmoreprominentlyto the attention of botanists in all parts of the world
his system will he applied extensively and with valuable results. For instance,
the investigationof the florasof larger or smaller areas fromthe points of view
here indicated would forman excellent branch of study forthe fieldbotanist,even
if the immediate outcomewere merelyan annotated list of the species, with the
determinationof the "biological type" to which each belongs. Raunkiaer has
himselfbroughtout a new edition of his Dansk Ekskursions-Flora,
(1906) in which
the biological type is indicated under the descriptionof each species. The very
fact that it is sometimesdifficultto decide to which type a species should be
assigned would doubtless lead to a nmoredetailed investigationof the life-histories
of many floweringplants-autecological studies of this kind are among the most
urgentlyneeded pieces of ecological work. On the other hand, the application of
Raunkiaer's principleswould also involve morethoroughand detailed observation
of climatic and other habitat factors than is usually made in phytogeographical
study.
For furtherdetails, referencemay be inade to the 1908 paper, which gives
tables based upon data fromvarious parts of the world,and an extensive bibliography.

LIST OF RAUNKIAER'S MEMOIRS REFERRED

TO

(1) " Om biologiskeTyper,med Hensyntil PlanternesTilpasning til at overleveugunstige

Aarstider." Bot. Tidsskrift,26, 1904.
(2) "Types biologiquespour la geographiebotanique." Btll. Acad. Roy. d. Sci. de Danemiark,1905, pp. 347-437, 41 figures.
(3) "Planterigets Livsformerog deres Betydningfor Geografien." Kjobenhavn, 1907,
132 pp., 1 plate, 77 figures.
(4) "LivsformernesStatistiksom Grundlagfor biologiskPlantegeografi." Bot. Tidsskr.,
29, 1908, pp. 42-83, 34 tables. (Translationby G. Toblerin Beih. Bot. Centralbl.,27, Abt. 2,
1910, pp. 171-206.)
Acad. Sci. de Danemanrk,
(5) "Livsformenhos Planterpaa ny Jord." Ml1ern.
8, 1909, 70 pp.,
29 figures.
(6) "Formnationsundersogelse
og' Formationsstatistik." Bot. Tidsskr.,30, 1909, 110 pp.,
20 figures.
(7) "Measuring apparatus for statisticalinvestigationsof plant-formations."Ibid., 33,
1912, pp. 45-48, 1 figure.

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