TAXON 63 (5) • October 2014: 1103–1111 Ronse De Craene & al.

• Floral formulae

ME THODS AND TECHNIQUES

Understanding the structure of flowers—The wonderful tool of floral

formulae: A response to Prenner & al.
Louis Ronse De Craene,1 Akitoshi Iwamoto,2 Kester Bull-Hereñu,3,4 Patricia Dos Santos,1 Javier A. Luna1,5
& Jennifer Farrar1,5

1 Royal Botanic Garden Edinburgh, Edinburgh EH3 5LR, Scotland, U.K.

2 Department of Biology, Tokyo Gakugei University, Tokyo, Japan
3 Escuela de Pedagogía en Biología y Ciencias, Universidad Central de Chile, Santiago, Chile
4 Departamento de Ecología, Pontificia Universidad Católica de Chile, Santiago, Chile
5 Institute of Molecular Plant Sciences, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JH, U.K.
Author for correspondence: Louis Ronse De Craene, l.ronsedecraene@rbge.ac.uk
ORCID: LRDC, http://orcid.org/0000-0002-8333-4596; AI, http://orcid.org/0000-0001-6492-495

DOI  http://dx.doi.org/10.12705/635.35

Abstract  This paper is a discussion and elaboration of a paper by Prenner & al. (2010), entitled “Floral formulae updated for
routine inclusion in formal taxonomic descriptions”. The aim of the Prenner paper was to promote the use of floral formulae
in botany and to reach a consensus among botanists for best practice. An important purpose of floral formulae is to induce
users to observe and describe flowers accurately. It is proposed that additional information on anther, ovule, style and stigma
should be included. Also, only visible organs should be included in a formula and theoretical speculations should be illustrated
with floral diagrams, which are complementary to formulae, unless there is good reason to include absent organs. We propose
a universal, standardized method to accurately shorthand a description of a flower. The level of detail given in the formula can
be highly flexible and depends on the intentions of the user.

Keywords  diagnostic characters; floral diagrams; floral formulae; floral morphology

INTRODUCTION
Flowers are remarkably conservative in their numbers
of parts. Whichever flower of Erodium (Geraniaceae) a keen
observer will look at (see Fig. 1A), he will always find the same
number of whorls and the same number of organs within each
whorl of the flower (five sepals and petals, two whorls of five
stamens, one of which is sterile, and five carpels: Fig. 1A).
Flower morphologists have sought for ways to succinctly
describe flowers, and a perfect tool was conceived in the form of
floral formulae. The first floral formulae were created in the first
decades of the nineteenth century and have been progressively
developed especially in Germany. For an overview of the history
of the use of floral formulae we refer to Prenner & al. (2010).
Although floral formulae (together with floral diagrams)
have been integral part of the basic teaching in botany, the
recent depreciation of botany at universities has not helped in
the recognition of the value of this methodology. Few recent
papers or textbooks use floral formulae, often in a very simpli-
fied and frequently inconsistent form, such as Judd & al. (2002),
Stützel (2006), Tsou & Mori (2007), Leins & Erbar (2010), or
Simpson (2010).
The value of floral formulae has been highlighted by
Prenner & al. (2010) who proposed an updated list of symbols
to describe flowers. Prenner & al. (2010: 241) also suggested
that floral formulae should be an inherent component of plant
diagnoses and taxonomic descriptions, “functioning as a logi-
cal phenotypic counterpart to the DNA barcode”. It is clear that
floral formulae have a clear advantage over lengthy descrip-
tions, as the structure of a flower can be presented in a suc-
cinct way, accessible to any reader. As such floral formulae
are a powerful incentive for consistent observations and also
a valuable teaching tool (Prenner & al., 2010). Prenner & al.
advocate the use of symbols that can be reproduced easily with
a non-sophisticated computer, enabling its universal use.
Floral diagrams, two-dimensional representations of the
ground-plan of the flower, are the major counterpart to flo-
ral formulae in floral descriptions. As discussed in Ronse De
Craene (2010), floral formulae have a disadvantage against
floral diagrams in that it is difficult to describe the position
of organs in the flower, and impossible to show any spatial
arrangements or any conspicuous special structures, such as
nectaries, coronas or appendages. However, floral formulae
have the advantage that they do not necessitate any pictorial
means. They also open possibilities for comparative research
and the understanding of trends in flower evolution.
We agree with Prenner & al. (2010) that floral formulae
(as well as floral diagrams) are under-used tools in systematic

Received: 10 Mar 2014 | returned for first revision: 15 Apr 2014 | last revision received: 9 Jun 2014 | accepted: 9 Jun 2014 | published online ahead
of inclusion in print and online issues: 24 Sep 2014 || © International Association for Plant Taxonomy (IAPT) 2014

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Ronse De Craene & al. • Floral formulae TAXON 63 (5) • October 2014: 1103–1111

Fig. 1. Examples of flowers to illustrate different floral formulae. A, flower of Erodium corsicum Léman (Geraniaceae) with five equal petals.
The abaxial side of the flower is below. B, flower of Lewisia cotyledon (S.Watson) B.L.Rob. (Montiaceae). The flower consists of two median
bracteoles and a variable number of tepals (sepals), consisting of two outer lateral sepals (asterisks) and a variable number of median sepals.
C, flower of Penstemon sp. (Plantaginaceae) laid open, showing the arrangement of different parts. The androecium consists of two pairs of sta-
mens of different length. D, lateral view of flower of Blumenbachia hieronymi Urb. (Loasaceae). Note the white petals alternating with coloured
staminode complexes, which consist of five staminodes. E, large complex flower of Nymphaea sp. (Nymphaeaceae) showing an arrangement of
alternating whorls that can be detected in the sepals (white numbers) and outer petals (black numbers); the inner whorls are less clear but androe-
cium and carpels are arranged in ten orthostichies. F, asymmetric flower of Vigna linearis (Kunth) Maréchal & al. (Leguminosae). The abaxial
(keel) petals are transformed into a coil that holds androecium and twisted style.

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TAXON 63 (5) • October 2014: 1103–1111 Ronse De Craene & al. • Floral formulae

research. Therefore, we believe that a broader use of these tools
should be encouraged. Ronse De Craene (2010: 39) came up
with a list of abbreviations comparable to Prenner & al. (2010),
although it was less elaborate. However, the advantages of flo-
ral formulae for educational purposes are clear and lecturers
may adapt a personal approach (E. Smets, pers. comm.). As
suggested by Prenner & al. (2010), their proposed system is
open for discussion, and Taxon is probably a good medium to
start this discussion. The ultimate aim is to reach consensus
among botanists for the use of a balanced and universal system.
A REEVALUATION OF FLORAL FORMULAE
The following points are a criticism of the paper by Prenner
& al. (2010) and suggestions for improvement. We also refer to
Table 1 for clarity.
The floral formulae proposed by Prenner & al. (2010) are
very detailed and contain more information than classical for-
mulae, including extrafloral organs associated with flowers,
such as bracts and bracteoles, and gynoecial characters, such
as the placentation and ovules. However, it is best to maintain
a hierarchical arrangement with subcategories. Treating parts
of organs (i.e., ovules) at the same level as the organs leads to
confusion and makes the floral formulae appear too complex.
A better approach includes a restriction of capital letters to
the main organs associated with the flower (see Table 1) Bract,
Bracteole, Epicalyx (i.e., agroup of structures below the flower
resembling an extra calyx; this can be a whorl of bracts or
structures of unknown/unclear origin), K for calyx, Corolla,
Perigon, Androecium, Gynoecium), and a use of lower case
letters for specific structures that are part of organs, such as
locules, ovules (including their position on the placenta), pollen
sacs, styles and stigmas.
The inclusion of details of the flower, such as the number
of pollen sacs or locules forces the user to accurately observe
the flower to record as many details as possible. Not including
these details in a consistent manner may lead to inconsisten-
cies in the recording of data, either by implying the presence
of floral characters, when these have never been checked,
or by assuming that the user will always know the relevant
characters. Several qualitative features can be shown through
superscript symbols and Prenner & al. use this to great effect.
For example, C stands for the number of petals, while c stands
for petaloidy, which can be used as a special feature of the
perigon. A calyx can often be pigmented, as, e.g., in Fuchsia
(Onagraceae) and this can be shown as Kc. Pc stands for a
petaloid perigon, when it is not possible to differentiate sepals
from petals, as in tulips (Weberling, 1989).
Inclusion of bracts and bracteoles in floral formulas is use-
ful, although strictly speaking they belong to the inflorescence.
Bracteoles are generally paired in eudicots and single in mono-
cots (Ronse De Craene, 2010). However, in some cases with a
multitude of bracts surrounding the flower a strict distinction
between bracts and bracteoles is not possible (e.g., Reaumuria
in Tamaricaceae: Ronse De Craene, 1990). Boundaries of flow-
ers can be vague, with no clear separation of bracts and sepals
(e.g., Strasburgeria in Strasburgeriaceae; Clusia in Clusi­aceae),

Table 1. Proposal for an improved use of symbols in floral formulae (partly based on Prenner & al., 2010 and Ronse De Craene, 2010).
Symmetry
  Median monosymmetry
  Transverse monosymmetry
    Oblique monosymmetry
 Disymmetry
 Polysymmetry
 Asymmetry
 Spiral flower
Organs
B Bract
Bt Bracteole
E Epicalyx (generally of bract origin but occasionally stipular)
K Calyx (number of sepals) (superscript = sepaloid organ)
C Corolla (number of petals) (superscript = petaloid organ)
P Perigon (number of tepals; no differentiation between calyx and corolla)
A Androecium (number of stamens) (superscript ∞ or definite number refers to stamen fascicles)
Aobd Obdiplostemony
Aobh Obhaplostemony
G Gynoecium (number of carpels)
G  -G-  G Position of ovary: superior, half-inferior, inferior

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or with shifts of bracts within the limits of flowers in the form
of an epicalyx (e.g., Hibiscus in Malv­aceae, Dipsacus in Cap-
rifoliaceae). In the case of Thunbergia (Acanthaceae) and
Calystegia (Convolvulaceae) the two large bracteoles behave
as an epicalyx enclosing the flower but they are still distinc-
tive organs (Weberling, 1989). In some cases the epicalyx may
even replace the original calyx (e.g., Thunbergia, Acanthaceae:
Schönenberger & Endress, 1998). An epicalyx can have differ-
ent origins (possibly stipular in Rosaceae and bract-derived
in Malvaceae: Eichler, 1878; Weberling, 1989), or the origin is
debatable (e.g., Lythraceae: Cadet, 1954; Mayr, 1969). In cases
where the epicalyx is clearly developed, the use of the letter E is
to be preferred. The symmetry symbol is placed before the epi-
calyx members to show that the epicalyx is clearly connected
to the flower (e.g., Lewisia, Potentilla: Fig. 1B; Table 2). This
is also shown for Meliosma, where bracteoles and sepals are
impossible to differentiate from each other (Table 2; Wanntorp
& Ronse De Craene, 2007).
Prenner & al. (2010) also include the placentation and
ovules in the floral formulae. This information is generally
missing from formulae, which are restricted to the organs and
their number, and in the case of carpels include their position
(superior, G; half inferior, -G-; inferior, G ). Prenner & al. use a
single symbol V for describing the number of ovules and refer

Table 1. Continued.
Elements of organs
lo Number of locules (allows for recognizing false septa)
ov Ovule number per ovary
ova Apical placentation
ovb Basal placentation
ovc Free-central placentation
ovl Laminar placentation
ovm Marginal placentation
ovp Parietal placentation
ovx Axile placentation
ovx-p Both axile and parietal present in the same ovary
ps Number of pollen sacs per anther
psl Longitudinal dehiscence
psp Poricidal dehiscence
pst Transversal dehiscence
psv Valvular dehiscence
sty Number of styles
stya Apical style(s)
styg Gynobasic style(s)
sti Stigma number (whether multilobed or single)
stia Apical stigma (not possible to define its position relative to the carpels)
stica Carinal stigma (opposite the carpels)
stico Commissural stigma (alternating with the carpels)
Additional information
+ Indicates when several whorls of the same organ can be distinguished
: Indicates differences within an organ whorl
Fusion of organs at three different levels: ( ), within the same whorl; [ ], involving two different whorls; { }, involving three differ-
( )  [ ]  { } ent whorls. Symbols used for organs reflecting mainly congenital fusion or hypanthial growth
^ Represents a calyptra or fusion of pollen sacs and stigmas or styles; e.g., C^
° Refers to organ reduction (can be used for the whole organ category, e.g. G°, or for part of the organs (e.g., A5° + 5)
– Refers to a range of organ number. The most common number is shown in bold (e.g., K4–5–6)
∞ Many organs, when definite numbers cannot be counted or are unknown
♀ Female (pistillate) flower
♂ Male (staminate) flower
® Resupination

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TAXON 63 (5) • October 2014: 1103–1111 Ronse De Craene & al. • Floral formulae

Table 2. Examples of floral formulae of flowers of diverse families encompassing variable structures. Note that the floral formulae proposed in this
table use a large number of possibilities for the description of the flower, but their ultimate application depends on the needs of users.
Species name Family Floral formula Reference
Amborella trichopoda Amborellaceae ♂ B1 Q Bt2 Pc9–11 A12–21 psl2:2 G0 Buzgo & al., 2004; Ronse De
Baill. ♀ B1 Q Bt2 Pc7–8 A°0–2 G4–5–6 lo1 sty0 stica1 ovb1 Craene, 2010

Nymphaea sp. Nymphaeaceae B1 Kc2+2 Cc4+8+4+4 A∞ psl2:2 -G-10–18 lo1 Fig. 1E; Ronse De Craene, 2010
sty0 stica1 ovl∞

Laurus nobilis L. Lauraceae ♂ B1 Pc2+2 A4+2+2+4 psv1:1 G0 L.P. Ronse De Craene, pers. obs.;
♀ B1 Pc2+2 A°2 G1 lo1 sty a1 stica1 ov a1 Watson & Dallwitz, 1992+

Canna indica L.* Cannaceae B1 Bt1 7 Kk(3) [Cc3 Ac1°–3°+11⁄2 :1⁄2°:2°] psl1/1°:0 Ronse De Craene, 2010; Almeida &
G(3) lo3 sty a1 stia1:1° ovx∞ al., 2013

Meliosma veitchiorum Sabiaceae B1 Bt2 Kk3 Cc2:3 A2:3° psv1:1 G(2) lo2 stya2 Wanntorp & Ronse De Craene, 2007;
Hemsl. stica2 ovm2 Ronse De Craene & Wanntorp, 2008

Lewisia cotyledon Montiaceae B1 Bt2 /n E2 Kc8–13 A6–8–11 psl2:2 G(2–3) lo3 Fig. 1B; Dos Santos & Ronse De
(S.Watson) B.L.Rob. sty a1 stico3 ov fc2 Craene, in prep.

Potentilla fruticosa L. Rosaceae B1 Bt2 Ek5 Kk5 Cc5 A10+5+5+10 psl2:2 G∞ lo1 Lindenhofer & Weber, 2000
sty a1 stica ov b1

Vigna linearis (Kunth) Papilionoid B1 Bt2 7 Kk(2:3)Cc2^7  :2:1 A(9:1) psl2:2 G1 lo1 Fig. 1F; Dwyer & al., 1980
Maréchal & al. Leguminosae sty a7 1 stica1 ov x10

Chamaecrysta desvauxii Caesalpinioid B1 Bt2 7 Kc5 Cc4:1 A5+5 psl2:2 G1 lo1 sty a1 stica1 Costa & al., 2007; A. Iwamoto,
(Colladi) Killip Leguminosae ov x11–13 pers. obs.

Mimusops elengi Wight Sapotaceae B1 Kc4+4 [Cc4+4 A8°+8] psl2:2 G(8) lo8 sty a1 Kümpers & al., subm.
stica8 ov x1

Fremontodendron califor- Malvaceae B1 Bt2 Kc5 C0 (A5°+52) pst2 G(5) lo5 sty a1 stica1 Von Balthazar & al., 2006; Van Heel,
nicum (Torr.) Coville** ov x∞ 1966

Blumenbachia hieronymi Loasaceae B1 Kk5 Cc5 A3°:2°:∞ psl2:2 G(3) lo1 sty a1 stia1 Fig. 1D; Hufford, 1990
Urb. ov p∞

Ceratophyllum Ceratophyllaceae ♂7 Q Pk7–12 A∞ psl2:2 Iwamoto & al., 2003; in prep.
demersum L.*** ♀ Pk5–9 G1 stya1 stia1 ov a1

* The fertile stamen consists of one theca, half of which is petaloid and the other half is fertile; the other theca is aborted (Almeida & al., 2013).
Therefore only one pollen sac is fertile.
** The sterile stamen is fused with two antepetalous stamens in a triplet resembling an ordinary, but highly fragmented stamen (von Balthazar &
al., 2006).
*** The perianth of Ceratophyllum is questionably present and could represent a series of bracts.

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Ronse De Craene & al. • Floral formulae
to the placentation as a secondary condition by an additional
lowercase character (e.g., Va, for apical placentation). They use
this symbol instead of the symbol O as they believe it avoids
confusion with a zero. However, the use of V for ovules is
counterintuitive. The use of lowercase and the symbol “ov”
removes this difficulty. Instead of describing the placentation
as a second lowercase character, we advocate the use of super-
script, such as ovp for parietal placentation.
The description of the flower is also incomplete and remains
inconsistent by only describing ovules without mentioning the
anthers with pollen sacs. Although anthers are generally dithe-
cal and tetrasporangiate in the majority of angiosperms with
two pollen sacs in each theca (symbol “ps”, shown as ps 2:2),
variation should be acknowledged, such as loss of one theca (ps
2:0) or loss of one pollen sac in each theca (ps 1:1). Therefore for
the sake of consistency, we propose the use of “ps” for pollen
sacs as opposed to “ov” for ovules. Dehiscence of pollen sacs
can also be shown, as this is an important diagnostic character:
psl for lateral dehiscence, psv for valvate dehiscence, pst for
transversal dehiscence, and psp for poricidal dehiscence.
Although details provided by Prenner & al. (2010) enrich
our understanding of the flower structure, the list is not com-
plete and there is room to add further details of style and
stigma. Style number (symbol “sty”) can be variable reflect-
ing the number of carpels or styles can be single. The vertical
insertion of the style (apical or gynobasic) can be shown with
the superscript characters stya and styg. While a style can
be either present or absent, stigmatic tissue (symbol “sti”) is
most often developed. The horizontal insertion of styles and
stigmas relative to the carpels can be carinal (i.e., opposite the
carpels, shown with stica) or commissural (i.e., alternating
with the carpels, shown with stico). When the stigma is apical
(i.e., terminal without clear association with the carpels), this
can be shown as stia.
Eichler (1875–1878) distinguished between empirical and
theoretical floral formulae; the former being a description of
what one actually sees in a flower and the latter can be a reflec-
tion of an evolutionary hypothesis. Stützel (2006) and Prenner
& al. (2010) also used theoretical formulae in their distinction
of organ reduction (with superscript r) and organ loss (with
superscript 0). Ronse De Craene (2010) used the symbol “ ° ”
for reduced or sterile organs, such as staminodes (A°) and car-
pellodes (G°). To include absent organs in a floral formula
makes sense only when there is clear evidence for this, such as
absence of petals in a group where petals are generally present
(e.g., Fremontodendron californicum in Malvaceae: Table 2).
In the example of orchids used by Prenner & al., the androe-
cium is presented as A3°+2:1°, following the hypothesis that
the androecium has been derived from the monocot ancestral
condition A3+3. While this is probably true, a medium such
as a floral diagram is more appropriate to show organ losses,
where an asterisk shows the position of the lost organ. Depend-
ing on the context, we advocate to show only organs that are
visible in the mature flower, even if they are present in early
developmental stages and are lost at maturity (e.g., the abaxial
petal of Melianthus: Ronse De Craene & al., 2001). Including
so-called phantom organs, viz. organs that may appear at a
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TAXON 63 (5) • October 2014: 1103–1111
certain developmental stage but are absent in mature flowers
(e.g., Choob, 1999), has meaning for developmental floral for-
mulae (see below), but not for descriptions of mature flowers.
Reduced organs obviously show a progression in their reduc-
tion, as staminodes can be sterile stamens with aborted anthers,
or small, barely developed stubs (Ronse De Craene & Smets,
2001; Ronse De Craene & al., 2002). If there is sufficient evi-
dence for a staminodial nature, any stubs should be referred
to as A°. Nectar leaves of Ranunculaceae are best described as
sterile stamens. In unisexual flowers the lost carpels or stamens
can be represented in the floral formulae, as done in Table 2.
However, this decision lies with the user in the context of floral
evolution.
A number of abbreviations in Prenner & al. (2010) are
misleading or insufficiently clear (see Table 1). For example,
oblique monosymmetry is shown by the symbol “ø”, but
this does not convey any information about the orientation
of monosymmetry. We advocate the use of different arrows
() reflecting the direction of oblique monosymmetry.
Stützel (2006) followed a similar approach. The same applies
for disymmetry, using “  ” instead of “ + ”. Prenner & al. (2010)
use a separate symmetry symbol for each type of organ in the
flower. Symmetry obviously varies in the flower, but several
reasons can be responsible for this, including gravity and loss
of organs (see Endress, 1999). For example, what can be done
with cases where stamens are inserted in a regular whorl, but
hang down on one side of the flower? Is this to be described
as a monosymmetric or a polysymmetric androecium as, e.g.,
in several Amaryllidaceae? Flowers may also change their
symmetry by late developmental processes (Endress, 1999;
Tucker, 1999). In the case of Vigna (Leguminosae) flowers
become asymmetric due to a coiling of the keel petals and
style (Fig. 1F). In Chamaecrista and Senna (Leguminosae)
the flower is asymmetric by unequal growth of petals and sta-
mens and a twist in the ovary, starting early in development
(Tucker, 1999). This regulates specific pollen deposition on
insects (Endress, 1999). A regular pentamerous flower often
has a trimerous gynoecium with a single symmetry plane as
in Polemoniaceae and Caryophyllaceae.
Describing a different monosymmetry for each whorl obvi-
ously adds to the complexity of the floral formula. We suggest
restricting the symbol of symmetry to the whole flower, as we
prefer to adhere to the overall perception of symmetry. How-
ever, it should be open to the user if they should wish to add a
different symbol for each whorl, especially in cases where there
is a clear switch in symmetry between different organs. There
are several cases where the flower has a basically symmetrical
groundplan, but where the flower is weakly monosymmetric
(e.g., Rhododendron, Epilobium, Erodium). It is best to describe
these flowers as polysymmetric. Any indications of monosym-
metry in different whorls can be shown by the use of a colon.
This is an important symbol, which shows differences in shape,
including partial sterility of the androecium and gynoecium.
This was done independently by Ronse De Craene (2010) and
Prenner & al. (2010) and is particularly useful to show the
orientation of petal lobes in a bilabiate flower. For example,
some Erodium show distinctive spots on the two adaxial petals
TAXON 63 (5) • October 2014: 1103–1111
(E. leucanthemum Boiss., E. ×willkommianum Sünd.) or a dif-
ference in size (E. foetidum (L.) L’Her.) and this can be shown
as C3:2 (L.P. Ronse De Craene, pers. obs.). Therefore C3:2
indicates an arrangement of a lower lip with three petal lobes
and an upper lip with 2 lobes (e.g., Penstemon, Fig. 1C).
Contrary to Donoghue & al. (1998) or Prenner & al. (2010)
we do not visualize the sequence from the adaxial (dorsal, pos-
terior) to the abaxial (ventral, anterior) side of the flower, but
from the abaxial to the adaxial, as this is the logical direction
for a pollinator approaching the flower and also reflects the
sequence of initiation of the petals in many monosymmetric
flowers (e.g., Tucker, 1999 for Fabaceae; Endress, 1998, 1999 for
Lamiales). “Anterior” linguistically also reflects a precedence
in time and space. A partly sterile androecial whorl inserted
on different levels can be shown as A2:2:1° as is a common
condition in Lamiales (e.g., Penstemon, Fig. 1C).
Pseudomonomery reflects the presence of a single fertile
carpel out of multiple carpels and refers to an incompletely
fertile ovary (see Eckardt, 1937; Ronse De Craene & Smets,
1998). This is an important diagnostic feature of some families
such as Anacardiaceae expressed as G(2°:1), where only a single
carpel out of three is fertile, while two locules are empty (e.g.,
Bachelier & Endress, 2009). A full gradation is possible, rang-
ing from three fully fertile carpels to a single carpel without
any traces of the other two. Apart from Anacardiaceae, this is
characteristic for Restionaceae (Ronse De Craene & al., 2002).
The representation of the position of stamens in the flower
is a feature of great systematic importance. In flowers with
two stamen whorls a distinction can be made between dip-
lostemony (the outer whorl inserted opposite the calyx) and
obdiplostemony (the outer whorl inserted in antepetalous or
alternisepalous position). In case of a single stamen whorl, it is
either inserted opposite the calyx (haplostemony) or opposite
the corolla (obhaplostemony; alternisepalous). Obhaplostemony
and obdiplostemony also have strong phylogenetic significance
as characters (see Ronse De Craene & Smets, 1995; Ronse De
Craene, 2010). Although we agree with Prenner & al. (2010) that
it is important to show the anomalous condition of obdiploste-
mony in the flower, a left-right arrow (n) does not clearly show
the number of whorls and can be confused with the symbol of
disymmetry; in addition, if other organs are arranged as a series
of whorls with “+”, this makes little sense. It is better to use
“OBH” for obhaplostemony and “OBD” for obdiplostemony in
superscript after the symbol for the stamens (Aobh and Aobd).
This allows for the use of stamen position to reconstruct the
spatial understanding of flowers, even in the absence of petals
or sepals.
An extreme case of a complex flower is Canna (Cann­
aceae, see Ronse De Craene, 2010; Glinos & Cocucci, 2011;
Almeida & al., 2013) with an androecium derived from two
whorls (A3+3): the outer whorl is restricted to a single sta-
minode and the inner whorl consists of two staminodes with
the fertile part of the androecium restricted to half of a theca:
A1°+11⁄2 :1⁄2°:2°.
In the case of Erodium (Fig. 1A), a complete floral formula
would be as follows: B1 Bt1 Kk5 Cc5 Aobd(5+5°) psl2:2 G(5)
lo5 stya1 stica5 ovx5. It is not necessary to include all possible
Version of Record (identical to print version).
Ronse De Craene & al. • Floral formulae
elements, although additional data complete the description of
the flower that can be used as a shortened diagnosis (cf. Prenner
& al., 2010). The complexity of the floral formulae depends
on the needs of the user. We refer to Table 2 for examples of
flowers with occasionally difficult structures.
ADDITIONAL COMPLICATIONS FOR
FLORAL FORMULAE
Floral formulae are not static and could be utilized for
developmental studies. Changes in the morphology of flowers
during development occur in several groups of plants, especially
in the symmetry of the flower (e.g., Endress, 1999; Tucker, 1999;
Ronse De Craene & Smets, 2001; Ronse De Craene & al., 2001;
Olson, 2003; Iwamoto & al., 2003; Patchell & al., 2011). This
can be highlighted in floral formulae, as has been done for the
floral diagram of Tropaeolum by Ronse De Craene & Smets
(2001). Tsou & Mori (2007) used floral formulae to describe
different stages in the development of the complex flowers of
Lecythidaceae. Leins & Erbar (2010) used symbols to show the
direction of stamen development in multistaminate flowers.
Indeed, the floral formulae are a useful tool to describe flow-
ers at different stages of their development and the sequence
of initiation of organs, although they should be used for spe-
cific comparative purposes, not involving mature flowers. For
example, a whorled centrifugal sequence of stamen initiation
can be presented and read in a reversed order in a floral for-
mula (e.g., Capparis: Ronse De Craene & Smets, 1997). The
gynoecium can also arise before the androecium, as described
in a number of families (e.g., Ge & al., 2007; Ronse De Craene,
2011; Wanntorp & al., 2011). Studies in floral development are
useful in understanding the structure of the flower, which can
be obscured at maturity. Especially with large flowers having
numerous organs an early developmental sequence can help
in understanding the number and arrangement of organs with
complex phyllotaxis, facilitating a depiction with floral formu-
lae (e.g., Nymphaea in Nymphaeaceae: Fig. 1E; Anaxagorea
in Annonaceae: Endress & Armstrong, 2011; Corbichonia in
Lophiocarpaceae: Ronse De Craene, 2010; Brockington & al.,
2013).
Another important issue with constructing floral formu-
lae is the interpretation of visually striking flower structures,
such as receptacular cups or hypanthia. There is an abundant
literature dealing with the controversy of floral cups being the
result of fusions of organs or the result of zonal growth lifting
organs into a common structure (see Weberling, 1989; Leins
& Erbar, 2010). The distinction between congenital fusion and
postgenital fusion of organs is intimately linked to this dis-
cussion (see Sattler, 1978; Leins & Erbar, 2010). As hypanthia
or receptacular outgrowths are highly variable in flowers, it
is preferable to limit the use of floral formulae to what can
actually be observed in the flower as a connection between
different organs. The use of three kinds of brackets allows for
additional detail although it does not necessarily reflect fusion
(see Table 1). Postgenital fusion can occasionally be shown
by the symbol “ ^ ”, as the connivent anthers in Asteraceae
1109
Ronse De Craene & al. • Floral formulae
and Asclepias (Asclepiadaceae) (A5^) or the development of a
calyptrate corolla in Vitaceae (C5^). Prenner & al. (2010) use the
brackets in subscript and superscript to show the difference in
the level of fusion, as in Asteraceae with the common stamen-
petal tube shown by [  ] and the fusion of the anthers by (  ).
Nectaries or other appendages (corona appendages, obvi-
ous concentrations of tufts of trichomes) are generally omitted
from floral formulae, although they can be highly important
visual cues in flowers (e.g., the corona of Narcissus: Waters
& al., 2013). Unfortunately, nectaries or coronas are too variable
in position and shape to be incorporated in purely descriptive
floral formulae, as they would add complexity to descriptions.
The use of floral diagrams is complementary in this way.
Another problem is the occasional loss of one of two peri-
anth whorls, leaving a single petaloid perianth, which can either
be derived from a calyx (e.g., in Daphne of Thymelaeaceae)
or from a corolla (as in Osyris of Santalaceae). This difficulty
can be solved in the context of the knowledge of the phylogeny
of plants in question, where Kc4 C0 and K0 Cc4 can be used
respectively, or a neutral approach can be advocated by using
the symbol Pc for tepals. We advocate the use of superscripts
c and k on a general basis, even if the organs are obviously
sepal-like or petal-like. With a broader use of floral formulae
in descriptions or diagnoses, this information is important.
Prenner & al. (2010) also advocate writing out the name of
a particular structure in full when it is highly distinctive in a
flower (e.g., labellum, pappus). This makes the floral formulae
longer and more complex and could be inconsistent due to the
great diversity of floral forms. It will be of little value to the
user if he is not aware of the structure and different authors
sometimes use different names for the same structure; at the
same time users familiar with the structure would find this
information superfluous.
Other complications arise when two types of flowers are
present (e.g., left and right-oriented). This would necessitate
the use of two different arrows, adding to the confusion. Also,
exaggerated contortions of organs cannot be shown fully, as for
example in Vigna (Fig. 1F) and Chamaecrista where the ovary
is curved for about 45 degrees.
CONCLUSIONS
We agree with Prenner & al. (2010) that floral formulae
should be given greater weight in botanical research, as a
valuable tool in comparative morphology for making concise
descriptions, identification keys, and even comparisons of flo-
ral developmental sequences. However, floral formulae cannot
replace floral diagrams in the spatial depiction of flowers and
in the description of special structures. At the same time floral
formulae must not be overcomplicated, as an accumulation
of too much information can be counterproductive. The use
of a hierarchical sequence in floral symbols allows for better
clarity in conveying the information. Floral formulae can be
as detailed as the user wishes it to be. The descriptors that we
propose here aim not to be dogmatic but to provide a clear
and standardized system that is fit for multiple purposes as a
1110 Version of Record (identical to print version).
TAXON 63 (5) • October 2014: 1103–1111
powerful tool to accurately describe and compare floral mor-
phologies. Presenting exact floral formulae requires a detailed
investigation of the flower by the user and is an excellent incen-
tive to observe flowers in the field and in the classroom.
Although there are several parallel systems for building
floral formulae (e.g., the American versus European system),
we emphasize the need of a universal language that is clear and
concise. A consensus among botanists is a necessary require-
ment to make the use of floral formulae really worthwhile. We
believe that this paper is a step in the right direction.
ACKNOWLEDGEMENTS
We thank RBGE for providing facilities to organize a Floral Mor-
phology Discussion Group, which led to the development of this paper.
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