Floral formula

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K5[C(5)A5]G(5)
FloralformulaofAnagallisarvensis.[1]:307Polysymmetricflower.The
perianthconsistsof5freesepalsand5joinedpetals,whicharefused
withandroecium.Flowerisbisexual,itcontains5stamens,pistilis
fusedoffivecarpels,ovaryissuperior.
Floral formula is a means to represent the structure of a flower using numbers, letters and various
symbols, presenting substantial information about the flower in a compact form. It can represent
particular species, or can be generalized to characterize higher taxa, usually giving ranges of organ
numbers. Floral formulae are one of the two ways of describing flower structure developed during the
19th century, the other being floral diagrams.[2] The format of floral formulae differs between authors, yet
they tend to convey the same information.[1]

Contents
[hide]

1History

2Contained information

2.1Organ numbers and fusion

2.2Ovary position

2.3Symmetry

2.4Sexuality
3Examples

4See also

5Notes

6References

7Bibliography

8External links

History[edit]
Floral formulae were developed at the beginning of the 19th century.[2] The first authors using them were
Cassel[3] (1820) and Martius[4] (1828). Grisebach[5] (1854) used them in his textbook to describe
characteristics of floral families, stating numbers of different organs separated by commas and
highlighting fusion. Sachs[6] (1873) used them together with floral diagrams, he noted their advantage of
being composed of "ordinary typeface". Although Eichler widely used floral diagrams in
his Blthendiagramme,[7][8] he used floral formualae sparingly, mainly for families with simple flowers.
Sattler's[9] Organogenesis of Flowers (1973) takes advantage of floral formulae and diagrams to describe
the ontogeny of 50 plant species. Newer books containing formulae include Plant Systematicsby Judd et
al.[10] (2002) and Simpson[11] (2010). Prenner et al. devised an extension of the existing model to broaden
the descriptive capability of the formula and argued that formulae should be included in
formal taxonomic descriptions.[2] Ronse De Craene (2010)[1]partially utilized their way of writing the
formulae in his book Floral Diagrams.

Contained information[edit]
Organ numbers and fusion[edit]
The formula expresses counts of different floral organs,[note 1] these are usually preceded by letters or
abbreviations according to the organ type. They are ordered corresponding to the arrangement of the
parts of the flower from the outside to the inside:

Bract
s

Bracteol
es

Tepals
(perigonorperiant
h),orsepals(calyx)
andpetals(corolla)

Stamens(androeciu
m)

Carpels(gynoeciu
m)

Ovule
s

V[2]or
O[9]

P[2]orCaCo[12]
B[2]

Bt[2]
K[2]or
Ca[12]

C[2]or
Co[12]

The labels with darker backgrounds are less common. "V" used by Prenner et al. for the number of
ovules per gynoecium is followed by lowercase letter describing the type of placentation.
For epicalyx/calyculus, the letter "k" is used.
The numbers are inserted after the labels, they may be formatted as sub- or superscript. If an organ is
absent, its number is written as "0" or it is omitted, if there are "many" (usually more than 1012)
instances, it can be written as "". Whorls of the same organ are separated by "+". Organ counts within

a whorl can be separated by ":", for example when part of the whorl is morphologically different. A range
can be given if the number is variable, e.g. when the formula summarizes a taxon.
K3+3 a calyx with six free sepals, arranged as two separate whorls
A many stamens
P312 perianth from three to twelve tepals
Groups of organs can be described by writing the number of instances in the group as
superscript.
A5 5 groups of 2 stamens
The formula can also express organ fusion. Fusion of one organ type can be shown
by enclosing the number in a circle, fusion of different organs can be represented
by ties, as e.g. in Judd et al. Prenner et al. state that this method is difficult to achieve
via standard typesetting.[2]:242 Joining of organs can be more readily written
using parentheses "()" if instances of the same organ are fused. Fusion between
different organs can be achieved by square "[]", eventually curly brackets "{}".
A(5) five fused stamens
[C(5) A5] corolla fused from 5 petals, fused to stamens
Prenner et al. propose superscript zero for a lost organ, and superscript "r"
for a reduced one. Ronse De Craene uses a degree symbol to mark
a staminode (infertile stamen) or pistillode (infertile carpel).
A3:2r+50 (Prenner et al.) androecium in two whorls, first contains 3 stamens and 2
staminodes, second whorl lost
A1+2 (Ronse De Craene) androecium in two whorls, first whorl containing a stamen, second
whorl containing two staminodes

Ovary position[edit]
Ovary position is shown by alternating the "G" label. Simpson
circumvents the intricate formatting by expressing the ovary
position by words.

superiorovary

Prenneretal.,[2]:243RonseDe
Craene[1]:39

ha

inferiorovary

G,inferior

G,

G
Sattler[9]:xviii

Simpson[11]

G,superior

Symmetry[edit]
Symmetry or arrangement may be described for the whole flower,
in such case the corresponding symbol is usually placed at the
beginning of the formula. It may be also outlined separately for
different organs, placing it after their labels or numbers, or it may

not be included in the formula at all. It is described by following

symbols:

polysymmetry
(actinomorphi
c)

Prenneretal.
[2]:242

RonseDe
Craene[1]:39

Sattler[2]:xviii

Juddetal.[10]:66

disymmetr
y

monosymmet
ry
(zygomorphic
)

asymmetr
y

,or,
dependingon
thesymmetry
plane
orientation

,arrow
orientation
dependingon
symmetry
plane
orientation

Subrahmanyam
[13]

Rosypal[14]

X
not
mentioned

%inmedian
plane,
in
lateralplane

Sexuality[edit]
Sexuality of the flower can be highlighted by or
for
hermaphrodite (bisexual), for male (staminate) and for
female (pistillate) flower. The symbols are usually placed at the
beginning of the formula, after or before the symmetry symbol.
Prenner et al.recommend to use the corresponding symbols (
and ) only for flowers of separate sexuality. Ronse De Craene
utilizes the words "pistillate" or "staminate" instead of the
symbols.
Floral formula can also incorporate the fruit type, Judd et al.
[10]
place it at the very end.

not
mentioned

Examples[edit]
K3 [C3 A13+:2] (3)[1]:39 the formula of Canna
edulis; asymmetric flower; calyx of three free sepals; corolla of
three free petals joined with androecium; androecium in two
whorls, the outer whorl contains 13 staminodes, the inner
contains of a stamen and 2 staminodes; gynoecium fused of 3
carpels, inferior ovary

B BtC K3:(2)C C3:2r A(3):2r+4r:10 G1 Vm810[2]:246 the

formula of Tamarindus indica; bract and petaloid bracteoles;
monosymmetric calyx of three and two petaloid sepals;
monosymmetric corolla of three and two reduced petals; two
whorls of stamens, the outer monosymmetric from three fused
and two reduced stamens, the inner of 4 reduced and 1 lost
stamen; monosymmetric gynoecium of 1 carpel with superior
ovary; marginal placentation with 810 ovules per gynoecium.

See also[edit]

Floral diagram

Notes[edit]
1.

Jump up^ Bracts and bracteoles are not considered

parts of the flower itself, yet they may be included in
the floral formula

References[edit]
1.

^ Jump up to:a b c d e f Ronse De Craene, Louis P.

(2010-02-04). Floral Diagrams: An Aid to
Understanding Flower Morphology and Evolution.
Cambridge: Cambridge University Press. ISBN 9780-521-49346-8.

2.

^ Jump up to:a b c d e f g h i j k l m n Prenner, Gerhard;

Richard M. Bateman; Paula J. Rudall (February
2010). "Floral formulae updated for routine inclusion
in formal taxonomic descriptions". Taxon. 59 (1):
241250. ISSN 0040-0262.

3.

Jump up^ Cassel, F. P. (1820). Morphonomia

botanica: sive observationes circa proportionem et
evolutionem partium plantarum. Colonia Agrippina
[Cologne]: M. DuMont-Schauberg.

4.

Jump up^ Martius, C. F. (1828). "ber die

Architectonik der Blthen". Isis (Oken) (21): 522
529.

5.

Jump up^ Grisebach, A. (1854). Grundriss der

systematischen Botanik. Gttingen: Verlag der
Dieterichschen Buchhandlung.

6.

Jump up^ Sachs, J. (1873). Lehrbuch der Botanik

nach dem gegenwaertigen Stand der Wissenschaft.
Leipzig: Engelmann.

7.

Jump up^ Eichler, August Wilhelm

(1875). Blthendiagramme, erster Theil: Enthaltend
Einleitung, Gymnospermen, Monocotylen und
sympetale Dicotylen. Leipzig: Wilhelm Engelmann.

8.

Jump up^ Eichler, August Wilhelm

(1878). Blthendiagramme, zweiter Theil: Enthaltend
die apetalen und choripetalen Dicotylen. Leipzig:
Wilhelm Engelmann.

9.

^ Jump up to:a b c Sattler, Rolf (1973). Organogenesis

of flowers; a photographic text-atlas. Toronto,
Buffalo: University of Toronto Press. ISBN 0-80201864-5.

10. ^ Jump up to:a b c Judd, Walter S.; Christopher S.

Campbell; Elizabeth A. Kellogg; Peter F. Stevens;
Michael J. Donoghue (2002). Plant Systematics: A
Phylogenetic Approach (2nd ed.). Sunderland,
Mass., U.S.A.: Sinauer
Associates. ISBN 0878934030.
11. ^ Jump up to:a b Simpson, Michael George
(2010). Plant Systematics. Oxford (Great Britain):
Academic Press. ISBN 978-0-12-374380-0.
12. ^ Jump up to:a b c http://www.cactus-art.biz/notebook/Dictionary/Dictionary_F/dictionary_floral_formul
a.htm
13. Jump up^ Subrahmanyam, N. S. (1997-0101). Modern Plant Taxonomy. Jangpura, New Delhi:
South Asia Books. ISBN 9780706993462.
14. Jump up^ Rosypal, Stanislav (2003). Nov pehled
biologie. Praha: Scientia. ISBN 80-7183-268-5.